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diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..6833f05 --- /dev/null +++ b/.gitattributes @@ -0,0 +1,3 @@ +* text=auto +*.txt text +*.md text diff --git a/34967-0.txt b/34967-0.txt new file mode 100644 index 0000000..0566e9e --- /dev/null +++ b/34967-0.txt @@ -0,0 +1,14522 @@ +The Project Gutenberg EBook of The Descent of Man and Selection in +Relation to Sex, Vol. I (1st editio, by Charles Darwin + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: The Descent of Man and Selection in Relation to Sex, Vol. I (1st edition) + +Author: Charles Darwin + +Release Date: January 15, 2011 [EBook #34967] + +Language: English + +Character set encoding: UTF-8 + +*** START OF THIS PROJECT GUTENBERG EBOOK THE DESCENT OF MAN AND *** + + + + +Produced by StevenGibbs, Turgut Dincer and the Online +Distributed Proofreading Team at http://www.pgdp.net + + + + + + + + + +THE DESCENT OF MAN, + +AND + +SELECTION IN RELATION TO SEX. + + + + +BY CHARLES DARWIN, M.A., F.R.S., &c. + +IN TWO VOLUMES.—VOL. I. + +WITH ILLUSTRATIONS. + +LONDON: + +JOHN MURRAY, ALBEMARLE STREET. + +1871. + +[_The right of Translation is reserved._] + +BY THE SAME AUTHOR. + + + ON THE ORIGIN OF SPECIES BY MEANS OF NATURAL SELECTION; or, THE + PRESERVATION OF FAVOURED RACES IN THE STRUGGLE FOR LIFE; Fifth + Edition (_Tenth Thousand_), with Additions and Corrections. + 1869. MURRAY. + + THE VARIATION OF ANIMALS AND PLANTS UNDER DOMESTICATION. In two vols. + With Illustrations. 1868. MURRAY. + + ON THE VARIOUS CONTRIVANCES BY WHICH BRITISH AND FOREIGN ORCHIDS ARE + FERTILISED BY INSECTS; and on the GOOD EFFECTS OF CROSSING. With + numerous Woodcuts. MURRAY. + + A NATURALIST’S VOYAGE ROUND THE WORLD; or, A JOURNAL OF RESEARCHES INTO + THE NATURAL HISTORY AND GEOLOGY OF THE COUNTRIES visited during the + voyage of H.M.S. ‘Beagle,’ under the command of Captain FITZROY, R.N. + _Eleventh Thousand_. MURRAY. + + ON THE STRUCTURE AND DISTRIBUTION OF CORAL REEFS. SMITH, ELDER, & CO. + + GEOLOGICAL OBSERVATIONS ON VOLCANIC ISLANDS. SMITH, ELDER, & CO. + + GEOLOGICAL OBSERVATIONS ON SOUTH AMERICA. SMITH, ELDER & CO. + + A MONOGRAPH OF THE CIRRIPEDIA. With numerous Illustrations. 2 vols. + 8vo. HARDWICKE. + + ON THE MOVEMENTS AND HABITS OF CLIMBING PLANTS. With Woodcuts. WILLIAMS + & NORGATE. + + +LONDON: PRINTED BY WILLIAM CLOWES AND SONS, STAMFORD STREET, AND CHARING +CROSS. + + + + + CONTENTS. + + + INTRODUCTION Page 1-5 + + + PART I. + + ON THE DESCENT OF MAN. + + + CHAPTER I. + + THE EVIDENCE OF THE DESCENT OF MAN FROM SOME LOWER FORM. + + Nature of the evidence bearing on the origin of man—Homologous + structures in man and the lower animals—Miscellaneous points + of correspondence—Development—Rudimentary structures, + muscles, sense-organs, hair, bones, reproductive organs, + &c.—The bearing of these three great classes of facts on the + origin of man 9-33 + + + CHAPTER II. + + COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER ANIMALS. + + The difference in mental power between the highest ape and the + lowest savage, immense—Certain instincts in common—The + emotions—Curiosity—Imitation—Attention—Memory—Imagination— + Reason—Progressive improvement—Tools and weapons used by + animals—Language—Self-consciousness—Sense of beauty—Belief + in God, spiritual agencies, superstitions 34-69 + + + CHAPTER III. + + COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER + ANIMALS—_continued_. + + The moral sense—Fundamental proposition—The qualities of + social animals—Origin of sociability—Struggle between opposed + instincts—Man and social animal—The more enduring social + instincts conquer other less persistent instincts—The social + virtues alone regarded by savages—The self-regarding virtues + acquired at a later stage of development—The importance of the + judgment of the members of the same community on + conduct—Transmission of moral tendencies—Summary 70-106 + + + CHAPTER IV. + + ON THE MANNER OF DEVELOPMENT OF MAN FROM SOME LOWER FORM. + + Variability of body and mind in man—Inheritance—Causes of + variability—Laws of variation the same in man as in the lower + animals—Direct action of the conditions of life—Effects of + the increased use and disuse of parts—Arrested + development—Reversion—Correlated variation—Rate of + increase—Checks to increase—Natural selection—Man the most + dominant animal in the world—Importance of his corporeal + structure—The causes which have led to his becoming + erect—Consequent changes of structure—Decrease in size of the + canine teeth—Increased size and altered shape of the + skull—Nakedness—Absence of a tail—Defenceless condition of + man 107-157 + + + CHAPTER V. + + ON THE DEVELOPMENT OF THE INTELLECTUAL AND MORAL FACULTIES + DURING PRIMEVAL AND CIVILISED TIMES. + + The advancement of the intellectual powers through natural + selection—Importance of imitation—Social and moral + faculties—Their development within the limits of the same + tribe—Natural selection as affecting civilised + nations—Evidence that civilised nations were once barbarous 158-184 + + + CHAPTER VI. + + ON THE AFFINITIES AND GENEALOGY OF MAN. + + Position of man in the animal series—The natural system + genealogical—Adaptive characters of slight value—Various + small points of resemblance between man and the + Quadrumana—Rank of man in the natural system—Birthplace and + antiquity of man—Absence of fossil connecting-links—Lower + stages in the genealogy of man, as inferred, firstly from his + affinities and secondly from his structure—Early androgynous + condition of the Vertebrata—Conclusion 185-213 + + + CHAPTER VII. + + ON THE RACES OF MAN. + + The nature and value of specific characters—Application to the + races of man—Arguments in favour of, and opposed to, ranking + the so-called races of man as distinct species—Sub-species— + Monogenists and polygenists—Convergence of character—Numerous + points of resemblance in body and mind between the most + distinct races of man—The state of man when he first spread + over the earth—Each race not descended from a single + pair—The extinction of races—The formation of races— + The effects of crossing—Slight influence of the direct + action of the conditions of life—Slight or no influence of + natural selection—Sexual selection. 214-250 + + + PART II. SEXUAL SELECTION. + + CHAPTER VIII. + + PRINCIPLES OF SEXUAL SELECTION. + + Secondary sexual characters—Sexual selection—Manner of + action—Excess of males—Polygamy—The male alone generally + modified through sexual selection—Eagerness of the + male—Variability of the male—Choice exerted by the + female—Sexual compared with natural selection—Inheritance at + corresponding periods of life, at corresponding seasons of the + year, and as limited by sex—Relations between the several + forms of inheritance—Causes why one sex and the young are not + modified through sexual selection—Supplement on the + proportional numbers of the two sexes throughout the animal + kingdom—On the limitation of the numbers of the two sexes + through natural selection 253-320 + + + CHAPTER IX. + + SECONDARY SEXUAL CHARACTERS IN THE LOWER CLASSES OF THE ANIMAL + KINGDOM. + + These characters absent in the lowest classes—Brilliant + colours—Mollusca—Annelids—Crustacea, secondary sexual + characters strongly developed; dimorphism; colour; characters + not acquired before maturity—Spiders, sexual colours of; + stridulation by the males—Myriapoda 321-340 + + + CHAPTER X. + + SECONDARY SEXUAL CHARACTERS OF INSECTS. + + Diversified structures possessed by the males for seizing the + females—Differences between the sexes, of which the meaning is + not understood—Difference in size between the + sexes—Thysanura—Diptera—Hemiptera—Homoptera, musical powers + possessed by the males alone—Orthoptera, musical instruments + of the males, much diversified in structure; pugnacity; + colours—Neuroptera, sexual differences in colour—Hymenoptera, + pugnacity and colours—Coleoptera, colours; furnished with + great horns, apparently as an ornament; battles; stridulating + organs generally common to both sexes 341-385 + + + CHAPTER XI. + + INSECTS, _continued_.—ORDER LEPIDOPTERA. + + Courtship of butterflies—Battles—Ticking noise—Colours + common to both sexes, or more brilliant in the + males—Examples—Not due to the direct action of the conditions + of life—Colours adapted for protection—Colours of + moths—Display—Perceptive powers of the + Lepidoptera—Variability—Causes of the difference in colour + between the males and females—Mimickry, female butterflies + more brilliantly coloured than the males—Bright colours of + caterpillars—Summary and concluding remarks on the secondary + sexual characters of insects—Birds and insects compared 386-423 + + + + +THE DESCENT OF MAN; + +AND ON + +SELECTION IN RELATION TO SEX. + + +INTRODUCTION. + +The nature of the following work will be best understood by a brief +account of how it came to be written. During many years I collected +notes on the origin or descent of man, without any intention of +publishing on the subject, but rather with the determination not to +publish, as I thought that I should thus only add to the prejudices +against my views. It seemed to me sufficient to indicate, in the first +edition of my ‘Origin of Species,’ that by this work “light would be +thrown on the origin of man and his history;” and this implies that man +must be included with other organic beings in any general conclusion +respecting his manner of appearance on this earth. Now the case wears a +wholly different aspect. When a naturalist like Carl Vogt ventures to +say in his address as President of the National Institution of Geneva +(1869), “personne, en Europe au moins, n’ose plus soutenir la création +indépendante et de toutes pièces, des espèces,” it is manifest that at +least a large number of naturalists must admit that species are the +modified descendants of other species; and this especially holds good +with the younger and rising naturalists. The greater number accept the +agency of natural selection; though some urge, whether with justice the +future must decide, that I have greatly overrated its importance. Of the +older and honoured chiefs in natural science, many unfortunately are +still opposed to evolution in every form. + +In consequence of the views now adopted by most naturalists, and which +will ultimately, as in every other case, be followed by other men, I +have been led to put together my notes, so as to see how far the general +conclusions arrived at in my former works were applicable to man. This +seemed all the more desirable as I had never deliberately applied these +views to a species taken singly. When we confine our attention to any +one form, we are deprived of the weighty arguments derived from the +nature of the affinities which connect together whole groups of +organisms—their geographical distribution in past and present times, +and their geological succession. The homological structure, +embryological development, and rudimentary organs of a species, whether +it be man or any other animal, to which our attention may be directed, +remain to be considered; but these great classes of facts afford, as it +appears to me, ample and conclusive evidence in favour of the principle +of gradual evolution. The strong support derived from the other +arguments should, however, always be kept before the mind. + +The sole object of this work is to consider, firstly, whether man, like +every other species, is descended from some pre-existing form; secondly, +the manner of his development; and thirdly, the value of the +differences between the so-called races of man. As I shall confine +myself to these points, it will not be necessary to describe in detail +the differences between the several races—an enormous subject which has +been fully discussed in many valuable works. The high antiquity of man +has recently been demonstrated by the labours of a host of eminent men, +beginning with M. Boucher de Perthes; and this is the indispensable +basis for understanding his origin. I shall, therefore, take this +conclusion for granted, and may refer my readers to the admirable +treatises of Sir Charles Lyell, Sir John Lubbock, and others. Nor shall +I have occasion to do more than to allude to the amount of difference +between man and the anthropomorphous apes; for Prof. Huxley, in the +opinion of most competent judges, has conclusively shewn that in every +single visible character man differs less from the higher apes than +these do from the lower members of the same order of Primates. + +This work contains hardly any original facts in regard to man; but as +the conclusions at which I arrived, after drawing up a rough draft, +appeared to me interesting, I thought that they might interest others. +It has often and confidently been asserted, that man’s origin can never +be known: but ignorance more frequently begets confidence than does +knowledge: it is those who know little, and not those who know much, who +so positively assert that this or that problem will never be solved by +science. The conclusion that man is the co-descendant with other species +of some ancient, lower, and extinct form, is not in any degree new. +Lamarck long ago came to this conclusion, which has lately been +maintained by several eminent naturalists and philosophers; for instance +by Wallace, Huxley, Lyell, Vogt, Lubbock, Büchner, Rolle, &c.,[1] and +especially by Häckel. This last naturalist, besides his great work, +‘Generelle Morphologie’ (1866), has recently (1868, with a second edit. +in 1870), published his ‘Natürliche Schöpfungsgeschichte,’ in which he +fully discusses the genealogy of man. If this work had appeared before +my essay had been written, I should probably never have completed it. +Almost all the conclusions at which I have arrived I find confirmed by +this naturalist, whose knowledge on many points is much fuller than +mine. Wherever I have added any fact or view from Prof. Häckel’s +writings, I give his authority in the text, other statements I leave as +they originally stood in my manuscript, occasionally giving in the +foot-notes references to his works, as a confirmation of the more +doubtful or interesting points. + +During many years it has seemed to me highly probable that sexual +selection has played an important part in differentiating the races of +man; but in my ‘Origin of Species’ (first edition, p. 199) I contented +myself by merely alluding to this belief. When I came to apply this view +to man, I found it indispensable to treat the whole subject in full +detail.[2] Consequently the second part of the present work, treating of +sexual selection, has extended to an inordinate length, compared with +the first part; but this could not be avoided. + +I had intended adding to the present volumes an essay on the expression +of the various emotions by man and the lower animals. My attention was +called to this subject many years ago by Sir Charles Bell’s admirable +work. This illustrious anatomist maintains that man is endowed with +certain muscles solely for the sake of expressing his emotions. As this +view is obviously opposed to the belief that man is descended from some +other and lower form, it was necessary for me to consider it. I likewise +wished to ascertain how far the emotions are expressed in the same +manner by the different races of man. But owing to the length of the +present work, I have thought it better to reserve my essay, which is +partially completed, for separate publication. + + + + +PART I. + +THE DESCENT OR ORIGIN OF MAN. + +PART I.—THE DESCENT OF MAN. + + + + +CHAPTER I. + +THE EVIDENCE OF THE DESCENT OF MAN FROM SOME LOWER FORM. + + + Nature of the evidence bearing on the origin of man—Homologous + structures in man and the lower animals—Miscellaneous points + of correspondence—Development—Rudimentary structures, + muscles, sense-organs, hair, bones, reproductive organs, + &c.—The bearing of these three great classes of facts on the + origin of man. + + +He who wishes to decide whether man is the modified descendant of some +pre-existing form, would probably first enquire whether man varies, +however slightly, in bodily structure and in mental faculties; and if +so, whether the variations are transmitted to his offspring in +accordance with the laws which prevail with the lower animals; such as +that of the transmission of characters to the same age or sex. Again, +are the variations the result, as far as our ignorance permits us to +judge, of the same general causes, and are they governed by the same +general laws, as in the case of other organisms; for instance by +correlation, the inherited effects of use and disuse, &c.? Is man +subject to similar malconformations, the result of arrested development, +of reduplication of parts, &c., and does he display in any of his +anomalies reversion to some former and ancient type of structure? It +might also naturally be enquired whether man, like so many other +animals, has given rise to varieties and sub-races, differing but +slightly from each other, or to races differing so much that they must +be classed as doubtful species? How are such races distributed over the +world; and how, when crossed, do they react on each other, both in the +first and succeeding generations? And so with many other points. + +The enquirer would next come to the important point, whether man tends +to increase at so rapid a rate, as to lead to occasional severe +struggles for existence, and consequently to beneficial variations, +whether in body or mind, being preserved, and injurious ones eliminated. +Do the races or species of men, whichever term may be applied, encroach +on and replace each other, so that some finally become extinct? We shall +see that all these questions, as indeed is obvious in respect to most of +them, must be answered in the affirmative, in the same manner as with +the lower animals. But the several considerations just referred to may +be conveniently deferred for a time; and we will first see how far the +bodily structure of man shows traces, more or less plain, of his descent +from some lower form. In the two succeeding chapters the mental powers +of man, in comparison with those of the lower animals, will be +considered. + + +_The Bodily Structure of Man._—It is notorious that man is constructed +on the same general type or model with other mammals. All the bones in +his skeleton can be compared with corresponding bones in a monkey, bat, +or seal. So it is with his muscles, nerves, blood-vessels and internal +viscera. The brain, the most important of all the organs, follows the +same law, as shewn by Huxley and other anatomists. Bischoff,[3] who is a +hostile witness, admits that every chief fissure and fold in the brain +of man has its analogy in that of the orang; but he adds that at no +period of development do their brains perfectly agree; nor could this be +expected, for otherwise their mental powers would have been the same. +Vulpian[4] remarks: “Les différences réelles qui existent entre +l’encéphale de l’homme et celui des singes supérieurs, sont bien +minimes. Il ne faut pas se faire d’illusions à cet égard. L’homme est +bien plus près des singes anthropomorphes par les caractères anatomiques +de son cerveau que ceux-ci ne le sont non-seulement des autres +mammifères, mais mêmes de certains quadrumanes, des guenons et des +macaques.” But it would be superfluous here to give further details on +the correspondence between man and the higher mammals in the structure +of the brain and all other parts of the body. + +It may, however, be worth while to specify a few points, not directly or +obviously connected with structure, by which this correspondence or +relationship is well shewn. + +Man is liable to receive from the lower animals, and to communicate to +them, certain diseases as hydrophobia, variola, the glanders, &c.; and +this fact proves the close similarity of their tissues and blood, both +in minute structure and composition, far more plainly than does their +comparison under the best microscope, or by the aid of the best chemical +analysis. Monkeys are liable to many of the same non-contagious diseases +as we are; thus Rengger,[5] who carefully observed for a long time the +_Cebus Azaræ_ in its native land, found it liable to catarrh, with the +usual symptoms, and which when often recurrent led to consumption. +These monkeys suffered also from apoplexy, inflammation of the bowels, +and cataract in the eye. The younger ones when shedding their milk-teeth +often died from fever. Medicines produced the same effect on them as on +us. Many kinds of monkeys have a strong taste for tea, coffee, and +spirituous liquors: they will also, as I have myself seen, smoke tobacco +with pleasure. Brehm asserts that the natives of north-eastern Africa +catch the wild baboons by exposing vessels with strong beer, by which +they are made drunk. He has seen some of these animals, which he kept in +confinement, in this state; and he gives a laughable account of their +behaviour and strange grimaces. On the following morning they were very +cross and dismal; they held their aching heads with both hands and wore +a most pitiable expression: when beer or wine was offered them, they +turned away with disgust, but relished the juice of lemons.[6] An +American monkey, an Ateles, after getting drunk on brandy, would never +touch it again, and thus was wiser than many men. These trifling facts +prove how similar the nerves of taste must be in monkeys and man, and +how similarly their whole nervous system is affected. + +Man is infested with internal parasites, sometimes causing fatal +effects, and is plagued by external parasites, all of which belong to +the same genera or families with those infesting other mammals. Man is +subject like other mammals, birds, and even insects, to that mysterious +law, which causes certain normal processes, such as gestation, as well +as the maturation and duration of various diseases, to follow lunar +periods.[7] His wounds are repaired by the same process of healing; and +the stumps left after the amputation of his limbs occasionally possess, +especially during an early embryonic period, some power of regeneration, +as in the lowest animals.[8] + +The whole process of that most important function, the reproduction of +the species, is strikingly the same in all mammals, from the first act +of courtship by the male[9] to the birth and nurturing of the young. +Monkeys are born in almost as helpless a condition as our own infants; +and in certain genera the young differ fully as much in appearance from +the adults, as do our children from their full-grown parents.[10] It has +been urged by some writers as an important distinction, that with man +the young arrive at maturity at a much later age than with any other +animal; but if we look to the races of mankind which inhabit tropical +countries the difference is not great, for the orang is believed not to +be adult till the age of from ten to fifteen years.[11] Man differs +from woman in size, bodily strength, hairyness, &c., as well as in mind, +in the same manner as do the two sexes of many mammals. It is, in short, +scarcely possible to exaggerate the close correspondence in general +structure, in the minute structure of the tissues, in chemical +composition and in constitution, between man and the higher animals, +especially the anthropomorphous apes. + + +_Embryonic Development._—Man is developed from an ovule, about the +125th of an inch in diameter, which differs in no respect from the +ovules of other animals. The embryo itself at a very early period can +hardly be distinguished from that of other members of the vertebrate +kingdom. At this period the arteries run in arch-like branches, as if to +carry the blood to branchiæ which are not present in the higher +vertebrata, though the slits on the sides of the neck still remain (_f_, +_g_, fig. 1), marking their former position. At a somewhat later period, +when the extremities are developed, “the feet of lizards and mammals,” +as the illustrious Von Baer remarks, “the wings and feet of birds, no +less than the hands and feet of man, all arise from the same fundamental +form.” It is, says Prof. Huxley,[12] “quite in the later stages of +development that the young human being presents marked differences from +the young ape, while the latter departs as much from the dog in its +developments, as the man does. Startling as this last assertion may +appear to be, it is demonstrably true.” + +As some of my readers may never have seen a drawing of an embryo, I have +given one of man and another of a dog, at about the same early stage of +development, carefully copied from two works of undoubted accuracy.[13] + +[Illustration: Fig. 1. Upper figure human embryo, from Ecker. Lower +figure that of a dog, from Bischoff. + + _a._ Fore-brain, cerebral hemispheres, &c. + _b._ Mid-brain, corpora quadrigemina. + _c._ Hind-brain, cerebellum, medulla oblongata. + _d._ Eye. + _e._ Ear. + _f._ First visceral arch. + _g._ Second visceral arch. + H. Vertebral columns and muscles in process of development. + _i._ Anterior ┐ + │ extremities. + K. Posterior ┘ + L. Tail or os coccyx.] + +After the foregoing statements made by such high authorities, it would +be superfluous on my part to give a number of borrowed details, shewing +that the embryo of man closely resembles that of other mammals. It may, +however, be added that the human embryo likewise resembles in various +points of structure certain low forms when adult. For instance, the +heart at first exists as a simple pulsating vessel; the excreta are +voided through a cloacal passage; and the os coccyx projects like a true +tail, “extending considerably beyond the rudimentary legs.”[14] In the +embryos of all air-breathing vertebrates, certain glands called the +corpora Wolffiana, correspond with and act like the kidneys of mature +fishes.[15] Even at a later embryonic period, some striking resemblances +between man and the lower animals may be observed. Bischoff says that +the convolutions of the brain in a human fœtus at the end of the +seventh month reach about the same stage of development as in a baboon +when adult.[16] The great toe, as Prof. Owen remarks,[17] “which forms +the fulcrum when standing or walking, is perhaps the most +characteristic peculiarity in the human structure;” but in an embryo, +about an inch in length, Prof. Wyman[18] found “that the great toe was +shorter than the others, and, instead of being parallel to them, +projected at an angle from the side of the foot, thus corresponding with +the permanent condition of this part in the quadrumana.” I will conclude +with a quotation from Huxley,[19] who after asking, does man originate +in a different way from a dog, bird, frog or fish? says, “the reply is +not doubtful for a moment; without question, the mode of origin and the +early stages of the development of man are identical with those of the +animals immediately below him in the scale: without a doubt in these +respects, he is far nearer to apes, than the apes are to the dog.” + + +_Rudiments._—This subject, though not intrinsically more important than +the two last, will for several reasons be here treated with more +fullness.[20] Not one of the higher animals can be named which does not +bear some part in a rudimentary condition; and man forms no exception to +the rule. Rudimentary organs must be distinguished from those that are +nascent; though in some cases the distinction is not easy. The former +are either absolutely useless, such as the mammæ of male quadrupeds, or +the incisor teeth of ruminants which never cut through the gums; or they +are of such slight service to their present possessors, that we cannot +suppose that they were developed under the conditions which now exist. +Organs in this latter state are not strictly rudimentary, but they are +tending in this direction. Nascent organs, on the other hand, though not +fully developed, are of high service to their possessors, and are +capable of further development. Rudimentary organs are eminently +variable; and this is partly intelligible, as they are useless or nearly +useless, and consequently are no longer subjected to natural selection. +They often become wholly suppressed. When this occurs, they are +nevertheless liable to occasional reappearance through reversion; and +this is a circumstance well worthy of attention. + +Disuse at that period of life, when an organ is chiefly used, and this +is generally during maturity, together with inheritance at a +corresponding period of life, seem to have been the chief agents in +causing organs to become rudimentary. The term “disuse” does not relate +merely to the lessened action of muscles, but includes a diminished flow +of blood to a part or organ, from being subjected to fewer alternations +of pressure, or from becoming in any way less habitually active. +Rudiments, however, may occur in one sex of parts normally present in +the other sex; and such rudiments, as we shall hereafter see, have often +originated in a distinct manner. In some cases organs have been reduced +by means of natural selection, from having become injurious to the +species under changed habits of life. The process of reduction is +probably often aided through the two principles of compensation and +economy of growth; but the later stages of reduction, after disuse has +done all that can fairly be attributed to it, and when the saving to be +effected by the economy of growth would be very small,[21] are difficult +to understand. The final and complete suppression of a part, already +useless and much reduced in size, in which case neither compensation nor +economy can come into play, is perhaps intelligible by the aid of the +hypothesis of pangenesis, and apparently in no other way. But as the +whole subject of rudimentary organs has been fully discussed and +illustrated in my former works,[22] I need here say no more on this +head. + +Rudiments of various muscles have been observed in many parts of the +human body;[23] and not a few muscles, which are regularly present in +some of the lower animals can occasionally be detected in man in a +greatly reduced condition. Every one must have noticed the power which +many animals, especially horses, possess of moving or twitching their +skin; and this is effected by the panniculus carnosus. Remnants of this +muscle in an efficient state are found in various parts of our bodies; +for instance, on the forehead, by which the eyebrows are raised. The +_platysma myoides_, which is well developed on the neck, belongs to this +system, but cannot be voluntarily brought into action. Prof. Turner, of +Edinburgh, has occasionally detected, as he informs me, muscular +fasciculi in five different situations, namely in the axillæ, near the +scapulæ, &c., all of which must be referred to the system of the +panniculus. He has also shewn[24] that the _musculus sternalis_ or +_sternalis brutorum_, which is not an extension of the _rectus +abdominalis_, but is closely allied to the panniculus, occurred in the +proportion of about 3 per cent. in upwards of 600 bodies: he adds, that +this muscle affords “an excellent illustration of the statement that +occasional and rudimentary structures are especially liable to variation +in arrangement.” + +Some few persons have the power of contracting the superficial muscles +on their scalps; and these muscles are in a variable and partially +rudimentary condition. M. A. de Candolle has communicated to me a +curious instance of the long-continued persistence or inheritance of +this power, as well as of its unusual development. He knows a family, in +which one member, the present head of a family, could, when a youth, +pitch several heavy books from his head by the movement of the scalp +alone; and he won wagers by performing this feat. His father, uncle, +grandfather, and all his three children possess the same power to the +same unusual degree. This family became divided eight generations ago +into two branches; so that the head of the above-mentioned branch is +cousin in the seventh degree to the head of the other branch. This +distant cousin resides in another part of France, and on being asked +whether he possessed the same faculty, immediately exhibited his power. +This case offers a good illustration how persistently an absolutely +useless faculty may be transmitted. + +The extrinsic muscles which serve to move the whole external ear, and +the intrinsic muscles which move the different parts, all of which +belong to the system of the panniculus, are in a rudimentary condition +in man; they are also variable in development, or at least in function. +I have seen one man who could draw his ears forwards, and another who +could draw them backwards;[25] and from what one of these persons told +me, it is probable that most of us by often touching our ears and thus +directing our attention towards them, could by repeated trials recover +some power of movement. The faculty of erecting the ears and of +directing them to different points of the compass, is no doubt of the +highest service to many animals, as they thus perceive the point of +danger; but I have never heard of a man who possessed the least power of +erecting his ears,—the one movement which might be of use to him. The +whole external shell of the ear may be considered a rudiment, together +with the various folds and prominences (helix and anti-helix, tragus and +anti-tragus, &c.) which in the lower animals strengthen and support the +ear when erect, without adding much to its weight. Some authors, +however, suppose that the cartilage of the shell serves to transmit +vibrations to the acoustic nerve; but Mr. Toynbee,[26] after collecting +all the known evidence on this head, concludes that the external shell +is of no distinct use. The ears of the chimpanzee and orang are +curiously like those of man, and I am assured by the keepers in the +Zoological Gardens that these animals never move or erect them; so that +they are in an equally rudimentary condition, as far as function is +concerned, as in man. Why these animals, as well as the progenitors of +man, should have lost the power of erecting their ears we cannot say. It +may be, though I am not quite satisfied with this view, that owing to +their arboreal habits and great strength they were but little exposed to +danger, and so during a lengthened period moved their ears but little, +and thus gradually lost the power of moving them. This would be a +parallel case with that of those large and heavy birds, which from +inhabiting oceanic islands have not been exposed to the attacks of +beasts of prey, and have consequently lost the power of using their +wings for flight. + +[Illustration: Fig. 2. Human Ear, modelled and drawn by Mr. Woolner. + +_a._ The projecting point.] + +The celebrated sculptor, Mr. Woolner, informs me of one little +peculiarity in the external ear, which he has often observed both in men +and women, and of which he perceived the full signification. His +attention was first called to the subject whilst at work on his figure +of Puck, to which he had given pointed ears. He was thus led to examine +the ears of various monkeys, and subsequently more carefully those of +man. The peculiarity consists in a little blunt point, projecting from +the inwardly folded margin, or helix. Mr. Woolner made an exact model of +one such case, and has sent me the accompanying drawing. (Fig. 2.) These +points not only project inwards, but often a little outwards, so that +they are visible when the head is viewed from directly in front or +behind. They are variable in size and somewhat in position, standing +either a little higher or lower; and they sometimes occur on one ear and +not on the other. Now the meaning of these projections is not, I think, +doubtful; but it may be thought that they offer too trifling a character +to be worth notice. This thought, however, is as false as it is natural. +Every character, however slight, must be the result of some definite +cause; and if it occurs in many individuals deserves consideration. The +helix obviously consists of the extreme margin of the ear folded +inwards; and this folding appears to be in some manner connected with +the whole external ear being permanently pressed backwards. In many +monkeys, which do not stand high in the order, as baboons and some +species of macacus,[27] the upper portion of the ear is slightly +pointed, and the margin is not at all folded inwards; but if the margin +were to be thus folded, a slight point would necessarily project inwards +and probably a little outwards. This could actually be observed in a +specimen of the _Ateles beelzebuth_ in the Zoological Gardens; and we +may safely conclude that it is a similar structure—a vestige of +formerly pointed ears—which occasionally reappears in man. + +The nictitating membrane, or third eyelid, with its accessory muscles +and other structures, is especially well developed in birds, and is of +much functional importance to them, as it can be rapidly drawn across +the whole eye-ball. It is found in some reptiles and amphibians, and in +certain fishes, as in sharks. It is fairly well developed in the two +lower divisions of the mammalian series, namely, in the monotremata and +marsupials, and in some few of the higher mammals, as in the walrus. But +in man, the quadrumana, and most other mammals, it exists, as is +admitted by all anatomists, as a mere rudiment, called the semilunar +fold.[28] + +The sense of smell is of the highest importance to the greater number of +mammals—to some, as the ruminants, in warning them of danger; to +others, as the carnivora, in finding their prey; to others, as the wild +boar, for both purposes combined. But the sense of smell is of extremely +slight service, if any, even to savages, in whom it is generally more +highly developed than in the civilised races. It does not warn them of +danger, nor guide them to their food; nor does it prevent the Esquimaux +from sleeping in the most fetid atmosphere, nor many savages from eating +half-putrid meat. Those who believe in the principle of gradual +evolution, will not readily admit that this sense in its present state +was originally acquired by man, as he now exists. No doubt he inherits +the power in _an_ enfeebled and so far rudimentary condition, from some +early progenitor, to whom it was highly serviceable and by whom it was +continually used. We can thus perhaps understand how it is, as Dr. +Maudsley has truly remarked,[29] that the sense of smell in man “is +singularly effective in recalling vividly the ideas and images of +forgotten scenes and places;” for we see in those animals, which have +this sense highly developed, such as dogs and horses, that old +recollections of persons and places are strongly associated with their +odour. + +Man differs conspicuously from all the other Primates in being almost +naked. But a few short straggling hairs are found over the greater part +of the body in the male sex, and fine down on that of the female sex, In +individuals belonging to the same race these hairs are highly variable, +not only in abundance, but likewise in position: thus the shoulders in +some Europeans are quite naked, whilst in others they bear thick tufts +of hair.[30] There can be little doubt that the hairs thus scattered +over the body are the rudiments of the uniform hairy coat of the lower +animals. This view is rendered all the more probable, as it is known +that fine, short, and pale-coloured hairs on the limbs and other parts +of the body occasionally become developed into “thickset, long, and +rather coarse dark hairs,” when abnormally nourished near old-standing +inflamed surfaces.[31] + +I am informed by Mr. Paget that persons belonging to the same family +often have a few hairs in their eyebrows much longer than the others; so +that this slight peculiarity seems to be inherited. These hairs +apparently represent the vibrissæ, which are used as organs of touch by +many of the lower animals. In a young chimpanzee I observed that a few +upright, rather long, hairs, projected above the eyes, where the true +eyebrows, if present, would have stood. + +The fine wool-like hair, or so-called lanugo, with which the human +fœtus during the sixth month is thickly covered, offers a more +curious case. It is first developed, during the fifth month, on the +eyebrows and face, and especially round the mouth, where it is much +longer than that on the head. A moustache of this kind was observed by +Eschricht[32] on a female fœtus; but this is not so surprising a +circumstance as it may at first appear, for the two sexes generally +resemble each other in all external characters during an early period of +growth. The direction and arrangement of the hairs on all parts of the +fœtal body are the same as in the adult, but are subject to much +variability. The whole surface, including even the forehead and ears, is +thus thickly clothed; but it is a significant fact that the palms of the +hands and the soles of the feet are quite naked, like the inferior +surfaces of all four extremities in most of the lower animals. As this +can hardly be an accidental coincidence, we must consider the woolly +covering of the fœtus to be the rudimental representative of the +first permanent coat of hair in those mammals which are born hairy. This +representation is much more complete, in accordance with the usual law +of embryological development, than that afforded by the straggling hairs +on the body of the adult. + +It appears as if the posterior molar or wisdom-teeth were tending to +become rudimentary in the more civilised races of man. These teeth are +rather smaller than the other molars, as is likewise the case with the +corresponding teeth in the chimpanzee and orang; and they have only two +separate fangs. They do not cut through the gums till about the +seventeenth year, and I am assured by dentists that they are much more +liable to decay, and are earlier lost, than the other teeth. It is also +remarkable that they are much more liable to vary both in structure and +in the period of their development than the other teeth.[33] In the +Melanian races, on the other hand, the wisdom-teeth are usually +furnished with three separate fangs, and are generally sound: they also +differ from the other molars in size less than in the Caucasian +races.[34] Prof. Schaaffhausen accounts for this difference between the +races by “the posterior dental portion of the jaw being always +shortened” in those that are civilised,[35] and this shortening may, I +presume, be safely attributed to civilised men habitually feeding on +soft, cooked food, and thus using their jaws less. I am informed by Mr. +Brace that it is becoming quite a common practice in the United States +to remove some of the molar teeth of children, as the jaw does not grow +large enough for the perfect development of the normal number. + +With respect to the alimentary canal I have met with an account of only +a single rudiment, namely the vermiform appendage of the cæcum. The +cæcum is a branch or diverticulum of the intestine, ending in a +cul-de-sac, and it is extremely long in many of the lower +vegetable-feeding mammals. In the marsupial koala it is actually more +than thrice as long as the whole body.[36] It is sometimes produced into +a long gradually-tapering point, and is sometimes constricted in parts. +It appears as if, in consequence of changed diet or habits, the cæcum +had become much shortened in various animals, the vermiform appendage +being left as a rudiment of the shortened part. That this appendage is a +rudiment, we may infer from its small size, and from the evidence which +Prof. Canestrini[37] has collected of its variability in man. It is +occasionally quite absent, or again is largely developed. The passage is +sometimes completely closed for half or two-thirds of its length, with +the terminal part consisting of a flattened solid expansion. In the +orang this appendage is long and convoluted: in man it arises from the +end of the short cæcum, and is commonly from four to five inches in +length, being only about the third of an inch in diameter. Not only is +it useless, but it is sometimes the cause of death, of which fact I have +lately heard two instances: this is due to small hard bodies, such as +seeds, entering the passage and causing inflammation.[38] + +In the Quadrumana and some other orders of mammals, especially in the +Carnivora, there is a passage near the lower end of the humerus, called +the supra-condyloid foramen, through which the great nerve of the fore +limb passes, and often the great artery. Now in the humerus of man, as +Dr. Struthers[39] and others have shewn, there is generally a trace of +this passage, and it is sometimes fairly well developed, being formed by +a depending hook-like process of bone, completed by a band of ligament. +When present the great nerve invariably passes through it, and this +clearly indicates that it is the homologue and rudiment of the +supra-condyloid foramen of the lower animals. Prof. Turner estimates, as +he informs me, that it occurs in about one per cent. of recent +skeletons; but during ancient times it appears to have been much more +common. Mr. Busk[40] has collected the following evidence on this head: +Prof. Broca “noticed the perforation in four and a half per cent. of the +arm-bones collected in the ‘Cimetière du Sud’ at Paris; and in the +Grotto of Orrony, the contents of which are referred to the Bronze +period, as many as eight humeri out of thirty-two were perforated; but +this extraordinary proportion, he thinks, might be due to the cavern +having been a sort of ‘family vault.’ Again, M. Dupont found 30 per +cent. of perforated bones in the caves of the Valley of the Lesse, +belonging to the Reindeer period; whilst M. Leguay, in a sort of +_dolmen_ at Argenteuil, observed twenty-five per cent. to be perforated; +and M. Pruner-Bey found twenty-six per cent. in the same condition in +bones from Vauréal. Nor should it be left unnoticed that M. Pruner-Bey +states that this condition is common in Guanche skeletons.” The fact +that ancient races, in this and several other cases, more frequently +present structures which resemble those of the lower animals than do the +modern races, is interesting. One chief cause seems to be that ancient +races stand somewhat nearer than modern races in the long line of +descent to their remote animal-like progenitors. + +The os coccyx in man, though functionless as a tail, plainly represents +this part in other vertebrate animals. At an early embryonic period it +is free, and, as we have seen, projects beyond the lower extremities. In +certain rare and anomalous cases it has been known, according to Isidore +Geoffroy St.-Hilaire and others,[41] to form a small external rudiment +of a tail. The os coccyx is short, usually including only four vertebræ: +and these are in a rudimental condition, for they consist, with the +exception of the basal one, of the centrum alone.[42] They are furnished +with some small muscles; one of which, as I am informed by Prof. Turner, +has been expressly described by Theile as a rudimentary repetition of +the extensor of the tail, which is so largely developed in many mammals. + +The spinal cord in man extends only as far downwards as the last dorsal +or first lumbar vertebra; but a thread-like structure (the _filum +terminale_) runs down the axis of the sacral part of the spinal canal, +and even along the back of the coccygeal bones. The upper part of this +filament, as Prof. Turner informs me, is undoubtedly homologous with the +spinal cord; but the lower part apparently consists merely of the _pia +mater_, or vascular investing membrane. Even in this case the os coccyx +may be said to possess a vestige of so important a structure as the +spinal cord, though no longer enclosed within a bony canal. The +following fact, for which I am also indebted to Prof. Turner, shews how +closely the os coccyx corresponds with the true tail in the lower +animals: Luschka has recently discovered at the extremity of the +coccygeal bones a very peculiar convoluted body, which is continuous +with the middle sacral artery; and this discovery led Krause and Meyer +to examine the tail of a monkey (Macacus) and of a cat, in both of which +they found, though not at the extremity, a similarly convoluted body. + +The reproductive system offers various rudimentary structures; but these +differ in one important respect from the foregoing cases. We are not +here concerned with a vestige of a part which does not belong to the +species in an efficient state; but with a part which is always present +and efficient in the one sex, being represented in the other by a mere +rudiment. Nevertheless, the occurrence of such rudiments is as difficult +to explain on the belief of the separate creation of each species, as in +the foregoing cases. Hereafter I shall have to recur to these rudiments, +and shall shew that their presence generally depends merely on +inheritance; namely, on parts acquired by one sex having been partially +transmitted to the other. Here I will only give some instances of such +rudiments. It is well known that in the males of all mammals, including +man, rudimentary mammæ exist. These in several instances have become +well developed, and have yielded a copious supply of milk. Their +essential identity in the two sexes is likewise shewn by their +occasional sympathetic enlargement in both during an attack of the +measles. The _vesicula prostatica_, which has been observed in many +male mammals, is now universally acknowledged to be the homologue of the +female uterus, together with the connected passage. It is impossible to +read Leuckart’s able description of this organ, and his reasoning, +without admitting the justness of his conclusion. This is especially +clear in the case of those mammals in which the true female uterus +bifurcates, for in the males of these the vesicula likewise +bifurcates.[43] Some additional rudimentary structures belonging to the +reproductive system might here have been adduced.[44] + + +The bearing of the three great classes of facts now given is +unmistakeable. But it would be superfluous here fully to recapitulate +the line of argument given in detail in my ‘Origin of Species.’ The +homological construction of the whole frame in the members of the same +class is intelligible, if we admit their descent from a common +progenitor, together with their subsequent adaptation to diversified +conditions. On any other view the similarity of pattern between the hand +of a man or monkey, the foot of a horse, the flipper of a seal, the wing +of a bat, &c., is utterly inexplicable. It is no scientific explanation +to assert that they have all been formed on the same ideal plan. With +respect to development, we can clearly understand, on the principle of +variations supervening at a rather late embryonic period, and being +inherited at a corresponding period, how it is that the embryos of +wonderfully different forms should still retain, more or less perfectly, +the structure of their common progenitor. No other explanation has ever +been given of the marvellous fact that the embryo of a man, dog, seal, +bat, reptile, &c., can at first hardly be distinguished from each other. +In order to understand the existence of rudimentary organs, we have only +to suppose that a former progenitor possessed the parts in question in a +perfect state, and that under changed habits of life they became greatly +reduced, either from simple disuse, or through the natural selection of +those individuals which were least encumbered with a superfluous part, +aided by the other means previously indicated. + +Thus we can understand how it has come to pass that man and all other +vertebrate animals have been constructed on the same general model, why +they pass through the same early stages of development, and why they +retain certain rudiments in common. Consequently we ought frankly to +admit their community of descent: to take any other view, is to admit +that our own structure and that of all the animals around us, is a mere +snare laid to entrap our judgment. This conclusion is greatly +strengthened, if we look to the members of the whole animal series, and +consider the evidence derived from their affinities or classification, +their geographical distribution and geological succession. It is only +our natural prejudice, and that arrogance which made our forefathers +declare that they were descended from demi-gods, which leads us to demur +to this conclusion. But the time will before long come when it will be +thought wonderful, that naturalists, who were well acquainted with the +comparative structure and development of man and other mammals, should +have believed that each was the work of a separate act of creation. + + + + +CHAPTER II. + +COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER ANIMALS. + + + The difference in mental power between the highest ape and the + lowest savage, immense—Certain instincts in common—The + emotions—Curiosity—Imitation—Attention—Memory—Imagination + —Reason—Progressive improvement—Tools and weapons used by + animals—Language—Self-consciousness—Sense of beauty—Belief + in God, spiritual agencies, superstitions. + + +We have seen in the last chapter that man bears in his bodily structure +clear traces of his descent from some lower form; but it may be urged +that, as man differs so greatly in his mental power from all other +animals, there must be some error in this conclusion. No doubt the +difference in this respect is enormous, even if we compare the mind of +one of the lowest savages, who has no words to express any number higher +than four, and who uses no abstract terms for the commonest objects or +affections,[45] with that of the most highly organised ape. The +difference would, no doubt, still remain immense, even if one of the +higher apes had been improved or civilised as much as a dog has been in +comparison with its parent-form, the wolf or jackal. The Fuegians rank +amongst the lowest barbarians; but I was continually struck with +surprise how closely the three natives on board H.M.S. “Beagle,” who had +lived some years in England and could talk a little English, resembled +us in disposition and in most of our mental faculties. If no organic +being excepting man had possessed any mental power, or if his powers had +been of a wholly different nature from those of the lower animals, then +we should never have been able to convince ourselves that our high +faculties had been gradually developed. But it can be clearly shewn that +there is no fundamental difference of this kind. We must also admit that +there is a much wider interval in mental power between one of the lowest +fishes, as a lamprey or lancelet, and one of the higher apes, than +between an ape and man; yet this immense interval is filled up by +numberless gradations. + +Nor is the difference slight in moral disposition between a barbarian, +such as the man described by the old navigator Byron, who dashed his +child on the rocks for dropping a basket of sea-urchins, and a Howard or +Clarkson; and in intellect, between a savage who does not use any +abstract terms, and a Newton or Shakspeare. Differences of this kind +between the highest men of the highest races and the lowest savages, are +connected by the finest gradations. Therefore it is possible that they +might pass and be developed into each other. + +My object in this chapter is solely to shew that there is no fundamental +difference between man and the higher mammals in their mental faculties. +Each division of the subject might have been extended into a separate +essay, but must here be treated briefly. As no classification of the +mental powers has been universally accepted, I shall arrange my remarks +in the order most convenient for my purpose; and will select those facts +which have most struck me, with the hope that they may produce some +effect on the reader. + +With respect to animals very low in the scale, I shall have to give some +additional facts under Sexual Selection, shewing that their mental +powers are higher than might have been expected. The variability of the +faculties in the individuals of the same species is an important point +for us, and some few illustrations will here be given. But it would be +superfluous to enter into many details on this head, for I have found on +frequent enquiry, that it is the unanimous opinion of all those who have +long attended to animals of many kinds, including birds, that the +individuals differ greatly in every mental characteristic. In what +manner the mental powers were first developed in the lowest organisms, +is as hopeless an enquiry as how life first originated. These are +problems for the distant future, if they are ever to be solved by man. + +As man possesses the same senses with the lower animals, his fundamental +intuitions must be the same. Man has also some few instincts in common, +as that of self-preservation, sexual love, the love of the mother for +her new-born offspring, the power possessed by the latter of sucking, +and so forth. But man, perhaps, has somewhat fewer instincts than those +possessed by the animals which come next to him in the series. The orang +in the Eastern islands, and the chimpanzee in Africa, build platforms on +which they sleep; and, as both species follow the same habit, it might +be argued that this was due to instinct, but we cannot feel sure that it +is not the result of both animals having similar wants and possessing +similar powers of reasoning. These apes, as we may assume, avoid the +many poisonous fruits of the tropics, and man has no such knowledge; but +as our domestic animals, when taken to foreign lands and when first +turned out in the spring, often eat poisonous herbs, which they +afterwards avoid, we cannot feel sure that the apes do not learn from +their own experience or from that of their parents what fruits to +select. It is however certain, as we shall presently see, that apes +have an instinctive dread of serpents, and probably of other dangerous +animals. + +The fewness and the comparative simplicity of the instincts in the +higher animals are remarkable in contrast with those of the lower +animals. Cuvier maintained that instinct and intelligence stand in an +inverse ratio to each other; and some have thought that the intellectual +faculties of the higher animals have been gradually developed from their +instincts. But Pouchet, in an interesting essay,[46] has shewn that no +such inverse ratio really exists. Those insects which possess the most +wonderful instincts are certainly the most intelligent. In the +vertebrate series, the least intelligent members, namely fishes and +amphibians, do not possess complex instincts; and amongst mammals the +animal most remarkable for its instincts, namely the beaver, is highly +intelligent, as will be admitted by every one who has read Mr. Morgan’s +excellent account of this animal.[47] + +Although the first dawnings of intelligence, according to Mr. Herbert +Spencer,[48] have been developed through the multiplication and +co-ordination of reflex actions, and although many of the simpler +instincts graduate into actions of this kind and can hardly be +distinguished from them, as in the case of young animals sucking, yet +the more complex instincts seem to have originated independently of +intelligence. I am, however, far from wishing to deny that instinctive +actions may lose their fixed and untaught character, and be replaced by +others performed by the aid of the free will. On the other hand, some +intelligent actions—as when, birds on oceanic islands first learn to +avoid man—after being performed during many generations, become +converted into instincts and are inherited. They may then be said to be +degraded in character, for they are no longer performed through reason +or from experience. But the greater number of the more complex instincts +appear to have been gained in a wholly different manner, through the +natural selection of variations of simpler instinctive actions. Such +variations appear to arise from the same unknown causes acting on the +cerebral organisation, which induce slight variations or individual +differences in other parts of the body; and these variations, owing to +our ignorance, are often said to arise spontaneously. We can, I think, +come to no other conclusion with respect to the origin of the more +complex instincts, when we reflect on the marvellous instincts of +sterile worker-ants and bees, which leave no offspring to inherit the +effects of experience and of modified habits. + +Although a high degree of intelligence is certainly compatible with the +existence of complex instincts, as we see in the insects just named and +in the beaver, it is not improbable that they may to a certain extent +interfere with each other’s development. Little is known about the +functions of the brain, but we can perceive that as the intellectual +powers become highly developed, the various parts of the brain must be +connected by the most intricate channels of intercommunication; and as a +consequence each separate part would perhaps tend to become less well +fitted to answer in a definite and uniform, that is instinctive, manner +to particular sensations or associations. + +I have thought this digression worth giving, because we may easily +underrate the mental powers of the higher animals, and especially of +man, when we compare their actions founded on the memory of past events, +on foresight, reason, and imagination, with exactly similar actions +instinctively performed by the lower animals; in this latter case the +capacity of performing such actions having been gained, step by step, +through the variability of the mental organs and natural selection, +without any conscious intelligence on the part of the animal during each +successive generation. No doubt, as Mr. Wallace has argued,[49] much of +the intelligent work done by man is due to imitation and not to reason; +but there is this great difference between his actions and many of those +performed by the lower animals, namely, that man cannot, on his first +trial, make, for instance, a stone hatchet or a canoe, through his power +of imitation. He has to learn his work by practice; a beaver, on the +other hand, can make its dam or canal, and a bird its nest, as well, or +nearly as well, the first time it tries, as when old and experienced. + +To return to our immediate subject: the lower animals, like man, +manifestly feel pleasure and pain, happiness and misery. Happiness is +never better exhibited than by young animals, such as puppies, kittens, +lambs, &c., when playing together, like our own children. Even insects +play together, as has been described by that excellent observer, P. +Huber,[50] who saw ants chasing and pretending to bite each other, like +so many puppies. + +The fact that the lower animals are excited by the same emotions as +ourselves is so well established, that it will not be necessary to weary +the reader by many details. Terror acts in the same manner on them as on +us, causing the muscles to tremble, the heart to palpitate, the +sphincters to be relaxed, and the hair to stand on end. Suspicion, the +offspring of fear, is eminently characteristic of most wild animals. +Courage and timidity are extremely variable qualities in the +individuals of the same species, as is plainly seen in our dogs. Some +dogs and horses are ill-tempered and easily turn sulky; others are +good-tempered; and these qualities are certainly inherited. Every one +knows how liable animals are to furious rage, and how plainly they show +it. Many anecdotes, probably true, have been published on the +long-delayed and artful revenge of various animals. The accurate Rengger +and Brehm[51] state that the American and African monkeys which they +kept tame, certainly revenged themselves. The love of a dog for his +master is notorious; in the agony of death he has been known to caress +his master, and every one has heard of the dog suffering under +vivisection, who licked the hand of the operator; this man, unless he +had a heart of stone, must have felt remorse to the last hour of his +life. As Whewell[52] has remarked, “who that reads the touching +instances of maternal affection, related so often of the women of all +nations, and of the females of all animals, can doubt that the principle +of action is the same in the two cases?” + +We see maternal affection exhibited in the most trifling details; thus +Rengger observed an American monkey (a Cebus) carefully driving away the +flies which plagued her infant; and Duvaucel saw a Hylobates washing the +faces of her young ones in a stream. So intense is the grief of female +monkeys for the loss of their young, that it invariably caused the death +of certain kinds kept under confinement by Brehm in N. Africa. +Orphan-monkeys were always adopted and carefully guarded by the other +monkeys, both males and females. One female baboon had so capacious a +heart that she not only adopted young monkeys of other species, but +stole young dogs and cats, which she continually carried about. Her +kindness, however, did not go so far as to share her food with her +adopted offspring, at which Brehm was surprised, as his monkeys always +divided everything quite fairly with their own young ones. An adopted +kitten scratched the above-mentioned affectionate baboon, who certainly +had a fine intellect, for she was much astonished at being scratched, +and immediately examined the kitten’s feet, and without more ado bit off +the claws. In the Zoological Gardens, I heard from the keeper that an +old baboon (_C. chacma_) had adopted a Rhesus monkey; but when a young +drill and mandrill were placed in the cage, she seemed to perceive that +these monkeys, though distinct species, were her nearer relatives, for +she at once rejected the Rhesus and adopted both of them. The young +Rhesus, as I saw, was greatly discontented at being thus rejected, and +it would, like a naughty child, annoy and attack the young drill and +mandrill whenever it could do so with safety; this conduct exciting +great indignation in the old baboon. Monkeys will also, according to +Brehm, defend their master when attacked by any one, as well as dogs to +whom they are attached, from the attacks of other dogs. But we here +trench on the subject of sympathy, to which I shall recur. Some of +Brehm’s monkeys took much delight in teasing, in various ingenious ways, +a certain old dog whom they disliked, as well as other animals. + +Most of the more complex emotions are common to the higher animals and +ourselves. Every one has seen how jealous a dog is of his master’s +affection, if lavished on any other creature; and I have observed the +same fact with monkeys. This shews that animals not only love, but have +the desire to be loved. Animals manifestly feel emulation. They love +approbation or praise; and a dog carrying a basket for his master +exhibits in a high degree self-complacency or pride. There can, I think, +be no doubt that a dog feels shame, as distinct from fear, and something +very like modesty when begging too often for food. A great dog scorns +the snarling of a little dog, and this may be called magnanimity. +Several observers have stated that monkeys certainly dislike being +laughed at; and they sometimes invent imaginary offences. In the +Zoological Gardens I saw a baboon who always got into a furious rage +when his keeper took out a letter or book and read it aloud to him; and +his rage was so violent that, as I witnessed on one occasion, he bit his +own leg till the blood flowed. + +We will now turn to the more intellectual emotions and faculties, which +are very important, as forming the basis for the development of the +higher mental powers. Animals manifestly enjoy excitement and suffer +from ennui, as may be seen with dogs, and, according to Rengger, with +monkeys. All animals feel Wonder, and many exhibit Curiosity. They +sometimes suffer from this latter quality, as when the hunter plays +antics and thus attracts them; I have witnessed this with deer, and so +it is with the wary chamois, and with some kinds of wild-ducks. Brehm +gives a curious account of the instinctive dread which his monkeys +exhibited towards snakes; but their curiosity was so great that they +could not desist from occasionally satiating their horror in a most +human fashion, by lifting up the lid of the box in which the snakes were +kept. I was so much surprised at his account, that I took a stuffed and +coiled-up snake into the monkey-house at the Zoological Gardens, and the +excitement thus caused was one of the most curious spectacles which I +ever beheld. Three species of Cercopithecus were the most alarmed; they +dashed about their cages and uttered sharp signal-cries of danger, which +were understood by the other monkeys. A few young monkeys and one old +Anubis baboon alone took no notice of the snake. I then placed the +stuffed specimen on the ground in one of the larger compartments. After +a time all the monkeys collected round it in a large circle, and staring +intently, presented a most ludicrous appearance. They became extremely +nervous; so that when a wooden ball, with which they were familiar as a +plaything, was accidently moved in the straw, under which it was partly +hidden, they all instantly started away. These monkeys behaved very +differently when a dead fish, a mouse, and some other new objects were +placed in their cages; for though at first frightened, they soon +approached, handled and examined them. I then placed a live snake in a +paper bag, with the mouth loosely closed, in one of the larger +compartments. One of the monkeys immediately approached, cautiously +opened the bag a little, peeped in, and instantly dashed away. Then I +witnessed what Brehm has described, for monkey after monkey, with head +raised high and turned on one side, could not resist taking momentary +peeps into the upright bag, at the dreadful object lying quiet at the +bottom. It would almost appear as if monkeys had some notion of +zoological affinities, for those kept by Brehm exhibited a strange, +though mistaken, instinctive dread of innocent lizards and frogs. An +orang, also, has been known to be much alarmed at the first sight of a +turtle.[53] + +The principle of _Imitation_ is strong in man, and especially in man in +a barbarous state. Desor[54] has remarked that no animal voluntarily +imitates an action performed by man, until in the ascending scale we +come to monkeys, which are well-known to be ridiculous mockers. Animals, +however, sometimes imitate each others’ actions: thus two species of +wolves, which had been reared by dogs, learned to bark, as does +sometimes the jackal,[55] but whether this can be called voluntary +imitation is another question. From one account which I have read, there +is reason to believe that puppies nursed by cats sometimes learn to lick +their feet and thus to clean their faces: it is at least certain, as I +hear from a perfectly trustworthy friend, that some dogs behave in this +manner. Birds imitate the songs of their parents, and sometimes those of +other birds; and parrots are notorious imitators of any sound which they +often hear. + +Hardly any faculty is more important for the intellectual progress of +man than the power of _Attention_. Animals clearly manifest this power, +as when a cat watches by a hole and prepares to spring on its prey. Wild +animals sometimes become so absorbed when thus engaged, that they may be +easily approached. Mr. Bartlett has given me a curious proof how +variable this faculty is in monkeys. A man who trains monkeys to act +used to purchase common kinds from the Zoological Society at the price +of five pounds for each; but he offered to give double the price, if he +might keep three or four of them for a few days, in order to select one. +When asked how he could possibly so soon learn whether a particular +monkey would turn out a good actor, he answered that it all depended on +their power of attention. If when he was talking and explaining anything +to a monkey, its attention was easily distracted, as by a fly on the +wall or other trifling object, the case was hopeless. If he tried by +punishment to make an inattentive monkey act, it turned sulky. On the +other hand, a monkey which carefully attended to him could always be +trained. + +It is almost superfluous to state that animals have excellent _Memories_ +for persons and places. A baboon at the Cape of Good Hope, as I have +been informed by Sir Andrew Smith, recognised him with joy after an +absence of nine months. I had a dog who was savage and averse to all +strangers, and I purposely tried his memory after an absence of five +years and two days. I went near the stable where he lived, and shouted +to him in my old manner; he showed no joy, but instantly followed me out +walking and obeyed me, exactly as if I had parted with him only +half-an-hour before. A train of old associations, dormant during five +years, had thus been instantaneously awakened in his mind. Even ants, as +P. Huber[56] has clearly shewn, recognised their fellow-ants belonging +to the same community after a separation of four months. Animals can +certainly by some means judge of the intervals of time between recurrent +events. + +The _Imagination_ is one of the highest prerogatives of man. By this +faculty he unites, independently of the will, former images and ideas, +and thus creates brilliant and novel results. A poet, as Jean Paul +Richter remarks,[57] “who must reflect whether he shall make a +character say yes or no—to the devil with him; he is only a stupid +corpse.” Dreaming gives us the best notion of this power; as Jean Paul +again says, “The dream is an involuntary art of poetry.” The value of +the products of our imagination depends of course on the number, +accuracy, and clearness of our impressions; on our judgment and taste in +selecting or rejecting the involuntary combinations, and to a certain +extent on our power of voluntarily combining them. As dogs, cats, +horses, and probably all the higher animals, even birds, as is stated on +good authority,[58] have vivid dreams, and this is shewn by their +movements and voice, we must admit that they possess some power of +imagination. + +Of all the faculties of the human mind, it will, I presume, be admitted +that _Reason_ stands at the summit. Few persons any longer dispute that +animals possess some power of reasoning. Animals may constantly be seen +to pause, deliberate, and resolve. It is a significant fact, that the +more the habits of any particular animal are studied by a naturalist, +the more he attributes to reason and the less to unlearnt instincts.[59] +In future chapters we shall see that some animals extremely low in the +scale apparently display a certain amount of reason. No doubt it is +often difficult to distinguish between the power of reason and that of +instinct. Thus Dr. Hayes, in his work on ‘The Open Polar Sea,’ +repeatedly remarks that his dogs, instead of continuing to draw the +sledges in a compact body, diverged and separated when they came to thin +ice, so that their weight might be more evenly distributed. This was +often the first warning and notice which the travellers received that +the ice was becoming thin and dangerous. Now, did the dogs act thus from +the experience of each individual, or from the example of the older and +wiser dogs, or from an inherited habit, that is from an instinct? This +instinct might possibly have arisen since the time, long ago, when dogs +were first employed by the natives in drawing their sledges; or the +Arctic wolves, the parent-stock of the Esquimaux dog, may have acquired +this instinct, impelling them not to attack their prey in a close pack +when on thin ice. Questions of this kind are most difficult to answer. + +So many facts have been recorded in various works shewing that animals +possess some degree of reason, that I will here give only two or three +instances, authenticated by Rengger, and relating to American monkeys, +which stand low in their order. He states that when he first gave eggs +to his monkeys, they smashed them and thus lost much of their contents; +afterwards they gently hit one end against some hard body, and picked +off the bits of shell with their fingers. After cutting themselves only +once with any sharp tool, they would not touch it again, or would handle +it with the greatest care. Lumps of sugar were often given them wrapped +up in paper; and Rengger sometimes put a live wasp in the paper, so that +in hastily unfolding it they got stung; after this had once happened, +they always first held the packet to their ears to detect any movement +within. Any one who is not convinced by such facts as these, and by what +he may observe with his own dogs, that animals can reason, would not be +convinced by anything that I could add. Nevertheless I will give one +case with respect to dogs, as it rests on two distinct observers, and +can hardly depend on the modification of any instinct. + +Mr. Colquhoun[60] winged two wild-ducks, which fell on the opposite side +of a stream; his retriever tried to bring over both at once, but could +not succeed; she then, though never before known to ruffle a feather, +deliberately killed one, brought over the other, and returned for the +dead bird. Col. Hutchinson relates that two partridges were shot at +once, one being killed, the other wounded; the latter ran away, and was +caught by the retriever, who on her return came across the dead bird; +“she stopped, evidently greatly puzzled, and after one or two trials, +finding she could not take it up without permitting the escape of the +winged bird, she considered a moment, then deliberately murdered it by +giving it a severe crunch, and afterwards brought away both together. +This was the only known instance of her ever having wilfully injured any +game.” Here we have reason, though not quite perfect, for the retriever +might have brought the wounded bird first and then returned for the dead +one, as in the case of the two wild-ducks. + +The muleteers in S. America say, “I will not give you the mule whose +step is easiest, but _la mas rational_,—the one that reasons best;” and +Humboldt[61] adds, “this popular expression, dictated by long +experience, combats the system of animated machines, better perhaps than +all the arguments of speculative philosophy.” + + +It has, I think, now been shewn that man and the higher animals, +especially the Primates, have some few instincts in common. All have the +same senses, intuitions and sensations—similar passions, affections, +and emotions, even the more complex ones; they feel wonder and +curiosity; they possess the same faculties of imitation, attention, +memory, imagination, and reason, though in very different degrees. +Nevertheless many authors have insisted that man is separated through +his mental faculties by an impassable barrier from all the lower +animals. I formerly made a collection of above a score of such +aphorisms, but they are not worth giving, as their wide difference and +number prove the difficulty, if not the impossibility, of the attempt. +It has been asserted that man alone is capable of progressive +improvement; that he alone makes use of tools or fire, domesticates +other animals, possesses property, or employs language; that no other +animal is self-conscious, comprehends itself, has the power of +abstraction, or possesses general ideas; that man alone has a sense of +beauty, is liable to caprice, has the feeling of gratitude, mystery, +&c.; believes in God, or is endowed with a conscience. I will hazard a +few remarks on the more important and interesting of these points. + +Archbishop Sumner formerly maintained[62] that man alone is capable of +progressive improvement. With animals, looking first to the individual, +every one who has had any experience in setting traps knows that young +animals can be caught much more easily than old ones; and they can be +much more easily approached by an enemy. Even with respect to old +animals, it is impossible to catch many in the same place and in the +same kind of trap, or to destroy them by the same kind of poison; yet it +is improbable that all should have partaken of the poison, and +impossible that all should have been caught in the trap. They must learn +caution by seeing their brethren caught or poisoned. In North America, +where the fur-bearing animals have long been pursued, they exhibit, +according to the unanimous testimony of all observers, an almost +incredible amount of sagacity, caution, and cunning; but trapping has +been there so long carried on that inheritance may have come into play. + +If we look to successive generations, or to the race, there is no doubt +that birds and other animals gradually both acquire and lose caution in +relation to man or other enemies;[63] and this caution is certainly in +chief part an inherited habit or instinct, but in part the result of +individual experience. A good observer, Leroy,[64] states that in +districts where foxes are much hunted, the young when they first leave +their burrows are incontestably much more wary than the old ones in +districts where they are not much disturbed. + +Our domestic dogs are descended from wolves and jackals,[65] and though +they may not have gained in cunning, and may have lost in waryness and +suspicion, yet they have progressed in certain moral qualities, such as +in affection, trust-worthiness, temper, and probably in general +intelligence. The common rat has conquered and beaten several other +species throughout Europe, in parts of North America, New Zealand, and +recently in Formosa, as well as on the mainland of China. Mr. +Swinhoe,[66] who describes these latter cases, attributes the victory of +the common rat over the large _Mus coninga_ to its superior cunning; and +this latter quality may be attributed to the habitual exercise of all +its faculties in avoiding extirpation by man, as well as to nearly all +the less cunning or weak-minded rats having been successively destroyed +by him. To maintain, independently of any direct evidence, that no +animal during the course of ages has progressed in intellect or other +mental faculties, is to beg the question of the evolution of species. +Hereafter we shall see that, according to Lartet, existing mammals +belonging to several orders have larger brains than their ancient +tertiary prototypes. + +It has often been said that no animal uses any tool; but the chimpanzee +in a state of nature cracks a native fruit, somewhat like a walnut, with +a stone.[67] Rengger[68] easily taught an American monkey thus to break +open hard palm-nuts, and afterwards of its own accord it used stones to +open other kinds of nuts, as well as boxes. It thus also removed the +soft rind of fruit that had a disagreeable flavour. Another monkey was +taught to open the lid of a large box with a stick, and afterwards it +used the stick as a lever to move heavy bodies; and I have myself seen a +young orang put a stick into a crevice, slip his hand to the other end, +and use it in the proper manner as a lever. In the cases just mentioned +stones and sticks were employed as implements; but they are likewise +used as weapons. Brehm[69] states, on the authority of the well-known +traveller Schimper, that in Abyssinia when the baboons belonging to one +species (_C. gelada_) descend in troops from the mountains to plunder +the fields, they sometimes encounter troops of another species (_C. +hamadryas_), and then a fight ensues. The Geladas roll down great +stones, which the Hamadryas try to avoid, and then, both species, +making a great uproar, rush furiously against each other. Brehm, when +accompanying the Duke of Coburg-Gotha, aided in an attack with fire-arms +on a troop of baboons in the pass of Mensa in Abyssinia. The baboons in +return rolled so many stones down the mountain, some as large as a man’s +head, that the attackers had to beat a hasty retreat; and the pass was +actually for a time closed against the caravan. It deserves notice that +these baboons thus acted in concert. Mr. Wallace[70] on three occasions +saw female orangs, accompanied by their young, “breaking off branches +and the great spiny fruit of the Durian tree, with every appearance of +rage; causing such a shower of missiles as effectually kept us from +approaching too near the tree.” + +In the Zoological Gardens a monkey which had weak teeth used to break +open nuts with a stone; and I was assured by the keepers that this +animal, after using the stone, hid it in the straw, and would not let +any other monkey touch it. Here, then, we have the idea of property; but +this idea is common to every dog with a bone, and to most or all birds +with their nests. + +The Duke of Argyll[71] remarks, that the fashioning of an implement for +a special purpose is absolutely peculiar to man; and he considers that +this forms an immeasurable gulf between him and the brutes. It is no +doubt a very important distinction, but there appears to me much truth +in Sir J. Lubbock’s suggestion,[72] that when primeval man first used +flint-stones for any purpose, he would have accidentally splintered +them, and would then have used the sharp fragments. From this step it +would be a small one to intentionally break the flints, and not a very +wide step to rudely fashion them. This latter advance, however, may have +taken long ages, if we may judge by the immense interval of time which +elapsed before the men of the neolithic period took to grinding and +polishing their stone tools. In breaking the flints, as Sir J. Lubbock +likewise remarks, sparks would have been emitted, and in grinding them +heat would have been evolved: “thus the two usual methods of obtaining +fire may have originated.” The nature of fire would have been known in +the many volcanic regions where lava occasionally flows through forests. +The anthropomorphous apes, guided probably by instinct, build for +themselves temporary platforms; but as many instincts are largely +controlled by reason, the simpler ones, such as this of building a +platform, might readily pass into a voluntary and conscious act. The +orang is known to cover itself at night with the leaves of the Pandanus; +and Brehm states that one of his baboons used to protect itself from the +heat of the sun by throwing a straw-mat over its head. In these latter +habits, we probably see the first steps towards some of the simpler +arts; namely rude architecture and dress, as they arose amongst the +early progenitors of man. + + +_Language._—This faculty has justly been considered as one of the chief +distinctions between man and the lower animals. But man, as a highly +competent judge, Archbishop Whately remarks, “is not the only animal +that can make use of language to express what is passing in his mind, +and can understand, more or less, what is so expressed by another.”[73] +In Paraguay the _Cebus Azaræ_ when excited utters at least six distinct +sounds, which excite in other monkeys similar emotions.[74] The +movements of the features and gestures of monkeys are understood by us, +and they partly understand ours, as Rengger and others declare. It is a +more remarkable fact that the dog, since being domesticated, has learnt +to bark[75] in at least four or five distinct tones. Although barking is +a new art, no doubt the wild species, the parents of the dog, expressed +their feelings by cries of various kinds. With the domesticated dog we +have the bark of eagerness, as in the chase; that of anger; the yelping +or howling bark of despair, as when shut up; that of joy, as when +starting on a walk with his master; and the very distinct one of demand +or supplication, as when wishing for a door or window to be opened. + +Articulate language is, however, peculiar to man; but he uses in common +with the lower animals inarticulate cries to express his meaning, aided +by gestures and the movements of the muscles of the face.[76] This +especially holds good with the more simple and vivid feelings, which are +but little connected with our higher intelligence. Our cries of pain, +fear, surprise, anger, together with their appropriate actions, and the +murmur of a mother to her beloved child, are more expressive than any +words. It is not the mere power of articulation that distinguishes man +from other animals, for as every one knows, parrots can talk; but it is +his large power of connecting definite sounds with definite ideas; and +this obviously depends on the development of the mental faculties. + +As Horne Tooke, one of the founders of the noble science of philology, +observes, language is an art, like brewing or baking; but writing would +have been a much more appropriate simile. It certainly is not a true +instinct, as every language has to be learnt. It differs, however, +widely from all ordinary arts, for man has an instinctive tendency to +speak, as we see in the babble of our young children; whilst no child +has an instinctive tendency to brew, bake, or write. Moreover, no +philologist now supposes that any language has been deliberately +invented; each has been slowly and unconsciously developed by many +steps. The sounds uttered by birds offer in several respects the nearest +analogy to language, for all the members of the same species utter the +same instinctive cries expressive of their emotions; and all the kinds +that have the power of singing exert this power instinctively; but the +actual song, and even the call-notes, are learnt from their parents or +foster-parents. These sounds, as Daines Barrington[77] has proved, “are +no more innate than language is in man.” The first attempts to sing “may +be compared to the imperfect endeavour in a child to babble.” The young +males continue practising, or, as the bird-catchers say, recording, for +ten or eleven months. Their first essays show hardly a rudiment of the +future song; but as they grow older we can perceive what they are aiming +at; and at last they are said “to sing their song round.” Nestlings +which have learnt the song of a distinct species, as with the +canary-birds educated in the Tyrol, teach and transmit their new song to +their offspring. The slight natural differences of song in the same +species inhabiting different districts may be appositely compared, as +Barrington remarks, “to provincial dialects;” and the songs of allied, +though distinct species may be compared with the languages of distinct +races of man. I have given the foregoing details to shew that an +instinctive tendency to acquire an art is not a peculiarity confined to +man. + +With respect to the origin of articulate language, after having read on +the one side the highly interesting works of Mr. Hensleigh Wedgwood, the +Rev. F. Farrar, and Prof. Schleicher,[78] and the celebrated lectures of +Prof. Max Müller on the other side, I cannot doubt that language owes +its origin to the imitation and modification, aided by signs and +gestures, of various natural sounds, the voices of other animals, and +man’s own instinctive cries. When we treat of sexual selection we shall +see that primeval man, or rather some early progenitor of man, probably +used his voice largely, as does one of the gibbon-apes at the present +day, in producing true musical cadences, that is in singing; we may +conclude from a widely-spread analogy that this power would have been +especially exerted during the courtship of the sexes, serving to express +various emotions, as love, jealousy, triumph, and serving as a challenge +to their rivals. The imitation by articulate sounds of musical cries +might have given rise to words expressive of various complex emotions. +As bearing on the subject of imitation, the strong tendency in our +nearest allies, the monkeys, in microcephalous idiots,[79] and in the +barbarous races of mankind, to imitate whatever they hear deserves +notice. As monkeys certainly understand much that is said to them by +man, and as in a state of nature they utter signal-cries of danger to +their fellows,[80] it does not appear altogether incredible, that some +unusually wise ape-like animal should have thought of imitating the +growl of a beast of prey, so as to indicate to his fellow monkeys the +nature of the expected danger. And this would have been a first step in +the formation of a language. + +As the voice was used more and more, the vocal organs would have been +strengthened and perfected through the principle of the inherited +effects of use; and this would have reacted on the power of speech. But +the relation between the continued use of language and the development +of the brain has no doubt been far more important. The mental powers in +some early progenitor of man must have been more highly developed than +in any existing ape, before even the most imperfect form of speech could +have come into use; but we may confidently believe that the continued +use and advancement of this power would have reacted on the mind by +enabling and encouraging it to carry on long trains of thought. A long +and complex train of thought can no more be carried on without the aid +of words, whether spoken or silent, than a long calculation without the +use of figures or algebra. It appears, also, that even ordinary trains +of thought almost require some form of language, for the dumb, deaf, and +blind girl, Laura Bridgman, was observed to use her fingers whilst +dreaming.[81] Nevertheless a long succession of vivid and connected +ideas, may pass through the mind without the aid of any form of +language, as we may infer from the prolonged dreams of dogs. We have, +also, seen that retriever-dogs are able to reason to a certain extent; +and this they manifestly do without the aid of language. The intimate +connection between the brain, as it is now developed in us, and the +faculty of speech, is well shewn by those curious cases of +brain-disease, in which speech is specially affected, as when the power +to remember substantives is lost, whilst other words can be correctly +used.[82] There is no more improbability in the effects of the continued +use of the vocal and mental organs being inherited, than in the case of +handwriting, which depends partly on the structure of the hand and +partly on the disposition of the mind; and handwriting is certainly +inherited.[83] + +Why the organs now used for speech should have been originally perfected +for this purpose, rather than any other organs, it is not difficult to +see. Ants have considerable powers of intercommunication by means of +their antennæ, as shewn by Huber, who devotes a whole chapter to their +language. We might have used our fingers as efficient instruments, for a +person with practice can report to a deaf man every word of a speech +rapidly delivered at a public meeting; but the loss of our hands, whilst +thus employed, would have been a serious inconvenience. As all the +higher mammals possess vocal organs constructed on the same general +plan with ours, and which are used as a means of communication, it was +obviously probable, if the power of communication had to be improved, +that these same organs would have been still further developed; and this +has been effected by the aid of adjoining and well-adapted parts, namely +the tongue and lips.[84] The fact of the higher apes not using their +vocal organs for speech, no doubt depends on their intelligence not +having been sufficiently advanced. The possession by them of organs, +which with long-continued practice might have been used for speech, +although not thus used, is paralleled by the case of many birds which +possess organs fitted for singing, though they never sing. Thus, the +nightingale and crow have vocal organs similarly constructed, these +being used by the former for diversified song, and by the latter merely +for croaking.[85] + +The formation of different languages and of distinct species, and the +proofs that both have been developed through a gradual process, are +curiously the same.[86] But we can trace the origin of many words +further back than in the case of species, for we can perceive that they +have arisen from the imitation of various sounds, as in alliterative +poetry. We find in distinct languages striking homologies due to +community of descent, and analogies due to a similar process of +formation. The manner in which certain letters or sounds change when +others change is very like correlated growth. We have in both cases the +reduplication of parts, the effects of long-continued use, and so forth. +The frequent presence of rudiments, both in languages and in species, is +still more remarkable. The letter _m_ in the word _am_, means _I_; so +that in the expression _I am_, a superfluous and useless rudiment has +been retained. In the spelling also of words, letters often remain as +the rudiments of ancient forms of pronunciation. Languages, like organic +beings, can be classed in groups under groups; and they can be classed +either naturally according to descent, or artificially by other +characters. Dominant languages and dialects spread widely and lead to +the gradual extinction of other tongues. A language, like a species, +when once extinct, never, as Sir C. Lyell remarks, reappears. The same +language never has two birthplaces. Distinct languages may be crossed +or blended together.[87] We see variability in every tongue, and new +words are continually cropping up; but as there is a limit to the powers +of the memory, single words, like whole languages, gradually become +extinct. As Max Müller[88] has well remarked:—“A struggle for life is +constantly going on amongst the words and grammatical forms in each +language. The better, the shorter, the easier forms are constantly +gaining the upper hand, and they owe their success to their own inherent +virtue.” To these more important causes of the survival of certain +words, mere novelty may, I think, be added; for there is in the mind of +man a strong love for slight changes in all things. The survival or +preservation of certain favoured words in the struggle for existence is +natural selection. + +The perfectly regular and wonderfully complex construction of the +languages of many barbarous nations has often been advanced as a proof, +either of the divine origin of these languages, or of the high art and +former civilisation of their founders. Thus F. von Schlegel writes: “In +those languages which appear to be at the lowest grade of intellectual +culture, we frequently observe a very high and elaborate degree of art +in their grammatical structure. This is especially the case with the +Basque and the Lapponian, and many of the American languages.”[89] But +it is assuredly an error to speak of any language as an art in the sense +of its having been elaborately and methodically formed. Philologists now +admit that conjugations, declensions, &c., originally existed as +distinct words, since joined together; and as such words express the +most obvious relations between objects and persons, it is not surprising +that they should have been used by the men of most races during the +earliest ages. With respect to perfection, the following illustration +will best shew how easily we may err: a Crinoid sometimes consists of no +less than 150,000 pieces of shell,[90] all arranged with perfect +symmetry in radiating lines; but a naturalist does not consider an +animal of this kind as more perfect than a bilateral one with +comparatively few parts, and with none of these alike, excepting on the +opposite sides of the body. He justly considers the differentiation and +specialisation of organs as the test of perfection. So with languages, +the most symmetrical and complex ought not to be ranked above irregular, +abbreviated, and bastardised languages, which have borrowed expressive +words and useful forms of construction from various conquering, or +conquered, or immigrant races. + +From these few and imperfect remarks I conclude that the extremely +complex and regular construction of many barbarous languages, is no +proof that they owe their origin to a special act of creation.[91] Nor, +as we have seen, does the faculty of articulate speech in itself offer +any insuperable objection to the belief that man has been developed from +some lower form. + + +_Self-consciousness, Individuality, Abstraction, General Ideas, &c._—It +would be useless to attempt discussing these high faculties, which, +according to several recent writers, make the sole and complete +distinction between man and the brutes, for hardly two authors agree in +their definitions. Such faculties could not have been fully developed in +man until his mental powers had advanced to a high standard, and this +implies the use of a perfect language. No one supposes that one of the +lower animals reflects whence he comes or whither he goes,—what is +death or what is life, and so forth. But can we feel sure that an old +dog with an excellent memory and some power of imagination, as shewn by +his dreams, never reflects on his past pleasures in the chase? and this +would be a form of self-consciousness. On the other hand, as Büchner[92] +has remarked, how little can the hard-worked wife of a degraded +Australian savage, who uses hardly any abstract words and cannot count +above four, exert her self-consciousness, or reflect on the nature of +her own existence. + +That animals retain their mental individuality is unquestionable. When +my voice awakened a train of old associations in the mind of the +above-mentioned dog, he must have retained his mental individuality, +although every atom of his brain had probably undergone change more than +once during the interval of five years. This dog might have brought +forward the argument lately advanced to crush all evolutionists, and +said, “I abide amid all mental moods and all material changes.... The +teaching that atoms leave their impressions as legacies to other atoms +falling into the places they have vacated is contradictory of the +utterance of consciousness, and is therefore false; but it is the +teaching necessitated by evolutionism, consequently the hypothesis is a +false one.”[93] + + +_Sense of Beauty._—This sense has been declared to be peculiar to man. +But when we behold male birds elaborately displaying their plumes and +splendid colours before the females, whilst other birds not thus +decorated make no such display, it is impossible to doubt that the +females admire the beauty of their male partners. As women everywhere +deck themselves with these plumes, the beauty of such ornaments cannot +be disputed. The Bower-birds by tastefully ornamenting their +playing-passages with gaily-coloured objects, as do certain +humming-birds their nests, offer additional evidence that they possess a +sense of beauty. So with the song of birds, the sweet strains poured +forth by the males during the season of love are certainly admired by +the females, of which fact evidence will hereafter be given. If female +birds had been incapable of appreciating the beautiful colours, the +ornaments, and voices of their male partners, all the labour and +anxiety exhibited by them in displaying their charms before the females +would have been thrown away; and this it is impossible to admit. Why +certain bright colours and certain sounds should excite pleasure, when +in harmony, cannot, I presume, be explained any more than why certain +flavours and scents are agreeable; but assuredly the same colours and +the same sounds are admired by us and by many of the lower animals. + +The taste for the beautiful, at least as far as female beauty is +concerned, is not of a special nature in the human mind; for it differs +widely in the different races of man, as will hereafter be shewn, and is +not quite the same even in the different nations of the same race. +Judging from the hideous ornaments and the equally hideous music admired +by most savages, it might be urged that their æsthetic faculty was not +so highly developed as in certain animals, for instance, in birds. +Obviously no animal would be capable of admiring such scenes as the +heavens at night, a beautiful landscape, or refined music; but such high +tastes, depending as they do on culture and complex associations, are +not enjoyed by barbarians or by uneducated persons. + +Many of the faculties, which have been of inestimable service to man for +his progressive advancement, such as the powers of the imagination, +wonder, curiosity, an undefined sense of beauty, a tendency to +imitation, and the love of excitement or novelty, could not fail to have +led to the most capricious changes of customs and fashions. I have +alluded to this point, because a recent writer[94] has oddly fixed on +Caprice “as one of the most remarkable and typical differences between +savages and brutes.” But not only can we perceive how it is that roan is +capricious, but the lower animals are, as we shall hereafter see, +capricious in their affections, aversions, and sense of beauty. There is +also good reason to suspect that they love novelty, for its own sake. + + +_Belief in God—Religion._—There is no evidence that man was +aboriginally endowed with the ennobling belief in the existence of an +Omnipotent God. On the contrary there is ample evidence, derived not +from hasty travellers, but from men who have long resided with savages, +that numerous races have existed and still exist, who have no idea of +one or more gods, and who have no words in their languages to express +such an idea.[95] The question is of course wholly distinct from that +higher one, whether there exists a Creator and Ruler of the universe; +and this has been answered in the affirmative by the highest intellects +that have ever lived. + +If, however, we include under the term “religion” the belief in unseen +or spiritual agencies, the case is wholly different; for this belief +seems to be almost universal with the less civilised races. Nor is it +difficult to comprehend how it arose. As soon as the important faculties +of the imagination, wonder, and curiosity, together with some power of +reasoning, had become partially developed, man would naturally have +craved to understand what was passing around him, and have vaguely +speculated on his own existence. As Mr. M’Lennan[96] has remarked, +“Some explanation of the phenomena of life, a man must feign for +himself; and to judge from the universality of it, the simplest +hypothesis, and the first to occur to men, seems to have been that +natural phenomena are ascribable to the presence in animals, plants, and +things, and in the forces of nature, of such spirits prompting to action +as men are conscious they themselves possess.” It is probable, as Mr. +Tylor has clearly shewn, that dreams may have first given rise to the +notion of spirits; for savages do not readily distinguish between +subjective and objective impressions. When a savage dreams, the figures +which appear before him are believed to have come from a distance and to +stand over him; or “the soul of the dreamer goes out on its travels, and +comes home with a remembrance of what it has seen.”[97] But until the +above-named faculties of imagination, curiosity, reason, &c., had been +fairly well developed in the mind of man, his dreams would not have led +him to believe in spirits, any more than in the case of a dog. + +The tendency in savages to imagine that natural objects and agencies +are animated by spiritual or living essences, is perhaps illustrated by +a little fact which I once noticed: my dog, a full-grown and very +sensible animal, was lying on the lawn during a hot and still day; but +at a little distance a slight breeze occasionally moved an open parasol, +which would have been wholly disregarded by the dog, had any one stood +near it. As it was, every time that the parasol slightly moved, the dog +growled fiercely and barked. He must, I think, have reasoned to himself +in a rapid and unconscious manner, that movement without any apparent +cause indicated the presence of some strange living agent, and no +stranger had a right to be on his territory. + +The belief in spiritual agencies would easily pass into the belief in +the existence of one or more gods. For savages would naturally attribute +to spirits the same passions, the same love of vengeance or simplest +form of justice, and the same affections which they themselves +experienced. The Fuegians appear to be in this respect in an +intermediate condition, for when the surgeon on board the “Beagle” shot +some young ducklings as specimens, York Minster declared in the most +solemn manner, “Oh! Mr. Bynoe, much rain, much snow, blow much;” and +this was evidently a retributive punishment for wasting human food. So +again he related how, when his brother killed a “wild man,” storms long +raged, much rain and snow fell. Yet we could never discover that the +Fuegians believed in what we should call a God, or practised any +religious rites; and Jemmy Button, with justifiable pride, stoutly +maintained that there was no devil in his land. This latter assertion is +the more remarkable, as with savages the belief in bad spirits is far +more common than the belief in good spirits. + +The feeling of religious devotion is a highly complex one, consisting of +love, complete submission to an exalted and mysterious superior, a +strong sense of dependence,[98] fear, reverence, gratitude, hope for the +future, and perhaps other elements. No being could experience so complex +an emotion until advanced in his intellectual and moral faculties to at +least a moderately high level. Nevertheless we see some distant approach +to this state of mind, in the deep love of a dog for his master, +associated with complete submission, some fear, and perhaps other +feelings. The behaviour of a dog when returning to his master after an +absence, and, as I may add, of a monkey to his beloved keeper, is widely +different from that towards their fellows. In the latter case the +transports of joy appear to be somewhat less, and the sense of equality +is shewn in every action. Professor Braubach[99] goes so far as to +maintain that a dog looks on his master as on a god. + +The same high mental faculties which first led man to believe in unseen +spiritual agencies, then in fetishism, polytheism, and ultimately in +monotheism, would infallibly lead him, as long as his reasoning powers +remained poorly developed, to various strange superstitions and customs. +Many of these are terrible to think of—such as the sacrifice of human +beings to a blood-loving god; the trial of innocent persons by the +ordeal of poison or fire; witchcraft, &c.—yet it is well occasionally +to reflect on these superstitions, for they shew us what an infinite +debt of gratitude we owe to the improvement of our reason, to science, +and our accumulated knowledge.[100] As Sir J. Lubbock has well +observed, “it is not too much to say that the horrible dread of unknown +evil hangs like a thick cloud over savage life, and embitters every +pleasure.” These miserable and indirect consequences of our highest +faculties may be compared with the incidental and occasional mistakes of +the instincts of the lower animals. + + + + +CHAPTER III. + +COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER +ANIMALS—_continued_. + + + The moral sense—Fundamental proposition—The qualities of + social animals—Origin of sociability—Struggle between opposed + instincts—Man a social animal—The more enduring social + instincts conquer other less persistent instincts—The social + virtues alone regarded by savages—The self-regarding virtues + acquired at a later stage of development—The importance of the + judgment of the members of the same community on + conduct—Transmission of moral tendencies—Summary. + + +I fully subscribe to the judgment of those writers[101] who maintain +that of all the differences between man and the lower animals, the moral +sense or conscience is by far the most important. This sense, as +Mackintosh[102] remarks, “has a rightful supremacy over every other +principle of human action;” it is summed up in that short but imperious +word _ought_, so full of high significance. It is the most noble of all +the attributes of man, leading him without a moment’s hesitation to risk +his life for that of a fellow-creature; or after due deliberation, +impelled simply by the deep feeling of right or duty, to sacrifice it in +some great cause. Immanuel Kant exclaims, “Duty! Wondrous thought, that +workest neither by fond insinuation, flattery, nor by any threat, but +merely by holding up thy naked law in the soul, and so extorting for +thyself always reverence, if not always obedience; before whom all +appetites are dumb, however secretly they rebel; whence thy +original?”[103] + +This great question has been discussed by many writers[104] of +consummate ability; and my sole excuse for touching on it is the +impossibility of here passing it over, and because, as far as I know, no +one has approached it exclusively from the side of natural history. The +investigation possesses, also, some independent interest, as an attempt +to see how far the study of the lower animals can throw light on one of +the highest psychical faculties of man. + +The following proposition seems to me in a high degree probable—namely, +that any animal whatever, endowed with well-marked social +instincts,[105] would inevitably acquire a moral sense or conscience, as +soon as its intellectual powers had become as well developed, or nearly +as well developed, as in man. For, _firstly_, the social instincts lead +an animal to take pleasure in the society of its fellows, to feel a +certain amount of sympathy with them, and to perform various services +for them. The services may be of a definite and evidently instinctive +nature; or there may be only a wish and readiness, as with most of the +higher social animals, to aid their fellows in certain general ways. But +these feelings and services are by no means extended to all the +individuals of the same species, only to those of the same association. +_Secondly_, as soon as the mental faculties had become highly developed, +images of all past actions and motives would be incessantly passing +through the brain of each individual; and that feeling of +dissatisfaction which invariably results, as we shall hereafter see, +from any unsatisfied instinct, would arise, as often as it was perceived +that the enduring and always present social instinct had yielded to some +other instinct, at the time stronger, but neither enduring in its +nature, nor leaving behind it a very vivid impression. It is clear that +many instinctive desires, such as that of hunger, are in their nature of +short duration; and after being satisfied are not readily or vividly +recalled. _Thirdly_, after the power of language had been acquired and +the wishes of the members of the same community could be distinctly +expressed, the common opinion how each member ought to act for the +public good, would naturally become to a large extent the guide to +action. But the social instincts would still give the impulse to act for +the good of the community, this impulse being strengthened, directed, +and sometimes even deflected by public opinion, the power of which +rests, as we shall presently see, on instinctive sympathy. _Lastly_, +habit in the individual would ultimately play a very important part in +guiding the conduct of each member; for the social instincts and +impulses, like all other instincts, would be greatly strengthened by +habit, as would obedience to the wishes and judgment of the community. +These several subordinate propositions must now be discussed; and some +of them at considerable length. + +It may be well first to premise that I do not wish to maintain that any +strictly social animal, if its intellectual faculties were to become as +active and as highly developed as in man, would acquire exactly the same +moral sense as ours. In the same manner as various animals have some +sense of beauty, though they admire widely different objects, so they +might have a sense of right and wrong, though led by it to follow widely +different lines of conduct. If, for instance, to take an extreme case, +men were reared under precisely the same conditions as hive-bees, there +can hardly be a doubt that our unmarried females would, like the +worker-bees, think it a sacred duty to kill their brothers, and mothers +would strive to kill their fertile daughters; and no one would think of +interfering. Nevertheless the bee, or any other social animal, would in +our supposed case gain, as it appears to me, some feeling of right and +wrong, or a conscience. For each individual would have an inward sense +of possessing certain stronger or more enduring instincts, and others +less strong or enduring; so that there would often be a struggle which +impulse should be followed; and satisfaction or dissatisfaction would be +felt, as past impressions were compared during their incessant passage +through the mind. In this case an inward monitor would tell the animal +that it would have been better to have followed the one impulse rather +than the other. The one course ought to have been followed: the one +would have been right and the other wrong; but to these terms I shall +have to recur. + + +_Sociability._—Animals of many kinds are social; we find even distinct +species living together, as with some American monkeys, and with the +united flocks of rooks, jackdaws, and starlings. Man shows the same +feeling in his strong love for the dog, which the dog returns with +interest. Every one must have noticed how miserable horses, dogs, sheep, +&c. are when separated from their companions; and what affection at +least the two former kinds show on their reunion. It is curious to +speculate on the feelings of a dog, who will rest peacefully for hours +in a room with his master or any of the family, without the least notice +being taken of him; but if left for a short time by himself, barks or +howls dismally. We will confine our attention to the higher social +animals, excluding insects, although these aid each other in many +important ways. The most common service which the higher animals perform +for each other, is the warning each other of danger by means of the +united senses of all. Every sportsman knows, as Dr. Jaeger remarks,[106] +how difficult it is to approach animals in a herd or troop. Wild horses +and cattle do not, I believe, make any danger-signal; but the attitude +of any one who first discovers an enemy, warns the others. Rabbits stamp +loudly on the ground with their hind-feet as a signal: sheep and chamois +do the same, but with their fore-feet, uttering likewise a whistle. Many +birds and some mammals post sentinels, which in the case of seals are +said[107] generally to be the females. The leader of a troop of monkeys +acts as the sentinel, and utters cries expressive both of danger and of +safety.[108] Social animals perform many little services for each +other: horses nibble, and cows lick each other, on any spot which +itches: monkeys search for each other’s external parasites; and Brehm +states that after a troop of the _Cercopithecus griseo-viridis_ has +rushed through a thorny brake, each monkey stretches itself on a branch, +and another monkey sitting by “conscientiously” examines its fur and +extracts every thorn or burr. + +Animals also render more important services to each other: thus wolves +and some other beasts of prey hunt in packs, and aid each other in +attacking their victims. Pelicans fish in concert. The Hamadryas baboons +turn over stones to find insects, &c.; and when they come to a large +one, as many as can stand round, turn it over together and share the +booty. Social animals mutually defend each other. The males of some +ruminants come to the front when there is danger and defend the herd +with their horns. I shall also in a future chapter give cases of two +young wild bulls attacking an old one in concert, and of two stallions +together trying to drive away a third stallion from a troop of mares. +Brehm encountered in Abyssinia a great troop of baboons which were +crossing a valley: some had already ascended the opposite mountain, and +some were still in the valley: the latter were attacked by the dogs, but +the old males immediately hurried down from the rocks, and with mouths +widely opened roared so fearfully, that the dogs precipitately +retreated. They were again encouraged to the attack; but by this time +all the baboons had reascended the heights, excepting a young one, about +six months old, who, loudly calling for aid, climbed on a block of rock +and was surrounded. Now one of the largest males, a true hero, came down +again from the mountain, slowly went to the young one, coaxed him, and +triumphantly led him away—the dogs being too much astonished to make an +attack. I cannot resist giving another scene which was witnessed by this +same naturalist; an eagle seized a young Cercopithecus, which, by +clinging to a branch, was not at once carried off; it cried loudly for +assistance, upon which the other members of the troop with much uproar +rushed to the rescue, surrounded the eagle, and pulled out so many +feathers, that he no longer thought of his prey, but only how to escape. +This eagle, as Brehm remarks, assuredly would never again attack a +monkey in a troop. + +It is certain that associated animals have a feeling of love for each +other which is not felt by adult and non-social animals. How far in most +cases they actually sympathise with each other’s pains and pleasures is +more doubtful, especially with respect to the latter. Mr. Buxton, +however, who had excellent means of observation,[109] states that his +macaws, which lived free in Norfolk, took “an extravagant interest” in a +pair with a nest, and whenever the female left it, she was surrounded by +a troop “screaming horrible acclamations in her honour.” It is often +difficult to judge whether animals have any feeling for each other’s +sufferings. Who can say what cows feel, when they surround and stare +intently on a dying or dead companion? That animals sometimes are far +from feeling any sympathy is too certain; for they will expel a wounded +animal from the herd, or gore or worry it to death. This is almost the +blackest fact in natural history, unless indeed the explanation which +has been suggested is true, that their instinct or reason leads them to +expel an injured companion, lest beasts of prey, including man, should +be tempted to follow the troop. In this case their conduct is not much +worse than that of the North American Indians who leave their feeble +comrades to perish on the plains, or the Feegeans, who, when their +parents get old or fall ill, bury them alive.[110] + +Many animals, however, certainly sympathise with each other’s distress +or danger. This is the case even with birds; Capt. Stansbury[111] found +on a salt lake in Utah an old and completely blind pelican, which was +very fat, and must have been long and well fed by his companions. Mr. +Blyth, as he informs me, saw Indian crows feeding two or three of their +companions which were blind; and I have heard of an analogous case with +the domestic cock. We may, if we choose, call these actions instinctive; +but such cases are much too rare for the development of any special +instinct.[112] I have myself seen a dog, who never passed a great friend +of his, a cat which lay sick in a basket, without giving her a few licks +with his tongue, the surest sign of kind feeling in a dog. + +It must be called sympathy that leads a courageous dog to fly at any one +who strikes his master, as he certainly will. I saw a person pretending +to beat a lady who had a very timid little dog on her lap, and the trial +had never before been made. The little creature instantly jumped away, +but after the pretended beating was over, it was really pathetic to see +how perseveringly he tried to lick his mistress’s face and comfort her. +Brehm[113] states that when a baboon in confinement was pursued to be +punished, the others tried to protect him. It must have been sympathy in +the cases above given which led the baboons and Cercopitheci to defend +their young comrades from the dogs and the eagle. I will give only one +other instance of sympathetic and heroic conduct in a little American +monkey. Several years ago a keeper at the Zoological Gardens, showed me +some deep and scarcely healed wounds on the nape of his neck, inflicted +on him whilst kneeling on the floor by a fierce baboon. The little +American monkey, who was a warm friend of this keeper, lived in the same +large compartment, and was dreadfully afraid of the great baboon. +Nevertheless, as soon as he saw his friend the keeper in peril, he +rushed to the rescue, and by screams and bites so distracted the baboon +that the man was able to escape, after running great risk, as the +surgeon who attended him thought, of his life. + +Besides love and sympathy, animals exhibit other qualities which in us +would be called moral; and I agree with Agassiz[114] that dogs possess +something very like a conscience. They certainly possess some power of +self-command, and this does not appear to be wholly the result of fear. +As Braubach[115] remarks, a dog will refrain from stealing food in the +absence of his master. Dogs have long been accepted as the very type of +fidelity and obedience. All animals living in a body which defend each +other or attack their enemies in concert, must be in some degree +faithful to each other; and those that follow a leader must be in some +degree obedient. When the baboons in Abyssinia[116] plunder a garden, +they silently follow their leader; and if an imprudent young animal +makes a noise, he receives a slap from the others to teach him silence +and obedience; but as soon as they are sure that there is no danger, all +show their joy by much clamour. + +With respect to the impulse which leads certain animals to associate +together, and to aid each other in many ways, we may infer that in most +cases they are impelled by the same sense of satisfaction or pleasure +which they experience in performing other instinctive actions; or by the +same sense of dissatisfaction, as in other cases of prevented +instinctive actions. We see this in innumerable instances, and it is +illustrated in a striking manner by the acquired instincts of our +domesticated animals; thus a young shepherd-dog delights in driving and +running round a flock of sheep, but not in worrying them; a young +foxhound delights in hunting a fox, whilst some other kinds of dogs as I +have witnessed, utterly disregard foxes. What a strong feeling of inward +satisfaction must impel a bird, so full of activity, to brood day after +day over her eggs. Migratory birds are miserable if prevented from +migrating, and perhaps they enjoy starting on their long flight. Some +few instincts are determined solely by painful feelings, as by fear, +which leads to self-preservation, or is specially directed against +certain enemies. No one, I presume, can analyse the sensations of +pleasure or pain. In many cases, however, it is probable that instincts +are persistently followed from the mere force of inheritance, without +the stimulus of either pleasure or pain. A young pointer, when it first +scents game, apparently cannot help pointing. A squirrel in a cage who +pats the nuts which it cannot eat, as if to bury them in the ground, can +hardly be thought to act thus either from pleasure or pain. Hence the +common assumption that men must be impelled to every action by +experiencing some pleasure or pain may be erroneous. Although a habit +may be blindly and implicitly followed, independently of any pleasure or +pain felt at the moment, yet if it be forcibly and abruptly checked, a +vague sense of dissatisfaction is generally experienced; and this is +especially true in regard to persons of feeble intellect. + +It has often been assumed that animals were in the first place rendered +social, and that they feel as a consequence uncomfortable when separated +from each other, and comfortable whilst together; but it is a more +probable view that these sensations were first developed, in order that +those animals which would profit by living in society, should be induced +to live together. In the same manner as the sense of hunger and the +pleasure of eating were, no doubt, first acquired in order to induce +animals to eat. The feeling of pleasure from society is probably an +extension of the parental or filial affections; and this extension may +be in chief part attributed to natural selection, but perhaps in part to +mere habit. For with those animals which were benefited by living in +close association, the individuals which took the greatest pleasure in +society would best escape various dangers; whilst those that cared least +for their comrades and lived solitary would perish in greater numbers. +With respect to the origin of the parental and filial affections, which +apparently lie at the basis of the social affections, it is hopeless to +speculate; but we may infer that they have been to a large extent +gained through natural selection. So it has almost certainly been with +the unusual and opposite feeling of hatred between the nearest +relations, as with the worker-bees which kill their brother-drones, and +with the queen-bees which kill their daughter-queens; the desire to +destroy, instead of loving, their nearest relations having been here of +service to the community. + +The all-important emotion of sympathy is distinct from that of love. A +mother may passionately love her sleeping and passive infant, but she +can then hardly be said to feel sympathy for it. The love of a man for +his dog is distinct from sympathy, and so is that of a dog for his +master. Adam Smith formerly argued, as has Mr. Bain recently, that the +basis of sympathy lies in our strong retentiveness of former states of +pain or pleasure. Hence, “the sight of another person enduring hunger, +cold, fatigue, revives in us some recollection of these states, which +are painful even in idea.” We are thus impelled to relieve the +sufferings of another, in order that our own painful feelings may be at +the same time relieved. In like manner we are led to participate in the +pleasures of others.[117] But I cannot see how this view explains the +fact that sympathy is excited in an immeasurably stronger degree by a +beloved than by an indifferent person. The mere sight of suffering, +independently of love, would suffice to call up in us vivid +recollections and associations. Sympathy may at first have originated in +the manner above suggested; but it seems now to have become an instinct, +which is especially directed towards beloved objects, in the same manner +as fear with animals is especially directed against certain enemies. As +sympathy is thus directed, the mutual love of the members of the same +community will extend its limits. No doubt a tiger or lion feels +sympathy for the sufferings of its own young, but not for any other +animal. With strictly social animals the feeling will be more or less +extended to all the associated members, as we know to be the case. With +mankind selfishness, experience, and imitation probably add, as Mr. Bain +has shewn, to the power of sympathy; for we are led by the hope of +receiving good in return to perform acts of sympathetic kindness to +others; and there can be no doubt that the feeling of sympathy is much +strengthened by habit. In however complex a manner this feeling may have +originated, as it is one of high importance to all those animals which +aid and defend each other, it will have been increased, through natural +selection; for those communities, which included the greatest number of +the most sympathetic members, would flourish best and rear the greatest +number of offspring. + +In many cases it is impossible to decide whether certain social +instincts have been acquired through natural selection, or are the +indirect result of other instincts and faculties, such as sympathy, +reason, experience, and a tendency to imitation; or again, whether they +are simply the result of long-continued habit. So remarkable an instinct +as the placing sentinels to warn the community of danger, can hardly +have been the indirect result of any other faculty; it must therefore +have been directly acquired. On the other hand, the habit followed by +the males of some social animals, of defending the community and of +attacking their enemies or their prey in concert, may perhaps have +originated from mutual sympathy; but courage, and in most cases +strength, must have been previously acquired, probably through natural +selection. + +Of the various instincts and habits, some are much stronger than others, +that is, some either give more pleasure in their performance and more +distress in their prevention than others; or, which is probably quite as +important, they are more persistently followed through inheritance +without exciting any special feeling of pleasure or pain. We are +ourselves conscious that some habits are much more difficult to cure or +change than others. Hence a struggle may often be observed in animals +between different instincts, or between an instinct and some habitual +disposition; as when a dog rushes after a hare, is rebuked, pauses, +hesitates, pursues again or returns ashamed to his master; or as between +the love of a female dog for her young puppies and for her master, for +she may be seen to slink away to them, as if half ashamed of not +accompanying her master. But the most curious instance known to me of +one instinct conquering another, is the migratory instinct conquering +the maternal instinct. The former is wonderfully strong; a confined bird +will at the proper season beat her breast against the wires of her cage, +until it is bare and bloody. It causes young salmon to leap out of the +fresh water, where they could still continue to live, and thus +unintentionally to commit suicide. Every one knows how strong the +maternal instinct is, leading even timid birds to face great danger, +though with hesitation and in opposition to the instinct of +self-preservation. Nevertheless the migratory instinct is so powerful +that late in the autumn swallows and house-martins frequently desert +their tender young, leaving them to perish miserably in their +nests.[118] + +We can perceive that an instinctive impulse, if it be in any way more +beneficial to a species than some other or opposed instinct, would be +rendered the more potent of the two through natural selection; for the +individuals which had it most strongly developed would survive in larger +numbers. Whether this is the case with the migratory in comparison with +the maternal instinct, may well be doubted. The great persistence or +steady action of the former at certain seasons of the year during the +whole day, may give it for a time paramount force. + + +_Man a social animal._—Most persons admit that man is a social being. +We see this in his dislike of solitude, and in his wish for society +beyond that of his own family. Solitary confinement is one of the +severest punishments which can be inflicted. Some authors suppose that +man primevally lived in single families; but at the present day, though +single families, or only two or three together, roam the solitudes of +some savage lands, they are always, as far as I can discover, friendly +with other families inhabiting the same district. Such families +occasionally meet in council, and they unite for their common defence. +It is no argument against savage man being a social animal, that the +tribes inhabiting adjacent districts are almost always at war with each +other; for the social instincts never extend to all the individuals of +the same species. Judging from the analogy of the greater number of the +Quadrumana, it is probable that the early ape-like progenitors of man +were likewise social; but this is not of much importance for us. +Although man, as he now exists, has few special instincts, having lost +any which his early progenitors may have possessed, this is no reason +why he should not have retained from an extremely remote period some +degree of instinctive love and sympathy for his fellows. We are indeed +all conscious that we do possess such sympathetic feelings;[119] but our +consciousness does not tell us whether they are instinctive, having +originated long ago in the same manner as with the lower animals, or +whether they have been acquired by each of us during our early years. As +man is a social animal, it is also probable that he would inherit a +tendency to be faithful to his comrades, for this quality is common to +most social animals. He would in like manner possess some capacity for +self-command, and perhaps of obedience to the leader of the community. +He would from an inherited tendency still be willing to defend, in +concert with others, his fellow-men, and would be ready to aid them in +any way which did not too greatly interfere with his own welfare or his +own strong desires. + +The social animals which stand at the bottom of the scale are guided +almost exclusively, and those which stand higher in the scale are +largely guided, in the aid which they give to the members of the same +community, by special instincts; but they are likewise in part impelled +by mutual love and sympathy, assisted apparently by some amount of +reason. Although man, as just remarked, has no special instincts to tell +him how to aid his fellow-men, he still has the impulse, and with his +improved intellectual faculties would naturally be much guided in this +respect by reason and experience. Instinctive sympathy would, also, +cause him to value highly the approbation of his fellow-men; for, as Mr. +Bain has clearly shewn,[120] the love of praise and the strong feeling +of glory, and the still stronger horror of scorn and infamy, “are due to +the workings of sympathy.” Consequently man would be greatly influenced +by the wishes, approbation, and blame of his fellow-men, as expressed by +their gestures and language. Thus the social instincts, which must have +been acquired by man in a very rude state, and probably even by his +early ape-like progenitors, still give the impulse to many of his best +actions; but his actions are largely determined by the expressed wishes +and judgment of his fellow-men, and unfortunately still oftener by his +own strong, selfish desires. But as the feelings of love and sympathy +and the power of self-command become strengthened by habit, and as the +power of reasoning becomes clearer so that man can appreciate the +justice of the judgments of his fellow-men, he will feel himself +impelled, independently of any pleasure or pain felt at the moment, to +certain lines of conduct. He may then say, I am the supreme judge of my +own conduct, and in the words of Kant, I will not in my own person +violate the dignity of humanity. + + +_The more enduring Social Instincts conquer the less Persistent +Instincts._—We have, however, not as yet considered the main point, on +which the whole question of the moral sense hinges. Why should a man +feel that he ought to obey one instinctive desire rather than another? +Why does he bitterly regret if he has yielded to the strong sense of +self-preservation, and has not risked his life to save that of a +fellow-creature; or why does he regret having stolen food from severe +hunger? + +It is evident in the first place, that with mankind the instinctive +impulses have different degrees of strength; a young and timid mother +urged by the maternal instinct will, without a moment’s hesitation, run +the greatest danger for her infant, but not for a mere fellow-creature. +Many a man, or even boy, who never before risked his life for another, +but in whom courage and sympathy were well developed, has, disregarding +the instinct of self-preservation, instantaneously plunged into a +torrent to save a drowning fellow-creature. In this case man is impelled +by the same instinctive motive, which caused the heroic little American +monkey, formerly described, to attack the great and dreaded baboon, to +save his keeper. Such actions as the above appear to be the simple +result of the greater strength of the social or maternal instincts than +of any other instinct or motive; for they are performed too +instantaneously for reflection, or for the sensation of pleasure or +pain; though if prevented distress would be caused. + +I am aware that some persons maintain that actions performed +impulsively, as in the above cases, do not come under the dominion of +the moral sense, and cannot be called moral. They confine this term to +actions done deliberately, after a victory over opposing desires, or to +actions prompted by some lofty motive. But it appears scarcely possible +to draw any clear line of distinction of this kind; though the +distinction may be real. As far as exalted motives are concerned, many +instances have been recorded of barbarians, destitute of any feeling of +general benevolence towards mankind, and not guided by any religious +motive, who have deliberately as prisoners sacrificed their lives,[121] +rather than betray their comrades; and surely their conduct ought to be +considered as moral. As far as deliberation and the victory over +opposing motives are concerned, animals may be seen doubting between +opposed instincts, as in rescuing their offspring or comrades from +danger; yet their actions, though done for the good of others, are not +called moral. Moreover, an action repeatedly performed by us, will at +last be done without deliberation or hesitation, and can then hardly be +distinguished from an instinct; yet surely no one will pretend that an +action thus done ceases to be moral. On the contrary, we all feel that +an act cannot be considered as perfect, or as performed in the most +noble manner, unless it be done impulsively, without deliberation or +effort, in the same manner as by a man in whom the requisite qualities +are innate. He who is forced to overcome his fear or want of sympathy +before he acts, deserves, however, in one way higher credit than the man +whose innate disposition leads him to a good act without effort. As we +cannot distinguish between motives, we rank all actions of a certain +class as moral, when they are performed by a moral being. A moral being +is one who is capable of comparing his past and future actions or +motives, and of approving or disapproving of them. We have no reason to +suppose that any of the lower animals have this capacity; therefore +when a monkey faces danger to rescue its comrade, or takes charge of an +orphan-monkey, we do not call its conduct moral. But in the case of man, +who alone can with certainty be ranked as a moral being, actions of a +certain class are called moral, whether performed deliberately after a +struggle with opposing motives, or from the effects of slowly-gained +habit, or impulsively through instinct. + +But to return to our more immediate subject; although some instincts are +more powerful than others, thus leading to corresponding actions, yet it +cannot be maintained that the social instincts are ordinarily stronger +in man, or have become stronger through long-continued habit, than the +instincts, for instance, of self-preservation, hunger, lust, vengeance, +&c. Why then does man regret, even though he may endeavour to banish any +such regret, that he has followed the one natural impulse, rather than +the other; and why does he further feel that he ought to regret his +conduct? Man in this respect differs profoundly from the lower animals. +Nevertheless we can, I think, see with some degree of clearness the +reason of this difference. + +Man, from the activity of his mental faculties, cannot avoid reflection: +past impressions and images are incessantly passing through his mind +with distinctness. Now with those animals which live permanently in a +body, the social instincts are ever present and persistent. Such animals +are always ready to utter the danger-signal, to defend the community, +and to give aid to their fellows in accordance with their habits; they +feel at all times, without the stimulus of any special passion or +desire, some degree of love and sympathy for them; they are unhappy if +long separated from them, and always happy to be in their company. So it +is with ourselves. A man who possessed no trace of such feelings would +be an unnatural monster. On the other hand, the desire to satisfy +hunger, or any passion, such as vengeance, is in its nature temporary, +and can for a time be fully satisfied. Nor is it easy, perhaps hardly +possible, to call up with complete vividness the feeling, for instance, +of hunger; nor indeed, as has often been remarked, of any suffering. The +instinct of self-preservation is not felt except in the presence of +danger; and many a coward has thought himself brave until he has met his +enemy face to face. The wish for another man’s property is perhaps as +persistent a desire as any that can be named; but even in this case the +satisfaction of actual possession is generally a weaker feeling than the +desire: many a thief, if not an habitual one, after success has wondered +why he stole some article. + +Thus, as man cannot prevent old impressions continually repassing +through his mind, he will be compelled to compare the weaker impressions +of, for instance, past hunger, or of vengeance satisfied or danger +avoided at the cost of other men, with the instinct of sympathy and +good-will to his fellows, which is still present and ever in some degree +active in his mind. He will then feel in his imagination that a stronger +instinct has yielded to one which now seems comparatively weak; and then +that sense of dissatisfaction will inevitably be felt with which man is +endowed, like every other animal, in order that his instincts may be +obeyed. The case before given, of the swallow, affords an illustration, +though of a reversed nature, of a temporary though for the time strongly +persistent instinct conquering another instinct which is usually +dominant over all others. At the proper season these birds seem all day +long to be impressed with the desire to migrate; their habits change; +they become restless, are noisy, and congregate in flocks. Whilst the +mother-bird is feeding or brooding over her nestlings, the maternal +instinct is probably stronger than the migratory; but the instinct which +is more persistent gains the victory, and at last, at a moment when her +young ones are not in sight, she takes flight and deserts them. When +arrived at the end of her long journey, and the migratory instinct +ceases to act, what an agony of remorse each bird would feel, if, from +being endowed with great mental activity, she could not prevent the +image continually passing before her mind of her young ones perishing in +the bleak north from cold and hunger. + +At the moment of action, man will no doubt be apt to follow the stronger +impulse; and though this may occasionally prompt him to the noblest +deeds, it will far more commonly lead him to gratify his own desires at +the expense of other men. But after their gratification, when past and +weaker impressions are contrasted with the ever-enduring social +instincts, retribution will surely come. Man will then feel dissatisfied +with himself, and will resolve with more or less force to act +differently for the future. This is conscience; for conscience looks +backwards and judges past actions, inducing that kind of +dissatisfaction, which if weak we call regret, and if severe remorse. + +These sensations are, no doubt, different from those experienced when +other instincts or desires are left unsatisfied; but every unsatisfied +instinct has its own proper prompting sensation, as we recognise with +hunger, thirst, &c. Man thus prompted, will through long habit acquire +such perfect self-command, that his desires and passions will at last +instantly yield to his social sympathies, and there will no longer be a +struggle between them. The still hungry, or the still revengeful man +will not think of stealing food, or of wreaking his vengeance. It is +possible, or, as we shall hereafter see, even probable, that the habit +of self-command may, like other habits, be inherited. Thus at last man +comes to feel, through acquired and perhaps inherited habit, that it is +best for him to obey his more persistent instincts. The imperious word +_ought_ seems merely to imply the consciousness of the existence of a +persistent instinct, either innate or partly acquired, serving him as a +guide, though liable to be disobeyed. We hardly use the word _ought_ in +a metaphorical sense, when we say hounds ought to hunt, pointers to +point, and retrievers to retrieve their game. If they fail thus to act, +they fail in their duty and act wrongly. + +If any desire or instinct, leading to an action opposed to the good of +others, still appears to a man, when recalled to mind, as strong as, or +stronger than, his social instinct, he will feel no keen regret at +having followed it; but he will be conscious that if his conduct were +known to his fellows, it would meet with their disapprobation; and few +are so destitute of sympathy as not to feel discomfort when this is +realised. If he has no such sympathy, and if his desires leading to bad +actions are at the time strong, and when recalled are not overmastered +by the persistent social instincts, then he is essentially a bad +man;[122] and the sole restraining motive left is the fear of +punishment, and the conviction that in the long run it would be best for +his own selfish interests to regard the good of others rather than his +own. + +It is obvious that every one may with an easy conscience gratify his own +desires, if they do not interfere with his social instincts, that is +with the good of others; but in order to be quite free from +self-reproach, or at least of anxiety, it is almost necessary for him to +avoid the disapprobation, whether reasonable or not, of his fellow men. +Nor must he break through the fixed habits of his life, especially if +these are supported by reason; for if he does, he will assuredly feel +dissatisfaction. He must likewise avoid the reprobation of the one God +or gods, in whom according to his knowledge or superstition he may +believe; but in this case the additional fear of divine punishment often +supervenes. + + +_The strictly Social Virtues at first alone regarded._—The above view +of the first origin and nature of the moral sense, which tells us what +we ought to do, and of the conscience which reproves us if we disobey +it, accords well with what we see of the early and undeveloped condition +of this faculty in mankind. The virtues which must be practised, at +least generally, by rude men, so that they may associate in a body, are +those which are still recognised as the most important. But they are +practised almost exclusively in relation to the men of the same tribe; +and their opposites are not regarded as crimes in relation to the men of +other tribes. No tribe could hold together if murder, robbery, +treachery, &c., were common; consequently such crimes within the limits +of the same tribe “are branded with everlasting infamy;”[123] but excite +no such sentiment beyond these limits. A North-American Indian is well +pleased with himself, and is honoured by others, when he scalps a man of +another tribe; and a Dyak cuts off the head of an unoffending person +and dries it as a trophy. The murder of infants has prevailed on the +largest scale throughout the world,[124] and has met with no reproach; +but infanticide, especially of females, has been thought to be good for +the tribe, or at least not injurious. Suicide during former times was +not generally considered as a crime,[125] but rather from the courage +displayed as an honourable act; and it is still largely practised by +some semi-civilised nations without reproach, for the loss to a nation +of a single individual is not felt: whatever the explanation may be, +suicide, as I hear from Sir J. Lubbock, is rarely practised by the +lowest barbarians. It has been recorded that an Indian Thug +conscientiously regretted that he had not strangled and robbed as many +travellers as did his father before him. In a rude state of civilisation +the robbery of strangers is, indeed, generally considered as honourable. + +The great sin of Slavery has been almost universal, and slaves have +often been treated in an infamous manner. As barbarians do not regard +the opinion of their women, wives are commonly treated like slaves. Most +savages are utterly indifferent to the sufferings of strangers, or even +delight in witnessing them. It is well known that the women and children +of the North-American Indians aided in torturing their enemies. Some +savages take a horrid pleasure in cruelty to animals,[126] and humanity +with them is an unknown virtue. Nevertheless, feelings of sympathy and +kindness are common, especially during sickness, between the members of +the same tribe, and are sometimes extended beyond the limits of the +tribe. Mungo Park’s touching account of the kindness of the negro women +of the interior to him is well known. Many instances could be given of +the noble fidelity of savages towards each other, but not to strangers; +common experience justifies the maxim of the Spaniard, “Never, never +trust an Indian.” There cannot be fidelity without truth; and this +fundamental virtue is not rare between the members of the same tribe: +thus Mungo Park heard the negro women teaching their young children to +love the truth. This, again, is one of the virtues which becomes so +deeply rooted in the mind that it is sometimes practised by savages even +at a high cost, towards strangers; but to lie to your enemy has rarely +been thought a sin, as the history of modern diplomacy too plainly +shews. As soon as a tribe has a recognised leader, disobedience becomes +a crime, and even abject submission is looked at as a sacred virtue. + +As during rude times no man can be useful or faithful to his tribe +without courage, this quality has universally been placed in the highest +rank; and although, in civilised countries, a good, yet timid, man may +be far more useful to the community than a brave one, we cannot help +instinctively honouring the latter above a coward, however benevolent. +Prudence, on the other hand, which does not concern the welfare of +others, though a very useful virtue, has never been highly esteemed. As +no man can practise the virtues necessary for the welfare of his tribe +without self-sacrifice, self-command, and the power of endurance, these +qualities have been at all times highly and most justly valued. The +American savage voluntarily submits without a groan to the most horrid +tortures to prove and strengthen his fortitude and courage; and we +cannot help admiring him, or even an Indian Fakir, who, from a foolish +religious motive, swings suspended by a hook buried in his flesh. + +The other self-regarding virtues, which do not obviously, though they +may really, affect the welfare of the tribe, have never been esteemed by +savages, though now highly appreciated by civilised nations. The +greatest intemperance with savages is no reproach. Their utter +licentiousness, not to mention unnatural crimes, is something +astounding.[127] As soon, however, as marriage, whether polygamous or +monogamous, becomes common, jealousy will lead to the inculcation of +female virtue; and this being honoured will tend to spread to the +unmarried females. How slowly it spreads to the male sex we see at the +present day. Chastity eminently requires self-command; therefore it has +been honoured from a very early period in the moral history of civilised +man. As a consequence of this, the senseless practice of celibacy has +been ranked from a remote period as a virtue.[128] The hatred of +indecency, which appears to us so natural as to be thought innate, and +which is so valuable an aid to chastity, is a modern virtue, +appertaining exclusively, as Sir G. Staunton remarks,[129] to civilised +life. This is shewn by the ancient religious rites of various nations, +by the drawings on the walls of Pompeii, and by the practices of many +savages. + +We have now seen that actions are regarded by savages, and were probably +so regarded by primeval man, as good or bad, solely as they affect in an +obvious manner the welfare of the tribe,—not that of the species, nor +that of man as an individual member of the tribe. This conclusion +agrees well with the belief that the so-called moral sense is +aboriginally derived from the social instincts, for both relate at first +exclusively to the community. The chief causes of the low morality of +savages, as judged by our standard, are, firstly, the confinement of +sympathy to the same tribe. Secondly, insufficient powers of reasoning, +so that the bearing of many virtues, especially of the self-regarding +virtues, on the general welfare of the tribe is not recognised. Savages, +for instance, fail to trace the multiplied evils consequent on a want of +temperance, chastity, &c. And, thirdly, weak power of self-command; for +this power has not been strengthened through long-continued, perhaps +inherited, habit, instruction and religion. + +I have entered into the above details on the immorality of savages,[130] +because some authors have recently taken a high view of their moral +nature, or have attributed most of their crimes to mistaken +benevolence.[131] These authors appear to rest their conclusion on +savages possessing, as they undoubtedly do possess, and often in a high +degree, those virtues which are serviceable, or even necessary, for the +existence of a tribal community. + + +_Concluding Remarks._—Philosophers of the derivative[132] school of +morals formerly assumed that the foundation of morality lay in a form of +Selfishness; but more recently in the “Greatest Happiness principle.” +According to the view given above, the moral sense is fundamentally +identical with the social instincts; and in the case of the lower +animals it would be absurd to speak of these instincts as having been +developed from selfishness, or for the happiness of the community. They +have, however, certainly been developed for the general good of the +community. The term, general good, may be defined as the means by which +the greatest possible number of individuals can be reared in full vigour +and health, with all their faculties perfect, under the conditions to +which they are exposed. As the social instincts both of man and the +lower animals have no doubt been developed by the same steps, it would +be advisable, if found practicable, to use the same definition in both +cases, and to take as the test of morality, the general good or welfare +of the community, rather than the general happiness; but this definition +would perhaps require some limitation on account of political ethics. + +When a man risks his life to save that of a fellow-creature, it seems +more appropriate to say that he acts for the general good or welfare, +rather than for the general happiness of mankind. No doubt the welfare +and the happiness of the individual usually coincide; and a contented, +happy tribe will flourish better than one that is discontented and +unhappy. We have seen that at an early period in the history of man, the +expressed wishes of the community will have naturally influenced to a +large extent the conduct of each member; and as all wish for happiness, +the “greatest happiness principle” will have become a most important +secondary guide and object; the social instincts, including sympathy, +always serving as the primary impulse and guide. Thus the reproach of +laying the foundation of the most noble part of our nature in the base +principle of selfishness is removed; unless indeed the satisfaction +which every animal feels when it follows its proper instincts, and the +dissatisfaction felt when prevented, be called selfish. + +The expression of the wishes and judgment of the members of the same +community, at first by oral and afterwards by written language, serves, +as just remarked, as a most important secondary guide of conduct, in aid +of the social instincts, but sometimes in opposition to them. This +latter fact is well exemplified by the _Law of Honour_, that is the law +of the opinion of our equals, and not of all our countrymen. The breach +of this law, even when the breach is known to be strictly accordant with +true morality, has caused many a man more agony than a real crime. We +recognise the same influence in the burning sense of shame which most of +us have felt even after the interval of years, when calling to mind some +accidental breach of a trifling though fixed rule of etiquette. The +judgment of the community will generally be guided by some rude +experience of what is best in the long run for all the members; but this +judgment will not rarely err from ignorance and from weak powers of +reasoning. Hence the strangest customs and superstitions, in complete +opposition to the true welfare and happiness of mankind, have become +all-powerful throughout the world. We see this in the horror felt by a +Hindoo who breaks his caste, in the shame of a Mahometan woman who +exposes her face, and in innumerable other instances. It would be +difficult to distinguish between the remorse felt by a Hindoo who has +eaten unclean food, from that felt after committing a theft; but the +former would probably be the more severe. + +How so many absurd rules of conduct, as well as so many absurd religious +beliefs, have originated we do not know; nor how it is that they have +become, in all quarters of the world, so deeply impressed on the mind +of men; but it is worthy of remark that a belief constantly inculcated +during the early years of life, whilst the brain is impressible, appears +to acquire almost the nature of an instinct; and the very essence of an +instinct is that it is followed independently of reason. Neither can we +say why certain admirable virtues, such as the love of truth, are much +more highly appreciated by some savage tribes than by others;[133] nor, +again, why similar differences prevail even amongst civilised nations. +Knowing how firmly fixed many strange customs and superstitions have +become, we need feel no surprise that the self-regarding virtues should +now appear to us so natural, supported as they are by reason, as to be +thought innate, although they were not valued by man in his early +condition. + +Notwithstanding many sources of doubt, man can generally and readily +distinguish between the higher and lower moral rules. The higher are +founded on the social instincts, and relate to the welfare of others. +They are supported by the approbation of our fellow-men and by reason. +The lower rules, though some of them when implying self-sacrifice hardly +deserve to be called lower, relate chiefly to self, and owe their origin +to public opinion, when matured by experience and cultivated; for they +are not practised by rude tribes. + +As man advances in civilisation, and small tribes are united into larger +communities, the simplest reason would tell each individual that he +ought to extend his social instincts and sympathies to all the members +of the same nation, though personally unknown to him. This point being +once reached, there is only an artificial barrier to prevent his +sympathies extending to the men of all nations and races. If, indeed, +such men are separated from him by great differences in appearance or +habits, experience unfortunately shews us how long it is before we look +at them as our fellow-creatures. Sympathy beyond the confines of man, +that is humanity to the lower animals, seems to be one of the latest +moral acquisitions. It is apparently unfelt by savages, except towards +their pets. How little the old Romans knew of it is shewn by their +abhorrent gladiatorial exhibitions. The very idea of humanity, as far as +I could observe, was new to most of the Gauchos of the Pampas. This +virtue, one of the noblest with which man is endowed, seems to arise +incidentally from our sympathies becoming more tender and more widely +diffused, until they are extended to all sentient beings. As soon as +this virtue is honoured and practised by some few men, it spreads +through instruction and example to the young, and eventually through +public opinion. + +The highest stage in moral culture at which we can arrive, is when we +recognise that we ought to control our thoughts, and “not even in inmost +thought to think again the sins that made the past so pleasant to +us.”[134] Whatever makes any bad action familiar to the mind, renders +its performance by so much the easier. As Marcus Aurelius long ago said, +“Such as are thy habitual thoughts, such also will be the character of +thy mind; for the soul is dyed by the thoughts.”[135] + +Our great philosopher, Herbert Spencer, has recently explained his views +on the moral sense. He says,[136] “I believe that the experiences of +utility organised and consolidated through all past generations of the +human race, have been producing corresponding modifications, which, by +continued transmission and accumulation, have become in us certain +faculties of moral intuition—certain emotions responding to right and +wrong conduct, which have no apparent basis in the individual +experiences of utility.” There is not the least inherent improbability, +as it seems to me, in virtuous tendencies being more or less strongly +inherited; for, not to mention the various dispositions and habits +transmitted by many of our domestic animals, I have heard of cases in +which a desire to steal and a tendency to lie appeared to run in +families of the upper ranks; and as stealing is so rare a crime in the +wealthy classes, we can hardly account by accidental coincidence for the +tendency occurring in two or three members of the same family. If bad +tendencies are transmitted, it is probable that good ones are likewise +transmitted. Excepting through the principle of the transmission of +moral tendencies, we cannot understand the differences believed to exist +in this respect between the various races of mankind. We have, however, +as yet, hardly sufficient evidence on this head. + +Even the partial transmission of virtuous tendencies would be an immense +assistance to the primary impulse derived directly from the social +instincts, and indirectly from the approbation of our fellow-men. +Admitting for the moment that virtuous tendencies are inherited, it +appears probable, at least in such cases as chastity, temperance, +humanity to animals, &c., that they become first impressed on the mental +organisation through habit, instruction, and example, continued during +several generations in the same family, and in a quite subordinate +degree, or not at all, by the individuals possessing such virtues, +having succeeded best in the struggle for life. My chief source of doubt +with respect to any such inheritance, is that senseless customs, +superstitions, and tastes, such as the horror of a Hindoo for unclean +food, ought on the same principle to be transmitted. Although this in +itself is perhaps not less probable than that animals should acquire +inherited tastes for certain kinds of food or fear of certain foes, I +have not met with any evidence in support of the transmission of +superstitious customs or senseless habits. + + +Finally, the social instincts which no doubt were acquired by man, as by +the lower animals, for the good of the community, will from the first +have given to him some wish to aid his fellows, and some feeling of +sympathy. Such impulses will have served him at a very early period as a +rude rule of right and wrong. But as man gradually advanced in +intellectual power and was enabled to trace the more remote consequences +of his actions; as he acquired sufficient knowledge to reject baneful +customs and superstitions; as he regarded more and more not only the +welfare but the happiness of his fellow-men; as from habit, following on +beneficial experience, instruction, and example, his sympathies became +more tender and widely diffused, so as to extend to the men of all +races, to the imbecile, the maimed, and other useless members of +society, and finally to the lower animals,—so would the standard of his +morality rise higher and higher. And it is admitted by moralists of the +derivative school and by some intuitionists, that the standard of +morality has risen since an early period in the history of man.[137] + +As a struggle may sometimes be seen going on between the various +instincts of the lower animals, it is not surprising that there should +be a struggle in man between his social instincts, with their derived +virtues, and his lower, though at the moment, stronger impulses or +desires. This, as Mr. Galton[138] has remarked, is all the less +surprising, as man has emerged from a state of barbarism within a +comparatively recent period. After having yielded to some temptation we +feel a sense of dissatisfaction, analogous to that felt from other +unsatisfied instincts, called in this case conscience; for we cannot +prevent past images and impressions continually passing through our +minds, and these in their weakened state we compare with the +ever-present social instincts, or with habits gained in early youth and +strengthened during our whole lives, perhaps inherited, so that they are +at last rendered almost as strong as instincts. Looking to future +generations, there is no cause to fear that the social instincts will +grow weaker, and we may expect that virtuous habits will grow stronger, +becoming perhaps fixed by inheritance. In this case the struggle between +our higher and lower impulses will be less severe, and virtue will be +triumphant. + + +_Summary of the two last Chapters._—There can be no doubt that the +difference between the mind of the lowest man and that of the highest +animal is immense. An anthropomorphous ape, if he could take a +dispassionate view of his own case, would admit that though he could +form an artful plan to plunder a garden—though he could use stones for +fighting or for breaking open nuts, yet that the thought of fashioning +a stone into a tool was quite beyond his scope. Still less, as he would +admit, could he follow out a train of metaphysical reasoning, or solve a +mathematical problem, or reflect on God, or admire a grand natural +scene. Some apes, however, would probably declare that they could and +did admire the beauty of the coloured skin and fur of their partners in +marriage. They would admit, that though they could make other apes +understand by cries some of their perceptions and simpler wants, the +notion of expressing definite ideas by definite sounds had never crossed +their minds. They might insist that they were ready to aid their +fellow-apes of the same troop in many ways, to risk their lives for +them, and to take charge of their orphans; but they would be forced to +acknowledge that disinterested love for all living creatures, the most +noble attribute of man, was quite beyond their comprehension. + +Nevertheless the difference in mind between man and the higher animals, +great as it is, is certainly one of degree and not of kind. We have seen +that the senses and intuitions, the various emotions and faculties, such +as love, memory, attention, curiosity, imitation, reason, &c., of which +man boasts, may be found in an incipient, or even sometimes in a +well-developed condition, in the lower animals. They are also capable of +some inherited improvement, as we see in the domestic dog compared with +the wolf or jackal. If it be maintained that certain powers, such as +self-consciousness, abstraction, &c., are peculiar to man, it may well +be that these are the incidental results of other highly-advanced +intellectual faculties; and these again are mainly the result of the +continued use of a highly developed language. At what age does the +new-born infant possess the power of abstraction, or become +self-conscious and reflect on its own existence? We cannot answer; nor +can we answer in regard to the ascending organic scale. The half-art and +half-instinct of language still bears the stamp of its gradual +evolution. The ennobling belief in God is not universal with man; and +the belief in active spiritual agencies naturally follows from his other +mental powers. The moral sense perhaps affords the best and highest +distinction between man and the lower animals; but I need not say +anything on this head, as I have so lately endeavoured to shew +that the social instincts,—the prime principle of man’s moral +constitution[139]—with the aid of active intellectual powers and the +effects of habit, naturally lead to the golden rule, “As ye would that +men should do to you, do ye to them likewise;” and this lies at the +foundation of morality. + +In a future chapter I shall make some few remarks on the probable steps +and means by which the several mental and moral faculties of man have +been gradually evolved. That this at least is possible ought not to be +denied, when we daily see their development in every infant; and when we +may trace a perfect gradation from the mind of an utter idiot, lower +than that of the lowest animal, to the mind of a Newton. + + + + +CHAPTER IV. + +ON THE MANNER OF DEVELOPMENT OF MAN FROM SOME LOWER FORM. + + + Variability of body and mind in man—Inheritance—Causes of + variability—Laws of variation the same in man as in the lower + animals—Direct action of the conditions of life—Effects of + the increased use and disuse of parts—Arrested + development—Reversion—Correlated variation—Rate of + increase—Checks to increase—Natural selection—Man the most + dominant animal in the world—Importance of his corporeal + structure—The causes which have led to his becoming + erect—Consequent changes of structure—Decrease in size of the + canine teeth—Increased size and altered shape of the + skull—Nakedness—Absence of a tail—Defenceless condition of + man. + + +We have seen in the first chapter that the homological structure of man, +his embryological development and the rudiments which he still retains, +all declare in the plainest manner that he is descended from some lower +form. The possession of exalted mental powers is no insuperable +objection to this conclusion. In order that an ape-like creature should +have been transformed into man, it is necessary that this early form, as +well as many successive links, should all have varied in mind and body. +It is impossible to obtain direct evidence on this head; but if it can +be shewn that man now varies—that his variations are induced by the +same general causes, and obey the same general laws, as in the case of +the lower animals—there can be little doubt that the preceding +intermediate links varied in a like manner. The variations at each +successive stage of descent must, also, have been in some manner +accumulated and fixed. + +The facts and conclusions to be given in this chapter relate almost +exclusively to the probable means by which the transformation of man has +been effected, as far as his bodily structure is concerned. The +following chapter will be devoted to the development of his intellectual +and moral faculties. But the present discussion likewise bears on the +origin of the different races or species of mankind, whichever term may +be preferred. + +It is manifest that man is now subject to much variability. No two +individuals of the same race are quite alike. We may compare millions of +faces, and each will be distinct. There is an equally great amount of +diversity in the proportions and dimensions of the various parts of the +body; the length of the legs being one of the most variable points.[140] +Although in some quarters of the world an elongated skull, and in other +quarters a short skull prevails, yet there is great diversity of shape +even within the limits of the same race, as with the aborigines of +America and South Australia,—the latter a race “probably as pure and +homogeneous in blood, customs, and language as any in existence”—and +even with the inhabitants of so confined an area as the Sandwich +Islands.[141] An eminent dentist assures me that there is nearly as much +diversity in the teeth, as in the features. The chief arteries so +frequently run in abnormal courses, that it has been found useful for +surgical purposes to calculate from 12,000 corpses how often each +course prevails.[142] The muscles are eminently variable: thus those of +the foot were found by Prof. Turner[143] not to be strictly alike in any +two out of fifty bodies; and in some the deviations were considerable. +Prof. Turner adds that the power of performing the appropriate movements +must have been modified in accordance with the several deviations. Mr. +J. Wood has recorded[144] the occurrence of 295 muscular variations in +thirty-six subjects, and in another set of the same number no less than +558 variations, reckoning both sides of the body as one. In the last +set, not one body out of the thirty-six was “found totally wanting in +departures from the standard descriptions of the muscular system given +in anatomical text-books.” A single body presented the extraordinary +number of twenty-five distinct abnormalities. The same muscle sometimes +varies in many ways: thus Prof. Macalister describes[145] no less than +twenty distinct variations in the _palmaris accessorius_. + +The famous old anatomist, Wolff,[146] insists that the internal viscera +are more variable than the external parts: _Nulla particula est quæ non +aliter et aliter in aliis se habeat hominibus._ He has even written a +treatise on the choice of typical examples of the viscera for +representation. A discussion on the beau-ideal of the liver, lungs, +kidneys, &c., as of the human face divine, sounds strange in our ears. + +The variability or diversity of the mental faculties in men of the same +race, not to mention the greater differences between the men of +distinct races, is so notorious that not a word need here be said. So it +is with the lower animals, as has been illustrated by a few examples in +the last chapter. All who have had charge of menageries admit this fact, +and we see it plainly in our dogs and other domestic animals. Brehm +especially insists that each individual monkey of those which he kept +under confinement in Africa had its own peculiar disposition and temper: +he mentions one baboon remarkable for its high intelligence; and the +keepers in the Zoological Gardens pointed out to me a monkey, belonging +to the New World division, equally remarkable for intelligence. Rengger, +also, insists on the diversity in the various mental characters of the +monkeys of the same species which he kept in Paraguay; and this +diversity, as he adds, is partly innate, and partly the result of the +manner in which they have been treated or educated.[147] + +I have elsewhere[148] so fully discussed the subject of Inheritance that +I need here add hardly anything. A greater number of facts have been +collected with respect to the transmission of the most trifling, as well +as of the most important characters in man than in any of the lower +animals; though the facts are copious enough with respect to the latter. +So in regard to mental qualities, their transmission is manifest in our +dogs, horses, and other domestic animals. Besides special tastes and +habits, general intelligence, courage, bad and good temper, &c., are +certainly transmitted. With man we see similar facts in almost every +family; and we now know through the admirable labours of Mr. Galton[149] +that genius, which implies a wonderfully complex combination of high +faculties, tends to be inherited; and, on the other hand, it is too +certain that insanity and deteriorated mental powers likewise run in the +same families. + +With respect to the causes of variability we are in all cases very +ignorant; but we can see that in man as in the lower animals, they stand +in some relation with the conditions to which each species has been +exposed during several generations. Domesticated animals vary more than +those in a state of nature; and this is apparently due to the +diversified and changing nature of their conditions. The different races +of man resemble in this respect domesticated animals, and so do the +individuals of the same race when inhabiting a very wide area, like that +of America. We see the influence of diversified conditions in the more +civilised nations, the members of which belong to different grades of +rank and follow different occupations, presenting a greater range of +character than the members of barbarous nations. But the uniformity of +savages has often been exaggerated, and in some cases can hardly be said +to exist.[150] It is nevertheless an error to speak of man, even if we +look only to the conditions to which he has been subjected, as “far more +domesticated”[151] than any other animal. Some savage races, such as +the Australians, are not exposed to more diversified conditions than are +many species which have very wide ranges. In another and much more +important respect, man differs widely from any strictly domesticated +animal; for his breeding has not been controlled, either through +methodical or unconscious selection. No race or body of men has been so +completely subjugated by other men, that certain individuals have been +preserved and thus unconsciously selected, from being in some way more +useful to their masters. Nor have certain male and female individuals +been intentionally picked out and matched, except in the well-known case +of the Prussian grenadiers; and in this case man obeyed, as might have +been expected, the law of methodical selection; for it is asserted that +many tall men were reared in the villages inhabited by the grenadiers +with their tall wives. + +If we consider all the races of man, as forming a single species, his +range is enormous; but some separate races, as the Americans and +Polynesians, have very wide ranges. It is a well-known law that +widely-ranging species are much more variable than species with +restricted ranges; and the variability of man may with more truth be +compared with that of widely-ranging species, than with that of +domesticated animals. + +Not only does variability appear to be induced in man and the lower +animals by the same general causes, but in both the same characters are +affected in a closely analogous manner. This has been proved in such +full detail by Godron and Quatrefages, that I need here only refer to +their works.[152] Monstrosities, which graduate into slight variations, +are likewise so similar in man and the lower animals, that the same +classification and the same terms can be used for both, as may be seen +in Isidore Geoffroy St.-Hilaire’s great work.[153] This is a necessary +consequence of the same laws of change prevailing throughout the animal +kingdom. In my work on the variation of domestic animals, I have +attempted to arrange in a rude fashion the laws of variation under the +following heads:—The direct and definite action of changed conditions, +as shewn by all or nearly all the individuals of the same species +varying in the same manner under the same circumstances. The effects of +the long-continued use or disuse of parts. The cohesion of homologous +parts. The variability of multiple parts. Compensation of growth; but of +this law I have found no good instances in the case of man. The effects +of the mechanical pressure of one part on another; as of the pelvis on +the cranium of the infant in the womb. Arrests of development, leading +to the diminution or suppression of parts. The reappearance of long-lost +characters through reversion. And lastly, correlated variation. All +these so-called laws apply equally to man and the lower animals; and +most of them even to plants. It would be superfluous here to discuss all +of them;[154] but several are so important for us, that they must be +treated at considerable length. + + +_The direct and definite action of changed conditions._—This is a most +perplexing subject. It cannot be denied that changed conditions produce +some effect, and occasionally a considerable effect, on organisms of all +kinds; and it seems at first probable that if sufficient time were +allowed this would be the invariable result. But I have failed to obtain +clear evidence in favour of this conclusion; and valid reasons may be +urged on the other side, at least as far as the innumerable structures +are concerned, which are adapted for special ends. There can, however, +be no doubt that changed conditions induce an almost indefinite amount +of fluctuating variability, by which the whole organisation is rendered +in some degree plastic. + +In the United States, above 1,000,000 soldiers, who served in the late +war, were measured, and the States in which they were born and reared +recorded.[155] From this astonishing number of observations it is proved +that local influences of some kind act directly on stature; and we +further learn that “the State where the physical growth has in great +measure taken place, and the State of birth, which indicates the +ancestry, seem to exert a marked influence on the stature.” For instance +it is established, “that residence in the Western States, during the +years of growth, tends to produce increase of stature.” On the other +hand, it is certain that with sailors, their manner of life delays +growth, as shewn “by the great difference between the statures of +soldiers and sailors at the ages of 17 and 18 years.” Mr. B. A. Gould +endeavoured to ascertain the nature of the influences which thus act on +stature; but he arrived only at negative results, namely, that they did +not relate to climate, the elevation of the land, soil, nor even “in any +controlling degree” to the abundance or need of the comforts of life. +This latter conclusion is directly opposed to that arrived at by +Villermé from the statistics of the height of the conscripts in +different parts of France. When we compare the differences in stature +between the Polynesian chiefs and the lower orders within the same +islands, or between the inhabitants of the fertile volcanic and low +barren coral islands of the same ocean,[156] or again between the +Fuegians on the eastern and western shores of their country, where the +means of subsistence are very different, it is scarcely possible to +avoid the conclusion that better food and greater comfort do influence +stature. But the preceding statements shew how difficult it is to arrive +at any precise result. Dr. Beddoe has lately proved that, with the +inhabitants of Britain, residence in towns and certain occupations have +a deteriorating influence on height; and he infers that the result is to +a certain extent inherited, as is likewise the case in the United +States. Dr. Beddoe further believes that wherever a “race attains its +maximum of physical development, it rises highest in energy and moral +vigour.”[157] + +Whether external conditions produce any other direct effect on man is +not known. It might have been expected that differences of climate would +have had a marked influence, as the lungs and kidneys are brought into +fuller activity under a low temperature, and the liver and skin under a +high one.[158] It was formerly thought that the colour of the skin and +the character of the hair were determined by light or heat; and +although it can hardly be denied that some effect is thus produced, +almost all observers now agree that the effect has been very small, even +after exposure during many ages. But this subject will be more properly +discussed when we treat of the different races of mankind. With our +domestic animals there are grounds for believing that cold and damp +directly affect the growth of the hair; but I have not met with any +evidence on this head in the case of man. + + +_Effects of the increased Use and Disuse of Parts._—It is well known +that use strengthens the muscles in the individual, and complete disuse, +or the destruction of the proper nerve, weakens them. When the eye is +destroyed the optic nerve often becomes atrophied. When an artery is +tied, the lateral channels increase not only in diameter, but in the +thickness and strength of their coats. When one kidney ceases acting +from disease, the other increases in size and does double work. Bones +increase not only in thickness, but in length, from carrying a greater +weight.[159] Different occupations habitually followed lead to changed +proportions in various parts of the body. Thus it was clearly +ascertained by the United States Commission[160] that the legs of the +sailors employed in the late war were longer by 0.217 of an inch than +those of the soldiers, though the sailors were on an average shorter +men; whilst their arms were shorter by 1.09 of an inch, and therefore +out of proportion shorter in relation to their lesser height. This +shortness of the arms is apparently due to their greater use, and is an +unexpected result; but sailors chiefly use their arms in pulling and not +in supporting weights. The girth of the neck and the depth of the instep +are greater, whilst the circumference of the chest, waist, and hips is +less in sailors than in soldiers. + +Whether the several foregoing modifications would become hereditary, if +the same habits of life were followed during many generations, is not +known, but is probable. Rengger[161] attributes the thin legs and thick +arms of the Payaguas Indians to successive generations having passed +nearly their whole lives in canoes, with their lower extremities +motionless. Other writers have come to a similar conclusion in other +analogous cases. According to Cranz,[162] who lived for a long time with +the Esquimaux, “the natives believe that ingenuity and dexterity in +seal-catching (their highest art and virtue) is hereditary; there is +really something in it, for the son of a celebrated seal-catcher will +distinguish himself though he lost his father in childhood.” But in this +case it is mental aptitude, quite as much as bodily structure, which +appears to be inherited. It is asserted that the hands of English +labourers are at birth larger than those of the gentry.[163] From the +correlation which exists, at least in some cases,[164] between the +development of the extremities and of the jaws, it is possible that in +those classes which do not labour much with their hands and feet, the +jaws would be reduced in size from this cause. That they are generally +smaller in refined and civilised men than in hard-working men or +savages, is certain. But with savages, as Mr. Herbert Spencer[165] has +remarked, the greater use of the jaws in chewing coarse, uncooked food, +would act in a direct manner on the masticatory muscles and on the bones +to which they are attached. In infants long before birth, the skin on +the soles of the feet is thicker than on any other part of the +body;[166] and it can hardly be doubted that this is due to the +inherited effects of pressure during a long series of generations. + +It is familiar to every one that watchmakers and engravers are liable to +become short-sighted, whilst sailors and especially savages are +generally long-sighted. Short-sight and long-sight certainly tend to be +inherited.[167] The inferiority of Europeans, in comparison with +savages, in eyesight and in the other senses, is no doubt the +accumulated and transmitted effect of lessened use during many +generations; for Rengger[168] states that he has repeatedly observed +Europeans, who had been brought up and spent their whole lives with the +wild Indians, who nevertheless did not equal them in the sharpness of +their senses. The same naturalist observes that the cavities in the +skull for the reception of the several sense-organs are larger in the +American aborigines than in Europeans; and this no doubt indicates a +corresponding difference in the dimensions of the organs themselves. +Blumenbach has also remarked on the large size of the nasal cavities in +the skulls of the American aborigines, and connects this fact with their +remarkably acute power of smell. The Mongolians of the plains of +Northern Asia, according to Pallas, have wonderfully perfect senses; and +Prichard believes that the great breadth of their skulls across the +zygomas follows from their highly-developed sense-organs.[169] + +The Quechua Indians inhabit the lofty plateaux of Peru, and Alcide +d’Orbigny states[170] that from continually breathing a highly rarefied +atmosphere they have acquired chests and lungs of extraordinary +dimensions. The cells, also, of the lungs are larger and more numerous +than in Europeans. These observations have been doubted; but Mr. D. +Forbes carefully measured many Aymaras, an allied race, living at the +height of between ten and fifteen thousand feet; and he informs me[171] +that they differ conspicuously from the men of all other races seen by +him, in the circumference and length of their bodies. In his table of +measurements, the stature of each man is taken at 1000, and the other +measurements are reduced to this standard. It is here seen that the +extended arms of the Aymaras are shorter than those of Europeans, and +much shorter than those of Negroes. The legs are likewise shorter, and +they present this remarkable peculiarity, that in every Aymara measured +the femur is actually shorter than the tibia. On an average the length +of the femur to that of the tibia is as 211 to 252; whilst in two +Europeans measured at the same time, the femora to the tibiæ were as +244 to 230; and in three Negroes as 258 to 241. The humerus is likewise +shorter relatively to the fore-arm. This shortening of that part of the +limb which is nearest to the body, appears to be, as suggested to me by +Mr. Forbes, a case of compensation in relation with the greatly +increased length of the trunk. The Aymaras present some other singular +points of structure, for instance, the very small projection of the +heel. + +These men are so thoroughly acclimatised to their cold and lofty abode, +that when formerly carried down by the Spaniards to the low Eastern +plains, and when now tempted down by high wages to the gold-washings, +they suffer a frightful rate of mortality. Nevertheless Mr. Forbes found +a few pure families which had survived during two generations; and he +observed that they still inherited their characteristic peculiarities. +But it was manifest, even without measurement, that these peculiarities +had all decreased; and on measurement their bodies were found not to be +so much elongated as those of the men on the high plateau; whilst their +femora had become somewhat lengthened, as had their tibiæ but in a less +degree. The actual measurements may be seen by consulting Mr. Forbes’ +memoir. From these valuable observations, there can, I think, be no +doubt that residence during many generations at a great elevation tends, +both directly and indirectly, to induce inherited modifications in the +proportions of the body.[172] + +Although man may not have been much modified during the latter stages of +his existence through the increased or decreased use of parts, the +facts now given shew that his liability in this respect has not been +lost; and we positively know that the same law holds good with the lower +animals. Consequently we may infer, that when at a remote epoch the +progenitors of man were in a transitional state, and were changing from +quadrupeds into bipeds, natural selection would probably have been +greatly aided by the inherited effects of the increased or diminished +use of the different parts of the body. + + +_Arrests of Development._—Arrested development differs from arrested +growth, as parts in the former state continue to grow whilst still +retaining their early condition. Various monstrosities come under this +head, and some are known to be occasionally inherited, as a +cleft-palate. It will suffice for our purpose to refer to the arrested +brain-development of microcephalous idiots, as described in Vogt’s great +memoir.[173] Their skulls are smaller, and the convolutions of the brain +are less complex than in normal men. The frontal sinus, or the +projection over the eyebrows, is largely developed, and the jaws are +prognathous to an “_effrayant_” degree; so that these idiots somewhat +resemble the lower types of mankind. Their intelligence and most of +their mental faculties are extremely feeble. They cannot acquire the +power of speech, and are wholly incapable of prolonged attention, but +are much given to imitation. They are strong and remarkably active, +continually gamboling and jumping about, and making grimaces. They often +ascend stairs on all-fours; and are curiously fond of climbing up +furniture or trees. We are thus reminded of the delight shewn by almost +all boys in climbing trees; and this again reminds us how lambs and +kids, originally alpine animals, delight to frisk on any hillock, +however small. + + +_Reversion._—Many of the cases to be here given might have been +introduced under the last heading. Whenever a structure is arrested in +its development, but still continues growing until it closely resembles +a corresponding structure in some lower and adult member of the same +group, we may in one sense consider it as a case of reversion. The lower +members in a group give us some idea how the common progenitor of the +group was probably constructed; and it is hardly credible that a part +arrested at an early phase of embryonic development should be enabled to +continue growing so as ultimately to perform its proper function, unless +it had acquired this power of continued growth during some earlier state +of existence, when the present exceptional or arrested structure was +normal. The simple brain of a microcephalous idiot, in as far as it +resembles that of an ape, may in this sense be said to offer a case of +reversion. There are other cases which come more strictly under our +present heading of reversion. Certain structures, regularly occurring in +the lower members of the group to which man belongs, occasionally make +their appearance in him, though not found in the normal human embryo; +or, if present in the normal human embryo, they become developed in an +abnormal manner, though this manner of development is proper to the +lower members of the same group. These remarks will be rendered clearer +by the following illustrations. + +In various mammals the uterus graduates from a double organ with two +distinct orifices and two passages, as in the marsupials, into a single +organ, showing no signs of doubleness except a slight internal fold, as +in the higher apes and man. The rodents exhibit a perfect series of +gradations between these two extreme states. In all mammals the uterus +is developed from two simple primitive tubes, the inferior portions of +which form the cornua; and it is in the words of Dr. Farre “by the +coalescence of the two cornua at their lower extremities that the body +of the uterus is formed in man; while in those animals in which no +middle portion or body exists, the cornua remain ununited. As the +development of the uterus proceeds, the two cornua become gradually +shorter, until at length they are lost, or, as it were, absorbed into +the body of the uterus.” The angles of the uterus are still produced +into cornua, even so high in the scale as in the lower apes, and their +allies the lemurs. + +Now in women anomalous cases are not very infrequent, in which the +mature uterus is furnished with cornua, or is partially divided into two +organs; and such cases, according to Owen, repeat “the grade of +concentrative development,” attained by certain rodents. Here perhaps we +have an instance of a simple arrest of embryonic development, with +subsequent growth and perfect functional development, for either side of +the partially double uterus is capable of performing the proper office +of gestation. In other and rarer cases, two distinct uterine cavities +are formed, each having its proper orifice and passage.[174] No such +stage is passed through during the ordinary development of the embryo, +and it is difficult to believe, though perhaps not impossible, that the +two simple, minute, primitive tubes could know how (if such an +expression may be used) to grow into two distinct uteri, each with a +well-constructed orifice and passage, and each furnished with numerous +muscles, nerves, glands and vessels, if they had not formerly passed +through a similar course of development, as in the case of existing +marsupials. No one will pretend that so perfect a structure as the +abnormal double uterus in woman could be the result of mere chance. But +the principle of reversion, by which long-lost dormant structures are +called back into existence, might serve as the guide for the full +development of the organ, even after the lapse of an enormous interval +of time. + +Professor Canestrini,[175] after discussing the foregoing and various +analogous cases, arrives at the same conclusion as that just given. He +adduces, as another instance, the malar bone, which, in some of the +Quadrumana and other mammals, normally consists of two portions. This is +its condition in the two-months-old human fœtus; and thus it +sometimes remains, through arrested development, in man when adult, more +especially in the lower prognathous races. Hence Canestrini concludes +that some ancient progenitor of man must have possessed this bone +normally divided into two portions, which subsequently became fused +together. In man the frontal bone consists of a single piece, but in the +embryo and in children, and in almost all the lower mammals, it consists +of two pieces separated by a distinct suture. This suture occasionally +persists, more or less distinctly, in man after maturity, and more +frequently in ancient than in recent crania, especially as Canestrini +has observed in those exhumed from the Drift and belonging to the +brachycephalic type. Here again he comes to the same conclusion as in +the analogous case of the malar bones. In this and other instances +presently to be given, the cause of ancient races approaching the lower +animals in certain characters more frequently than do the modern races, +appears to be that the latter stand at a somewhat greater distance in +the long line of descent from their early semi-human progenitors. + +Various other anomalies in man, more or less analogous with the +foregoing, have been advanced by different authors[176] as cases of +reversion; but these seem not a little doubtful, for we have to descend +extremely low in the mammalian series before we find such structures +normally present.[177] + +In man the canine teeth are perfectly efficient instruments for +mastication. But their true canine character, as Owen[178] remarks, “is +indicated by the conical form of the crown, which terminates in an +obtuse point, is convex outward and flat or sub-concave within, at the +base of which surface there is a feeble prominence. The conical form is +best expressed in the Melanian races, especially the Australian. The +canine is more deeply implanted, and by a stronger fang than the +incisors.” Nevertheless this tooth no longer serves man as a special +weapon for tearing his enemies or prey; it may, therefore, as far as its +proper function is concerned, be considered as rudimentary. In every +large collection of human skulls some may be found, as Häckel[179] +observes, with the canine teeth projecting considerably beyond the +others in the same manner, but in a less degree, as in the +anthropomorphous apes. In these cases, open spaces between the teeth in +the one jaw are left for the reception of the canines belonging to the +opposite jaw. An interspace of this kind in a Kaffir skull, figured by +Wagner, is surprisingly wide.[180] Considering how few ancient skulls +have been examined in comparison with recent skulls, it is an +interesting fact that in at least three cases the canines project +largely; and in the Naulette jaw they are spoken of as enormous.[181] + +The males alone of the anthropomorphous apes have their canines fully +developed; but in the female gorilla, and in a less degree in the female +orang, these teeth project considerably beyond the others; therefore the +fact that women sometimes have, as I have been assured, considerably +projecting canines, is no serious objection to the belief that their +occasional great development in man is a case of reversion to an +ape-like progenitor. He who rejects with scorn the belief that the shape +of his own canines, and their occasional great development in other men, +are due to our early progenitors having been provided with these +formidable weapons, will probably reveal by sneering the line of his +descent. For though he no longer intends, nor has the power, to use +these teeth as weapons, he will unconsciously retract his “snarling +muscles” (thus named by Sir C. Bell)[182] so as to expose them ready for +action, like a dog prepared to fight. + +Many muscles are occasionally developed in man, which are proper to the +Quadrumana or other mammals. Professor Vlacovich[183] examined forty +male subjects, and found a muscle, called by him the ischiopubic, in +nineteen of them; in three others there was a ligament which represented +this muscle; and in the remaining eighteen no trace of it. Out of thirty +female subjects this muscle was developed on both sides in only two, but +in three others the rudimentary ligament was present. This muscle, +therefore, appears to be much more common in the male than in the female +sex; and on the principle of the descent of man from some lower form, +its presence can be understood; for it has been detected in several of +the lower animals, and in all of these it serves exclusively to aid the +male in the act of reproduction. + +Mr. J. Wood, in his valuable series of papers,[184] has minutely +described a vast number of muscular variations in man, which resemble +normal structures in the lower animals. Looking only to the muscles +which closely resemble those regularly present in our nearest allies, +the Quadrumana, they are too numerous to be here even specified. In a +single male subject, having a strong bodily frame and well-formed skull, +no less than seven muscular variations were observed, all of which +plainly represented muscles proper to various kinds of apes. This man, +for instance, had on both sides of his neck a true and powerful +“_levator claviculæ_,” such as is found in all kinds of apes, and which +is said to occur in about one out of sixty human subjects.[185] Again, +this man had “a special abductor of the metatarsal bone of the fifth +digit, such as Professor Huxley and Mr. Flower have shewn to exist +uniformly in the higher and lower apes.” The hands and arms of man are +eminently characteristic structures, but their muscles are extremely +liable to vary, so as to resemble the corresponding muscles in the lower +animals.[186] Such resemblances are either complete and perfect or +imperfect, yet in this latter case manifestly of a transitional nature. +Certain variations are more common in man, and others in woman, without +our being able to assign any reason. Mr. Wood, after describing numerous +cases, makes the following pregnant remark: “Notable departures from the +ordinary type of the muscular structures run in grooves or directions, +which must be taken to indicate some unknown factor, of much importance +to a comprehensive knowledge of general and scientific anatomy.”[187] + +That this unknown factor is reversion to a former state of existence may +be admitted as in the highest degree probable. It is quite incredible +that a man should through mere accident abnormally resemble, in no less +than seven of his muscles, certain apes, if there had been no genetic +connection between them. On the other hand, if man is descended from +some ape-like creature, no valid reason can be assigned why certain +muscles should not suddenly reappear after an interval of many thousand +generations, in the same manner as with horses, asses, and mules, +dark-coloured stripes suddenly reappear on the legs and shoulders, after +an interval of hundreds, or more probably thousands, of generations. + +These various cases of reversion are so closely related to those of +rudimentary organs given in the first chapter, that many of them might +have been indifferently introduced in either chapter. Thus a human +uterus furnished with cornua may be said to represent in a rudimentary +condition the same organ in its normal state in certain mammals. Some +parts which are rudimental in man, as the os coccyx in both sexes and +the mammæ in the male sex, are always present; whilst others, such as +the supra-condyloid foramen, only occasionally appear, and therefore +might have been introduced under the head of reversion. These several +reversionary, as well as the strictly rudimentary, structures reveal the +descent of man from some lower form in an unmistakeable manner. + + +_Correlated Variation._—In man, as in the lower animals, many +structures are so intimately related, that when one part varies so does +another, without our being able, in most cases, to assign any reason. We +cannot say whether the one part governs the other, or whether both are +governed by some earlier developed part. Various monstrosities, as I. +Geoffroy repeatedly insists, are thus intimately connected. Homologous +structures are particularly liable to change together, as we see on the +opposite sides of the body, and in the upper and lower extremities. +Meckel long ago remarked that when the muscles of the arm depart from +their proper type, they almost always imitate those of the leg; and so +conversely with the muscles of the legs. The organs of sight and +hearing, the teeth and hair, the colour of the skin and hair, colour and +constitution, are more or less correlated.[188] Professor Schaaffhausen +first drew attention to the relation apparently existing between a +muscular frame and strongly-pronounced supra-orbital ridges, which are +so characteristic of the lower races of man. + +Besides the variations which can be grouped with more or less +probability under the foregoing heads, there is a large class of +variations which may be provisionally called spontaneous, for they +appear, owing to our ignorance, to arise without any exciting cause. It +can, however, be shewn that such variations, whether consisting of +slight individual differences, or of strongly-marked and abrupt +deviations of structure, depend much more on the constitution of the +organism than on the nature of the conditions to which it has been +subjected.[189] + + +_Rate of Increase._—Civilised populations have been known under +favourable conditions, as in the United States, to double their number +in twenty-five years; and according to a calculation by Euler, this +might occur in a little over twelve years.[190] At the former rate the +present population of the United States, namely, thirty millions, would +in 657 years cover the whole terraqueous globe so thickly, that four men +would have to stand on each square yard of surface. The primary or +fundamental check to the continued increase of man is the difficulty of +gaining subsistence and of living in comfort. We may infer that this is +the case from what we see, for instance, in the United States, where +subsistence is easy and there is plenty of room. If such means were +suddenly doubled in Great Britain, our number would be quickly doubled. +With civilised nations the above primary check acts chiefly by +restraining marriages. The greater death-rate of infants in the poorest +classes is also very important; as well as the greater mortality at all +ages, and from various diseases, of the inhabitants of crowded and +miserable houses. The effects of severe epidemics and wars are soon +counterbalanced, and more than counterbalanced, in nations placed under +favourable conditions. Emigration also comes in aid as a temporary +check, but not to any great extent with the extremely poor classes. + +There is reason to suspect, as Malthus has remarked, that the +reproductive power is actually less in barbarous than in civilised +races. We know nothing positively on this head, for with savages no +census has been taken; but from the concurrent testimony of +missionaries, and of others who have long resided with such people, it +appears that their families are usually small, and large ones rare. This +may be partly accounted for, as it is believed, by the women suckling +their infants for a prolonged period; but it is highly probable that +savages, who often suffer much hardship, and who do not obtain so much +nutritious food as civilised men, would be actually less prolific. I +have shewn in a former work,[191] that all our domesticated quadrupeds +and birds, and all our cultivated plants, are more fertile than the +corresponding species in a state of nature. It is no valid objection to +this conclusion that animals suddenly supplied with an excess of food, +or when rendered very fat, and that most plants when suddenly removed +from very poor to very rich soil, are rendered more or less sterile. We +might, therefore, expect that civilised men, who in one sense are highly +domesticated, would be more prolific than wild men. It is also probable +that the increased fertility of civilised nations would become, as with +our domestic animals, an inherited character: it is at least known that +with mankind a tendency to produce twins runs in families.[192] + +Notwithstanding that savages appear to be less prolific than civilised +people, they would no doubt rapidly increase if their numbers were not +by some means rigidly kept down. The Santali, or hill-tribes of India, +have recently afforded a good illustration of this fact; for they have +increased, as shewn by Mr. Hunter,[193] at an extraordinary rate since +vaccination has been introduced, other pestilences mitigated, and war +sternly repressed. This increase, however, would not have been possible +had not these rude people spread into the adjoining districts and worked +for hire. Savages almost always marry; yet there is some prudential +restraint, for they do not commonly marry at the earliest possible age. +The young men are often required to show that they can support a wife, +and they generally have first to earn the price with which to purchase +her from her parents. With savages the difficulty of obtaining +subsistence occasionally limits their number in a much more direct +manner than with civilised people, for all tribes periodically suffer +from severe famines. At such times savages are forced to devour much bad +food, and their health can hardly fail to be injured. Many accounts have +been published of their protruding stomachs and emaciated limbs after +and during famines. They are then, also, compelled to wander much about, +and their infants, as I was assured in Australia, perish in large +numbers. As famines are periodical, depending chiefly on extreme +seasons, all tribes must fluctuate in number. They cannot steadily and +regularly increase, as there is no artificial increase in the supply of +food. Savages when hardly pressed encroach on each other’s territories, +and war is the result; but they are indeed almost always at war with +their neighbours. They are liable to many accidents on land and water in +their search for food; and in some countries they must suffer much from +the larger beasts of prey. Even in India, districts have been +depopulated by the ravages of tigers. + +Malthus has discussed these several checks, but he does not lay stress +enough on what is probably the most important of all, namely +infanticide, especially of female infants, and the habit of procuring +abortion. These practices now prevail in many quarters of the world, and +infanticide seems formerly to have prevailed, as Mr. M’Lennan[194] has +shewn, on a still more extensive scale. These practices appear to have +originated in savages recognising the difficulty, or rather the +impossibility of supporting all the infants that are born. +Licentiousness may also be added to the foregoing checks; but this does +not follow from failing means of subsistence; though there is reason to +believe that in some cases (as in Japan) it has been intentionally +encouraged as a means of keeping down the population. + +If we look back to an extremely remote epoch, before man had arrived at +the dignity of manhood, he would have been guided more by instinct and +less by reason than are savages at the present time. Our early +semi-human progenitors would not have practised infanticide, for the +instincts of the lower animals are never so perverted as to lead them +regularly to destroy their own offspring. There would have been no +prudential restraint from marriage, and the sexes would have freely +united at an early age. Hence the progenitors of man would have tended +to increase rapidly, but checks of some kind, either periodical or +constant, must have kept down their numbers, even more severely than +with existing savages. What the precise nature of these checks may have +been, we cannot say, any more than with most other animals. We know that +horses and cattle, which are not highly prolific animals, when first +turned loose in South America, increased at an enormous rate. The +slowest breeder of all known animals, namely the elephant, would in a +few thousand years stock the whole world. The increase of every species +of monkey must be checked by some means; but not, as Brehm remarks, by +the attacks of beasts of prey. No one will assume that the actual power +of reproduction in the wild horses and cattle of America, was at first +in any sensible degree increased; or that, as each district became fully +stocked, this same power was diminished. No doubt in this case and in +all others, many checks concur, and different checks under different +circumstances; periodical dearths, depending on unfavourable seasons, +being probably the most important of all. So it will have been with the +early progenitors of man. + + +_Natural Selection._—We have now seen that man is variable in body and +mind; and that the variations are induced, either directly or +indirectly, by the same general causes, and obey the same general laws, +as with the lower animals. Man has spread widely over the face of the +earth, and must have been exposed, during his incessant migrations,[195] +to the most diversified conditions. The inhabitants of Tierra del +Fuego, the Cape of Good Hope, and Tasmania in the one hemisphere, and of +the Arctic regions in the other, must have passed through many climates +and changed their habits many times, before they reached their present +homes.[196] The early progenitors of man must also have tended, like all +other animals, to have increased beyond their means of subsistence; they +must therefore occasionally have been exposed to a struggle for +existence, and consequently to the rigid law of natural selection. +Beneficial variations of all kinds will thus, either occasionally or +habitually, have been preserved, and injurious ones eliminated. I do not +refer to strongly-marked deviations of structure, which occur only at +long intervals of time, but to mere individual differences. We know, for +instance, that the muscles of our hands and feet, which determine our +powers of movement, are liable, like those of the lower animals,[197] to +incessant variability. If then the ape-like progenitors of man which +inhabited any district, especially one undergoing some change in its +conditions, were divided into two equal bodies, the one half which +included all the individuals best adapted by their powers of movement +for gaining subsistence or for defending themselves, would on an average +survive in greater number and procreate more offspring than the other +and less well endowed half. + +Man in the rudest state in which he now exists is the most dominant +animal that has ever appeared on the earth. He has spread more widely +than any other highly organised form; and all others have yielded +before him. He manifestly owes this immense superiority to his +intellectual faculties, his social habits, which lead him to aid and +defend his fellows, and to his corporeal structure. The supreme +importance of these characters has been proved by the final arbitrament +of the battle for life. Through his powers of intellect, articulate +language has been evolved; and on this his wonderful advancement has +mainly depended. He has invented and is able to use various weapons, +tools, traps, &c., with which he defends himself, kills or catches prey, +and otherwise obtains food. He has made rafts or canoes on which to fish +or cross over to neighbouring fertile islands. He has discovered the art +of making fire, by which hard and stringy roots can be rendered +digestible, and poisonous roots or herbs innocuous. This last discovery, +probably the greatest, excepting language, ever made by man, dates from +before the dawn of history. These several inventions, by which man in +the rudest state has become so preeminent, are the direct result of the +development of his powers of observation, memory, curiosity, +imagination, and reason. I cannot, therefore, understand how it is that +Mr. Wallace[198] maintains, that “natural selection could only have +endowed the savage with a brain a little superior to that of an ape.” + +Although the intellectual powers and social habits of man are of +paramount importance to him, we must not underrate the importance of his +bodily structure, to which subject the remainder of this chapter will be +devoted. The development of the intellectual and social or moral +faculties will be discussed in the following chapter. + +Even to hammer with precision is no easy matter, as every one who has +tried to learn carpentry will admit. To throw a stone with as true an +aim as can a Fuegian in defending himself, or in killing birds, requires +the most consummate perfection in the correlated action of the muscles +of the hand, arm, and shoulder, not to mention a fine sense of touch. In +throwing a stone or spear, and in many other actions, a man must stand +firmly on his feet; and this again demands the perfect coadaptation of +numerous muscles. To chip a flint into the rudest tool, or to form a +barbed spear or hook from a bone, demands the use of a perfect hand; +for, as a most capable judge, Mr. Schoolcraft,[199] remarks, the shaping +fragments of stone into knives, lances, or arrow-heads, shews +“extraordinary ability and long practice.” We have evidence of this in +primeval men having practised a division of labour; each man did not +manufacture his own flint tools or rude pottery; but certain individuals +appear to have devoted themselves to such work, no doubt receiving in +exchange the produce of the chase. Archæologists are convinced that an +enormous interval of time elapsed before our ancestors thought of +grinding chipped flints into smooth tools. A man-like animal who +possessed a hand and arm sufficiently perfect to throw a stone with +precision or to form a flint into a rude tool, could, it can hardly be +doubted, with sufficient practice make almost anything, as far as +mechanical skill alone is concerned, which a civilised man can make. The +structure of the hand in this respect may be compared with that of the +vocal organs, which in the apes are used for uttering various +signal-cries, or, as in one species, musical cadences; but in man +closely similar vocal organs have become adapted through the inherited +effects of use for the utterance of articulate language. + +Turning now to the nearest allies of man, and therefore to the best +representatives of our early progenitors, we find that the hands in the +Quadrumana are constructed on the same general pattern as in us, but are +far less perfectly adapted for diversified uses. Their hands do not +serve so well as the feet of a dog for locomotion; as may be seen in +those monkeys which walk on the outer margins of the palms, or on the +backs of their bent fingers, as in the chimpanzee and orang.[200] Their +hands, however, are admirably adapted for climbing trees. Monkeys seize +thin branches or ropes, with the thumb on one side and the fingers and +palm on the other side, in the same manner as we do. They can thus also +carry rather large objects, such as the neck of a bottle, to their +mouths. Baboons turn over stones and scratch up roots with their hands. +They seize nuts, insects, or other small objects with the thumb in +opposition to the fingers, and no doubt they thus extract eggs and the +young from the nests of birds. American monkeys beat the wild oranges on +the branches until the rind is cracked, and then tear it off with the +fingers of the two hands. Other monkeys open mussel-shells with the two +thumbs. With their fingers they pull out thorns and burrs, and hunt for +each other’s parasites. In a state of nature they break open hard fruits +with the aid of stones. They roll down stones or throw them at their +enemies; nevertheless, they perform these various actions clumsily, and +they are quite unable, as I have myself seen, to throw a stone with +precision. + +It seems to me far from true that because “objects are grasped clumsily” +by monkeys, “a much less specialised organ of prehension” would have +served them[201] as well as their present hands. On the contrary, I see +no reason to doubt that a more perfectly constructed hand would have +been an advantage to them, provided, and it is important to note this, +that their hands had not thus been rendered less well adapted for +climbing trees. We may suspect that a perfect hand would have been +disadvantageous for climbing; as the most arboreal monkeys in the world, +namely Ateles in America and Hylobates in Asia, either have their thumbs +much reduced in size and even rudimentary, or their fingers +partially coherent, so that their hands are converted into mere +grasping-hooks.[202] + +As soon as some ancient member in the great series of the Primates came, +owing to a change in its manner of procuring subsistence, or to a change +in the conditions of its native country, to live somewhat less on trees +and more on the ground, its manner of progression would have been +modified; and in this case it would have had to become either more +strictly quadrupedal or bipedal. Baboons frequent hilly and rocky +districts, and only from necessity climb up high trees;[203] and they +have acquired almost the gait of a dog. Man alone has become a biped; +and we can, I think, partly see how he has come to assume his erect +attitude, which forms one of the most conspicuous differences between +him and his nearest allies. Man could not have attained his present +dominant position in the world without the use of his hands which are so +admirably adapted to act in obedience to his will. As Sir C. Bell[204] +insists “the hand supplies all instruments, and by its correspondence +with the intellect gives him universal dominion.” But the hands and arms +could hardly have become perfect enough to have manufactured weapons, or +to have hurled stones and spears with a true aim, as long as they were +habitually used for locomotion and for supporting the whole weight of +the body, or as long as they were especially well adapted, as previously +remarked, for climbing trees. Such rough treatment would also have +blunted the sense of touch, on which their delicate use largely depends. +From these causes alone it would have been an advantage to man to have +become a biped; but for many actions it is almost necessary that both +arms and the whole upper part of the body should be free; and he must +for this end stand firmly on his feet. To gain this great advantage, the +feet have been rendered flat, and the great toe peculiarly modified, +though this has entailed the loss of the power of prehension. It accords +with the principle of the division of physiological labour, which +prevails throughout the animal kingdom, that as the hands became +perfected for prehension, the feet should have become perfected for +support and locomotion. With some savages, however, the foot has not +altogether lost its prehensile power, as shewn by their manner of +climbing trees and of using them in other ways.[205] + +If it be an advantage to man to have his hands and arms free and to +stand firmly on his feet, of which there can be no doubt from his +preeminent success in the battle of life, then I can see no reason why +it should not have been advantageous to the progenitors of man to have +become more and more erect or bipedal. They would thus have been better +able to have defended themselves with stones or clubs, or to have +attacked their prey, or otherwise obtained food. The best constructed +individuals would in the long run have succeeded best, and have survived +in larger numbers. If the gorilla and a few allied forms had become +extinct, it might have been argued with great force and apparent truth, +that an animal could not have been gradually converted from a quadruped +into a biped; as all the individuals in an intermediate condition would +have been miserably ill-fitted for progression. But we know (and this is +well worthy of reflection) that several kinds of apes are now actually +in this intermediate condition; and no one doubts that they are on the +whole well adapted for their conditions of life. Thus the gorilla runs +with a sidelong shambling gait, but more commonly progresses by resting +on its bent hands. The long-armed apes occasionally use their arms like +crutches, swinging their bodies forward between them, and some kinds of +Hylobates, without having been taught, can walk or run upright with +tolerable quickness; yet they move awkwardly, and much less securely +than man. We see, in short, with existing monkeys various gradations +between a form of progression strictly like that of a quadruped and that +of a biped or man. + +As the progenitors of man became more and more erect, with their hands +and arms more and more modified for prehension and other purposes, with +their feet and legs at the same time modified for firm support and +progression, endless other changes of structure would have been +necessary. The pelvis would have had to be made broader, the spine +peculiarly curved and the head fixed in an altered position, and all +these changes have been attained by man. Prof. Schaaffhausen[206] +maintains that “the powerful mastoid processes of the human skull are +the result of his erect position;” and these processes are absent in the +orang, chimpanzee, &c., and are smaller in the gorilla than in man. +Various other structures might here have been specified, which appear +connected with man’s erect position. It is very difficult to decide how +far all these correlated modifications are the result of natural +selection, and how far of the inherited effects of the increased use of +certain parts, or of the action of one part on another. No doubt these +means of change act and react on each other: thus when certain muscles, +and the crests of bone to which they are attached, become enlarged by +habitual use, this shews that certain fictions are habitually performed +and must be serviceable. Hence the individuals which performed them +best, would tend to survive in greater numbers. + +The free use of the arms and hands, partly the cause and partly the +result of man’s erect position, appears to have led in an indirect +manner to other modifications of structure. The early male progenitors +of man were, as previously stated, probably furnished with great canine +teeth; but as they gradually acquired the habit of using stones, clubs, +or other weapons, for fighting with their enemies, they would have used +their jaws and teeth less and less. In this case, the jaws, together +with the teeth, would have become reduced in size, as we may feel sure +from innumerable analogous cases. In a future chapter we shall meet with +a closely-parallel case, in the reduction or complete disappearance of +the canine teeth in male ruminants, apparently in relation with the +development of their horns; and in horses, in relation with their habit +of fighting with their incisor teeth and hoofs. + +In the adult male anthropomorphous apes, as Rütimeyer,[207] and others +have insisted, it is precisely the effect which the jaw-muscles by their +great development have produced on the skull, that causes it to differ +so greatly in many respects from that of man, and has given to it “a +truly frightful physiognomy.” Therefore as the jaws and teeth in the +progenitors of man gradually become reduced in size, the adult skull +would have presented nearly the same characters which it offers in the +young of the anthropomorphous apes, and would thus have come to resemble +more nearly that of existing man. A great reduction of the canine teeth +in the males would almost certainly, as we shall hereafter see, have +affected through inheritance the teeth of the females. + +As the various mental faculties were gradually developed, the brain +would almost certainly have become larger. No one, I presume, doubts +that the large size of the brain in man, relatively to his body, in +comparison with that of the gorilla or orang, is closely connected with +his higher mental powers. We meet with closely analogous facts with +insects, in which the cerebral ganglia are of extraordinary dimensions +in ants; these ganglia in all the Hymenoptera being many times larger +than in the less intelligent orders, such as beetles.[208] On the other +hand, no one supposes that the intellect of any two animals or of any +two men can be accurately gauged by the cubic contents of their skulls. +It is certain that there may be extraordinary mental activity with an +extremely small absolute mass of nervous matter: thus the wonderfully +diversified instincts, mental powers, and affections of ants are +generally known, yet their cerebral ganglia are not so large as the +quarter of a small pin’s head. Under this latter point of view, the +brain of an ant is one of the most marvellous atoms of matter in the +world, perhaps more marvellous than the brain of man. + +The belief that there exists in man some close relation between the size +of the brain and the development of the intellectual faculties is +supported by the comparison of the skulls of savage and civilised races, +of ancient and modern people, and by the analogy of the whole +vertebrate series. Dr. J. Barnard Davis has proved[209] by many careful +measurements, that the mean internal capacity of the skull in Europeans +is 92·3 cubic inches; in Americans 87·5; in Asiatics 87·1; and in +Australians only 81·9 inches. Professor Broca[210] found that skulls +from graves in Paris of the nineteenth century, were larger than those +from vaults of the twelfth century, in the proportion of 1484 to 1426; +and Prichard is persuaded that the present inhabitants of Britain have +“much more capacious brain-cases” than the ancient inhabitants. +Nevertheless it must be admitted that some skulls of very high +antiquity, such as the famous one of Neanderthal, are well developed and +capacious. With respect to the lower animals, M. E. Lartet,[211] by +comparing the crania of tertiary and recent mammals, belonging to the +same groups, has come to the remarkable conclusion that the brain is +generally larger and the convolutions more complex in the more recent +form. On the other hand I have shewn[212] that the brains of domestic +rabbits are considerably reduced in bulk, in comparison with those of +the wild rabbit or hare; and this may be attributed to their having been +closely confined during many generations, so that they have exerted but +little their intellect, instincts, senses, and voluntary movements. + +The gradually increasing weight of the brain and skull in man must have +influenced the development of the supporting spinal column, more +especially whilst he was becoming erect. As this change of position was +being brought about, the internal pressure of the brain, will, also, +have influenced the form of the skull; for many facts shew how easily +the skull is thus affected. Ethnologists believe that it is modified by +the kind of cradle in which infants sleep. Habitual spasms of the +muscles and a cicatrix from a severe burn have permanently modified the +facial bones. In young persons whose heads from disease have become +fixed either sideways or backwards, one of the eyes has changed its +position, and the bones of the skull have been modified; and this +apparently results from the brain pressing in a new direction.[213] I +have shewn that with long-eared rabbits, even so trifling a cause as the +lopping forward of one ear drags forward on that side almost every bone +of the skull; so that the bones on the opposite sides no longer strictly +correspond. Lastly, if any animal were to increase or diminish much in +general size, without any change in its mental powers; or if the mental +powers were to be much increased or diminished without any great change +in the size of the body; the shape of the skull would almost certainly +be altered. I infer this from my observations on domestic rabbits, some +kinds of which have become very much larger than the wild animal, whilst +others have retained nearly the same size, but in both cases the brain +has been much reduced relatively to the size of the body. Now I was at +first much surprised by finding that in all these rabbits the skull had +become elongated or dolichocephalic; for instance, of two skulls of +nearly equal breadth, the one from a wild rabbit and the other from a +large domestic kind, the former was only 3·15 and the latter 4·3 inches +in length.[214] One of the most marked distinctions in different races +of man is that the skull in some is elongated, and in others rounded; +and here the explanation suggested by the case of the rabbits may +partially hold good; for Welcker finds that short “men incline more to +brachycephaly, and tall men to dolichocephaly;”[215] and tall men may be +compared with the larger and longer-bodied rabbits, all of which have +elongated skulls, or are dolichocephalic. + +From these several facts we can to a certain extent understand the means +through which the great size and more or less rounded form of the skull +has been acquired by man; and these are characters eminently distinctive +of him in comparison with the lower animals. + +Another most conspicuous difference between man and the lower animals is +the nakedness of his skin. Whales and dolphins (Cetacea), dugongs +(Sirenia) and the hippopotamus are naked; and this may be advantageous +to them for gliding through the water; nor would it be injurious to them +from the loss of warmth, as the species which inhabit the colder regions +are protected by a thick layer of blubber, serving the same purpose as +the fur of seals and otters. Elephants and rhinoceroses are almost +hairless; and as certain extinct species which formerly lived under an +arctic climate were covered with long wool or hair, it would almost +appear as if the existing species of both genera had lost their hairy +covering from exposure to heat. This appears the more probable, as the +elephants in India which live on elevated and cool districts are more +hairy[216] than those on the lowlands. May we then infer that man became +divested of hair from having aboriginally inhabited some tropical land? +The fact of the hair being chiefly retained in the male sex on the chest +and face, and in both sexes at the junction of all four limbs with the +trunk, favours this inference, assuming that the hair was lost before +man became erect; for the parts which now retain most hair would then +have been most protected from the heat of the sun. The crown of the +head, however, offers a curious exception, for at all times it must have +been one of the most exposed parts, yet it is thickly clothed with hair. +In this respect man agrees with the great majority of quadrupeds, which +generally have their upper and exposed surfaces more thickly clothed +than the lower surface. Nevertheless, the fact that the other members of +the order of Primates, to which man belongs, although inhabiting various +hot regions, are well clothed with hair, generally thickest on the upper +surface,[217] is strongly opposed to the supposition that man became +naked through the action of the sun. I am inclined to believe, as we +shall see under sexual selection, that man, or rather primarily woman, +became divested of hair for ornamental purposes; and according to this +belief it is not surprising that man should differ so greatly in +hairiness from all his lower brethren, for characters gained through +sexual selection often differ in closely-related forms to an +extraordinary degree. + +According to a popular impression, the absence of a tail is eminently +distinctive of man; but as those apes which come nearest to man are +destitute of this organ, its disappearance does not especially concern +us. Nevertheless it may be well to own that no explanation, as far as I +am aware, has ever been given of the loss of the tail by certain apes +and man. Its loss, however, is not surprising, for it sometimes differs +remarkably in length in species of the same genera: thus in some species +of Macacus the tail is longer than the whole body, consisting of +twenty-four vertebræ; in others it consists of a scarcely visible stump, +containing only three or four vertebræ. In some kinds of baboons there +are twenty-five, whilst in the mandrill there are ten very small stunted +caudal vertebræ, or, according to Cuvier,[218] sometimes only five. This +great diversity in the structure and length of the tail in animals +belonging to the same genera, and following nearly the same habits of +life, renders it probable that the tail is not of much importance to +them; and if so, we might have expected that it would sometimes have +become more or less rudimentary, in accordance with what we incessantly +see with other structures. The tail almost always tapers towards the end +whether it be long or short; and this, I presume, results from the +atrophy, through disuse, of the terminal muscles together with their +arteries and nerves, leading to the atrophy of the terminal bones. With +respect to the os coccyx, which in man and the higher apes manifestly +consists of the few basal and tapering segments of an ordinary tail, I +have heard it asked how could these have become completely embedded +within the body; but there is no difficulty in this respect, for in many +monkeys the basal segments of the true tail are thus embedded. For +instance, Mr. Murie informs me that in the skeleton of a not full-grown +_Macacus inornatus_, he counted nine or ten caudal vertebræ, which +altogether were only 1·8 inch in length. Of these the three basal ones +appeared to have been embedded; the remainder forming the free part of +the tail, which was only one inch in length, and half an inch in +diameter. Here, then, the three embedded caudal vertebræ plainly +correspond with the four coalesced vertebræ of the human os coccyx. + + +I have now endeavoured to shew that some of the most distinctive +characters of man have in all probability been acquired, either +directly, or more commonly indirectly, through natural selection. We +should bear in mind that modifications in structure or constitution, +which are of no service to an organism in adapting it to its habits of +life, to the food which it consumes, or passively to the surrounding +conditions, cannot have been thus acquired. We must not, however, be too +confident in deciding what modifications are of service to each being: +we should remember how little we know about the use of many parts, or +what changes in the blood or tissues may serve to fit an organism for a +new climate or some new kind of food. Nor must we forget the principle +of correlation, by which, as Isidore Geoffroy has shewn in the case of +man, many fit-range deviations of structure are tied together. +Independently of correlation, a change in one part often leads through +the increased or decreased use of other parts, to other changes of a +quite unexpected nature. It is also well to reflect on such facts, as +the wonderful growth of galls on plants caused by the poison of an +insect, and on the remarkable changes of colour in the plumage of +parrots when fed on certain fishes, or inoculated with the poison of +toads;[219] for we can thus see that the fluids of the system, if +altered for some special purpose, might induce other strange changes. We +should especially bear in mind that modifications acquired and +continually used during past ages for some useful purpose would probably +become firmly fixed and might be long inherited. + +Thus a very large yet undefined extension may safely be given to the +direct and indirect results of natural selection; but I now admit, after +reading the essay by Nägeli on plants, and the remarks by various +authors with respect to animals, more especially those recently made by +Professor Broca, that in the earlier editions of my ‘Origin of Species’ +I probably attributed too much to the action of natural selection or the +survival of the fittest. I have altered the fifth edition of the Origin +so as to confine my remarks to adaptive changes of structure. I had not +formerly sufficiently considered the existence of many structures which +appear to be, as far as we can judge, neither beneficial nor injurious; +and this I believe to be one of the greatest oversights as yet detected +in my work. I may be permitted to say as some excuse, that I had two +distinct objects in view, firstly, to shew that species had not been +separately created, and secondly, that natural selection had been the +chief agent of change, though largely aided by the inherited effects of +habit, and slightly by the direct action of the surrounding conditions. +Nevertheless I was not able to annul the influence of my former belief, +then widely prevalent, that each species had been purposely created; and +this led to my tacitly assuming that every detail of structure, +excepting rudiments, was of some special, though unrecognised, service. +Any one with this assumption in his mind would naturally extend the +action of natural selection, either during past or present times, too +far. Some of those who admit the principle of evolution, but reject +natural selection, seem to forget, when criticising my book, that I had +the above two objects in view; hence if I have erred in giving to +natural selection great power, which I am far from admitting, or in +having exaggerated its power, which is in itself probable, I have at +least, as I hope, done good service in aiding to overthrow the dogma of +separate creations. + +That all organic beings, including man, present many modifications of +structure which are of no service to them at present, nor have been +formerly, is, as I can now see, probable. We know not what produces the +numberless slight differences between the individuals of each species, +for reversion only carries the problem a few steps backwards; but each +peculiarity must have had its own efficient cause. If these causes, +whatever they may be, were to act more uniformly and energetically +during a lengthened period (and no reason can be assigned why this +should not sometimes occur), the result would probably be not mere +slight individual differences, but well-marked, constant modifications. +Modifications which are in no way beneficial cannot have been kept +uniform through natural selection, though any which were injurious would +have been thus eliminated. Uniformity of character would, however, +naturally follow from, the assumed uniformity of the exciting causes, +and likewise from the free intercrossing of many individuals. The same +organism might acquire in this manner during successive periods +successive modifications, and these would be transmitted in a nearly +uniform state as long as the exciting causes remained the same and there +was free intercrossing. With respect to the exciting causes we can only +say, as when speaking of so-called spontaneous variations, that they +relate much more closely to the constitution of the varying organism, +than to the nature of the conditions to which it has been subjected. + + +_Conclusion._—In this chapter we have seen that as man at the present +day is liable, like every other animal, to multiform individual +differences or slight variations, so no doubt were the early progenitors +of man; the variations being then as now induced by the same general +causes, and governed by the same general and complex laws. As all +animals tend to multiply beyond their means of subsistence, so it must +have been with the progenitors of man; and this will inevitably have led +to a struggle for existence and to natural selection. This latter +process will have been greatly aided by the inherited effects of the +increased use of parts; these two processes incessantly reacting on each +other. It appears, also, as we shall hereafter see, that various +unimportant characters have been acquired by man through sexual +selection. An unexplained residuum of change, perhaps a large one, must +be left to the assumed uniform action of those unknown agencies, which +occasionally induce strongly-marked and abrupt deviations of structure +in our domestic productions. + +Judging from the habits of savages and of the greater number of the +Quadrumana, primeval men, and even the ape-like progenitors of man, +probably lived in society. With strictly social animals, natural +selection sometimes acts indirectly on the individual, through the +preservation of variations which are beneficial only to the community. A +community including a large number of well-endowed individuals increases +in number and is victorious over other and less well-endowed +communities; although each separate member may gain no advantage over +the other members of the same community. With associated insects many +remarkable structures, which are of little or no service to the +individual or its own offspring, such as the pollen-collecting +apparatus, or the sting of the worker-bee, or the great jaws of +soldier-ants, have been thus acquired. With the higher social animals, I +am not aware that any structure has been modified solely for the good of +the community, though some are of secondary service to it. For instance, +the horns of ruminants and the great canine teeth of baboons appear to +have been acquired by the males as weapons for sexual strife, but they +are used in defence of the herd or troop. In regard to certain mental +faculties the case, as we shall see in the following chapter, is wholly +different; for these faculties have been chiefly, or even exclusively, +gained for the benefit of the community; the individuals composing the +community being at the same time indirectly benefited. + +It has often been objected to such views as the foregoing, that man is +one of the most helpless and defenceless creatures in the world; and +that during his early and less well-developed condition he would have +been still more helpless. The Duke of Argyll, for instance, insists[220] +that “the human frame has diverged from the structure of brutes, in the +direction of greater physical helplessness and weakness. That is to say, +it is a divergence which of all others it is most impossible to ascribe +to mere natural selection.” He adduces the naked and unprotected state +of the body, the absence of great teeth or claws for defence, the little +strength of man, his small speed in running, and his slight power of +smell, by which to discover food or to avoid danger. To these +deficiencies there might have been added the still more serious loss of +the power of quickly climbing trees, so as to escape from enemies. +Seeing that the unclothed Fuegians can exist under their wretched +climate, the loss of hair would not have been a great injury to primeval +man, if he inhabited a warm country. When we compare defenceless man +with the apes, many of which are provided with formidable canine teeth, +we must remember that these in their fully-developed condition are +possessed by the males alone, being chiefly used by them for fighting +with their rivals; yet the females which are not thus provided, are able +to survive. + +In regard to bodily size or strength, we do not know whether man is +descended from some comparatively small species, like the chimpanzee, or +from one as powerful as the gorilla; and, therefore, we cannot say +whether man has become larger and stronger, or smaller and weaker, in +comparison with his progenitors. We should, however, bear in mind that +an animal possessing great size, strength, and ferocity, and which, like +the gorilla, could defend itself from all enemies, would probably, +though not necessarily, have failed to become social; and this would +most effectually have checked the acquirement by man of his higher +mental qualities, such as sympathy and the love of his fellow-creatures. +Hence it might have been an immense advantage to man to have sprung +from some comparatively weak creature. + +The slight corporeal strength of man, his little speed, his want of +natural weapons, &c., are more than counterbalanced, firstly by his +intellectual powers, through which he has, whilst still remaining in a +barbarous state, formed for himself weapons, tools, &c., and secondly by +his social qualities which lead him to give aid to his fellow-men and to +receive it in return. No country in the world abounds in a greater +degree with dangerous beasts than Southern Africa; no country presents +more fearful physical hardships than the Arctic regions; yet one of the +puniest races, namely, the Bushmen, maintain themselves in Southern +Africa, as do the dwarfed Esquimaux in the Arctic regions. The early +progenitors of man were, no doubt, inferior in intellect, and probably +in social disposition, to the lowest existing savages; but it is quite +conceivable that they might have existed, or even flourished, if, whilst +they gradually lost their brute-like powers, such as climbing trees, +&c., they at the same time advanced in intellect. But granting that the +progenitors of man were far more helpless and defenceless than any +existing savages, if they had inhabited some warm continent or large +island, such as Australia or New Guinea, or Borneo (the latter island +being now tenanted by the orang), they would not have been exposed to +any special danger. In an area as large as one of these islands, the +competition between tribe and tribe would have been sufficient, under +favourable conditions, to have raised man, through the survival of the +fittest, combined with the inherited effects of habit, to his present +high position in the organic scale. + + + + +CHAPTER V. + +ON THE DEVELOPMENT OF THE INTELLECTUAL AND MORAL FACULTIES DURING +PRIMEVAL AND CIVILISED TIMES. + + + The advancement of the intellectual powers through natural + selection—Importance of imitation—Social and moral + faculties—Their development within the limits of the same + tribe—Natural selection as affecting civilised + nations—Evidence that civilised nations were once barbarous. + + +The subjects to be discussed in this chapter are of the highest +interest, but are treated by me in a most imperfect and fragmentary +manner. Mr. Wallace, in an admirable paper before referred to,[221] +argues that man after he had partially acquired those intellectual and +moral faculties which distinguish him from the lower animals, would have +been but little liable to have had his bodily structure modified through +natural selection or any other means. For man is enabled through his +mental faculties “to keep with an unchanged body in harmony with the +changing universe.” He has great power of adapting his habits to new +conditions of life. He invents weapons, tools and various stratagems, by +which he procures food and defends himself. When he migrates into a +colder climate he uses clothes, builds sheds, and makes fires; and, by +the aid of fire, cooks food otherwise indigestible. He aids his +fellow-men in many ways, and anticipates future events. Even at a remote +period he practised some subdivision of labour. + +The lower animals, on the other hand, must have their bodily structure +modified in order to survive under greatly changed conditions. They must +be rendered stronger, or acquire more effective teeth or claws, in order +to defend themselves from new enemies; or they must be reduced in size +so as to escape detection and danger. When they migrate into a colder +climate they must become clothed with thicker fur, or have their +constitutions altered. If they fail to be thus modified, they will cease +to exist. + +The case, however, is widely different, as Mr. Wallace has with justice +insisted, in relation to the intellectual and moral faculties of man. +These faculties are variable; and we have every reason to believe that +the variations tend to be inherited. Therefore, if they were formerly of +high importance to primeval man and to his ape-like progenitors, they +would have been perfected or advanced through natural selection. Of the +high importance of the intellectual faculties there can be no doubt, for +man mainly owes to them his preeminent position in the world. We can +see that, in the rudest state of society, the individuals who were the +most sagacious, who invented and used the best weapons or traps, and who +were best able to defend themselves, would rear the greatest number of +offspring. The tribes which included the largest number of men thus +endowed would increase in number and supplant other tribes. Numbers +depend primarily on the means of subsistence, and this, partly on the +physical nature of the country, but in a much higher degree on the arts +which are there practised. As a tribe increases and is victorious, it is +often still further increased by the absorption of other tribes.[222] +The stature and strength of the men of a tribe are likewise of some +importance for its success, and these depend in part on the nature and +amount of the food which can be obtained. In Europe the men of the +Bronze period were supplanted by a more powerful and, judging from their +sword-handles, larger-handed race;[223] but their success was probably +due in a much higher degree to their superiority in the arts. + +All that we know about savages, or may infer from their traditions and +from old monuments, the history of which is quite forgotten by the +present inhabitants, shew that from the remotest times successful tribes +have supplanted other tribes. Relics of extinct or forgotten tribes have +been discovered throughout the civilised regions of the earth, on the +wild plains of America, and on the isolated islands in the Pacific +Ocean. At the present day civilised nations are everywhere supplanting +barbarous nations, excepting where the climate opposes a deadly barrier; +and they succeed mainly, though not exclusively, through their arts, +which are the products of the intellect. It is, therefore, highly +probable that with mankind the intellectual faculties have been +gradually perfected through natural selection; and this conclusion is +sufficient for our purpose. Undoubtedly it would have been very +interesting to have traced the development of each separate faculty from +the state in which it exists in the lower animals to that in which it +exists in man; but neither my ability nor knowledge permit the attempt. + +It deserves notice that as soon as the progenitors of man became social +(and this probably occurred at a very early period), the advancement of +the intellectual faculties will have been aided and modified in an +important manner, of which we see only traces in the lower animals, +namely, through the principle of imitation, together with reason and +experience. Apes are much given to imitation, as are the lowest savages; +and the simple fact previously referred to, that after a time no animal +can be caught in the same place by the same sort of trap, shews that +animals learn by experience, and imitate each others’ caution. Now, if +some one man in a tribe, more sagacious than the others, invented a new +snare or weapon, or other means of attack or defence, the plainest +self-interest, without the assistance of much reasoning power, would +prompt the other members to imitate him; and all would thus profit. The +habitual practice of each new art must likewise in some slight degree +strengthen the intellect. If the new invention were an important one, +the tribe would increase in number, spread, and supplant other tribes. +In a tribe thus rendered more numerous there would always be a rather +better chance of the birth of other superior and inventive members. If +such men left children to inherit their mental superiority, the chance +of the birth of still more ingenious members would be somewhat better, +and in a very small tribe decidedly better. Even if they left no +children, the tribe would still include their blood-relations; and it +has been ascertained by agriculturists[224] that by preserving and +breeding from the family of an animal, which when slaughtered was found +to be valuable, the desired character has been obtained. + + +Turning now to the social and moral faculties. In order that primeval +men, or the ape-like progenitors of man, should have become social, they +must have acquired the same instinctive feelings which impel other +animals to live in a body; and they no doubt exhibited the same general +disposition. They would have felt uneasy when separated from their +comrades, for whom they would have felt some degree of love; they would +have warned each other of danger, and have given mutual aid in attack or +defence. All this implies some degree of sympathy, fidelity, and +courage. Such social qualities, the paramount importance of which to the +lower animals is disputed by no one, were no doubt acquired by the +progenitors of man in a similar manner, namely, through natural +selection, aided by inherited habit. When two tribes of primeval man, +living in the same country, came into competition, if the one tribe +included (other circumstances being equal) a greater number of +courageous, sympathetic, and faithful members, who were always ready to +warn each other of danger, to aid and defend each other, this tribe +would without doubt succeed best and conquer the other. Let it be borne +in mind how all-important, in the never-ceasing wars of savages, +fidelity and courage must be. The advantage which disciplined soldiers +have over undisciplined hordes follows chiefly from the confidence which +each man feels in his comrades. Obedience, as Mr. Bagehot has well +shewn,[225] is of the highest value, for any form of government is +better than none. Selfish and contentious people will not cohere, and +without coherence nothing can be effected. A tribe possessing the above +qualities in a high degree would spread and be victorious over other +tribes; but in the course of time it would, judging from all past +history, be in its turn overcome by some other and still more highly +endowed tribe. Thus the social and moral qualities would tend slowly to +advance and be diffused throughout the world. + +But it may be asked, how within the limits of the same tribe did a large +number of members first become endowed with these social and moral +qualities, and how was the standard of excellence raised? It is +extremely doubtful whether the offspring of the more sympathetic and +benevolent parents, or of those which were the most faithful to their +comrades, would be reared in greater number than the children of selfish +and treacherous parents of the same tribe. He who was ready to sacrifice +his life, as many a savage has been, rather than betray his comrades, +would often leave no offspring to inherit his noble nature. The bravest +men, who were always willing to come to the front in war, and who freely +risked their lives for others, would on an average perish in larger +number than other men. Therefore it seems scarcely possible (bearing in +mind that we are not here speaking of one tribe being victorious over +another) that the number of men gifted with such virtues, or that the +standard of their excellence, could be increased through natural +selection, that is, by the survival of the fittest. + +Although the circumstances which lead to an increase in the number of +men thus endowed within the same tribe are too complex to be clearly +followed out, we can trace some of the probable steps. In the first +place, as the reasoning powers and foresight of the members became +improved, each man would soon learn from experience that if he aided his +fellow-men, he would commonly receive aid in return. From this low +motive he might acquire the habit of aiding his fellows; and the habit +of performing benevolent actions certainly strengthens the feeling of +sympathy, which gives the first impulse to benevolent actions. Habits, +moreover, followed during many generations probably tend to be +inherited. + +But there is another and much more powerful stimulus to the development +of the social virtues, namely, the praise and the blame of our +fellow-men. The love of approbation and the dread of infamy, as well as +the bestowal of praise or blame, are primarily due, as we have seen in +the third chapter, to the instinct of sympathy; and this instinct no +doubt was originally acquired, like all the other social instincts, +through natural selection. At how early a period the progenitors of man, +in the course of their development, became capable of feeling and being +impelled by the praise or blame of their fellow-creatures, we cannot, of +course, say. But it appears that even dogs appreciate encouragement, +praise, and blame. The rudest savages feel the sentiment of glory, as +they clearly show by preserving the trophies of their prowess, by their +habit of excessive boasting, and even by the extreme care which they +take of their personal appearance and decorations; for unless they +regarded the opinion of their comrades, such habits would be senseless. + +They certainly feel shame at the breach of some of their lesser rules; +but how far they experience remorse is doubtful. I was at first +surprised that I could not recollect any recorded instances of this +feeling in savages; and Sir J. Lubbock[226] states that he knows of +none. But if we banish from our minds all cases given in novels and +plays and in death-bed confessions made to priests, I doubt whether many +of us have actually witnessed remorse; though we may have often seen +shame and contrition for smaller offences. Remorse is a deeply hidden +feeling. It is incredible that a savage, who will sacrifice his life +rather than betray his tribe, or one who will deliver himself up as a +prisoner rather than break his parole,[227] would not feel remorse in +his inmost soul, though he might conceal it, if he had failed in a duty +which he held sacred. + +We may therefore conclude that primeval man, at a very remote period, +would have been influenced by the praise and blame of his fellows. It is +obvious, that the members of the same tribe would approve of conduct +which appeared to them to be for the general good, and would reprobate +that which appeared evil. To do good unto others—to do unto others as +ye would they should do unto you,—is the foundation-stone of morality. +It is, therefore, hardly possible to exaggerate the importance during +rude times of the love of praise and the dread of blame. A man who was +not impelled by any deep, instinctive feeling, to sacrifice his life for +the good of others, yet was roused to such actions by a sense of glory, +would by his example excite the same wish for glory in other men, and +would strengthen by exercise the noble feeling of admiration. He might +thus do far more good to his tribe than by begetting offspring with a +tendency to inherit his own high character. + +With increased experience and reason, man perceives the more remote +consequences of his actions, and the self-regarding virtues, such as +temperance, chastity, &c., which during early times are, as we have +before seen, utterly disregarded, come to be highly esteemed or even +held sacred. I need not, however, repeat what I have said on this head +in the third chapter. Ultimately a highly complex sentiment, having its +first origin in the social instincts, largely guided by the approbation +of our fellow-men, ruled by reason, self-interest, and in later times by +deep religious feelings, confirmed by instruction and habit, all +combined, constitute our moral sense or conscience. + +It must not be forgotten that although a high standard of morality gives +but a slight or no advantage to each individual man and his children +over the other men of the same tribe, yet that an advancement in the +standard of morality and an increase in the number of well-endowed men +will certainly give an immense advantage to one tribe over another. +There can be no doubt that a tribe including many members who, from +possessing in a high degree the spirit of patriotism, fidelity, +obedience, courage, and sympathy, were always ready to give aid to each +other and to sacrifice themselves for the common good, would be +victorious over most other tribes; and this would be natural selection. +At all times throughout the world tribes have supplanted other tribes; +and as morality is one element in their success, the standard of +morality and the number of well-endowed men will thus everywhere tend to +rise and increase. + +It is, however, very difficult to form any judgment why one particular +tribe and not another has been successful and has risen in the scale of +civilisation. Many savages are in the same condition as when first +discovered several centuries ago. As Mr. Bagehot has remarked, we are +apt to look at progress as the normal rule in human society; but history +refutes this. The ancients did not even entertain the idea; nor do the +oriental nations at the present day. According to another high +authority, Mr. Maine,[228] “the greatest part of mankind has never +shewn a particle of desire that its civil institutions should be +improved.” Progress seems to depend on many concurrent favourable +conditions, far too complex to be followed out. But it has often been +remarked, that a cool climate from leading to industry and the various +arts has been highly favourable, or even indispensable for this end. The +Esquimaux, pressed by hard necessity, have succeeded in many ingenious +inventions, but their climate has been too severe for continued +progress. Nomadic habits, whether over wide plains, or through the dense +forests of the tropics, or along the shores of the sea, have in every +case been highly detrimental. Whilst observing the barbarous inhabitants +of Tierra del Fuego, it struck me that the possession of some property, +a fixed abode, and the union of many families under a chief, were the +indispensable requisites for civilisation. Such habits almost +necessitate the cultivation of the ground; and the first steps in +cultivation would probably result, as I have elsewhere shewn,[229] from +some such accident as the seeds of a fruit-tree falling on a heap of +refuse and producing an unusually fine variety. The problem, however, of +the first advance of savages towards civilisation is at present much too +difficult to be solved. + + +_Natural Selection as affecting Civilised Nations._—In the last and +present chapters I have considered the advancement of man from a former +semi-human condition to his present state as a barbarian. But some +remarks on the agency of natural selection on civilised nations may be +here worth adding. This subject has been ably discussed by Mr. W. R. +Greg,[230] and previously by Mr. Wallace and Mr. Galton.[231] Most of +my remarks are taken from these three authors. With savages, the weak in +body or mind are soon eliminated; and those that survive commonly +exhibit a vigorous state of health. We civilised men, on the other hand, +do our utmost to check the process of elimination; we build asylums for +the imbecile, the maimed, and the sick; we institute poor-laws; and our +medical men exert their utmost skill to save the life of every one to +the last moment. There is reason to believe that vaccination has +preserved thousands, who from a weak constitution would formerly have +succumbed to small-pox. Thus the weak members of civilised societies +propagate their kind. No one who has attended to the breeding of +domestic animals will doubt that this must be highly injurious to the +race of man. It is surprising how soon a want of care, or care wrongly +directed, leads to the degeneration of a domestic race; but excepting in +the case of man himself, hardly any one is so ignorant as to allow his +worst animals to breed. + +The aid which we feel impelled to give to the helpless is mainly an +incidental result of the instinct of sympathy, which was originally +acquired as part of the social instincts, but subsequently rendered, in +the manner previously indicated, more tender and more widely diffused. +Nor could we check our sympathy, if so urged by hard reason, without +deterioration in the noblest part of our nature. The surgeon may harden +himself whilst performing an operation, for he knows that he is acting +for the good of his patient; but if we were intentionally to neglect the +weak and helpless, it could only be for a contingent benefit, with a +certain and great present evil. Hence we must bear without complaining +the undoubtedly bad effects of the weak surviving and propagating their +kind; but there appears to be at least one check in steady action, +namely the weaker and inferior members of society not marrying so freely +as the sound; and this check might be indefinitely increased, though +this is more to be hoped for than expected, by the weak in body or mind +refraining from marriage. + +In all civilised countries man accumulates property and bequeaths it to +his children. So that the children in the same country do not by any +means start fair in the race for success. But this is far from an +unmixed evil; for without the accumulation of capital the arts could not +progress; and it is chiefly through their power that the civilised races +have extended, and are now everywhere extending, their range, so as to +take the place of the lower races. Nor does the moderate accumulation of +wealth interfere with the process of selection. When a poor man becomes +rich, his children enter trades or professions in which there is +struggle enough, so that the able in body and mind succeed best. The +presence of a body of well-instructed men, who have not to labour for +their daily bread, is important to a degree which cannot be +over-estimated; as all high intellectual work is carried on by them, and +on such work material progress of all kinds mainly depends, not to +mention other and higher advantages. No doubt wealth when very great +tends to convert men into useless drones, but their number is never +large; and some degree of elimination here occurs, as we daily see rich +men, who happen to be fools or profligate, squandering away all their +wealth. + +Primogeniture with entailed estates is a more direct evil, though it may +formerly have been a great advantage by the creation of a dominant +class, and any government is better than anarchy. The eldest sons, +though they may be weak in body or mind, generally marry, whilst the +younger sons, however superior in these respects, do not so generally +marry. Nor can worthless eldest sons with entailed estates squander +their wealth. But here, as elsewhere, the relations of civilised life +are so complex that some compensatory checks intervene. The men who are +rich through primogeniture are able to select generation after +generation the more beautiful and charming women; and these must +generally be healthy in body and active in mind. The evil consequences, +such as they may be, of the continued preservation of the same line of +descent, without any selection, are checked by men of rank always +wishing to increase their wealth and power; and this they effect by +marrying heiresses. But the daughters of parents who have produced +single children, are themselves, as Mr. Galton has shewn,[232] apt to be +sterile; and thus noble families are continually cut off in the direct +line, and their wealth flows into some side channel; but unfortunately +this channel is not determined by superiority of any kind. + +Although civilisation thus checks in many ways the action of natural +selection, it apparently favours, by means of improved food and the +freedom from occasional hardships, the better development of the body. +This may be inferred from civilised men having been found, wherever +compared, to be physically stronger than savages. They appear also to +have equal powers of endurance, as has been proved in many adventurous +expeditions. Even the great luxury of the rich can be but little +detrimental; for the expectation of life of our aristocracy, at all ages +and of both sexes, is very little inferior to that of healthy English +lives in the lower classes.[233] + +We will now look to the intellectual faculties alone. If in each grade +of society the members were divided into two equal bodies, the one +including the intellectually superior and the other the inferior, there +can be little doubt that the former would succeed best in all +occupations and rear a greater number of children. Even in the lowest +walks of life, skill and ability must be of some advantage, though in +many occupations, owing to the great division of labour, a very small +one. Hence in civilised nations there will be some tendency to an +increase both in the number and in the standard of the intellectually +able. But I do not wish to assert that this tendency may not be more +than counterbalanced in other ways, as by the multiplication of the +reckless and improvident; but even to such as these, ability must be +some advantage. + +It has often been objected to views like the foregoing, that the most +eminent men who have ever lived have left no offspring to inherit their +great intellect. Mr. Galton says,[234] “I regret I am unable to solve +the simple question whether, and how far, men and women who are +prodigies of genius are infertile. I have, however, shewn that men of +eminence are by no means so.” + +Great lawgivers, the founders of beneficent religions, great +philosophers and discoverers in science, aid the progress of mankind in +a far higher degree by their works than by leaving a numerous progeny. +In the case of corporeal structures, it is the selection of the slightly +better-endowed and the elimination of the slightly less well-endowed +individuals, and not the preservation of strongly-marked and rare +anomalies, that leads to the advancement of a species.[235] So it will +be with the intellectual faculties, namely from the somewhat more able +men in each grade of society succeeding rather better than the less +able, and consequently increasing in number, if not otherwise prevented. +When in any nation the standard of intellect and the number of +intellectual men have increased, we may expect from the law of the +deviation from an average, as shewn by Mr. Galton, that prodigies of +genius will appear somewhat more frequently than before. + +In regard to the moral qualities, some elimination of the worst +dispositions is always in progress even in the most civilised nations. +Malefactors are executed, or imprisoned for long periods, so that they +cannot freely transmit their bad qualities. Melancholic and insane +persons are confined, or commit suicide. Violent and quarrelsome men +often come to a bloody end. Restless men who will not follow any steady +occupation—and this relic of barbarism is a great check to +civilisation[236]—emigrate to newly-settled countries, where they prove +useful pioneers. Intemperance is so highly destructive, that the +expectation of life of the intemperate, at the age, for instance, of +thirty, is only 13.8 years; whilst for the rural labourers of England at +the same age it is 40·59 years.[237] Profligate women bear few children, +and profligate men rarely marry; both suffer from disease. In the +breeding of domestic animals, the elimination of those individuals, +though few in number, which are in any marked manner inferior, is by no +means an unimportant element towards success. This especially holds good +with injurious characters which tend to reappear through reversion, such +as blackness in sheep; and with mankind some of the worst dispositions, +which occasionally without any assignable cause make their appearance in +families, may perhaps be reversions to a savage state, from which we are +not removed by very many generations. This view seems indeed recognised +in the common expression that such men are the black sheep of the +family. + +With civilised nations, as far as an advanced standard of morality, and +an increased number of fairly well-endowed men are concerned, natural +selection apparently effects but little; though the fundamental social +instincts were originally thus gained. But I have already said enough, +whilst treating of the lower races, on the causes which lead to the +advance of morality, namely, the approbation of our fellow-men—the +strengthening of our sympathies by habit—example and imitation— +reason—experience and even self-interest—instruction during +youth, and religious feelings. + +A most important obstacle in civilised countries to an increase in the +number of men of a superior class has been strongly urged by Mr. Greg +and Mr. Galton,[238] namely, the fact that the very poor and reckless, +who are often degraded by vice, almost invariably marry early, whilst +the careful and frugal, who are generally otherwise virtuous, marry late +in life, so that they may be able to support themselves and their +children in comfort. Those who marry early produce within a given period +not only a greater number of generations, but, as shewn by Dr. +Duncan,[239] they produce many more children. The children, moreover, +that are born by mothers during the prime of life are heavier and +larger, and therefore probably more vigorous, than those born at other +periods. Thus the reckless, degraded, and often vicious members of +society, tend to increase at a quicker rate than the provident and +generally virtuous members. Or as Mr. Greg puts the case: “The careless, +squalid, unaspiring Irishman multiplies like rabbits: the frugal, +foreseeing, self-respecting, ambitious Scot, stern in his morality, +spiritual in his faith, sagacious and disciplined in his intelligence, +passes his best years in struggle and in celibacy, marries late, and +leaves few behind him. Given a land originally peopled by a thousand +Saxons and a thousand Celts—and in a dozen generations five-sixths of +the population would be Celts, but five-sixths of the property, of the +power, of the intellect, would belong to the one-sixth of Saxons that +remained. In the eternal ‘struggle for existence,’ it would be the +inferior and _less_ favoured race that had prevailed—and prevailed by +virtue not of its good qualities but of its faults.” + +There are, however, some checks to this downward tendency. We have seen +that the intemperate suffer from a high rate of mortality, and the +extremely profligate leave few offspring. The poorest classes crowd into +towns, and it has been proved by Dr. Stark from the statistics of ten +years in Scotland,[240] that at all ages the death-rate is higher in +towns than in rural districts, “and during the first five years of life +the town death-rate is almost exactly double that of the rural +districts.” As these returns include both the rich and the poor, no +doubt more than double the number of births would be requisite to keep +up the number of the very poor inhabitants in the towns, relatively to +those in the country. With women, marriage at too early an age is highly +injurious; for it has been found in France that, “twice as many wives +under twenty die in the year, as died out of the same number of the +unmarried.” The mortality, also, of husbands under twenty is +“excessively high,”[241] but what the cause of this may be seems +doubtful. Lastly, if the men who prudently delay marrying until they can +bring up their families in comfort, were to select, as they often do, +women in the prime of life, the rate of increase in the better class +would be only slightly lessened. + +It was established from an enormous body of statistics, taken during +1853, that the unmarried men throughout France, between the ages of +twenty and eighty, die in a much larger proportion than the married: for +instance, out of every 1000 unmarried men, between the ages of twenty +and thirty, 11·3 annually died, whilst of the married only 6·5 +died.[242] A similar law was proved to hold good, during the years 1863 +and 1864, with the entire population above the age of twenty in +Scotland: for instance, out of every 1000 unmarried men, between the +ages of twenty and thirty, 14·97 annually died, whilst of the married +only 7·24 died, that is less than half.[243] Dr. Stark remarks on this, +“Bachelorhood is more destructive to life than the most unwholesome +trades, or than residence in an unwholesome house or district where +there has never been the most distant attempt at sanitary improvement.” +He considers that the lessened mortality is the direct result of +“marriage, and the more regular domestic habits which attend that +state.” He admits, however, that the intemperate, profligate, and +criminal classes, whose duration of life is low, do not commonly marry; +and it must likewise be admitted that men with a weak constitution, +ill health, or any great infirmity in body or mind, will often not wish +to marry, or will be rejected. Dr. Stark seems to have come to the +conclusion that marriage in itself is a main cause of prolonged life, +from finding that aged married men still have a considerable advantage +in this respect over the unmarried of the same advanced age; but every +one must have known instances of men, who with weak health during youth +did not marry, and yet have survived to old age, though remaining weak +and therefore always with a lessened chance of life. There is another +remarkable circumstance which seems to support Dr. Stark’s conclusion, +namely, that widows and widowers in France suffer in comparison with the +married a very heavy rate of mortality; but Dr. Farr attributes this to +the poverty and evil habits consequent on the disruption of the family, +and to grief. On the whole we may conclude with Dr. Farr that the lesser +mortality of married than of unmarried men, which seems to be a general +law, “is mainly due to the constant elimination of imperfect types, and +to the skilful selection of the finest individuals out of each +successive generation;” the selection relating only to the marriage +state, and acting on all corporeal, intellectual, and moral qualities. +We may, therefore, infer that sound and good men who out of prudence +remain for a time unmarried do not suffer a high rate of mortality. + +If the various checks specified in the two last paragraphs, and perhaps +others as yet unknown, do not prevent the reckless, the vicious and +otherwise inferior members of society from increasing at a quicker rate +than the better class of men, the nation will retrograde, as has +occurred too often in the history of the world. We must remember that +progress is no invariable rule. It is most difficult to say why one +civilised nation rises, becomes more powerful, and spreads more widely, +than another; or why the same nation progresses more at one time than at +another. We can only say that it depends on an increase in the actual +number of the population, on the number of the men endowed with high +intellectual and moral faculties, as well as on their standard of +excellence. Corporeal structure, except so far as vigour of body leads +to vigour of mind, appears to have little influence. + +It has been urged by several writers that as high intellectual powers +are advantageous to a nation, the old Greeks, who stood some grades +higher in intellect than any race that has ever existed,[244] ought to +have risen, if the power of natural selection were real, still higher +in the scale, increased in number, and stocked the whole of Europe. Here +we have the tacit assumption, so often made with respect to corporeal +structures, that there is some innate tendency towards continued +development in mind and body. But development of all kinds depends on +many concurrent favourable circumstances. Natural selection acts only in +a tentative manner. Individuals and races may have acquired certain +indisputable advantages, and yet have perished from failing in other +characters. The Greeks may have retrograded from a want of coherence +between the many small states, from the small size of their whole +country, from the practice of slavery, or from extreme sensuality; for +they did not succumb until “they were enervated and corrupt to the very +core.”[245] The western nations of Europe, who now so immeasurably +surpass their former savage progenitors and stand at the summit of +civilisation, owe little or none of their superiority to direct +inheritance from the old Greeks; though they owe much to the written +works of this wonderful people. + +Who can positively say why the Spanish nation, so dominant at one time, +has been distanced in the race. The awakening of the nations of Europe +from the dark ages is a still more perplexing problem. At this early +period, as Mr. Galton[246] has remarked, almost all the men of a gentle +nature, those given to meditation or culture of the mind, had no refuge +except in the bosom of the Church which demanded celibacy; and this +could hardly fail to have had a deteriorating influence on each +successive generation. During this same period the Holy Inquisition +selected with extreme care the freest and boldest men in order to burn +or imprison them. In Spain alone some of the best men—those who doubted +and questioned, and without doubting there can be no progress—were +eliminated during three centuries at the rate of a thousand a year. The +evil which the Catholic Church has thus effected, though no doubt +counterbalanced to a certain, perhaps large extent in other ways, is +incalculable; nevertheless, Europe has progressed at an unparalleled +rate. + +The remarkable success of the English as colonists over other European +nations, which is well illustrated by comparing the progress of the +Canadians of English and French extraction, has been ascribed to their +“daring and persistent energy;” but who can say how the English gained +their energy. There is apparently much truth in the belief that the +wonderful progress of the United States, as well as the character of the +people, are the results of natural selection; the more energetic, +restless, and courageous men from all parts of Europe having emigrated +during the last ten or twelve generations to that great country, and +having there succeeded best.[247] Looking to the distant future, I do +not think that the Rev. Mr. Zincke takes an exaggerated view when he +says:[248] “All other series of events—as that which resulted in the +culture of mind in Greece, and that which resulted in the empire of +Rome—only appear to have purpose and value when viewed in connection +with, or rather as subsidiary to ... the great stream of Anglo-Saxon +emigration to the west.” + +Obscure as is the problem of the advance of civilisation, we can at +least see that a nation which produced during a lengthened period the +greatest number of highly intellectual, energetic, brave, patriotic, and +benevolent men, would generally prevail over less favoured nations. + +Natural selection follows from the struggle for existence; and this from +a rapid rate of increase. It is impossible not bitterly to regret, but +whether wisely is another question, the rate at which man tends to +increase; for this leads in barbarous tribes to infanticide and many +other evils, and in civilised nations to abject poverty, celibacy, and +to the late marriages of the prudent. But as man suffers from the same +physical evils with the lower animals, he has no right to expect an +immunity from the evils consequent on the struggle for existence. Had he +not been subjected to natural selection, assuredly he would never have +attained to the rank of manhood. When we see in many parts of the world +enormous areas of the most fertile land peopled by a few wandering +savages, but which are capable of supporting numerous happy homes, it +might be argued that the struggle for existence had not been +sufficiently severe to force man upwards to his highest standard. +Judging from all that we know of man and the lower animals, there has +always been sufficient variability in the intellectual and moral +faculties, for their steady advancement through natural selection. No +doubt such advancement demands many favourable concurrent circumstances; +but it may well be doubted whether the most favourable would have +sufficed, had not the rate of increase been rapid, and the consequent +struggle for existence severe to an extreme degree. + + +_On the evidence that all civilised nations were once barbarous._—As we +have had to consider the steps by which some semi-human creature has +been gradually raised to the rank of man in his most perfect state, the +present subject cannot be quite passed over. But it has been treated in +so full and admirable a manner by Sir J. Lubbock,[249] Mr. Tylor, Mr. +M’Lennan, and others, that I need here give only the briefest summary of +their results. The arguments recently advanced by the Duke of +Argyll[250] and formerly by Archbishop Whately, in favour of the belief +that man came into the world as a civilised being and that all savages +have since undergone degradation, seem to me weak in comparison with +those advanced on the other side. Many nations, no doubt, have fallen +away in civilisation, and some may have lapsed into utter barbarism, +though on this latter head I have not met with any evidence. The +Fuegians were probably compelled by other conquering hordes to settle in +their inhospitable country, and they may have become in consequence +somewhat more degraded; but it would be difficult to prove that they +have fallen much below the Botocudos who inhabit the finest parts of +Brazil. + +The evidence that all civilised nations are the descendants of +barbarians, consists, on the one side, of clear traces of their former +low condition in still-existing customs, beliefs, language, &c.; and on +the other side, of proofs that savages are independently able to raise +themselves a few steps in the scale of civilisation, and have actually +thus risen. The evidence on the first head is extremely curious, but +cannot be here given: I refer to such cases as that, for instance, of +the art of enumeration, which, as Mr. Tylor clearly shews by the words +still used in some places, originated in counting the fingers, first of +one hand and then of the other, and lastly of the toes. We have traces +of this in our own decimal system, and in the Roman numerals, which +after reaching to the number V., change into VI., &c., when the other +hand no doubt was used. So again, “when we speak of three-score and ten, +we are counting by the vigesimal system, each score thus ideally made, +standing for 20—for ‘one man’ as a Mexican or Carib would put it.”[251] +According to a large and increasing school of philologists, every +language bears the marks of its slow and gradual evolution. So it is +with the art of writing, as letters are rudiments of pictorial +representations. It is hardly possible to read Mr. M’Lennan’s work[252] +and not admit that almost all civilised nations still retain some traces +of such rude habits as the forcible capture of wives. What ancient +nation, as the same author asks, can be named that was originally +monogamous? The primitive idea of justice, as shewn by the law of battle +and other customs of which traces still remain, was likewise most rude. +Many existing superstitions are the remnants of former false religious +beliefs. The highest form of religion—the grand idea of God hating sin +and loving righteousness—was unknown during primeval times. + +Turning to the other kind of evidence: Sir J. Lubbock has shewn that +some savages have recently improved a little in some of their simpler +arts. From the extremely curious account which he gives of the weapons, +tools, and arts, used or practised by savages in various parts of the +world, it cannot be doubted that these have nearly all been independent +discoveries, excepting perhaps the art of making fire.[253] The +Australian boomerang is a good instance of one such independent +discovery. The Tahitians when first visited had advanced in many +respects beyond the inhabitants of most of the other Polynesian islands. +There are no just grounds for the belief that the high culture of the +native Peruvians and Mexicans was derived from any foreign source;[254] +many native plants were there cultivated, and a few native animals +domesticated. We should bear in mind that a wandering crew from some +semi-civilised land, if washed to the shores of America, would not, +judging from the small influence of most missionaries, have produced any +marked effect on the natives, unless they had already become somewhat +advanced. Looking to a very remote period in the history of the world, +we find, to use Sir J. Lubbock’s well-known terms, a paleolithic and +neolithic period; and no one will pretend that the art of grinding rough +flint tools was a borrowed one. In all parts of Europe, as far east as +Greece, in Palestine, India, Japan, New Zealand, and Africa, including +Egypt, flint tools have been discovered in abundance; and of their use +the existing inhabitants retain no tradition. There is also indirect +evidence of their former use by the Chinese and ancient Jews. Hence +there can hardly be a doubt that the inhabitants of these many +countries, which include nearly the whole civilised world, were once in +a barbarous condition. To believe that man was aboriginally civilised +and then suffered utter degradation in so many regions, is to take a +pitiably low view of human nature. It is apparently a truer and more +cheerful view that progress has been much more general than +retrogression; that man has risen, though by slow and interrupted steps, +from a lowly condition to the highest standard as yet attained by him in +knowledge, morals, and religion. + + + + +CHAPTER VI. + +ON THE AFFINITIES AND GENEALOGY OF MAN. + + + Position of man in the animal series—The natural system + genealogical—Adaptive characters of slight value—Various + small points of resemblance between man and the + Quadrumana—Rank of man in the natural system—Birthplace and + antiquity of man—Absence of fossil connecting-links—Lower + stages in the genealogy of man, as inferred, firstly from his + affinities and secondly from his structure—Early androgynous + condition of the Vertebrata—Conclusion. + + +Even if it be granted that the difference between man and his nearest +allies is as great in corporeal structure as some naturalists maintain, +and although we must grant that the difference between them is immense +in mental power, yet the facts given in the previous chapters declare, +as it appears to me, in the plainest manner, that man is descended from +some lower form, notwithstanding that connecting-links have not hitherto +been discovered. + +Man is liable to numerous, slight, and diversified variations, which are +induced by the same general causes, are governed and transmitted in +accordance with the same general laws, as in the lower animals. Man +tends to multiply at so rapid a rate that his offspring are necessarily +exposed to a struggle for existence, and consequently to natural +selection. He has given rise to many races, some of which are so +different that they have often been ranked by naturalists as distinct +species. His body is constructed on the same homological plan as that of +other mammals, independently of the uses to which the several parts may +be put. He passes through the same phases of embryological development. +He retains many rudimentary and useless structures, which no doubt were +once serviceable. Characters occasionally make their reappearance in +him, which we have every reason to believe were possessed by his early +progenitors. If the origin of man had been wholly different from that of +all other animals, these various appearances would be mere empty +deceptions; but such an admission is incredible. These appearances, on +the other hand, are intelligible, at least to a large extent, if man is +the co-descendant with other mammals of some unknown and lower form. + +Some naturalists, from being deeply impressed with the mental and +spiritual powers of man, have divided the whole organic world into three +kingdoms, the Human, the Animal, and the Vegetable, thus giving to man a +separate kingdom.[255] Spiritual powers cannot be compared or classed by +the naturalist; but he may endeavour to shew, as I have done, that the +mental faculties of man and the lower animals do not differ in kind, +although immensely in degree. A difference in degree, however great, +does not justify us in placing man in a distinct kingdom, as will +perhaps be best illustrated by comparing the mental powers of two +insects, namely, a coccus or scale-insect and an ant, which undoubtedly +belong to the same class. The difference is here greater, though of a +somewhat different kind, than that between man and the highest mammal. +The female coccus, whilst young, attaches itself by its proboscis to a +plant; sucks the sap but never moves again; is fertilised and lays eggs; +and this is its whole history. On the other hand, to describe the habits +and mental powers of a female ant, would require, as Pierre Huber has +shewn, a large volume; I may, however, briefly specify a few points. +Ants communicate information to each other, and several unite for the +same work, or games of play. They recognise their fellow-ants after +months of absence. They build great edifices, keep them clean, close the +doors in the evening, and post sentries. They make roads, and even +tunnels under rivers. They collect food for the community, and when an +object, too large for entrance, is brought to the nest, they enlarge the +door, and afterwards build it up again.[256] They go out to battle in +regular bands, and freely sacrifice their lives for the common weal. +They emigrate in accordance with a preconcerted plan. They capture +slaves. They keep Aphides as milch-cows. They move the eggs of their +aphides, as well as their own eggs and cocoons, into warm parts of the +nest, in order that they may be quickly hatched; and endless similar +facts could be given. On the whole, the difference in mental power +between an ant and a coccus is immense; yet no one has ever dreamed of +placing them in distinct classes, much less in distinct kingdoms. No +doubt this interval is bridged over by the intermediate mental powers of +many other insects; and this is not the case with man and the higher +apes. But we have every reason to believe that breaks in the series are +simply the result of many forms having become extinct. + +Professor Owen, relying chiefly on the structure of the brain, has +divided the mammalian series into four sub-classes. One of these he +devotes to man; in another he places both the marsupials and the +monotremata; so that he makes man as distinct from all other mammals as +are these two latter groups conjoined. This view has not been accepted, +as far as I am aware, by any naturalist capable of forming an +independent judgment, and therefore need not here be further considered. + +We can understand why a classification founded on any single character +or organ—even an organ so wonderfully complex and important as the +brain—or on the high development of the mental faculties, is almost +sure to prove unsatisfactory. This principle has indeed been tried with +hymenopterous insects; but when thus classed by their habits or +instincts, the arrangement proved thoroughly artificial.[257] +Classifications may, of course, be based on any character whatever, as +on size, colour, or the element inhabited; but naturalists have long +felt a profound conviction that there is a natural system. This system, +it is now generally admitted, must be, as far as possible, genealogical +in arrangement,—that is, the co-descendants of the same form must be +kept together in one group, separate from the co-descendants of any +other form; but if the parent-forms are related, so will be their +descendants, and the two groups together will form a larger group. The +amount of difference between the several groups—that is the amount of +modification which each has undergone—will be expressed by such terms +as genera, families, orders, and classes. As we have no record of the +lines of descent, these lines can be discovered only by observing the +degrees of resemblance between the beings which are to be classed. For +this object numerous points of resemblance are of much more importance +than the amount of similarity or dissimilarity in a few points. If two +languages were found to resemble each other in a multitude of words and +points of construction, they would be universally recognised as having +sprung from a common source, notwithstanding that they differed greatly +in some few words or points of construction. But with organic beings the +points of resemblance must not consist of adaptations to similar habits +of life: two animals may, for instance, have had their whole frames +modified for living in the water, and yet they will not be brought any +nearer to each other in the natural system. Hence we can see how it is +that resemblances in unimportant structures, in useless and rudimentary +organs, and in parts not as yet fully developed or functionally active, +are by far the most serviceable for classification; for they can hardly +be due to adaptations within a late period; and thus they reveal the old +lines of descent or of true affinity. + +We can further see why a great amount of modification in some one +character ought not to lead us to separate widely any two organisms. A +part which already differs much from the same part in other allied forms +has already, according to the theory of evolution, varied much; +consequently it would (as long as the organism remained exposed to the +same exciting conditions) be liable to further variations of the same +kind; and these, if beneficial, would be preserved, and thus continually +augmented. In many cases the continued development of a part, for +instance, of the beak of a bird, or of the teeth of a mammal, would not +be advantageous to the species for gaining its food, or for any other +object; but with man we can see no definite limit, as far as advantage +is concerned, to the continued development of the brain and mental +faculties. Therefore in determining the position of man in the natural +or genealogical system, the extreme development of his brain ought not +to outweigh a multitude of resemblances in other less important or +quite unimportant points. + +The greater number of naturalists who have taken into consideration the +whole structure of man, including his mental faculties, have followed +Blumenbach and Cuvier, and have placed man in a separate Order, under +the title of the Bimana, and therefore on an equality with the Orders of +the Quadrumana, Carnivora, &c. Recently many of our best naturalists +have recurred to the view first propounded by Linnæus, so remarkable for +his sagacity, and have placed man in the same Order with the Quadrumana, +under the title of the Primates. The justice of this conclusion will be +admitted if, in the first place, we bear in mind the remarks just made +on the comparatively small importance for classification of the great +development of the brain in man; bearing, also, in mind that the +strongly-marked differences between the skulls of man and the Quadrumana +(lately insisted upon by Bischoff, Aeby, and others) apparently follow +from their differently developed brains. In the second place, we must +remember that nearly all the other and more important differences +between man and the Quadrumana are manifestly adaptive in their nature, +and relate chiefly to the erect position of man; such as the structure +of his hand, foot, and pelvis, the curvature of his spine, and the +position of his head. The family of seals offers a good illustration of +the small importance of adaptive characters for classification. These +animals differ from all other Carnivora in the form of their bodies and +in the structure of their limbs, far more than does man from the higher +apes; yet in every system, from that of Cuvier to the most recent one by +Mr. Flower,[258] seals are ranked as a mere family in the Order of the +Carnivora. If man had not been his own classifier, he would never have +thought of founding a separate order for his own reception. + +It would be beyond my limits, and quite beyond my knowledge, even to +name the innumerable points of structure in which man agrees with the +other Primates. Our great anatomist and philosopher, Prof. Huxley, has +fully discussed this subject,[259] and has come to the conclusion that +man in all parts of his organisation differs less from the higher apes, +than these do from the lower members of the same group. Consequently +there “is no justification for placing man in a distinct order.” + +In an early part of this volume I brought forward various facts, shewing +how closely man agrees in constitution with the higher mammals; and this +agreement, no doubt, depends on our close similarity in minute structure +and chemical composition. I gave, as instances, our liability to the +same diseases, and to the attacks of allied parasites; our tastes in +common for the same stimulants, and the similar effects thus produced, +as well as by various drugs; and other such facts. + +As small unimportant points of resemblance between man and the higher +apes are not commonly noticed in systematic works, and as, when +numerous, they clearly reveal our relationship, I will specify a few +such points. The relative position of the features are manifestly the +same in man and the Quadrumana; and the various emotions are displayed +by nearly similar movements of the muscles and skin, chiefly above the +eyebrows and round the mouth. Some few expressions are, indeed, almost +the same, as in the weeping of certain kinds of monkeys, and in the +laughing noise made by others, during which the corners of the mouth are +drawn backwards, and the lower eyelids wrinkled. The external ears are +curiously alike. In man the nose is much more prominent than in most +monkeys; but we may trace the commencement of an aquiline curvature in +the nose of the Hoolock Gibbon; and this in the _Semnopithecus nasica_ +is carried to a ridiculous extreme. + +The faces of many monkeys are ornamented with beards, whiskers, or +moustaches. The hair on the head grows to a great length in some species +of Semnopithecus;[260] and in the Bonnet monkey (_Macacus radiatus_) it +radiates from a point on the crown, with a parting down the middle, as +in man. It is commonly said that the forehead gives to man his noble and +intellectual appearance; but the thick hair on the head of the Bonnet +monkey terminates abruptly downwards, and is succeeded by such short and +fine hair, or down, that at a little distance the forehead, with the +exception of the eyebrows, appears quite naked. It has been erroneously +asserted that eyebrows are not present in any monkey. In the species +just named the degree of nakedness of the forehead differs in different +individuals; and Eschricht states[261] that in our children the limit +between the hairy scalp and the naked forehead is sometimes not well +defined; so that here we seem to have a trifling case of reversion to a +progenitor, in whom the forehead had not as yet become quite naked. + +It is well known that the hair on our arms tends to converge from above +and below to a point at the elbow. This curious arrangement, so unlike +that in most of the lower mammals, is common to the gorilla, chimpanzee, +orang, some species of Hylobates, and even to some few American monkeys. +But in _Hylobates agilis_ the hair on the fore-arm is directed +downwards or towards the wrist in the ordinary manner; and in _H. lar_ +it is nearly erect, with only a very slight forward inclination; so that +in this latter species it is in a transitional state. It can hardly be +doubted that with most mammals the thickness of the hair and its +direction on the back is adapted to throw off the rain; even the +transverse hairs on the fore-legs of a dog may serve for this end when +he is coiled up asleep. Mr. Wallace remarks that the convergence of the +hair towards the elbow on the arms of the orang (whose habits he has so +carefully studied) serves to throw off the rain, when, as is the custom +of this animal, the arms are bent, with the hands clasped round a branch +or over its own head. We should, however, bear in mind that the attitude +of an animal may perhaps be in part determined by the direction of the +hair; and not the direction of the hair by the attitude. If the above +explanation is correct in the case of the orang, the hair on our +fore-arms offers a curious record of our former state; for no one +supposes that it is now of any use in throwing off the rain, nor in our +present erect condition is it properly directed for this purpose. + +It would, however, be rash to trust too much to the principle of +adaptation in regard to the direction of the hair in man or his early +progenitors; for it is impossible to study the figures given by +Eschricht of the arrangement of the hair on the human fœtus (this +being the same as in the adult) and not agree with this excellent +observer that other and more complex causes have intervened. The points +of convergence seem to stand in some relation to those points in the +embryo which are last closed in during development. There appears, also, +to exist some relation between the arrangement of the hair on the +limbs, and the course of the medullary arteries.[262] + +It must not be supposed that the resemblances between man and certain +apes in the above and many other points—such as in having a naked +forehead, long tresses on the head, &c.—are all necessarily the result +of unbroken inheritance from a common progenitor thus characterised, or +of subsequent reversion. Many of these resemblances are more probably +due to analogous variation, which follows, as I have elsewhere attempted +to shew,[263] from co-descended organisms having a similar constitution +and having been acted on by similar causes inducing variability. With +respect to the similar direction of the hair on the fore-arms of man and +certain monkeys, as this character is common to almost all the +anthropomorphous apes, it may probably be attributed to inheritance; but +not certainly so, as some very distinct American monkeys are thus +characterised. The same remark is applicable to the tailless condition +of man; for the tail is absent in all the anthropomorphous apes. +Nevertheless this character cannot with certainty be attributed to +inheritance, as the tail, though not absent, is rudimentary in several +other Old World and in some New World species, and is quite absent in +several species belonging to the allied group of Lemurs. + +Although, as we have now seen, man has no just right to form a separate +Order for his own reception, he may perhaps claim a distinct Sub-order +or Family. Prof. Huxley, in his last work,[264] divides the Primates +into three Sub-orders; namely, the Anthropidæ with man alone, the +Simiadæ including monkeys of all kinds, and the Lemuridæ with the +diversified genera of lemurs. As far as differences in certain important +points of structure are concerned, man may no doubt rightly claim the +rank of a Sub-order; and this rank is too low, if we look chiefly to his +mental faculties. Nevertheless, under a genealogical point of view it +appears that this rank is too high, and that man ought to form merely a +Family, or possibly even only a Sub-family. If we imagine three lines of +descent proceeding from a common source, it is quite conceivable that +two of them might after the lapse of ages be so slightly changed as +still to remain as species of the same genus; whilst the third line +might become so greatly modified as to deserve to rank as a distinct +Sub-family, Family, or even Order. But in this case it is almost certain +that the third line would still retain through inheritance numerous +small points of resemblance with the other two lines. Here then would +occur the difficulty, at present insoluble, how much weight we ought to +assign in our classifications to strongly-marked differences in some few +points,—that is to the amount of modification undergone; and how much +to close resemblance in numerous unimportant points, as indicating the +lines of descent or genealogy. The former alternative is the most +obvious, and perhaps the safest, though the latter appears the most +correct as giving a truly natural classification. + +To form a judgment on this head, with reference to man we must glance at +the classification of the Simiadæ. This family is divided by almost all +naturalists into the Catarhine group, or Old World monkeys, all of which +are characterised (as their name expresses) by the peculiar structure of +their nostrils and by having four premolars in each jaw; and into the +Platyrhine group or New World monkeys (including two very distinct +sub-groups), all of which are characterised by differently-constructed +nostrils and by having six premolars in each jaw. Some other small +differences might be mentioned. Now man unquestionably belongs in his +dentition, in the structure of his nostrils, and some other respects, to +the Catarhine or Old World division; nor does he resemble the +Platyrhines more closely than the Catarhines in any characters, +excepting in a few of not much importance and apparently of an adaptive +nature. Therefore it would be against all probability to suppose that +some ancient New World species had varied, and had thus produced a +man-like creature with all the distinctive characters proper to the Old +World division; losing at the same time all its own distinctive +characters. There can consequently hardly be a doubt that man is an +offshoot from the Old World Simian stem; and that under a genealogical +point of view, he must be classed with the Catarhine division.[265] + +The anthropomorphous apes, namely the gorilla, chimpanzee, orang, and +hylobates, are separated as a distinct sub-group from the other Old +World monkeys by most naturalists. I am aware that Gratiolet, relying on +the structure of the brain, does not admit the existence of this +sub-group, and no doubt it is a broken one; thus the orang, as Mr. St. +G. Mivart remarks,[266] “is one of the most peculiar and aberrant forms +to be found in the Order.” The remaining, non-anthropomorphous, Old +World monkeys, are again divided by some naturalists into two or three +smaller sub-groups; the genus Semnopithecus, with its peculiar +sacculated stomach, being the type of one such sub-group. But it appears +from M. Gaudry’s wonderful discoveries in Attica, that during the +Miocene period a form existed there, which connected Semnopithecus and +Macacus; and this probably illustrates the manner in which the other and +higher groups were once blended together. + +If the anthropomorphous apes be admitted to form a natural sub-group, +then as man agrees with them, not only in all those characters which he +possesses in common with the whole Catarhine group, but in other +peculiar characters, such as the absence of a tail and of callosities +and in general appearance, we may infer that some ancient member of the +anthropomorphous sub-group gave birth to man. It is not probable that a +member of one of the other lower sub-groups should, through the law of +analogous variation, have given rise to a man-like creature, resembling +the higher anthropomorphous apes in so many respects. No doubt man, in +comparison with most of his allies, has undergone an extraordinary +amount of modification, chiefly in consequence of his greatly developed +brain and erect position; nevertheless we should bear in mind that he +“is but one of several exceptional forms of Primates.”[267] + +Every naturalist, who believes in the principle of evolution, will +grant that the two main divisions of the Simiadæ, namely the Catarhine +and Platyrhine monkeys, with their sub-groups, have all proceeded from +some one extremely ancient progenitor. The early descendants of this +progenitor, before they had diverged to any considerable extent from +each other, would still have formed a single natural group; but some of +the species or incipient genera would have already begun to indicate by +their diverging characters the future distinctive marks of the Catarhine +and Platyrhine divisions. Hence the members of this supposed ancient +group would not have been so uniform in their dentition or in the +structure of their nostrils, as are the existing Catarhine monkeys in +one way and the Platyrhines in another way, but would have resembled in +this respect the allied Lemuridæ which differ greatly from each other in +the form of their muzzles,[268] and to an extraordinary degree in their +dentition. + +The Catarhine and Platyrhine monkeys agree in a multitude of characters, +as is shewn by their unquestionably belonging to one and the same Order. +The many characters which they possess in common can hardly have been +independently acquired by so many distinct species; so that these +characters must have been inherited. But an ancient form which possessed +many characters common to the Catarhine and Platyrhine monkeys, and +others in an intermediate condition, and some few perhaps distinct from +those now present in either group, would undoubtedly have been ranked, +if seen by a naturalist, as an ape or monkey. And as man under a +genealogical point of view belongs to the Catarhine or Old World stock, +we must conclude, however much the conclusion may revolt our pride, +that our early progenitors would have been properly thus +designated.[269] But we must not fall into the error of supposing that +the early progenitor of the whole Simian stock, including man, was +identical with, or even closely resembled, any existing ape or monkey. + + +_On the Birthplace and Antiquity of Man._—We are naturally led to +enquire where was the birthplace of man at that stage of descent when +our progenitors diverged from the Catarhine stock. The fact that they +belonged to this stock clearly shews that they inhabited the Old World; +but not Australia nor any oceanic island, as we may infer from the laws +of geographical distribution. In each great region of the world the +living mammals are closely related to the extinct species of the same +region. It is therefore probable that Africa was formerly inhabited by +extinct apes closely allied to the gorilla and chimpanzee; and as these +two species are now man’s nearest allies, it is somewhat more probable +that our early progenitors lived on the African continent than +elsewhere. But it is useless to speculate on this subject, for an ape +nearly as large as a man, namely the Dryopithecus of Lartet, which was +closely allied to the anthropomorphous Hylobates, existed in Europe +during the Upper Miocene period; and since so remote a period the earth +has certainly undergone many great revolutions, and there has been ample +time for migration on the largest scale. + +At the period and place, whenever and wherever it may have been, when +man first lost his hairy covering, he probably inhabited a hot country; +and this would have been favourable for a frugiferous diet, on which, +judging from analogy, he subsisted. We are far from knowing how long ago +it was when man first diverged from the Catarhine stock; but this may +have occurred at an epoch as remote as the Eocene period; for the higher +apes had diverged from the lower apes as early as the Upper Miocene +period, as shewn by the existence of the Dryopithecus. We are also quite +ignorant at how rapid a rate organisms, whether high or low in the +scale, may under favourable circumstances be modified: we know, however, +that some have retained the same form during an enormous lapse of time. +From what we see going on under domestication, we learn that within the +same period some of the co-descendants of the same species may be not at +all changed, some a little, and some greatly changed. Thus it may have +been with man, who has undergone a great amount of modification in +certain characters in comparison with the higher apes. + +The great break in the organic chain between man and his nearest allies, +which cannot be bridged over by any extinct or living species, has often +been advanced as a grave objection to the belief that man is descended +from some lower form; but this objection will not appear of much weight +to those who, convinced by general reasons, believe in the general +principle of evolution. Breaks incessantly occur in all parts of the +series, some being wide, sharp and defined, others less so in various +degrees; as between the orang and its nearest allies—between the +Tarsius and the other Lemuridæ—between the elephant and in a more +striking manner between the Ornithorhynchus or Echidna, and other +mammals. But all these breaks depend merely on the number of related +forms which have become extinct. At some future period, not very distant +as measured by centuries, the civilised races of man will almost +certainly exterminate and replace throughout the world the savage races. +At the same time the anthropomorphous apes, as Professor Schaaffhausen +has remarked,[270] will no doubt be exterminated. The break will then be +rendered wider, for it will intervene between man in a more civilised +state, as we may hope, than the Caucasian, and some ape as low as a +baboon, instead of as at present between the negro or Australian and the +gorilla. + +With respect to the absence of fossil remains, serving to connect man +with his ape-like progenitors, no one will lay much stress on this fact, +who will read Sir C. Lyell’s discussion,[271] in which he shews that in +all the vertebrate classes the discovery of fossil remains has been an +extremely slow and fortuitous process. Nor should it be forgotten that +those regions which are the most likely to afford remains connecting man +with some extinct ape-like creature, have not as yet been searched by +geologists. + + +_Lower Stages in the Genealogy of Man._—We have seen that man appears +to have diverged from the Catarhine or Old World division of the +Simiadæ, after these had diverged from the New World division. We will +now endeavour to follow the more remote traces of his genealogy, +trusting in the first place to the mutual affinities between the various +classes and orders, with some slight aid from the periods, as far as +ascertained, of their successive appearance on the earth. The Lemuridæ +stand below and close to the Simiadæ, constituting a very distinct +family of the Primates, or, according to Häckel, a distinct Order. This +group is diversified and broken to an extraordinary degree, and includes +many aberrant forms. It has, therefore, probably suffered much +extinction. Most of the remnants survive on islands, namely in +Madagascar and in the islands of the Malayan archipelago, where they +have not been exposed to such severe competition as they would have been +on well-stocked continents. This group likewise presents many +gradations, leading, as Huxley remarks,[272] “insensibly from the crown +and summit of the animal creation down to creatures from which there is +but a step, as it seems, to the lowest, smallest, and least intelligent +of the placental mammalia.” From these various considerations it is +probable that the Simiadæ were originally developed from the progenitors +of the existing Lemuridæ; and these in their turn from forms standing +very low in the mammalian series. + +The Marsupials stand in many important characters below the placental +mammals. They appeared at an earlier geological period, and their range +was formerly much more extensive than what it now is. Hence the +Placentata are generally supposed to have been derived from the +Implacentata or Marsupials; not, however, from forms closely like the +existing Marsupials, but from their early progenitors. The Monotremata +are plainly allied to the Marsupials; forming a third and still lower +division in the great mammalian series. They are represented at the +present day solely by the Ornithorhynchus and Echidna; and these two +forms may be safely considered as relics of a much larger group which +have been preserved in Australia through some favourable concurrence of +circumstances. The Monotremata are eminently interesting, as in several +important points of structure they lead towards the class of reptiles. + +In attempting to trace the genealogy of the Mammalia, and therefore of +man, lower down in the series, we become involved in greater and greater +obscurity. He who wishes to see what ingenuity and knowledge can effect, +may consult Prof. Häckel’s works.[273] I will content myself with a few +general remarks. Every evolutionist will admit that the five great +vertebrate classes, namely, mammals, birds, reptiles, amphibians, and +fishes, are all descended from some one prototype; for they have much in +common, especially during their embryonic state. As the class of fishes +is the most lowly organised and appeared before the others, we may +conclude that all the members of the vertebrate kingdom are derived from +some fish-like animal, less highly organised than any as yet found in +the lowest known formations. The belief that animals so distinct as a +monkey or elephant and a humming-bird, a snake, frog, and fish, &c., +could all have sprung from the same parents, will appear monstrous to +those who have not attended to the recent progress of natural history. +For this belief implies the former existence of links closely binding +together all these forms, now so utterly unlike. + +Nevertheless it is certain that groups of animals have existed, or do +now exist, which serve to connect more or less closely the several great +vertebrate classes. We have seen that the Ornithorhynchus graduates +towards reptiles; and Prof. Huxley has made the remarkable discovery, +confirmed by Mr. Cope and others, that the old Dinosaurians are +intermediate in many important respects between certain reptiles and +certain birds—the latter consisting of the ostrich-tribe (itself +evidently a widely-diffused remnant of a larger group) and of the +Archeopteryx, that strange Secondary bird having a long tail like that +of the lizard. Again, according to Prof. Owen,[274] the +Ichthyosaurians—great sea-lizards furnished with paddles—present many +affinities with fishes, or rather, according to Huxley, with amphibians. +This latter class (including in its highest division frogs and toads) is +plainly allied to the Ganoid fishes. These latter fishes swarmed during +the earlier geological periods, and were constructed on what is called a +highly generalised type, that is they presented diversified affinities +with other groups of organisms. The amphibians and fishes are also so +closely united by the Lepidosiren, that naturalists long disputed in +which of these two classes it ought to be placed. The Lepidosiren and +some few Ganoid fishes have been preserved from utter extinction by +inhabiting our rivers, which are harbours of refuge, bearing the same +relation to the great waters of the ocean that islands bear to +continents. + +Lastly, one single member of the immense and diversified class of +fishes, namely the lancelet or amphioxus, is so different from all other +fishes, that Häckel maintains that it ought to form a distinct class in +the vertebrate kingdom. This fish is remarkable for its negative +characters; it can hardly be said to possess a brain, vertebral column, +or heart, &c.; so that it was classed by the older naturalists amongst +the worms. Many years ago Prof. Goodsir perceived that the lancelet +presented some affinities with the Ascidians, which are invertebrate, +hermaphrodite, marine creatures permanently attached to a support. They +hardly appear like animals, and consist of a simple, tough, leathery +sack, with two small projecting orifices. They belong to the Molluscoida +of Huxley—a lower division of the great kingdom of the Mollusca; but +they have recently been placed by some naturalists amongst the Vermes or +worms. Their larvæ somewhat resemble tadpoles in shape,[275] and have +the power of swimming freely about. Some observations lately made by M. +Kowalevsky,[276] since confirmed by Prof. Kuppfer, will form a discovery +of extraordinary interest, if still further extended, as I hear from M. +Kowalevsky in Naples he has now effected. The discovery is that the +larvæ of Ascidians are related to the Vertebrata, in their manner of +development, in the relative position of the nervous system, and in +possessing a structure closely like the _chorda dorsalis_ of vertebrate +animals. It thus appears, if we may rely on embryology, which has always +proved the safest guide in classification, that we have at last gained a +clue to the source whence the Vertebrata have been derived. We should +thus be justified in believing that at an extremely remote period a +group of animals existed, resembling in many respects the larvæ of our +present Ascidians, which diverged into two great branches—the one +retrograding in development and producing the present class of +Ascidians, the other rising to the crown and summit of the animal +kingdom by giving birth to the Vertebrata. + + +We have thus far endeavoured rudely to trace the genealogy of the +Vertebrata by the aid of their mutual affinities. We will now look to +man as he exists; and we shall, I think, be able partially to restore +during successive periods, but not in due order of time, the structure +of our early progenitors. This can be effected by means of the rudiments +which man still retains, by the characters which occasionally make their +appearance in him through reversion, and by the aid of the principles of +morphology and embryology. The various facts, to which I shall here +allude, have been given in the previous chapters. The early progenitors +of man were no doubt once covered with hair, both sexes having beards; +their ears were pointed and capable of movement; and their bodies were +provided with a tail, having the proper muscles. Their limbs and bodies +were also acted on by many muscles which now only occasionally reappear, +but are normally present in the Quadrumana. The great artery and nerve +of the humerus ran through a supra-condyloid foramen. At this or some +earlier period, the intestine gave forth a much larger diverticulum or +cæcum than that now existing. The foot, judging from the condition of +the great toe in the fœtus, was then prehensile; and our progenitors, +no doubt, were arboreal in their habits, frequenting some warm, +forest-clad land. The males were provided with great canine teeth, +which served them as formidable weapons. + +At a much earlier period the uterus was double; the excreta were voided +through a cloaca; and the eye was protected by a third eyelid or +nictitating membrane. At a still earlier period the progenitors of man +must have been aquatic in their habits; for morphology plainly tells us +that our lungs consist of a modified swim-bladder, which once served as +a float. The clefts on the neck in the embryo of man show where the +branchiæ once existed. At about this period the true kidneys were +replaced by the corpora Wolffiana. The heart existed as a simple +pulsating vessel; and the chorda dorsalis took the place of a vertebral +column. These early predecessors of man, thus seen in the dim recesses +of time, must have been as lowly organised as the lancelet or amphioxus, +or even still more lowly organised. + +There is one other point deserving a fuller notice. It has long been +known that in the vertebrate kingdom one sex bears rudiments of various +accessory parts, appertaining to the reproductive system, which properly +belong to the opposite sex; and it has now been ascertained that at a +very early embryonic period both sexes possess true male and female +glands. Hence some extremely remote progenitor of the whole vertebrate +kingdom appears to have been hermaphrodite or androgynous.[277] But here +we encounter a singular difficulty. In the mammalian class the males +possess in their vesiculæ prostaticæ rudiments of a uterus with the +adjacent passage; they bear also rudiments of mammæ, and some male +marsupials have rudiments of a marsupial sack.[278] Other analogous +facts could be added. Are we, then, to suppose that some extremely +ancient mammal possessed organs proper to both sexes, that is, continued +androgynous after it had acquired the chief distinctions of its proper +class, and therefore after it had diverged from the lower classes of the +vertebrate kingdom? This seems improbable in the highest degree; for had +this been the case, we might have expected that some few members of the +two lower classes, namely fishes[279] and amphibians, would still have +remained androgynous. We must, on the contrary, believe that when the +five vertebrate classes diverged from their common progenitor the sexes +had already become separated. To account, however, for male mammals +possessing rudiments of the accessory female organs, and for female +mammals possessing rudiments of the masculine organs, we need not +suppose that their early progenitors were still androgynous after they +had assumed their chief mammalian characters. It is quite possible that +as the one sex gradually acquired the accessory organs proper to it, +some of the successive steps or modifications were transmitted to the +opposite sex. When we treat of sexual selection, we shall meet with +innumerable instances of this form of transmission,—as in the case of +the spurs, plumes, and brilliant colours, acquired by male birds for +battle or ornament, and transferred to the females in an imperfect or +rudimentary condition. + +The possession by male mammals of functionally imperfect mammary organs +is, in some respects, especially curious. The Monotremata have the +proper milk-secreting glands with orifices, but no nipples; and as these +animals stand at the very base of the mammalian series, it is probable +that the progenitors of the class possessed, in like manner, the +milk-secreting glands, but no nipples. This conclusion is supported by +what is known of their manner of development; for Professor Turner +informs me, on the authority of Kölliker and Lauger, that in the embryo +the mammary glands can be distinctly traced before the nipples are in +the least visible; and it should be borne in mind that the development +of successive parts in the individual generally seems to represent and +accord with the development of successive beings in the same line of +descent. The Marsupials differ from the Monotremata by possessing +nipples; so that these organs were probably first acquired by the +Marsupials after they had diverged from, and risen above, the +Monotremata, and were then transmitted to the placental mammals. No one +will suppose that after the Marsupials had approximately acquired their +present structure, and therefore at a rather late period in the +development of the mammalian series, any of its members still remained +androgynous. We seem, therefore, compelled to recur to the foregoing +view, and to conclude that the nipples were first developed in the +females of some very early marsupial form, and were then, in accordance +with a common law of inheritance, transferred in a functionally +imperfect condition to the males. + +Nevertheless a suspicion has sometimes crossed my mind that long after +the progenitors of the whole mammalian class had ceased to be +androgynous, both sexes might have yielded milk and thus nourished their +young; and in the case of the Marsupials, that both sexes might have +carried their young in marsupial sacks. This will not appear utterly +incredible, if we reflect that the males of syngnathous fishes receive +the eggs of the females in their abdominal pouches, hatch them, and +afterwards, as some believe, nourish the young;[280]—that certain other +male fishes hatch the eggs within their mouths or branchial +cavities;—that certain male toads take the chaplets of eggs from the +females and wind them round their own thighs, keeping them there until +the tadpoles are born;—that certain male birds undertake the whole duty +of incubation, and that male pigeons, as well as the females, feed their +nestlings with a secretion from their crops. But the above suspicion +first occurred to me from the mammary glands in male mammals being +developed so much more perfectly than the rudiments of those other +accessory reproductive parts, which are found in the one sex though +proper to the other. The mammary glands and nipples, as they exist in +male mammals, can indeed hardly be called rudimentary; they are simply +not fully developed and not functionally active. They are +sympathetically affected under the influence of certain diseases, like +the same organs in the female. At birth they often secrete a few drops +of milk; and they have been known occasionally in man and other mammals +to become well developed, and to yield a fair supply of milk. Now if we +suppose that during a former prolonged period male mammals aided the +females in nursing their offspring, and that afterwards from some cause, +as from a smaller number of young being produced, the males ceased +giving this aid, disuse of the organs during maturity would lead to +their becoming inactive; and from two well-known principles of +inheritance this state of inactivity would probably be transmitted to +the males at the corresponding age of maturity. But at all earlier ages +these organs would be left unaffected, so that they would be equally +well developed in the young of both sexes. + + +_Conclusion._—The best definition of advancement or progress in the +organic scale ever given, is that by Von Baer; and this rests on the +amount of differentiation and specialisation of the several parts of the +same being, when arrived, as I should be inclined to add, at maturity. +Now as organisms have become slowly adapted by means of natural +selection for diversified lines of life, their parts will have become, +from the advantage gained by the division of physiological labour, more +and more differentiated and specialised for various functions. The same +part appears often to have been modified first for one purpose, and then +long afterwards for some other and quite distinct purpose; and thus all +the parts are rendered more and more complex. But each organism will +still retain the general type of structure of the progenitor from which +it was aboriginally derived. In accordance with this view it seems, if +we turn to geological evidence, that organisation on the whole has +advanced throughout the world by slow and interrupted steps. In the +great kingdom of the Vertebrata it has culminated in man. It must not, +however, be supposed that groups of organic beings are always supplanted +and disappear as soon as they have given birth to other and more perfect +groups. The latter, though victorious over their predecessors, may not +have become better adapted for all places in the economy of nature. Some +old forms appear to have survived from inhabiting protected sites, where +they have not been exposed to very severe competition; and these often +aid us in constructing our genealogies, by giving us a fair idea of +former and lost populations. But we must not fall into the error of +looking at the existing members of any lowly-organised group as perfect +representatives of their ancient predecessors. + +The most ancient progenitors in the kingdom of the Vertebrata, at which +we are able to obtain an obscure glance, apparently consisted of a group +of marine animals,[281] resembling the larvæ of existing Ascidians. +These animals probably gave rise to a group of fishes, as lowly +organised as the lancelet; and from these the Ganoids, and other fishes +like the Lepidosiren, must have been developed. From such fish a very +small advance would carry us on to the amphibians. We have seen that +birds and reptiles were once intimately connected together; and the +Monotremata now, in a slight degree, connect mammals with reptiles. But +no one can at present say by what line of descent the three higher and +related classes, namely, mammals, birds, and reptiles, were derived from +either of the two lower vertebrate classes, namely amphibians and +fishes. In the class of mammals the steps are not difficult to conceive +which led from the ancient Monotremata to the ancient Marsupials; and +from these to the early progenitors of the placental mammals. We may +thus ascend to the Lemuridæ; and the interval is not wide from these to +the Simiadæ. The Simiadæ then branched off into two great stems, the New +World and Old World monkeys; and from the latter, at a remote period, +Man, the wonder and glory of the Universe, proceeded. + +Thus we have given to man a pedigree of prodigious length, but not, it +may be said, of noble quality. The world, it has often been remarked, +appears as if it had long been preparing for the advent of man; and +this, in one sense is strictly true, for he owes his birth to a long +line of progenitors. If any single link in this chain had never existed, +man would not have been exactly what he now is. Unless we wilfully close +our eyes, we may, with our present knowledge, approximately recognise +our parentage; nor need we feel ashamed of it. The most humble organism +is something much higher than the inorganic dust under our feet; and no +one with an unbiassed mind can study any living creature, however +humble, without being struck with enthusiasm at its marvellous structure +and properties. + + + + +CHAPTER VII. + +ON THE RACES OF MAN. + + + The nature and value of specific characters—Application to the + races of man—Arguments in favour of, and opposed to, ranking + the so-called races of man as distinct + species—Sub-species—Monogenists and polygenists—Convergence + of character—Numerous points of resemblance in body and mind + between the most distinct races of man—The state of man when + he first spread over the earth—Each race not descended from a + single pair—The extinction of races—The formation of + races—The effects of crossing—Slight influence of the direct + action of the conditions of life—Slight or no influence of + natural selection—Sexual selection. + + +It is not my intention here to describe the several so-called races of +men; but to inquire what is the value of the differences between them +under a classificatory point of view, and how they have originated. In +determining whether two or more allied forms ought to be ranked as +species or varieties, naturalists are practically guided by the +following considerations; namely, the amount of difference between them, +and whether such differences relate to few or many points of structure, +and whether they are of physiological importance; but more especially +whether they are constant. Constancy of character is what is chiefly +valued and sought for by naturalists. Whenever it can be shewn, or +rendered probable, that the forms in question have remained distinct for +a long period, this becomes an argument of much weight in favour of +treating them as species. Even a slight degree of sterility between any +two forms when first crossed, or in their offspring, is generally +considered as a decisive test of their specific distinctness; and their +continued persistence without blending within the same area, is usually +accepted as sufficient evidence, either of some degree of mutual +sterility, or in the case of animals of some repugnance to mutual +pairing. + +Independently of blending from intercrossing, the complete absence, in a +well-investigated region, of varieties linking together any two +closely-allied forms, is probably the most important of all the +criterions of their specific distinctness; and this is a somewhat +different consideration from mere constancy of character, for two forms +may be highly variable and yet not yield intermediate varieties. +Geographical distribution is often unconsciously and sometimes +consciously brought into play; so that forms living in two widely +separated areas, in which most of the other inhabitants are specifically +distinct, are themselves usually looked at as distinct; but in truth +this affords no aid in distinguishing geographical races from so-called +good or true species. + +Now let us apply these generally-admitted principles to the races of +man, viewing him in the same spirit as a naturalist would any other +animal. In regard to the amount of difference between the races, we must +make some allowance for our nice powers of discrimination gained by the +long habit of observing ourselves. In India, as Elphinstone +remarks,[282] although a newly-arrived European cannot at first +distinguish the various native races, yet they soon appear to him +extremely dissimilar; and the Hindoo cannot at first perceive any +difference between the several European nations. Even the most distinct +races of man, with the exception of certain negro tribes, are much more +like each other in form than would at first be supposed. This is well +shewn by the French photographs in the Collection Anthropologique du +Muséum of the men belonging to various races, the greater number of +which, as many persons to whom I have shown them have remarked, might +pass for Europeans. Nevertheless, these men if seen alive would +undoubtedly appear very distinct, so that we are clearly much influenced +in our judgment by the mere colour of the skin and hair, by slight +differences in the features, and by expression. + +There is, however, no doubt that the various races, when carefully +compared and measured, differ much from each other,—as in the texture +of the hair, the relative proportions of all parts of the body,[283] the +capacity of the lungs, the form and capacity of the skull, and even in +the convolutions of the brain.[284] But it would be an endless task to +specify the numerous points of structural difference. The races differ +also in constitution, in acclimatisation, and in liability to certain +diseases. Their mental characteristics are likewise very distinct; +chiefly as it would appear in their emotional, but partly in their +intellectual, faculties. Every one who has had the opportunity of +comparison, must have been struck with the contrast between the +taciturn, even morose, aborigines of S. America and the light-hearted, +talkative negroes. There is a nearly similar contrast between the Malays +and the Papuans,[285] who live under the same physical conditions, and +are separated from each other only by a narrow space of sea. + +We will first consider the arguments which may be advanced in favour of +classing the races of man as distinct species, and then those on the +other side. If a naturalist, who had never before seen such beings, were +to compare a Negro, Hottentot, Australian, or Mongolian, he would at +once perceive that they differed in a multitude of characters, some of +slight and some of considerable importance. On inquiry he would find +that they were adapted to live under widely different climates, and that +they differed somewhat in bodily constitution and mental disposition. If +he were then told that hundreds of similar specimens could be brought +from the same countries, he would assuredly declare that they were as +good species as many to which he had been in the habit of affixing +specific names. This conclusion would be greatly strengthened as soon as +he had ascertained that these forms had all retained the same character +for many centuries; and that negroes, apparently identical with existing +negroes, had lived at least 4000 years ago.[286] He would also hear from +an excellent observer, Dr. Lund,[287] that the human skulls found in +the caves of Brazil, entombed with many extinct mammals, belonged to the +same type as that now prevailing throughout the American Continent. + +Our naturalist would then perhaps turn to geographical distribution, and +he would probably declare that forms differing not only in appearance, +but fitted for the hottest and dampest or driest countries, as well as +for the arctic regions, must be distinct species. He might appeal to the +fact that no one species in the group next to man, namely the +Quadrumana, can resist a low temperature or any considerable change of +climate; and that those species which come nearest to man have never +been reared to maturity, even under the temperate climate of Europe. He +would be deeply impressed with the fact, first noticed by Agassiz,[288] +that the different races of man are distributed over the world in the +same zoological provinces, as those inhabited by undoubtedly distinct +species and genera of mammals. This is manifestly the case with the +Australian, Mongolian, and Negro races of man; in a less well-marked +manner with the Hottentots; but plainly with the Papuans and Malays, who +are separated, as Mr. Wallace has shewn, by nearly the same line which +divides the great Malayan and Australian zoological provinces. The +aborigines of America range throughout the Continent; and this at first +appears opposed to the above rule, for most of the productions of the +Southern and Northern halves differ widely; yet some few living forms, +as the opossum, range from the one into the other, as did formerly some +of the gigantic Edentata. The Esquimaux, like other Arctic animals, +extend round the whole polar regions. It should be observed that the +mammalian forms which inhabit the several zoological provinces, do not +differ from each other in the same degree; so that it can hardly be +considered as an anomaly that the Negro differs more, and the American +much less, from the other races of man than do the mammals of the same +continents from those of the other provinces. Man, it may be added, does +not appear to have aboriginally inhabited any oceanic island; and in +this respect he resembles the other members of his class. + +In determining whether the varieties of the same kind of domestic animal +should be ranked as specifically distinct, that is, whether any of them +are descended from distinct wild species, every naturalist would lay +much stress on the fact, if established, of their external parasites +being specifically distinct. All the more stress would be laid on this +fact, as it would be an exceptional one, for I am informed by Mr. Denny +that the most different kinds of dogs, fowls, and pigeons, in England, +are infested by the same species of Pediculi or lice. Now Mr. A. Murray +has carefully examined the Pediculi collected in different countries +from the different races of man;[289] and he finds that they differ, not +only in colour, but in the structure of their claws and limbs. In every +case in which numerous specimens were obtained the differences were +constant. The surgeon of a whaling ship in the Pacific assured me that +when the Pediculi, with which some Sandwich Islanders on board swarmed, +strayed on to the bodies of the English sailors, they died in the course +of three or four days. These Pediculi were darker coloured and appeared +different from those proper to the natives of Chiloe in South America, +of which he gave me specimens. These, again, appeared larger and much +softer than European lice. Mr. Murray procured four kinds from Africa, +namely from the Negroes of the Eastern and Western coasts, from the +Hottentots and Caffres; two kinds from the natives of Australia; two +from North, and two from South America. In these latter cases it may be +presumed that the Pediculi came from natives inhabiting different +districts. With insects slight structural differences, if constant, are +generally esteemed of specific value: and the fact of the races of man +being infested by parasites, which appear to be specifically distinct, +might fairly be urged as an argument that the races themselves ought to +be classed as distinct species. + +Our supposed naturalist having proceeded thus far in his investigation, +would next inquire whether the races of men, when crossed, were in any +degree sterile. He might consult the work[290] of a cautious and +philosophical observer, Professor Broca; and in this he would find good, +evidence that some races were quite fertile together; but evidence of an +opposite nature in regard to other races. Thus it has been asserted that +the native women of Australia and Tasmania rarely produce children to +European men; the evidence, however, on this head has now been shewn to +be almost valueless. The half-castes are killed by the pure blacks; and +an account has lately been published of eleven half-caste youths +murdered and burnt at the same time, whose remains were found by the +police.[291] Again, it has often been said that when mulattoes +intermarry they produce few children; on the other hand, Dr. Bachman of +Charlestown[292] positively asserts that he has known mulatto families +which have intermarried for several generations, and have continued on +an average as fertile as either pure whites or pure blacks. Inquiries +formerly made by Sir C. Lyell on this subject led him, as he informs me, +to the same conclusion. In the United States the census for the year +1854 included, according to Dr. Bachman, 405,751 mulattoes; and this +number, considering all the circumstances of the case, seems small; but +it may partly be accounted for by the degraded and anomalous position of +the class, and by the profligacy of the women. A certain amount of +absorption of mulattoes into negroes must always be in progress; and +this would lead to an apparent diminution of the former. The inferior +vitality of mulattoes is spoken of in a trustworthy work[293] as a +well-known phenomenon; but this is a different consideration from their +lessened fertility; and can hardly be advanced as a proof of the +specific distinctness of the parent races. No doubt both animal and +vegetable hybrids, when produced from extremely distinct species, are +liable to premature death; but the parents of mulattoes cannot be put +under the category of extremely distinct species. The common Mule, so +notorious for long life and vigour, and yet so sterile, shews how little +necessary connection there is in hybrids between lessened fertility and +vitality: other analogous cases could be added. + +Even if it should hereafter be proved that all the races of men were +perfectly fertile together, he who was inclined from other reasons to +rank them as distinct species, might with justice argue that fertility +and sterility are not safe criterions of specific distinctness. We know +that these qualities are easily affected by changed conditions of life +or by close inter-breeding, and that they are governed by highly complex +laws, for instance that of the unequal fertility of reciprocal crosses +between the same two species. With forms which must be ranked as +undoubted species, a perfect series exists from those which are +absolutely sterile when crossed, to those which are almost or quite +fertile. The degrees of sterility do not coincide strictly with the +degrees of difference in external structure or habits of life. Man in +many respects may be compared with those animals which have long been +domesticated, and a large body of evidence can be advanced in favour of +the Pallasian doctrine[294] that domestication tends to eliminate the +sterility which is so general a result of the crossing of species in a +state of nature. From these several considerations, it may be justly +urged that the perfect fertility of the intercrossed races of man, if +established, would not absolutely preclude us from ranking them as +distinct species. + +Independently of fertility, the character of the offspring from a cross +has sometimes been thought to afford evidence whether the parent-forms +ought to be ranked as species or varieties; but after carefully studying +the evidence, I have come to the conclusion that no general rules of +this kind can be trusted. Thus with mankind the offspring of distinct +races resemble in all respects the offspring of true species and of +varieties. This is shewn, for instance, by the manner in which the +characters of both parents are blended, and by one form absorbing +another through repeated crosses. In this latter case the progeny both +of crossed species and varieties retain for a long period a tendency to +revert to their ancestors, especially to that one which is prepotent in +transmission. When any character has suddenly appeared in a race or +species as the result of a single act of variation, as is general with +monstrosities,[295] and this race is crossed with another not thus +characterised, the characters in question do not commonly appear in a +blended condition in the young, but are transmitted to them either +perfectly developed or not at all. As with the crossed races of man +cases of this kind rarely or never occur, this may be used as an +argument against the view suggested by some ethnologists, namely that +certain characters, for instance the blackness of the negro, first +appeared as a sudden variation or sport. Had this occurred, it is +probable that mulattoes would often have been born, either completely +black or completely white. + +We have now seen that a naturalist might feel himself fully justified in +ranking the races of man as distinct species; for he has found that they +are distinguished by many differences in structure and constitution, +some being of importance. These differences have, also, remained nearly +constant for very long periods of time. He will have been in some degree +influenced by the enormous range of man, which is a great anomaly in the +class of mammals, if mankind be viewed as a single species. He will have +been struck with the distribution of the several so-called races, in +accordance with that of other undoubtedly distinct species of mammals. +Finally he might urge that the mutual fertility of all the races has not +as yet been fully proved; and even if proved would not be an absolute +proof of their specific identity. + + +On the other side of the question, if our supposed naturalist were to +enquire whether the forms of man kept distinct like ordinary species, +when mingled together in large numbers in the same country, he would +immediately discover that this was by no means the case. In Brazil he +would behold an immense mongrel population of Negroes and Portuguese; in +Chiloe and other parts of South America, he would behold the whole +population consisting of Indians and Spaniards blended in various +degrees.[296] In many parts of the same continent he would meet with the +most complex crosses between Negroes, Indians, and Europeans; and such +triple crosses afford the severest test, judging from the vegetable +kingdom, of the mutual fertility of the parent-forms. In one island of +the Pacific he would find a small population of mingled Polynesian and +English blood; and in the Viti Archipelago a population of Polynesians +and Negritos crossed in all degrees. Many analogous cases could be +added, for instance, in South Africa. Hence the races of man are not +sufficiently distinct to co-exist without fusion; and this it is, which +in all ordinary cases affords the usual test of specific distinctness. + +Our naturalist would likewise be much disturbed as soon as he perceived +that the distinctive characters of every race of man were highly +variable. This strikes every one when he first beholds the negro-slaves +in Brazil, who have been imported from all parts of Africa. The same +remark holds good with the Polynesians, and with many other races. It +may be doubted whether any character can be named which is distinctive +of a race and is constant. Savages, even within the limits of the same +tribe, are not nearly so uniform in character, as has often been said. +Hottentot women offer certain peculiarities, more strongly marked than +those occurring in any other race, but these are known not to be of +constant occurrence. In the several American tribes, colour and +hairyness differ considerably; as does colour to a certain degree, and +the shape of the features greatly, in the Negroes of Africa. The shape +of the skull varies much in some races;[297] and so it is with every +other character. Now all naturalists have learnt by dearly-bought +experience, how rash it is to attempt to define species by the aid of +inconstant characters. + +But the most weighty of all the arguments against treating the races of +man as distinct species, is that they graduate into each other, +independently in many cases, as far as we can judge, of their having +intercrossed. Man has been studied more carefully than any other organic +being, and yet there is the greatest possible diversity amongst capable +judges whether he should be classed as a single species or race, or as +two (Virey), as three (Jacquinot), as four (Kant), five (Blumenbach), +six (Buffon), seven (Hunter), eight (Agassiz), eleven (Pickering), +fifteen (Bory St. Vincent), sixteen (Desmoulins), twenty-two (Morton), +sixty (Crawfurd), or as sixty-three, according to Burke.[298] This +diversity of judgment does not prove that the races ought not to be +ranked as species, but it shews that they graduate into each other, and +that it is hardly possible to discover clear distinctive characters +between them. + +Every naturalist who has had the misfortune to undertake the +description of a group of highly varying organisms, has encountered +cases (I speak after experience) precisely like that of man; and if of a +cautious disposition, he will end by uniting all the forms which +graduate into each other as a single species; for he will say to himself +that he has no right to give names to objects which he cannot define. +Cases of this kind occur in the Order which includes man, namely in +certain genera of monkeys; whilst in other genera, as in Cercopithecus, +most of the species can be determined with certainty. In the American +genus Cebus, the various forms are ranked by some naturalists as +species, by others as mere geographical races. Now if numerous specimens +of Cebus were collected from all parts of South America, and those forms +which at present appear to be specifically distinct, were found to +graduate into each other by close steps, they would be ranked by most +naturalists as mere varieties or races; and thus the greater number of +naturalists have acted with respect to the races of man. Nevertheless it +must be confessed that there are forms, at least in the vegetable +kingdom,[299] which we cannot avoid naming as species, but which are +connected together, independently of intercrossing, by numberless +gradations. + +Some naturalists have lately employed the term “sub-species” to +designate forms which possess many of the characteristics of true +species, but which hardly deserve so high a rank. Now if we reflect on +the weighty arguments, above given, for raising the races of man to the +dignity of species, and the insuperable difficulties on the other side +in defining them, the term “sub-species” might here be used with much +propriety. But from long habit the term “race” will perhaps always be +employed. The choice of terms is only so far important as it is highly +desirable to use, as far as that may be possible, the same terms for the +same degrees of difference. Unfortunately this is rarely possible; for +within the same family the larger genera generally include +closely-allied forms, which can be distinguished only with much +difficulty, whilst the smaller genera include forms that are perfectly +distinct; yet all must equally be ranked as species. So again the +species within the same large genus by no means resemble each other to +the same degree: on the contrary, in most cases some of them can be +arranged in little groups round other species, like satellites round +planets.[300] + + +The question whether mankind consists of one or several species has of +late years been much agitated by anthropologists, who are divided into +two schools of monogenists and polygenists. Those who do not admit the +principle of evolution, must look at species either as separate +creations or as in some manner distinct entities; and they must decide +what forms to rank as species by the analogy of other organic beings +which are commonly thus received. But it is a hopeless endeavour to +decide this point on sound grounds, until some definition of the term +“species” is generally accepted; and the definition must not include an +element which cannot possibly be ascertained, such as an act of +creation. We might as well attempt without any definition to decide +whether a certain number of houses should be called a village, or town, +or city. We have a practical illustration of the difficulty in the +never-ending doubts whether many closely-allied mammals, birds, +insects, and plants, which represent each other in North America and +Europe, should be ranked species or geographical races; and so it is +with the productions of many islands situated at some little distance +from the nearest continent. + +Those naturalists, on the other hand, who admit the principle of +evolution, and this is now admitted by the greater number of rising men, +will feel no doubt that all the races of man are descended from a single +primitive stock; whether or not they think fit to designate them as +distinct species, for the sake of expressing their amount of +difference.[301] With our domestic animals the question whether the +various races have arisen from one or more species is different. +Although all such races, as well as all the natural species within the +same genus, have undoubtedly sprung from the same primitive stock, yet +it is a fit subject for discussion, whether, for instance, all the +domestic races of the dog have acquired their present differences since +some one species was first domesticated and bred by man; or whether they +owe some of their characters to inheritance from distinct species, which +had already been modified in a state of nature. With mankind no such +question can arise, for he cannot be said to have been domesticated at +any particular period. + +When the races of man diverged at an extremely remote epoch from their +common progenitor, they will have differed but little from each other, +and been few in number; consequently they will then, as far as their +distinguishing characters are concerned, have had less claim to rank as +distinct species, than the existing so-called races. Nevertheless such +early races would perhaps have been ranked by some naturalists as +distinct species, so arbitrary is the term, if their differences, +although extremely slight, had been more constant than at present, and +had not graduated into each other. + +It is, however, possible, though far from probable, that the early +progenitors of man might at first have diverged much in character, until +they became more unlike each other than are any existing races; but that +subsequently, as suggested by Vogt,[302] they converged in character. +When man selects for the same object the offspring of two distinct +species, he sometimes induces, as far as general appearance is +concerned, a considerable amount of convergence. This is the case, as +shewn by Von Nathusius,[303] with the improved breeds of pigs, which are +descended from two distinct species; and in a less well-marked manner +with the improved breeds of cattle. A great anatomist, Gratiolet, +maintains that the anthropomorphous apes do not form a natural +sub-group; but that the orang is a highly developed gibbon or +semnopithecus; the chimpanzee a highly developed macacus; and the +gorilla a highly developed mandrill. If this conclusion, which rests +almost exclusively on brain-characters, be admitted, we should have a +case of convergence at least in external characters, for the +anthropomorphous apes are certainly more like each other in many points +than they are to other apes. All analogical resemblances, as of a whale +to a fish, may indeed be said to be cases of convergence; but this term +has never been applied to superficial and adaptive resemblances. It +would be extremely rash in most cases to attribute to convergence close +similarity in many points of structure in beings which had once been +widely different. The form of a crystal is determined solely by the +molecular forces, and it is not surprising that dissimilar substances +should sometimes assume the same form; but with organic beings we should +bear in mind that the form of each depends on an infinitude of complex +relations, namely on the variations which have arisen, these being due +to causes far too intricate to be followed out,—on the nature of the +variations which have been preserved, and this depends on the +surrounding physical conditions, and in a still higher degree on the +surrounding organisms with which each has come into competition,—and +lastly, on inheritance (in itself a fluctuating element) from +innumerable progenitors, all of which have had their forms determined +through equally complex relations. It appears utterly incredible that +two organisms, if differing in a marked manner, should ever afterwards +converge so closely as to lead to a near approach to identity throughout +their whole organisation. In the case of the convergent pigs above +referred to, evidence of their descent from two primitive stocks is +still plainly retained, according to Von Nathusius, in certain bones of +their skulls. If the races of man were descended, as supposed by some +naturalists, from two or more distinct species, which had differed as +much, or nearly as much, from each other, as the orang differs from the +gorilla, it can hardly be doubted that marked differences in the +structure of certain bones would still have been discoverable in man as +he now exists. + + +Although the existing races of man differ in many respects, as in +colour, hair, shape of skull, proportions of the body, &c., yet if their +whole organisation be taken into consideration they are found to +resemble each other closely in a multitude of points. Many of these +points are of so unimportant or of so singular a nature, that it is +extremely improbable that they should have been independently acquired +by aboriginally distinct species or races. The same remark holds good +with equal or greater force with respect to the numerous points of +mental similarity between the most distinct races of man. The American +aborigines, Negroes and Europeans differ as much from each other in mind +as any three races that can be named; yet I was incessantly struck, +whilst living with the Fuegians on board the “Beagle,” with the many +little traits of character, shewing how similar their minds were to +ours; and so it was with a full-blooded negro with whom I happened once +to be intimate. + +He who will carefully read Mr. Tylor’s and Sir J. Lubbock’s interesting +works[304] can hardly fail to be deeply impressed with the close +similarity between the men of all races in tastes, dispositions and +habits. This is shewn by the pleasure which they all take in dancing, +rude music, acting, painting, tattooing, and otherwise decorating +themselves,—in their mutual comprehension of gesture-language—and, as +I shall be able to shew in a future essay, by the same expression in +their features, and by the same inarticulate cries, when they are +excited by various emotions. This similarity, or rather identity, is +striking, when contrasted with the different expressions which may be +observed in distinct species of monkeys. There is good evidence that the +art of shooting with bows and arrows has not been handed down from any +common progenitor of mankind, yet the stone arrow-heads, brought from +the most distant parts of the world and manufactured at the most remote +periods, are, as Nilsson has shewn,[305] almost identical; and this fact +can only be accounted for by the various races having similar inventive +or mental powers. The same observation has been made by +archæologists[306] with respect to certain widely-prevalent ornaments, +such as zigzags, &c.; and with respect to various simple beliefs and +customs, such as the burying of the dead under megalithic structures. I +remember observing in South America,[307] that there, as in so many +other parts of the world, man has generally chosen the summits of lofty +hills, on which to throw up piles of stones, either for the sake of +recording some remarkable event, or for burying his dead. + +Now when naturalists observe a close agreement in numerous small details +of habits, tastes and dispositions between two or more domestic races, +or between nearly-allied natural forms, they use this fact as an +argument that all are descended from a common progenitor who was thus +endowed; and consequently that all should be classed under the same +species. The same argument may be applied with much force to the races +of man. + +As it is improbable that the numerous and unimportant points of +resemblance between the several races of man in bodily structure and +mental faculties (I do not here refer to similar customs) should all +have been independently acquired, they must have been inherited from +progenitors who were thus characterised. We thus gain some insight into +the early state of man, before he had spread step by step over the face +of the earth. The spreading of man to regions widely separated by the +sea, no doubt, preceded any considerable amount of divergence of +character in the several races; for otherwise we should sometimes meet +with the same race in distinct continents; and this is never the case. +Sir J. Lubbock, after comparing the arts now practised by savages in all +parts of the world, specifies those which man could not have known, when +he first wandered from his original birthplace; for if once learnt they +would never have been forgotten.[308] He thus shews that “the spear, +which is but a development of the knife-point, and the club, which is +but a long hammer, are the only things left.” He admits, however, that +the art of making fire probably had already been discovered, for it is +common to all the races now existing, and was known to the ancient +cave-inhabitants of Europe. Perhaps the art of making rude canoes or +rafts was likewise known; but as man existed at a remote epoch, when the +land in many places stood at a very different level, he would have been +able, without the aid of canoes, to have spread widely. Sir J. Lubbock +further remarks how improbable it is that our earliest ancestors could +have “counted as high as ten, considering that so many races now in +existence cannot get beyond four.” Nevertheless, at this early period, +the intellectual and social faculties of man could hardly have been +inferior in any extreme degree to those now possessed by the lowest +savages; otherwise primeval man could not have been so eminently +successful in the struggle for life, as proved by his early and wide +diffusion. + +From the fundamental differences between certain languages, some +philologists have inferred that when man first became widely diffused he +was not a speaking animal; but it may be suspected that languages, far +less perfect than any now spoken, aided by gestures, might have been +used, and yet have left no traces on subsequent and more +highly-developed tongues. Without the use of some language, however +imperfect, it appears doubtful whether man’s intellect could have risen +to the standard implied by his dominant position at an early period. + +Whether primeval man, when he possessed very few arts of the rudest +kind, and when his power of language was extremely imperfect, would have +deserved to be called man, must depend on the definition which we +employ. In a series of forms graduating insensibly from some ape-like +creature to man as he now exists, it would be impossible to fix on any +definite point when the term “man” ought to be used. But this is a +matter of very little importance. So again it is almost a matter of +indifference whether the so-called races of man are thus designated, or +are ranked as species or sub-species; but the latter term appears the +most appropriate. Finally, we may conclude that when the principles of +evolution are generally accepted, as they surely will be before long, +the dispute between the monogenists and the polygenists will die a +silent and unobserved death. + + +One other question ought not to be passed over without notice, namely, +whether, as is sometimes assumed, each sub-species or race of man has +sprung from a single pair of progenitors. With our domestic animals a +new race can readily be formed from a single pair possessing some new +character, or even from a single individual thus characterised, by +carefully matching the varying offspring; but most of our races have +been formed, not intentionally from a selected pair, but unconsciously +by the preservation of many individuals which have varied, however +slightly, in some useful or desired manner. If in one country stronger +and heavier horses, and in another country lighter and fleeter horses, +were habitually preferred, we may feel sure that two distinct sub-breeds +would, in the course of time, be produced, without any particular pairs +or individuals having been separated and bred from in either country. +Many races have been thus formed, and their manner of formation is +closely analogous with that of natural species. We know, also, that the +horses which have been brought to the Falkland Islands have become, +during successive generations, smaller and weaker, whilst those which +have run wild on the Pampas have acquired larger and coarser heads; and +such changes are manifestly due, not to any one pair, but to all the +individuals having been subjected to the same conditions, aided, +perhaps, by the principle of reversion. The new sub-breeds in none of +these cases are descended from any single pair, but from many +individuals which have varied in different degrees, but in the same +general manner; and we may conclude that the races of man have been +similarly produced, the modifications being either the direct result of +exposure to different conditions, or the indirect result of some form of +selection. But to this latter subject we shall presently return. + + +_On the Extinction of the Races of Man._—The partial and complete +extinction of many races and sub-races of man are historically known +events. Humboldt saw in South America a parrot which was the sole living +creature that could speak the language of a lost tribe. Ancient +monuments and stone implements found in all parts of the world, of which +no tradition is preserved by the present inhabitants, indicate much +extinction. Some small and broken tribes, remnants of former races, +still survive in isolated and generally mountainous districts. In Europe +the ancient races were all, according to Schaaffhausen,[309] “lower in +the scale than the rudest living savages;” they must therefore have +differed, to a certain extent, from any existing race. The remains +described by Professor Broca[310] from Les Eyzies, though they +unfortunately appear to have belonged to a single family, indicate a +race with a most singular combination of low or simious and high +characteristics, and is “entirely different from any other race, ancient +or modern, that we have ever heard of.” It differed, therefore, from the +quaternary race of the caverns of Belgium. + +Unfavourable physical conditions appear to have had but little effect in +the extinction of races.[311] Man has long lived in the extreme regions +of the North, with no wood wherewith to make his canoes or other +implements, and with blubber alone for burning and giving him warmth, +but more especially for melting the snow. In the Southern extremity of +America the Fuegians survive without the protection of clothes, or of +any building worthy to be called a hovel. In South Africa the aborigines +wander over the most arid plains, where dangerous beasts abound. Man can +withstand the deadly influence of the Terai at the foot of the Himalaya, +and the pestilential shores of tropical Africa. + +Extinction follows chiefly from the competition of tribe with tribe, +and race with race. Various checks are always in action, as specified in +a former chapter, which serve to keep down the numbers of each savage +tribe,—such as periodical famines, the wandering of the parents and the +consequent deaths of infants, prolonged suckling, the stealing of women, +wars, accidents, sickness, licentiousness, especially infanticide, and, +perhaps, lessened fertility from less nutritious food, and many +hardships. If from any cause any one of these checks is lessened, even +in a slight degree, the tribe thus favoured will tend to increase; and +when one of two adjoining tribes becomes more numerous and powerful than +the other, the contest is soon settled by war, slaughter, cannibalism, +slavery, and absorption. Even when a weaker tribe is not thus abruptly +swept away, if it once begins to decrease, it generally goes on +decreasing until it is extinct.[312] + +When civilised nations come into contact with barbarians the struggle is +short, except where a deadly climate gives its aid to the native race. +Of the causes which lead to the victory of civilised nations, some are +plain and some very obscure. We can see that the cultivation of the land +will be fatal in many ways to savages, for they cannot, or will not, +change their habits. New diseases and vices are highly destructive; and +it appears that in every nation a new disease causes much death, until +those who are most susceptible to its destructive influence are +gradually weeded out;[313] and so it may be with the evil effects from +spirituous liquors, as well as with the unconquerably strong taste for +them shewn by so many savages. It further appears, mysterious as is the +fact, that the first meeting of distinct and separated people generates +disease.[314] Mr. Sproat, who in Vancouver Island closely attended to +the subject of extinction, believes that changed habits of life, which +always follow from the advent of Europeans, induces much ill-health. He +lays, also, great stress on so trifling a cause as that the natives +become “bewildered and dull by the new life around them; they lose the +motives for exertion, and get no new ones in their place.”[315] + +The grade of civilisation seems a most important element in the success +of nations which come in competition. A few centuries ago Europe feared +the inroads of Eastern barbarians; now, any such fear would be +ridiculous. It is a more curious fact, that savages did not formerly +waste away, as Mr. Bagehot has remarked, before the classical nations, +as they now do before modern civilised nations; had they done so, the +old moralists would have mused over the event; but there is no lament in +any writer of that period over the perishing barbarians.[316] + +Although the gradual decrease and final extinction of the races of man +is an obscure problem, we can see that it depends on many causes, +differing in different places and at different times. It is the same +difficult problem as that presented by the extinction of one of the +higher animals—of the fossil horse, for instance, which disappeared +from South America, soon afterwards to be replaced, within the same +districts, by countless troops of the Spanish horse. The New Zealander +seems conscious of this parallelism, for he compares his future fate +with that of the native rat almost exterminated by the European rat. The +difficulty, though great to our imagination, and really great if we wish +to ascertain the precise causes, ought not to be so to our reason, as +long as we keep steadily in mind that the increase of each species and +each race is constantly hindered by various checks; so that if any new +check, or cause of destruction, even a slight one, be superadded, the +race will surely decrease in number; and as it has everywhere been +observed that savages are much opposed to any change of habits, by which +means injurious checks could be counterbalanced, decreasing numbers will +sooner or later lead to extinction; the end, in most cases, being +promptly determined by the inroads of increasing and conquering tribes. + + +_On the Formation of the Races of Man._—It may be premised that when we +find the same race, though broken up into distinct tribes, ranging over +a great area, as over America, we may attribute their general +resemblance to descent from a common stock. In some cases the crossing +of races already, distinct has led to the formation of new races. The +singular fact that Europeans and Hindoos, who belong to the same Aryan +stock and speak a language fundamentally the same, differ widely in +appearance, whilst Europeans differ but little from Jews, who belong to +the Semitic stock and speak quite another language, has been accounted +for by Broca[317] through the Aryan branches having been largely crossed +during their wide diffusion by various indigenous tribes. When two races +in close contact cross, the first result is a heterogeneous mixture: +thus Mr. Hunter, in describing the Santali or hill-tribes of India, says +that hundreds of imperceptible gradations may be traced “from the black, +squat tribes of the mountains to the tall olive-coloured Brahman, with +his intellectual brow, calm eyes, and high but narrow head;” so that it +is necessary in courts of justice to ask the witnesses whether they are +Santalis or Hindoos.[318] Whether a heterogeneous people, such as the +inhabitants of some of the Polynesian islands, formed by the crossing of +two distinct races, with few or no pure members left, would ever become +homogeneous, is not known from direct evidence. But as with our +domesticated animals, a crossed breed can certainly, in the course of a +few generations, be fixed and made uniform by careful selection,[319] we +may infer that the free and prolonged intercrossing during many +generations of a heterogeneous mixture would supply the place of +selection, and overcome any tendency to reversion, so that a crossed +race would ultimately become homogeneous, though it might not partake in +an equal degree of the characters of the two parent-races. + +Of all the differences between the races of man, the colour of the skin +is the most conspicuous and one of the best marked. Differences of this +kind, it was formerly thought, could be accounted for by long exposure +under different climates; but Pallas first shewed that this view is not +tenable, and he has been followed by almost all anthropologists.[320] +The view has been rejected chiefly because the distribution of the +variously coloured races, most of whom must have long inhabited their +present homes, does not coincide with corresponding differences of +climate. Weight must also be given to such cases as that of the Dutch +families, who, as we hear on excellent authority,[321] have not +undergone the least change of colour, after residing for three centuries +in South Africa. The uniform appearance in various parts of the world of +gypsies and Jews, though the uniformity of the latter has been somewhat +exaggerated,[322] is likewise an argument on the same side. A very damp +or a very dry atmosphere has been supposed to be more influential in +modifying the colour of the skin than mere heat; but as D’Orbigny in +South America, and Livingstone in Africa, arrived at diametrically +opposite conclusions with respect to dampness and dryness, any +conclusion on this head must be considered as very doubtful.[323] + +Various facts, which I have elsewhere given, prove that the colour of +the skin and hair is sometimes correlated in a surprising manner with a +complete immunity from the action of certain vegetable poisons and from +the attacks of certain parasites. Hence it occurred to me, that negroes +and other dark races might have acquired their dark tints by the darker +individuals escaping during a long series of generations from the deadly +influence of the miasmas of their native countries. + +I afterwards found that the same idea had long ago occurred to Dr. +Wells.[324] That negroes, and even mulattoes, are almost completely +exempt from the yellow-fever, which is so destructive in tropical +America, has long been known.[325] They likewise escape to a large +extent the fatal intermittent fevers that prevail along, at least, 2600 +miles of the shores of Africa, and which annually cause one-fifth of the +white settlers to die, and another fifth to return home invalided.[326] +This immunity in the negro seems to be partly inherent, depending on +some unknown peculiarity of constitution, and partly the result of +acclimatisation. Pouchet[327] states that the negro regiments, borrowed +from the Viceroy of Egypt for the Mexican war, which had been recruited +near the Soudan, escaped the yellow-fever almost equally well with the +negroes originally brought from various parts of Africa, and accustomed +to the climate of the West Indies. That acclimatisation plays a part is +shewn by the many cases in which negroes, after having resided for some +time in a colder climate, have become to a certain extent liable to +tropical fevers.[328] The nature of the climate under which the white +races have long resided, likewise has some influence on them; for during +the fearful epidemic of yellow-fever in Demerara during 1837, Dr. Blair +found that the death-rate of the immigrants was proportional to the +latitude of the country whence they had come. With the negro the +immunity, as far as it is the result of acclimatisation, implies +exposure during a prodigious length of time; for the aborigines of +tropical America, who have resided there from time immemorial, are not +exempt from yellow-fever; and the Rev. B. Tristram states, that there +are districts in Northern Africa which the native inhabitants are +compelled annually to leave, though the negroes can remain with safety. + +That the immunity of the negro is in any degree correlated with the +colour of his skin is a mere conjecture: it may be correlated with some +difference in his blood, nervous system, or other tissues. Nevertheless, +from the facts above alluded to, and from some connection apparently +existing between complexion and a tendency to consumption, the +conjecture seemed to me not improbable. Consequently I endeavoured, with +but little success,[329] to ascertain how far it held good. The late +Dr. Daniell, who had long lived on the West Coast of Africa, told me +that he did not believe in any such relation. He was himself unusually +fair, and had withstood the climate in a wonderful manner. When he first +arrived as a boy on the coast, an old and experienced negro chief +predicted from his appearance that this would prove the case. Dr. +Nicholson, of Antigua, after having attended to this subject, wrote to +me that he did not think that dark-coloured Europeans escaped the +yellow-fever better than those that were light-coloured. Mr. J. M. +Harris altogether denies[330] that Europeans with dark hair withstand a +hot climate better than other men; on the contrary, experience has +taught him in making a selection of men for service on the coast of +Africa, to choose those with red hair. As far, therefore, as these +slight indications serve, there seems no foundation for the hypothesis, +which has been accepted by several writers, that the colour of the black +races may have resulted from darker and darker individuals having +survived in greater numbers, during their exposure to the +fever-generating miasmas of their native countries. + +Although with our present knowledge we cannot account for the +strongly-marked differences in colour between the races of man, either +through correlation with constitutional peculiarities, or through the +direct action of climate; yet we must not quite ignore the latter +agency, for there is good reason to believe that some inherited effect +is thus produced.[331] + +We have seen in our third chapter that the conditions of life, such as +abundant food and general comfort, affect in a direct manner the +development of the bodily frame, the effects being transmitted. Through +the combined influences of climate and changed habits of life, European +settlers, in the United States undergo, as is generally admitted, a +slight but extraordinarily rapid change of appearance. There is, also, a +considerable body of evidence shewing that in the Southern States the +house-slaves of the third generation present a markedly different +appearance from the field-slaves.[332] + +If, however, we look to the races of man, as distributed over the world, +we must infer that their characteristic differences cannot be accounted +for by the direct action of different conditions of life, even after +exposure to them for an enormous period of time. The Esquimaux live +exclusively on animal food; they are clothed in thick fur, and are +exposed to intense cold and to prolonged darkness; yet they do not +differ in any extreme degree from the inhabitants of Southern China, who +live entirely on vegetable food and are exposed almost naked to a hot, +glaring climate. The unclothed Fuegians live on the marine productions +of their inhospitable shores; the Botocudos of Brazil wander about the +hot forests of the interior and live chiefly on vegetable productions; +yet these tribes resemble each other so closely that the Fuegians on +board the “Beagle” were mistaken by some Brazilians for Botocudos. The +Botocudos again, as well as the other inhabitants of tropical America, +are wholly different from the Negroes who inhabit the opposite shores of +the Atlantic, are exposed to a nearly similar climate, and follow nearly +the same habits of life. + +Nor can the differences between the races of man be accounted for, +except to a quite insignificant degree, by the inherited effects of the +increased or decreased use of parts. Men who habitually live in canoes, +may have their legs somewhat stunted; those who inhabit lofty regions +have their chests enlarged; and those who constantly use certain +sense-organs have the cavities in which they are lodged somewhat +increased in size, and their features consequently a little modified. +With civilised nations, the reduced size of the jaws from lessened use, +the habitual play of different muscles serving to express different +emotions, and the increased size of the brain from greater intellectual +activity, have together produced a considerable effect on their general +appearance in comparison with savages.[333] It is also possible that +increased bodily stature, with no corresponding increase in the size of +the brain, may have given to some races (judging from the previously +adduced cases of the rabbits) an elongated skull of the dolichocephalic +type. + +Lastly, the little-understood principle of correlation will almost +certainly have come into action, as in the case of great muscular +development and strongly projecting supra-orbital ridges. It is not +improbable that the texture of the hair, which differs much in the +different races, may stand in some kind of correlation with the +structure of the skin; for the colour of the hair and skin are certainly +correlated, as is its colour and texture with the Mandans.[334] The +colour of the skin and the odour emitted by it are likewise in some +manner connected. With the breeds of sheep the number of hairs within a +given space and the number of the excretory pores stand in some relation +to each other.[335] If we may judge from the analogy of our domesticated +animals, many modifications of structure in man probably come under this +principle of correlated growth. + + +We have now seen that the characteristic differences between the races +of man cannot be accounted for in a satisfactory manner by the direct +action of the conditions of life, nor by the effects of the continued +use of parts, nor through the principle of correlation. We are therefore +led to inquire whether slight individual differences, to which man is +eminently liable, may not have been preserved and augmented during a +long series of generations through natural selection. But here we are at +once met by the objection that beneficial variations alone can be thus +preserved; and as far as we are enabled to judge (although always liable +to error on this head) not one of the external differences between the +races of man are of any direct or special service to him. The +intellectual and moral or social faculties must of course be excepted +from this remark; but differences in these faculties can have had little +or no influence on external characters. The variability of all the +characteristic differences between the races, before referred to, +likewise indicates that these differences cannot be of much importance; +for, had they been important, they would long ago have been either fixed +and preserved, or eliminated. In this respect man resembles those forms, +called by naturalists protean or polymorphic, which have remained +extremely variable, owing, as it seems, to their variations being of an +indifferent nature, and consequently to their having escaped the action +of natural selection. + +We have thus far been baffled in all our attempts to account for the +differences between the races of man; but there remains one important +agency, namely Sexual Selection, which appears to have acted as +powerfully on man, as on many other animals. I do not intend to assert +that sexual selection will account for all the differences between the +races. An unexplained residuum is left, about which we can in our +ignorance only say, that as individuals are continually born with, for +instance, heads a little rounder or narrower, and with noses a little +longer or shorter, such slight differences might become fixed and +uniform, if the unknown agencies which induced them were to act in a +more constant manner, aided by long-continued intercrossing. Such +modifications come under the provisional class, alluded to in our fourth +chapter, which for the want of a better term have been called +spontaneous variations. Nor do I pretend that the effects of sexual +selection can be indicated with scientific precision; but it can be +shewn that it would be an inexplicable fact if man had not been modified +by this agency, which has acted so powerfully on innumerable animals, +both high and low in the scale. It can further be shewn that the +differences between the races of man, as in colour, hairyness, form of +features, &c., are of the nature which it might have been expected would +have been acted on by sexual selection. But in order to treat this +subject in a fitting manner, I have found it necessary to pass the whole +animal kingdom in review; I have therefore devoted to it the Second Part +of this work. At the close I shall return to man, and, after attempting +to shew how far he has been modified through sexual selection, will give +a brief summary of the chapters in this First Part. + + + + +PART II.—SEXUAL SELECTION. + + +CHAPTER VIII. + +PRINCIPLES OF SEXUAL SELECTION. + + + Secondary sexual characters—Sexual selection—Manner of + action—Excess of males—Polygamy—The male alone generally + modified through sexual selection—Eagerness of the + male—Variability of the male—Choice exerted by the + female—Sexual compared with natural selection—Inheritance, at + corresponding periods of life, at corresponding seasons of the + year, and as limited by sex—Relations between the several + forms of inheritance—Causes why one sex and the young are not + modified through sexual selection—Supplement on the + proportional numbers of the two sexes throughout the animal + kingdom—On the limitation of the numbers of the two sexes + through natural selection. + + +With animals which have their sexes separated, the males necessarily +differ from the females in their organs of reproduction; and these +afford the primary sexual characters. But the sexes often differ in what +Hunter has called secondary sexual characters, which are not directly +connected with the act of reproduction; for instance, in the male +possessing certain organs of sense or locomotion, of which the female is +quite destitute, or in having them more highly-developed, in order that +he may readily find or reach her; or again, in the male having special +organs of prehension so as to hold her securely. These latter organs of +infinitely diversified kinds graduate into, and in some cases can hardly +be distinguished from, those which are commonly ranked as primary, such +as the complex appendages at the apex of the abdomen in male insects. +Unless indeed we confine the term “primary” to the reproductive glands, +it is scarcely possible to decide, as far as the organs of prehension +are concerned, which ought to be called primary and which secondary. + +The female often differs from the male in having organs for the +nourishment or protection of her young, as the mammary glands of +mammals, and the abdominal sacks of the marsupials. The male, also, in +some few cases differs from the female in possessing analogous organs, +as the receptacles for the ova possessed by the males of certain fishes, +and those temporarily developed in certain male frogs. Female bees have +a special apparatus for collecting and carrying pollen, and their +ovipositor is modified into a sting for the defence of their larvæ and +the community. In the females of many insects the ovipositor is modified +in the most complex manner for the safe placing of the eggs. Numerous +similar cases could be given, but they do not here concern us. There +are, however, other sexual differences quite disconnected with the +primary organs with which we are more especially concerned—such as the +greater size, strength, and pugnacity of the male, his weapons of +offence or means of defence against rivals, his gaudy colouring and +various ornaments, his power of song, and other such characters. + +Besides the foregoing primary and secondary sexual differences, the male +and female sometimes differ in structures connected with different +habits of life, and not at all, or only indirectly, related to the +reproductive functions. Thus the females of certain flies (Culicidæ and +Tabanidæ) are blood-suckers, whilst the males live on flowers and have +their mouths destitute of mandibles.[336] The males alone of certain +moths and of some crustaceans (_e.g._ Tanais) have imperfect, closed +mouths, and cannot feed. The Complemental males of certain cirripedes +live like epiphytic plants either on the female or hermaphrodite form, +and are destitute of a mouth and prehensile limbs. In these cases it is +the male which has been modified and has lost certain important organs, +which the other members of the same group possess. In other cases it is +the female which has lost such parts; for instance, the female glow-worm +is destitute of wings, as are many female moths, some of which never +leave their cocoons. Many female parasitic crustaceans have lost their +natatory legs. In some weevil-beetles (Curculionidæ) there is a great +difference between the male and female in the length of the rostrum or +snout;[337] but the meaning of this and of many analogous differences, +is not at all understood. Differences of structure between the two sexes +in relation to different habits of life are generally confined to the +lower animals; but with some few birds the beak of the male differs from +that of the female. No doubt in most, but apparently not in all these +cases, the differences are indirectly connected with the propagation of +the species: thus a female which has to nourish a multitude of ova will +require more food than the male, and consequently will require special +means for procuring it. A male animal which lived for a very short time +might without detriment lose through disuse its organs for procuring +food; but he would retain his locomotive organs in a perfect state, so +that he might reach the female. The female, on the other hand, might +safely lose her organs for flying, swimming, or walking, if she +gradually acquired habits which rendered such powers useless. + +We are, however, here concerned only with that kind of selection, which +I have called sexual selection. This depends on the advantage which +certain individuals have over other individuals of the same sex and +species, in exclusive relation to reproduction. When the two sexes +differ in structure in relation to different habits of life, as in the +cases above mentioned, they have no doubt been modified through natural +selection, accompanied by inheritance limited to one and the same sex. +So again the primary sexual organs, and those for nourishing or +protecting the young, come under this same head; for those individuals +which generated or nourished their offspring best, would leave, _cæteris +paribus_, the greatest number to inherit their superiority; whilst those +which generated or nourished their offspring badly, would leave but few +to inherit their weaker powers. As the male has to search for the +female, he requires for this purpose organs of sense and locomotion, but +if these organs are necessary for the other purposes of life, as is +generally the case, they will have been developed through natural +selection. When the male has found the female he sometimes absolutely +requires prehensile organs to hold her; thus Dr. Wallace informs me that +the males of certain moths cannot unite with the females if their tarsi +or feet are broken. The males of many oceanic crustaceans have their +legs and antennæ modified in an extraordinary manner for the prehension +of the female; hence we may suspect that owing to these animals being +washed about by the waves of the open sea, they absolutely require these +organs in order to propagate their kind, and if so their development +will have been the result of ordinary or natural selection. + +When the two sexes follow exactly the same habits of life, and the male +has more highly developed sense or locomotive organs than the female, it +may be that these in their perfected state are indispensable to the male +for finding the female; but in the vast majority of cases, they serve +only to give one male an advantage over another, for the less +well-endowed males, if time were allowed them, would succeed in pairing +with the females; and they would in all other respects, judging from the +structure of the female, be equally well adapted for their ordinary +habits of life. In such cases sexual selection must have come into +action, for the males have acquired their present structure, not from +being better fitted to survive in the struggle for existence, but from +having gained an advantage over other males, and from having transmitted +this advantage to their male offspring alone. It was the importance of +this distinction which led me to designate this form of selection as +sexual selection. So again, if the chief service rendered to the male by +his prehensile organs is to prevent the escape of the female before the +arrival of other males, or when assaulted by them, these organs will +have been perfected through sexual selection, that is by the advantage +acquired by certain males over their rivals. But in most cases it is +scarcely possible to distinguish between the effects of natural and +sexual selection. Whole chapters could easily be filled with details on +the differences between the sexes in their sensory, locomotive, and +prehensile organs. As, however, these structures are not more +interesting than others adapted for the ordinary purposes of life, I +shall almost pass them over, giving only a few instances under each +class. + +There are many other structures and instincts which must have been +developed through sexual selection—such as the weapons of offence and +the means of defence possessed by the males for fighting with and +driving away their rivals—their courage and pugnacity—their ornaments +of many kinds—their organs for producing vocal or instrumental +music—and their glands for emitting odours; most of these latter +structures serving only to allure or excite the female. That these +characters are the result of sexual and not of ordinary selection is +clear, as unarmed, unornamented, or unattractive males would succeed +equally well in the battle for life and in leaving a numerous progeny, +if better endowed males were not present. We may infer that this would +be the case, for the females, which are unarmed and unornamented, are +able to survive and procreate their kind. Secondary sexual characters of +the kind just referred to, will be fully discussed in the following +chapters, as they are in many respects interesting, but more especially +as they depend on the will, choice, and rivalry of the individuals of +either sex. When we behold two males fighting for the possession of the +female, or several male birds displaying their gorgeous plumage, and +performing the strangest antics before an assembled body of females, we +cannot doubt that, though led by instinct, they know what they are +about, and consciously exert their mental and bodily powers. + +In the same manner as man can improve the breed of his game-cocks by the +selection of those birds which are victorious in the cockpit, so it +appears that the strongest and most vigorous males, or those provided +with the best weapons, have prevailed under nature, and have led to the +improvement of the natural breed or species. Through repeated deadly +contests, a slight degree of variability, if it led to some advantage, +however slight, would suffice for the work of sexual selection; and it +is certain that secondary sexual characters are eminently variable. In +the same manner as man can give beauty, according to his standard of +taste, to his male poultry—can give to the Sebright bantam a new and +elegant plumage, an erect and peculiar carriage—so it appears that in a +state of nature female birds, by having long selected the more +attractive males, have added to their beauty. No doubt this implies +powers of discrimination and taste on the part of the female which will +at first appear extremely improbable; but I hope hereafter to shew that +this is not the case. + +From our ignorance on several points, the precise manner in which sexual +selection acts is to a certain extent uncertain. Nevertheless if those +naturalists who already believe in the mutability of species, will read +the following chapters, they will, I think, agree with me that sexual +selection has played an important part in the history of the organic +world. It is certain that with almost all animals there is a struggle +between the males for the possession of the female. This fact is so +notorious that it would be superfluous to give instances. Hence the +females, supposing that their mental capacity sufficed for the exertion +of a choice, could select one out of several males. But in numerous +cases it appears as if it had been specially arranged that there should +be a struggle between many males. Thus with migratory birds, the males +generally arrive before the females at their place of breeding, so that +many males are ready to contend for each female. The bird-catchers +assert that this is invariably the case with the nightingale and +blackcap, as I am informed by Mr. Jenner Weir, who confirms the +statement with respect to the latter species. + +Mr. Swaysland of Brighton, who has been in the habit, during the last +forty years, of catching our migratory birds on their first arrival, +writes to me that he has never known the females of any species to +arrive before their males. During one spring he shot thirty-nine males +of Ray’s wagtail (_Budytes Raii_) before he saw a single female. Mr. +Gould has ascertained by dissection, as he informs me, that male snipes +arrive in this country before the females. In the case of fish, at the +period when the salmon ascend our rivers, the males in large numbers are +ready to breed before the females. So it apparently is with frogs and +toads. Throughout the great class of insects the males almost always +emerge from the pupal state before the other sex, so that they generally +swarm for a time before any females can be seen.[338] The cause of this +difference between the males and females in their periods of arrival and +maturity is sufficiently obvious. Those males which annually first +migrated into any country, or which in the spring were first ready to +breed, or were the most eager, would leave the largest number of +offspring; and these would tend to inherit similar instincts and +constitutions. On the whole there can be no doubt that with almost all +animals, in which the sexes are separate, there is a constantly +recurrent struggle between the males for the possession of the females. + +Our difficulty in regard to sexual selection lies in understanding how +it is that the males which conquer other males, or those which prove the +most attractive to the females, leave a greater number of offspring to +inherit their superiority than the beaten and less attractive males. +Unless this result followed, the characters which gave to certain males +an advantage over others, could not be perfected and augmented through +sexual selection. When the sexes exist in exactly equal numbers, the +worst-endowed males will ultimately find females (excepting where +polygamy prevails), and leave as many offspring, equally well fitted for +their general habits of life, as the best-endowed males. From various +facts and considerations, I formerly inferred that with most animals, in +which secondary sexual characters were well developed, the males +considerably exceeded the females in number; and this does hold good in +some few cases. If the males were to the females as two to one, or as +three to two, or even in a somewhat lower ratio, the whole affair would +be simple; for the better-armed or more attractive males would leave the +largest number of offspring. But after investigating, as far as +possible, the numerical proportions of the sexes, I do not believe that +any great inequality in number commonly exists. In most cases sexual +selection appears to have been effective in the following manner. + +Let us take any species, a bird for instance, and divide the females +inhabiting a district into two equal bodies: the one consisting of the +more vigorous and better-nourished individuals, and the other of the +less vigorous and healthy. The former, there can be little doubt, would +be ready to breed in the spring before the others; and this is the +opinion of Mr. Jenner Weir, who has during many years carefully attended +to the habits of birds. There can also be no doubt that the most +vigorous, healthy, and best-nourished females would on an average +succeed in rearing the largest number of offspring. The males, as we +have seen, are generally ready to breed before the females; of the males +the strongest, and with some species the best armed, drive away the +weaker males; and the former would then unite with the more vigorous and +best-nourished females, as these are the first to breed. Such vigorous +pairs would surely rear a larger number of offspring than the retarded +females, which would be compelled, supposing the sexes to be numerically +equal, to unite with the conquered and less powerful males; and this is +all that is wanted to add, in the course of successive generations, to +the size, strength and courage of the males, or to improve their +weapons. + +But in a multitude of cases the males which conquer other males, do not +obtain possession of the females, independently of choice on the part of +the latter. The courtship of animals is by no means so simple and short +an affair as might be thought. The females are most excited by, or +prefer pairing with, the more ornamented males, or those which are the +best songsters, or play the best antics; but it is obviously probable, +as has been actually observed in some cases, that they would at the same +time prefer the more vigorous and lively males.[339] Thus the more +vigorous females, which are the first to breed, will have the choice of +many males; and though they may not always select the strongest or best +armed, they will select those which are vigorous and well armed, and in +other respects the most attractive. Such early pairs would have the same +advantage in rearing offspring on the female side as above explained, +and nearly the same advantage on the male side. And this apparently has +sufficed during a long course of generations to add not only to the +strength and fighting-powers of the males, but likewise to their +various ornaments or other attractions. + +In the converse and much rarer case of the males selecting particular +females, it is plain that those which were the most vigorous and had +conquered others, would have the freest choice; and it is almost certain +that they would select vigorous as well as attractive females. Such +pairs would have an advantage in rearing offspring, more especially if +the male had the power to defend the female during the pairing-season, +as occurs with some of the higher animals, or aided in providing for the +young. The same principles would apply if both sexes mutually preferred +and selected certain individuals of the opposite sex; supposing that +they selected not only the more attractive, but likewise the more +vigorous individuals. + + +_Numerical Proportion of the Two Sexes._—I have remarked that sexual +selection would be a simple affair if the males considerably exceeded in +number the females. Hence I was led to investigate, as far as I could, +the proportions between the two sexes of as many animals as possible; +but the materials are scanty. I will here give only a brief abstract of +the results, retaining the details for a supplementary discussion, so as +not to interfere with the course of my argument. Domesticated animals +alone afford the opportunity of ascertaining the proportional numbers at +birth; but no records have been specially kept for this purpose. By +indirect means, however, I have collected a considerable body of +statistical data, from which it appears that with most of our domestic +animals the sexes are nearly equal at birth. Thus with race-horses, +25,560 births have been recorded during twenty-one years, and the male +births have been to the female births as 99·7 to 100. With greyhounds +the inequality is greater than with any other animal, for during twelve +years, out of 6878 births, the male births have been as 110·1 to 100 +female births. It is, however, in some degree doubtful whether it is +safe to infer that the same proportional numbers would hold good under +natural conditions as under domestication; for slight and unknown +differences in the conditions affect to a certain extent the proportion +of the sexes. Thus with mankind, the male births in England are as +104·5, in Russia as 108·9, and with the Jews of Livornia as 120 _to_ 100 +females. The proportion is also mysteriously affected by the +circumstance of the births being legitimate or illegitimate. + +For our present purpose we are concerned with the proportion of the +sexes, not at birth, but at maturity, and this adds another element of +doubt; for it is a well ascertained fact that with man a considerably +larger proportion of males than of females die before or during birth, +and during the first few years of infancy. So it almost certainly is +with male lambs, and so it may be with the males of other animals. The +males of some animals kill each other by fighting; or they drive each +other about until they become greatly emaciated. They must, also, whilst +wandering about in eager search for the females, be often exposed to +various dangers. With many kinds of fish the males are much smaller than +the females, and they are believed often to be devoured by the latter or +by other fishes. With some birds the females appear to die in larger +proportion than the males: they are also liable to be destroyed on their +nests, or whilst in charge of their young. With insects the female larvæ +are often larger than those of the males, and would consequently be more +likely to be devoured: in some cases the mature females are less active +and less rapid in their movements than the males, and would not be so +well able to escape from danger. Hence, with animals in a state of +nature, in order to judge of the proportions of the sexes at maturity, +we must rely on mere estimation; and this, except perhaps when the +inequality is strongly marked, is but little trustworthy. Nevertheless, +as far as a judgment can be formed, we may conclude from the facts given +in the supplement, that the males of some few mammals, of many birds, of +some fish and insects, considerably exceed in number the females. + +The proportion between the sexes fluctuates slightly during successive +years: thus with race-horses, for every 100 females born, the males +varied from 107.1 in one year to 92.6 in another year, and with +greyhounds from 116.3 to 95.3. But had larger numbers been tabulated +throughout a more extensive area than England, these fluctuations would +probably have disappeared; and such as they are, they would hardly +suffice to lead under a state of nature to the effective action of +sexual selection. Nevertheless with some few wild animals, the +proportions seem, as shewn in the supplement, to fluctuate either during +different seasons or in different localities in a sufficient degree to +lead to such action. For it should be observed that any advantage gained +during certain years or in certain localities by those males which were +able to conquer other males, or were the most attractive to the females, +would probably be transmitted to the offspring and would not +subsequently be eliminated. During the succeeding seasons, when from the +equality of the sexes every male was everywhere able to procure a +female, the stronger or more attractive males previously produced would +still have at least as good a chance of leaving offspring as the less +strong or less attractive. + + +_Polygamy._—The practice of polygamy leads to the same results as +would follow from an actual inequality in the number of the sexes; for +if each male secures two or more females, many males will not be able to +pair; and the latter assuredly will be the weaker or less attractive +individuals. Many mammals and some few birds are polygamous, but with +animals belonging to the lower classes I have found no evidence of this +habit. The intellectual powers of such animals are, perhaps, not +sufficient to lead them to collect and guard a harem of females. That +some relation exists between polygamy and the development of secondary +sexual characters, appears nearly certain; and this supports the view +that a numerical preponderance of males would be eminently favourable to +the action of sexual selection. Nevertheless many animals, especially +birds, which are strictly monogamous, display strongly-marked secondary +sexual characters; whilst some few animals, which are polygamous, are +not thus characterised. + +We will first briefly run through the class of mammals, and then turn to +birds. The gorilla seems to be a polygamist, and the male differs +considerably from the female; so it is with some baboons which live in +herds containing twice as many adult females as males. In South America +the _Mycetes caraya_ presents well-marked sexual differences in colour, +beard, and vocal organs, and the male generally lives with two or three +wives: the male of the _Cebus capucinus_ differs somewhat from the +female, and appears to be polygamous.[340] Little is known on this head +with respect to most other monkeys, but some species are strictly +monogamous. The ruminants are eminently polygamous, and they more +frequently present sexual differences than almost any other group of +mammals, especially in their weapons, but likewise in other characters. +Most deer, cattle, and sheep are polygamous; as are most antelopes, +though some of the latter are monogamous. Sir Andrew Smith, in speaking +of the antelopes of South Africa, says that in herds of about a dozen +there was rarely more than one mature male. The Asiatic _Antilope saiga_ +appears to be the most inordinate polygamist in the world; for +Pallas[341] states that the male drives away all rivals, and collects a +herd of about a hundred, consisting of females and kids: the female is +hornless and has softer hair, but does not otherwise differ much from +the male. The horse is polygamous, but, except in his greater size and +in the proportions of his body, differs but little from the mare. The +wild boar, in his great tusks and some other characters, presents +well-marked sexual characters; in Europe and in India he leads a +solitary life, except during the breeding-season; but at this season he +consorts in India with several females, as Sir W. Elliot, who has had +large experience in observing this animal, believes: whether this holds +good in Europe is doubtful, but is supported by some statements. The +adult male Indian elephant, like the boar, passes much of his time in +solitude; but when associating with others, “it is rare to find,” as Dr. +Campbell states, “more than one male with a whole herd of females.” The +larger males expel or kill the smaller and weaker ones. The male differs +from the female by his immense tusks and greater size, strength, and +endurance; so great is the difference in these latter respects, that +the males when caught are valued at twenty per cent. above the +females.[342] With other pachydermatous animals the sexes differ very +little or not at all, and they are not, as far as known, polygamists. +Hardly a single species amongst the Cheiroptera and Edentata, or in the +great Orders of the Rodents and Insectivora, presents well-developed +secondary sexual differences; and I can find no account of any species +being polygamous, excepting, perhaps, the common rat, the males of +which, as some rat-catchers affirm, live with several females. + +The lion in South Africa, as I hear from Sir Andrew Smith, sometimes +lives with a single female, but generally with more than one, and, in +one case, was found with as many as five females, so that he is +polygamous. He is, as far as I can discover, the sole polygamist in the +whole group of the terrestrial Carnivora, and he alone presents +well-marked sexual characters. If, however, we turn to the marine +Carnivora, the case is widely different; for many species of seals +offer, as we shall hereafter see, extraordinary sexual differences, and +they are eminently polygamous. Thus the male sea-elephant of the +Southern Ocean, always possesses, according to Péron, several females, +and the sea-lion of Forster is said to be surrounded by from twenty to +thirty females. In the North, the male sea-bear of Steller is +accompanied by even a greater number of females. + +With respect to birds, many species, the sexes of which differ greatly +from each other, are certainly monogamous. In Great Britain we see +well-marked sexual differences in, for instance, the wild-duck which +pairs with a single female, with the common blackbird, and with the +bullfinch which is said to pair for life. So it is, as I am informed by +Mr. Wallace, with the Chatterers or Cotingidæ of South America, and +numerous other birds. In several groups I have not been able to discover +whether the species are polygamous or monogamous. Lesson says that birds +of paradise, so remarkable for their sexual differences, are polygamous, +but Mr. Wallace doubts whether he had sufficient evidence. Mr. Salvin +informs me that he has been led to believe that humming-birds are +polygamous. The male widow-bird; remarkable for his caudal plumes, +certainly seems to be a polygamist.[343] I have been assured by Mr. +Jenner Weir and by others, that three starlings not rarely frequent the +same nest; but whether this is a case of polygamy or polyandry has not +been ascertained. + +The Gallinaceæ present almost as strongly marked sexual differences as +birds of paradise or humming-birds, and many of the species are, as is +well known, polygamous; others being strictly monogamous. What a +contrast is presented between the sexes by the polygamous peacock or +pheasant, and the monogamous guinea-fowl or partridge! Many similar +cases could be given, as in the grouse tribe, in which the males of the +polygamous capercailzie and black-cock differ greatly from the females; +whilst the sexes of the monogamous red grouse and ptarmigan differ very +little. Amongst the Cursores, no great number of species offer +strongly-marked sexual differences, except the bustards, and the great +bustard (_Otis tarda_), is said to be polygamous. With the Grallatores, +extremely few species differ sexually, but the ruff (_Machetes pugnax_) +affords a strong exception, and this species is believed by Montagu to +be a polygamist. Hence it appears that with birds there often exists a +close relation between polygamy and the development of strongly-marked +sexual differences. On asking Mr. Bartlett, at the Zoological Gardens, +who has had such large experience with birds, whether the male tragopan +(one of the Gallinaceæ) was polygamous, I was struck by his answering, +“I do not know, but should think so from his splendid colours.” + +It deserves notice that the instinct of pairing with a single female is +easily lost under domestication. The wild-duck is strictly monogamous, +the domestic-duck highly polygamous. The Rev. W. D. Fox informs me that +with some half-tamed wild-ducks, kept on a large pond in his +neighbourhood, so many mallards were shot by the gamekeeper that only +one was left for every seven or eight females; yet unusually large +broods were reared. The guinea-fowl is strictly monogamous; but Mr. Fox +finds that his birds succeed best when he keeps one cock to two or three +hens.[344] Canary-birds pair in a state of nature, but the breeders in +England successfully put one male to four or five females; nevertheless +the first female, as Mr. Fox has been assured, is alone treated as the +wife, she and her young ones being fed by him; the others are treated as +concubines. I have noticed these cases, as it renders it in some degree +probable that monogamous species, in a state of nature, might readily +become either temporarily or permanently polygamous. + +With respect to reptiles and fishes, too little is known of their habits +to enable us to speak of their marriage arrangements. The stickle-back +Gasterosteus, however, is said to be a polygamist;[345] and the male +during the breeding-season differs conspicuously from the female. + +To sum up on the means through which, as far as we can judge, sexual +selection has led to the development of secondary sexual characters. It +has been shewn that the largest number of vigorous offspring will be +reared from the pairing of the strongest and best-armed males, which +have conquered other males, with the most vigorous and best-nourished +females, which are the first to breed in the spring. Such females, if +they select the more attractive, and at the same time vigorous, males, +will rear a larger number of offspring than the retarded females, which +must pair with the less vigorous and less attractive males. So it will +be if the more vigorous males select the more attractive and at the same +time healthy and vigorous females; and this will especially hold good if +the male defends the female, and aids in providing food for the young. +The advantage thus gained by the more vigorous pairs in rearing a larger +number of offspring has apparently sufficed to render sexual selection +efficient. But a large preponderance in number of the males over the +females would be still more efficient; whether the preponderance was +only occasional and local, or permanent; whether it occurred at birth, +or subsequently from the greater destruction of the females; or whether +it indirectly followed from the practice of polygamy. + + +_The Male generally more modified than the Female._—Throughout the +animal kingdom, when the sexes differ from each other in external +appearance, it is the male which, with rare exceptions, has been chiefly +modified; for the female still remains more like the young of her own +species, and more like the other members of the same group. The cause of +this seems to lie in the males of almost all animals having stronger +passions than the females. Hence it is the males that fight together and +sedulously display their charms before the females; and those which are +victorious transmit their superiority to their male offspring. Why the +males do not transmit their characters to both sexes will hereafter be +considered. That the males of all mammals eagerly pursue the females is +notorious to every one. So it is with birds; but many male birds do not +so much pursue the female, as display their plumage, perform strange +antics, and pour forth their song, in her presence. With the few fish +which have been observed, the male seems much more eager than the +female; and so it is with alligators, and apparently with Batrachians. +Throughout the enormous class of insects, as Kirby remarks,[346] “the +law is, that the male shall seek the female.” With spiders and +crustaceans, as I hear from two great authorities, Mr. Blackwall and Mr. +C. Spence Bate, the males are more active and more erratic in their +habits than the females. With insects and crustaceans, when the organs +of sense or locomotion are present in the one sex and absent in the +other, or when, as is frequently the case, they are more highly +developed in the one than the other, it is almost invariably the male, +as far as I can discover, which retains such organs, or has them most +developed; and this shews that the male is the more active member in the +courtship of the sexes.[347] + +The female, on the other hand, with the rarest exception, is less eager +than the male. As the illustrious Hunter[348] long ago observed, she +generally “requires to be courted;” she is coy, and may often be seen +endeavouring for a long time to escape from the male. Every one who has +attended to the habits of animals will be able to call to mind instances +of this kind. Judging from various facts, hereafter to be given, and +from the results which may fairly be attributed to sexual selection, the +female, though comparatively passive, generally exerts some choice and +accepts one male in preference to others. Or she may accept, as +appearances would sometimes lead us to believe, not the male which is +the most attractive to her, but the one which is the least distasteful. +The exertion of some choice on the part of the female seems almost as +general a law as the eagerness of the male. + +We are naturally led to enquire why the male in so many and such widely +distinct classes has been rendered more eager than the female, so that +he searches for her and plays the more active part in courtship. It +would be no advantage and some loss of power if both sexes were mutually +to search for each other; but why should the male almost always be the +seeker? With plants, the ovules after fertilisation have to be nourished +for a time; hence the pollen is necessarily brought to the female +organs—being placed on the stigma, through the agency of insects or of +the wind, or by the spontaneous movements of the stamens; and with the +Algæ, &c., by the locomotive power of the antherozooids. With +lowly-organised animals permanently affixed to the same spot and having +their sexes separate, the male element is invariably brought to the +female; and we can see the reason; for the ova, even if detached before +being fertilised and not requiring subsequent nourishment or protection, +would be, from their larger relative size, less easily transported than +the male element. Hence plants[349] and many of the lower animals are, +in this respect, analogous. In the case of animals not affixed to the +same spot, but enclosed within a shell with no power of protruding any +part of their bodies, and in the case of animals having little power of +locomotion, the males must trust the fertilising element to the risk of +at least a short transit through the waters of the sea. It would, +therefore, be a great advantage to such animals, as their organisation +became perfected, if the males when ready to emit the fertilising +element, were to acquire the habit of approaching the female as closely +as possible. The males of various lowly-organised animals having thus +aboriginally acquired the habit of approaching and seeking the females, +the same habit would naturally be transmitted to their more highly +developed male descendants; and in order that they should become +efficient seekers, they would have to be endowed with strong passions. +The acquirement of such passions would naturally follow from the more +eager males leaving a larger number of offspring than the less eager. + +The great eagerness of the male has thus indirectly led to the much +more frequent development of secondary sexual characters in the male +than in the female. But the development of such characters will have +been much aided, if the conclusion at which I arrived after studying +domesticated animals, can be trusted, namely, that the male is more +liable to vary than the female. I am aware how difficult it is to verify +a conclusion of this kind. Some slight evidence, however, can be gained +by comparing the two sexes in mankind, as man has been more carefully +observed than any other animal. During the Novara Expedition[350] a vast +number of measurements of various parts of the body in different races +were made, and the men were found in almost every case to present a +greater range of variation than the women; but I shall have to recur to +this subject in a future chapter. Mr. J. Wood,[351] who has carefully +attended to the variation of the muscles in man, puts in italics the +conclusion that “the greatest number of abnormalities in each subject is +found in the males.” He had previously remarked that “altogether in 102 +subjects the varieties of redundancy were found to be half as many again +as in females, contrasting widely with the greater frequency of +deficiency in females before described.” Professor Macalister like wise +remarks[352] that variations in the muscles “are probably more common in +males than females.” Certain muscles which are not normally present in +mankind are also more frequently developed in the male than in the +female sex, although exceptions to this rule are said to occur. Dr. +Burt Wilder[353] has tabulated the cases of 152 individuals with +supernumerary digits, of which 86 were males, and 39, or less than half, +females; the remaining 27 being of unknown sex. It should not, however, +be overlooked that women would more frequently endeavour to conceal a +deformity of this kind than men. Whether the large proportional number +of deaths of the male offspring of man and apparently of sheep, compared +with the female offspring, before, during, and shortly after birth (see +supplement), has any relation to a stronger tendency in the organs of +the male to vary and thus to become abnormal in structure or function, I +will not pretend to conjecture. + +In various classes of animals a few exceptional cases occur, in which +the female instead of the male has acquired well pronounced secondary +sexual characters, such as brighter colours, greater size, strength, or +pugnacity. With birds, as we shall hereafter see, there has sometimes +been a complete transposition of the ordinary characters proper to each +sex; the females having become the more eager in courtship, the males +remaining comparatively passive, but apparently selecting, as we may +infer from the results, the more attractive females. Certain female +birds have thus been rendered more highly coloured or otherwise +ornamented, as well as more powerful and pugnacious than the males, +these characters being transmitted to the female offspring alone. + +It may be suggested that in some cases a double process of selection has +been carried on; the males having selected the more attractive females, +and the latter the more attractive males. This process however, though +it might lead to the modification of both sexes, would not make the one +sex different from the other, unless indeed their taste for the +beautiful differed; but this is a supposition too improbable in the case +of any animal, excepting man, to be worth considering. There are, +however, many animals, in which the sexes resemble each other, both +being furnished with the same ornaments, which analogy would lead us to +attribute to the agency of sexual selection. In such cases it may be +suggested with more plausibility, that there has been a double or mutual +process of sexual selection; the more vigorous and precocious females +having selected the more attractive and vigorous males, the latter +having rejected all except the more attractive females. But from what we +know of the habits of animals, this view is hardly probable, the male +being generally eager to pair with any female. It is more probable that +the ornaments common to both sexes were acquired by one sex, generally +the male, and then transmitted to the offspring of both sexes. If, +indeed, during a lengthened period the males of any species were greatly +to exceed the females in number, and then during another lengthened +period under different conditions the reverse were to occur, a double, +but not simultaneous, process of sexual selection might easily be +carried on, by which the two sexes might be rendered widely different. + +We shall hereafter see that many animals exist, of which neither sex is +brilliantly coloured or provided with special ornaments, and yet the +members of both sexes or of one alone have probably been modified +through sexual selection. The absence of bright tints or other ornaments +may be the result of variations of the right kind never having occurred, +or of the animals themselves preferring simple colours, such as plain +black or white. Obscure colours have often been acquired through natural +selection for the sake of protection, and the acquirement through +sexual selection of conspicuous colours, may have been checked from the +danger thus incurred. But in other cases the males have probably +struggled together during long ages, through brute force, or by the +display of their charms, or by both means combined, and yet no effect +will have been produced unless a larger number of offspring were left by +the more successful males to inherit their superiority, than by the less +successful males; and this, as previously shewn, depends on various +complex contingencies. + +Sexual selection acts in a less rigorous manner than natural selection. +The latter produces its effects by the life or death at all ages of the +more or less successful individuals. Death, indeed, not rarely ensues +from the conflicts of rival males. But generally the less successful +male merely fails to obtain a female, or obtains later in the season a +retarded and less vigorous female, or, if polygamous, obtains fewer +females; so that they leave fewer, or less vigorous, or no offspring. In +regard to structures acquired through ordinary or natural selection, +there is in most cases, as long as the conditions of life remain the +same, a limit to the amount of advantageous modification in relation to +certain special ends; but in regard to structures adapted to make one +male victorious over another, either in fighting or in charming the +female, there is no definite limit to the amount of advantageous +modification; so that as long as the proper variations arise the work of +sexual selection will go on. This circumstance may partly account for +the frequent and extraordinary amount of variability presented by +secondary sexual characters. Nevertheless, natural selection will +determine that characters of this kind shall not be acquired by the +victorious males, which would be injurious to them in any high degree, +either by expending too much of their vital powers, or by exposing them +to any great danger. The development, however, of certain structures—of +the horns, for instance, in certain stags—has been carried to a +wonderful extreme; and in some instances to an extreme which, as far as +the general conditions of life are concerned, must be slightly injurious +to the male. From this fact we learn that the advantages which favoured +males have derived from conquering other males in battle or courtship, +and thus leaving a numerous progeny, have been in the long run greater +than those derived from rather more perfect adaptation to the external +conditions of life. We shall further see, and this could never have been +anticipated, that the power to charm the female has been in some few +instances more important than the power to conquer other males in +battle. + + +LAWS OF INHERITANCE. + +In order to understand how sexual selection has acted, and in the course +of ages has produced conspicuous results with many animals of many +classes, it is necessary to bear in mind the laws of inheritance, as far +as they are known. Two distinct elements are included under the term +“inheritance,” namely the transmission and the development of +characters; but as these generally go together, the distinction is often +overlooked. We see this distinction in those characters which are +transmitted through the early years of life, but are developed only at +maturity or during old age. We see the same distinction more clearly +with secondary sexual characters, for these are transmitted through both +sexes, though developed in one alone. That they are present in both +sexes, is manifest when two species, having strongly-marked sexual +characters, are crossed, for each transmits the characters proper to +its own male and female sex to the hybrid offspring of both sexes. The +same fact is likewise manifest, when characters proper to the male are +occasionally developed in the female when she grows old or becomes +diseased; and so conversely with the male. Again, characters +occasionally appear, as if transferred from the male to the female, as +when, in certain breeds of the fowl, spurs regularly appear in the young +and healthy females; but in truth they are simply developed in the +female; for in every breed each detail in the structure of the spur is +transmitted through the female to her male offspring. In all cases of +reversion, characters are transmitted through two, three, or many +generations, and are then under certain unknown favourable conditions +developed. This important distinction between transmission and +development will be easiest kept in mind by the aid of the hypothesis of +pangenesis, whether or not it be accepted as true. According to this +hypothesis, every unit or cell of the body throws off gemmules or +undeveloped atoms, which are transmitted to the offspring of both sexes, +and are multiplied by self-division. They may remain undeveloped during +the early years of life or during successive generations; their +development into units or cells, like those from which they were +derived, depending on their affinity for, and union with, other units or +cells previously developed in the due order of growth. + + +_Inheritance at Corresponding Periods of Life._—This tendency is well +established. If a new character appears in an animal whilst young, +whether it endures throughout life or lasts only for a time, it will +reappear, as a general rule, at the same age and in the same manner in +the offspring. If, on the other hand, a new character appears at +maturity, or even during old age, it tends to reappear in the offspring +at the same advanced age. When deviations from this rule occur, the +transmitted characters much oftener appear before than after the +corresponding age. As I have discussed this subject at sufficient length +in another work,[354] I will here merely give two or three instances, +for the sake of recalling the subject to the reader’s mind. In several +breeds of the Fowl, the chickens whilst covered with down, in their +first true plumage, and in their adult plumage, differ greatly from each +other, as well as from their common parent-form, the _Gallus bankiva_; +and these characters are faithfully transmitted by each breed to their +offspring at the corresponding period of life. For instance, the +chickens of spangled Hamburghs, whilst covered with down, have a few +dark spots on the head and rump, but are not longitudinally striped, as +in many other breeds; in their first true plumage, “they are beautifully +pencilled,” that is each feather is transversely marked by numerous dark +bars; but in their second plumage the feathers all become spangled or +tipped with a dark round spot.[355] Hence in this breed variations have +occurred and have been transmitted at three distinct periods of life. +The Pigeon offers a more remarkable case, because the aboriginal +parent-species does not undergo with advancing age any change of +plumage, excepting that at maturity the breast becomes more iridescent; +yet there are breeds which do not acquire their characteristic colours +until they have moulted two, three, or four times; and these +modifications of plumage are regularly transmitted. + + +_Inheritance at Corresponding Seasons of the Year._—With animals in a +state of nature innumerable instances occur of characters periodically +appearing at different seasons. We see this with the horns of the stag, +and with the fur of arctic animals which becomes thick and white during +the winter. Numerous birds acquire bright colours and other decorations +during the breeding-season alone. I can throw but little light on this +form of inheritance from facts observed under domestication. Pallas +states,[356] that in Siberia domestic cattle and horses periodically +become lighter-coloured during the winter; and I have observed a similar +marked change of colour in certain ponies in England. Although I do not +know that this tendency to assume a differently coloured coat during +different seasons of the year is transmitted, yet it probably is so, as +all shades of colour are strongly inherited by the horse. Nor is this +form of inheritance, as limited by season, more remarkable than +inheritance as limited by age or sex. + + +_Inheritance as Limited by Sex._—The equal transmission of characters +to both, sexes is the commonest form of inheritance, at least with those +animals which do not present strongly-marked sexual differences, and +indeed with many of these. But characters are not rarely transferred +exclusively to that sex, in which they first appeared. Ample evidence on +this head has been advanced in my work on Variation under +Domestication; but a few instances may here be given. There are breeds +of the sheep and goat, in which the horns of the male differ greatly in +shape from those of the female; and these differences, acquired under +domestication, are regularly transmitted to the same sex. With +tortoise-shell cats the females alone, as a general rule, are thus +coloured, the males being rusty-red. With most breeds of the fowl, the +characters proper to each sex are transmitted to the same sex alone. So +general is this form of transmission that it is an anomaly when we see +in certain breeds variations transmitted equally to both sexes. There +are also certain sub-breeds of the fowl in which the males can hardly be +distinguished from each other, whilst the females differ considerably in +colour. With the pigeon the sexes of the parent-species do not differ in +any external character; nevertheless in certain domesticated breeds the +male is differently coloured from the female.[357] The wattle in the +English Carrier pigeon and the crop in the Pouter are more highly +developed in the male than in the female; and although these characters +have been gained through long-continued selection by man, the difference +between the two sexes is wholly due to the form of inheritance which has +prevailed; for it has arisen, not from, but rather in opposition to, the +wishes of the breeder. + +Most of our domestic races have been formed by the accumulation of many +slight variations; and as some of the successive steps have been +transmitted to one sex alone, and some to both sexes, we find in the +different breeds of the same species all gradations between great sexual +dissimilarity and complete similarity. Instances have already been +given with the breeds of the fowl and pigeon; and under nature analogous +cases are of frequent occurrence. With animals under domestication, but +whether under nature I will not venture to say, one sex may lose +characters proper to it, and may thus come to resemble to a certain +extent the opposite sex; for instance, the males of some breeds of the +fowl have lost their masculine plumes and hackles. On the other hand the +differences between the sexes may be increased under domestication, as +with merino sheep, in which the ewes have lost their horns. Again, +characters proper to one sex may suddenly appear in the other sex; as +with those sub-breeds of the fowl in which the hens whilst young acquire +spurs; or, as in certain Polish sub-breeds, in which the females, as +there is reason to believe, originally acquired a crest, and +subsequently transferred it to the males. All these cases are +intelligible on the hypothesis of pangenesis; for they depend on the +gemmules of certain units of the body, although present in both sexes, +becoming through the influence of domestication dormant in the one sex; +or if naturally dormant, becoming developed. + +There is one difficult question which it will be convenient to defer to +a future chapter; namely, whether a character at first developed in both +sexes, can be rendered through selection limited in its development to +one sex alone. If, for instance, a breeder observed that some of his +pigeons (in which species characters are usually transferred in an equal +degree to both sexes) varied into pale blue; could he by long-continued +selection make a breed, in which the males alone should be of this tint, +whilst the females remained unchanged? I will here only say, that this, +though perhaps not impossible, would be extremely difficult; for the +natural result of breeding from the pale-blue males would be to change +his whole stock, including both sexes, into this tint. If, however, +variations of the desired tint appeared, which were from the first +limited in their development to the male sex, there would not be the +least difficulty in making a breed characterised by the two sexes being +of a different colour, as indeed has been effected with a Belgian breed, +in which the males alone are streaked with black. In a similar manner, +if any variation appeared in a female pigeon, which was from the first +sexually limited in its development, it would be easy to make a breed +with the females alone thus characterised; but if the variation was not +thus originally limited, the process would be extremely difficult, +perhaps impossible. + + +_On the Relation between the period of Development of a Character and +its transmission to one sex or to both sexes._—Why certain characters +should be inherited by both sexes, and other characters by one sex +alone, namely by that sex in which the character first appeared, is in +most cases quite unknown. We cannot even conjecture why with certain +sub-breeds of the pigeon, black striæ, though transmitted through the +female, should be developed in the male alone, whilst every other +character is equally transferred to both sexes. Why, again, with cats, +the tortoise-shell colour should, with rare exceptions, be developed in +the female alone. The very same characters, such as deficient or +supernumerary digits, colour-blindness, &c., may with mankind be +inherited by the males alone of one family, and in another family by the +females alone, though in both cases transmitted through the opposite as +well as the same sex.[358] Although we are thus ignorant, two rules +often hold good, namely that variations which, first appear in either +sex at a late period of life, tend to be developed in the same sex +alone; whilst variations which first appear early in life in either sex +tend to be developed in both sexes. I am, however, far from supposing +that this is the sole determining cause. As I have not elsewhere +discussed this subject, and as it has an important bearing on sexual +selection, I must here enter into lengthy and somewhat intricate +details. + +It is in itself probable that any character appearing at an early age +would tend to be inherited equally by both sexes, for the sexes do not +differ much in constitution, before the power of reproduction is gained. +On the other hand, after this power has been gained and the sexes have +come to differ in constitution, the gemmules (if I may again use the +language of pangenesis) which are cast off from each varying part in the +one sex would be much more likely to possess the proper affinities for +uniting with the tissues of the same sex, and thus becoming developed, +than with those of the opposite sex. + +I was first led to infer that a relation of this kind exists, from the +fact that whenever and in whatever manner the adult male has come to +differ from the adult female, he differs in the same manner from the +young of both sexes. The generality of this fact is quite remarkable: it +holds good with almost all mammals, birds, amphibians, and fishes; also +with many crustaceans, spiders and some few insects, namely certain +orthoptera and libellulæ. In all these cases the variations, through the +accumulation of which the male acquired his proper masculine characters, +must have occurred at a somewhat late period of life; otherwise the +young males would have been similarly characterised; and conformably +with our rule, they are transmitted to and developed in the adult males +alone. When, on the other hand, the adult male closely resembles the +young of both sexes (these, with rare exceptions, being alike), he +generally resembles the adult female; and in most of these cases the +variations through which the young and old acquired their present +characters, probably occurred in conformity with our rule during youth. +But there is here room for doubt, as characters are sometimes +transferred to the offspring at an earlier age than that at which they +first appeared in the parents, so that the parents may have varied when +adult, and have transferred their characters to their offspring whilst +young. There are, moreover, many animals, in which the two sexes closely +resemble each other, and yet both differ from their young; and here the +characters of the adults must have been acquired late in life; +nevertheless, these characters in apparent contradiction to our rule, +are transferred to both sexes. We must not, however, overlook the +possibility or even probability of successive variations of the same +nature sometimes occurring, under exposure to similar conditions, +simultaneously in both sexes at a rather late period of life; and in +this case the variations would be transferred to the offspring of both +sexes at a corresponding late age; and there would be no real +contradiction to our rule of the variations which occur late in life +being transferred exclusively to the sex in which they first appeared. +This latter rule seems to hold true more generally than the second rule, +namely, that variations which occur in either sex early in life tend to +be transferred to both sexes. As it was obviously impossible even to +estimate in how large a number of cases throughout the animal kingdom +these two propositions hold good, it occurred to me to investigate some +striking or crucial instances, and to rely on the result. + +An excellent case for investigation is afforded by the Deer Family. In +all the species, excepting one, the horns are developed in the male +alone, though certainly transmitted through the female, and capable of +occasional abnormal development in her. In the reindeer, on the other +hand, the female is provided with horns; so that in this species, the +horns ought, according to our rule, to appear early in life, long before +the two sexes had arrived at maturity and had come to differ much in +constitution. In all the other species of deer the horns ought to appear +later in life, leading to their development in that sex alone, in which +they first appeared in the progenitor of the whole Family. Now in seven +species, belonging to distinct sections of the family and inhabiting +different regions, in which the stags alone bear horns, I find that the +horns first appear at periods varying from nine months after birth in +the roebuck to ten or twelve or even more months in the stags of the six +other larger species.[359] But with the reindeer the case is widely +different, for as I hear from Prof. Nilsson, who kindly made special +enquiries for me in Lapland, the horns appear in the young animals +within four or five weeks after birth, and at the same time in both +sexes. So that here we have a structure, developed at a most unusually +early age in one species of the family, and common to both sexes in this +one species. + +In several kinds of antelopes the males alone are provided with horns, +whilst in the greater number both sexes have horns. With respect to the +period of development, Mr. Blyth informs me that there lived at one time +in the Zoological Gardens a young koodoo (_Ant. strepsiceros_), in which +species the males alone are horned, and the young of a closely-allied +species, viz. the eland (_Ant. oreas_), in which both sexes are horned. +Now in strict conformity with our rule, in the young male koodoo, +although arrived at the age of ten months, the horns were remarkably +small considering the size ultimately attained by them: whilst in the +young male eland, although only three months old, the horns were already +very much larger than in the koodoo. It is also worth notice that in the +prong-horned antelope,[360] in which species the horns, though present +in both sexes, are almost rudimentary in the female, they do not appear +until about five or six months after birth. With sheep, goats, and +cattle, in which the horns are well developed in both sexes, though not +quite equal in size, they can be felt, or even seen, at birth or soon +afterwards.[361] Our rule, however, fails in regard to some breeds of +sheep, for instance merinos, in which the rams alone are horned; for I +cannot find on enquiry,[362] that the horns are developed later in life +in this breed than in ordinary sheep in which both sexes are horned. But +with domesticated sheep the presence or absence of horns is not a firmly +fixed character; a certain proportion of the merino ewes bearing small +horns, and some of the rams being hornless; whilst with ordinary sheep +hornless ewes are occasionally produced. + +In most of the species of the splendid family of the Pheasants, the +males differ conspicuously from the females, and they acquire their +ornaments at a rather late period of life. The eared pheasant +(_Crossoptilon auritum_), however, offers a remarkable exception, for +both sexes possess the fine caudal plumes, the large ear-tufts and the +crimson velvet about the head; and I find on enquiry in the Zoological +Gardens that all these characters, in accordance with our rule, appear +very early in life. The adult male can, however, be distinguished from +the adult female by one character, namely by the presence of spurs; and +conformably with our rule, these do not begin to be developed, as I am +assured by Mr. Bartlett, before the age of six months, and even at this +age, can hardly be distinguished in the two sexes.[363] The male and +female Peacock differ conspicuously from each other in almost every +part of their plumage, except in the elegant head-crest, which is common +to both sexes; and this is developed very early in life, long before the +other ornaments which are confined to the male. The wild-duck offers an +analogous case, for the beautiful green speculum on the wings is common +to both sexes, though duller and somewhat smaller in the female, and it +is developed early in life, whilst the curled tail-feathers and other +ornaments peculiar to the male are developed later.[364] Between such +extreme cases of close sexual resemblance and wide dissimilarity, as +those of the Crossoptilon and peacock, many intermediate ones could be +given, in which the characters follow in their order of development our +two rules. + +As most insects emerge from their pupal state in a mature condition, it +is doubtful whether the period of development determines the +transference of their characters to one or both sexes. But we do not +know that the coloured scales, for instance, in two species of +butterflies, in one of which the sexes differ in colour, whilst in the +other they are alike, are developed at the same relative age in the +cocoon. Nor do we know whether all the scales are simultaneously +developed on the wings of the same species of butterfly, in which +certain coloured marks are confined to one sex, whilst other marks are +common to both sexes. A difference of this kind in the period of +development is not so improbable as it may at first appear; for with the +Orthoptera, which assume their adult state, not by a single +metamorphosis, but by a succession of moults, the young males of some +species at first resemble the females, and acquire their distinctive +masculine characters only during a later moult. Strictly analogous cases +occur during the successive moults of certain male crustaceans. + +We have as yet only considered the transference of characters, +relatively to their period of development, with species in a natural +state; we will now turn to domesticated animals; first touching on +monstrosities and diseases. The presence of supernumerary digits, and +the absence of certain phalanges, must be determined at an early +embryonic period—the tendency to profuse bleeding is at least +congenital, as is probably colour-blindness—yet these peculiarities, +and other similar ones, are often limited in their transmission to one +sex; so that the rule that characters which are developed at an early +period tend to be transmitted to both sexes, here wholly fails. But this +rule, as before remarked, does not appear to be nearly so generally true +as the converse proposition, namely, that characters which appear late +in life in one sex are transmitted exclusively to the same sex. From the +fact of the above abnormal peculiarities becoming attached to one sex, +long before the sexual functions are active, we may infer that there +must be a difference of some kind between the sexes at an extremely +early age. With respect to sexually-limited diseases, we know too little +of the period at which they originate, to draw any fair conclusion. +Gout, however, seems to fall under our rule; for it is generally caused +by intemperance after early youth, and is transmitted from the father to +his sons in a much more marked manner than to his daughters. + +In the various domestic breeds of sheep, goats, and cattle, the males +differ from their respective females in the shape or development of +their horns, forehead, mane, dewlap, tail, and hump on the shoulders; +and these peculiarities, in accordance with our rule, are not fully +developed until rather late in life. With dogs, the sexes do not differ, +except that in certain breeds, especially in the Scotch deer-hound, the +male is much larger and heavier than the female; and as we shall see in +a future chapter, the male goes on increasing in size to an unusually +late period of life, which will account, according to our rule, for his +increased size being transmitted to his male offspring alone. On the +other hand, the tortoise-shell colour of the hair, which is confined to +female cats, is quite distinct at birth, and this case violates our +rule. There is a breed of pigeons in which the males alone are streaked +with black, and the streaks can be detected even in the nestlings; but +they become more conspicuous at each successive moult, so that this case +partly opposes and partly supports the rule. With the English Carrier +and Pouter pigeon the full development of the wattle and the crop occurs +rather late in life, and these characters, conformably with our rule, +are transmitted in full perfection to the males alone. The following +cases perhaps come within the class previously alluded to, in which the +two sexes have varied in the same manner at a rather late period of +life, and have consequently transferred their new characters to both +sexes at a corresponding late period; and if so, such cases are not +opposed to our rule. Thus there are sub-breeds of the pigeon, described +by Neumeister,[365] both sexes of which change colour after moulting +twice or thrice, as does likewise the Almond Tumbler; nevertheless these +changes, though occurring rather late in life, are common to both sexes. +One variety of the Canary-bird, namely the London Prize, offers a nearly +analogous case. + +With the breeds of the Fowl the inheritance of various characters by one +sex or by both sexes, seems generally determined by the period at which +such characters are developed. Thus in all the many breeds in which the +adult male differs greatly in colour from the female and from the adult +male parent-species, he differs from the young male, so that the newly +acquired characters must have appeared at a rather late period of life. +On the other hand with most of the breeds in which the two sexes +resemble each other, the young are coloured in nearly the same manner as +their parents, and this renders it probable that their colours first +appeared early in life. We have instances of this fact in all black and +white breeds, in which the young and old of both sexes are alike; nor +can it be maintained that there is something peculiar in a black or +white plumage, leading to its transference to both sexes; for the males +alone of many natural species are either black or white, the females +being very differently coloured. With the so-called Cuckoo sub-breeds of +the fowl, in which the feathers are transversely pencilled with dark +stripes, both sexes and the chickens are coloured in nearly the same +manner. The laced plumage of the Sebright bantam is the same in both +sexes, and in the chickens the feathers are tipped with black, which +makes a near approach to lacing. Spangled Hamburghs, however, offer a +partial exception, for the two sexes, though not quite alike, resemble +each other more closely than do the sexes of the aboriginal +parent-species, yet they acquire their characteristic plumage late in +life, for the chickens are distinctly pencilled. Turning to other +characters besides colour: the males alone of the wild parent-species +and of most domestic breeds possess a fairly well developed comb, but in +the young of the Spanish fowl it is largely developed at a very early +age, and apparently in consequence of this it is of unusual size in the +adult females. In the Game breeds pugnacity is developed at a +wonderfully early age, of which curious proofs could be given; and this +character is transmitted to both sexes, so that the hens, from their +extreme pugnacity, are now generally exhibited in separate pens. With +the Polish breeds the bony protuberance of the skull which supports the +crest is partially developed even before the chickens are hatched, and +the crest itself soon begins to grow, though at first feebly;[366] and +in this breed a great bony protuberance and an immense crest +characterise the adults of both sexes. + +Finally, from what we have now seen of the relation which exists in many +natural species and domesticated races, between the period of the +development of their characters and the manner of their +transmission—for example the striking fact of the early growth of the +horns in the reindeer, in which both sexes have horns, in comparison +with their much later growth in the other species in which the male +alone bears horns—we may conclude that one cause, though not the sole +cause, of characters being exclusively inherited by one sex, is their +development at a late age. And secondly, that one, though apparently a +less efficient, cause of characters being inherited by both sexes is +their development at an early age, whilst the sexes differ but little in +constitution. It appears, however, that some difference must exist +between the sexes even during an early embryonic period, for characters +developed at this age not rarely become attached to one sex. + + +_Summary and concluding remarks._—From the foregoing discussion on the +various laws of inheritance, we learn that characters often or even +generally tend to become developed in the same sex, at the same age, and +periodically at the same season of the year, in which they first +appeared in the parents. But these laws, from unknown causes, are very +liable to change. Hence the successive steps in the modification of a +species might readily be transmitted in different ways; some of the +steps being transmitted to one sex, and some to both; some to the +offspring at one age, and some at all ages. Not only are the laws of +inheritance extremely complex, but so are the causes which induce and +govern variability. The variations thus caused are preserved and +accumulated by sexual selection, which is in itself an extremely complex +affair, depending, as it does, on ardour in love, courage, and the +rivalry of the males, and on the powers of perception, taste, and will +of the female. Sexual selection will also be dominated by natural +selection for the general welfare of the species. Hence the manner in +which the individuals of either sex or of both sexes are affected +through sexual selection cannot fail to be complex in the highest +degree. + +When variations occur late in life in one sex, and are transmitted to +the same sex at the same age, the other sex and the young are +necessarily left unmodified. When they occur late in life, but are +transmitted to both sexes at the same age, the young alone are left +unmodified. Variations, however, may occur at any period of life in one +sex or in both, and be transmitted to both sexes at all ages, and then +all the individuals of the species will be similarly modified. In the +following chapters it will be seen that all these cases frequently occur +under nature. + +Sexual selection can never act on any animal whilst young, before the +age for reproduction has arrived. From the great eagerness of the male +it has generally acted on this sex and not on the females. The males +have thus become provided with weapons for fighting with their rivals, +or with organs for discovering and securely holding the female, or for +exciting and charming her. When the sexes differ in these respects, it +is also, as we have seen, an extremely general law that the adult male +differs more or less from the young male; and we may conclude from this +fact that the successive variations, by which the adult male became +modified, cannot have occurred much before the age for reproduction. How +then are we to account for this general and remarkable coincidence +between the period of variability and that of sexual selection,— +principles which are quite independent of each other? I +think we can see the cause: it is not that the males have never varied +at an early age, but that such variations have commonly been lost, +whilst those occurring at a later age have been preserved. + +All animals produce more offspring than can survive to maturity; and we +have every reason to believe that death falls heavily on the weak and +inexperienced young. If then a certain proportion of the offspring were +to vary at birth or soon afterwards, in some manner which at this age +was of no service to them, the chance of the preservation of such +variations would be small. We have good evidence under domestication how +soon variations of all kinds are lost, if not selected. But variations +which occurred at or near maturity, and which were of immediate service +to either sex, would probably be preserved; as would similar variations +occurring at an earlier period in any individuals which happened to +survive. As this principle has an important bearing on sexual selection, +it may be advisable to give an imaginary illustration. We will take a +pair of animals, neither very fertile nor the reverse, and assume that +after arriving at maturity they live on an average for five years, +producing each year five young. They would thus produce 25 offspring; +and it would not, I think, be an unfair estimate to assume that 18 or 20 +out of the 25 would perish before maturity, whilst still young and +inexperienced; the remaining seven or five sufficing to keep up the +stock of mature individuals. If so, we can see that variations which +occurred during youth, for instance in brightness, and which were not of +the least service to the young, would run a good chance of being utterly +lost. Whilst similar variations, which occurring at or near maturity in +the comparatively few individuals surviving to this age, and which +immediately gave an advantage to certain males, by rendering them more +attractive to the females, would be likely to be preserved. No doubt +some of the variations in brightness which occurred at an earlier age +would by chance be preserved, and eventually give to the male the same +advantage as those which appeared later; and this will account for the +young males commonly partaking to a certain extent (as may be observed +with many birds) of the bright colours of their adult male parents. If +only a few of the successive variations in brightness were to occur at a +late age, the adult male would be only a little brighter than the young +male; and such cases are common. + +In this illustration I have assumed that the young varied in a manner +which was of no service to them; but many characters proper to the adult +male would be actually injurious to the young,—as bright colours from +making them conspicuous, or horns of large size from expending much +vital force. Such variations in the young would promptly be eliminated +through natural selection. With the adult and experienced males, on the +other hand, the advantage thus derived in their rivalry with other males +would often more than counterbalance exposure to some degree of danger. +Thus we can understand how it is that variations which must originally +have appeared rather late in life have alone or in chief part been +preserved for the development of secondary sexual characters; and the +remarkable coincidence between the periods of variability and of sexual +selection is intelligible. + +As variations which give to the male an advantage in lighting with other +males, or in finding, securing, or charming the female, would be of no +use to the female, they will not have been preserved in this sex either +during youth or maturity. Consequently such variations would be +extremely liable to be lost; and the female, as far as these characters +are concerned, would be left unmodified, excepting in so far as she may +have received them by transference from the male. No doubt if the female +varied and transferred serviceable characters to her male offspring, +these would be favoured through sexual selection; and then both sexes +would thus far be modified in the same manner. But I shall hereafter +have to recur to these more intricate contingencies. + +In the following chapters, I shall treat of the secondary sexual +characters in animals of all classes, and shall endeavour in each case +to apply the principles explained in the present chapter. The lowest +classes will detain us for a very short time, but the higher animals, +especially birds, must be treated at considerable length. It should be +borne in mind that for reasons already assigned, I intend to give only a +few illustrative instances of the innumerable structures by the aid of +which the male finds the female, or, when found, holds her. On the other +hand, all structures and instincts by which the male conquers other +males, and by which he allures or excites the female, will be fully +discussed, as these are in many ways the most interesting. + + +_Supplement on the proportional numbers of the two sexes in animals +belonging to various classes._ + + +As no one, as far as I can discover, has paid attention to the relative +numbers of the two sexes throughout the animal kingdom, I will here give +such materials as I have been able to collect, although they are +extremely imperfect. They consist in only a few instances of actual +enumeration, and the numbers are not very large. As the proportions are +known with certainty on a large scale in the case of man alone, I will +first give them, as a standard of comparison. + + +_Man._—In England during ten years (from 1857 to 1866) 707,120 children +on an annual average have been born alive, in the proportion of 104.5 +males to 100 females. But in 1857 the male births throughout England +were as 105.2, and in 1865 as 104.0 to 100. Looking to separate +districts, in Buckinghamshire (where on an average 5000 children are +annually born) the _mean_ proportion of male to female births, during +the whole period of the above ten years, was as 102.8 to 100; whilst in +N. Wales (where the average annual births are 12,873) it was as high as +106.2 to 100. Taking a still smaller district, viz., Rutlandshire (where +the annual births average only 739), in 1864 the male births were as +114.6, and in 1862 as 97.0 to 100; but even in this small district the +average of the 7385 births during the whole ten years was as 104.5 to +100; that is in the same ratio as throughout England.[367] The +proportions are sometimes slightly disturbed by unknown causes; thus +Prof. Faye states “that in some districts of Norway there has been +during a decennial period a steady deficiency of boys, whilst in others +the opposite condition has existed.” In France during forty-four years +the male to the female births have been as 106.2 to 100; but during this +period it has occurred five times in one department, and six times in +another, that the female births have exceeded the males. In Russia the +average proportion is as high as 108.9 to 100.[368] It is a singular +fact that with Jews the proportion of male births is decidedly larger +than with Christians: thus in Prussia the proportion is as 113, in +Breslau as 114, and in Livonia as 120 to 100; the Christian births in +these countries being the same as usual, for instance, in Livonia as 104 +to 100.[369] It is a still more singular fact that in different nations, +under different conditions and climates, in Naples, Prussia, Westphalia, +France and England, the excess of male over female births is less when +they are illegitimate than when legitimate.[370] + +In various parts of Europe, according to Prof. Faye and other authors, +“a still greater preponderance of males would be met with, if death +struck both sexes in equal proportion in the womb and during birth. But +the fact is, that for every 100 still-born females, we have in several +countries from 134.6 to 144.9 still-born males.” Moreover during the +first four or five years of life more male children die than females; +“for example in England, during the first year, 126 boys die +for every 100 girls,—a proportion which in France is still more +unfavourable.”[371] As a consequence of this excess in the death-rate of +male children, and of the exposure of men when adult to various dangers, +and of their tendency to emigrate, the females in all old-settled +countries, where statistical records have been kept,[372] are found to +preponderate considerably over the males. + +It has often been supposed that the relative ages of the parents +determine the sex of the offspring; and Prof. Leuckart[373] has advanced +what he considers sufficient evidence, with respect to man and certain +domesticated animals, to shew that this is one important factor in the +result. So again the period of impregnation has been thought to be the +efficient cause; but recent observations discountenance this belief. +Again, with mankind polygamy has been supposed to lead to the birth of a +greater proportion of female infants; but Dr. J. Campbell[374] carefully +attended to this subject in the harems of Siam, and he concludes that +the proportion of male to female births is the same as from monogamous +unions. Hardly any animal has been rendered so highly polygamous as our +English race-horses, and we shall immediately see that their male and +female offspring are almost exactly equal in number. + + + _Horses._—Mr. Tegetmeier has been so kind as to tabulate for + me from the ‘Racing Calendar’ the births of race-horses during + a period of twenty-one years, viz. from 1846 to 1867; 1849 + being omitted, as no returns were that year published. The + total births have been 25,560,[375] consisting of 12,763 males + and 12,797 females, or in the proportion of 99.7 males to 100 + females. As these numbers are tolerably large, and as they are + drawn from all parts of England, during several years, we may + with much confidence conclude that with the domestic horse, or + at least with the race-horse, the two sexes are produced in + almost equal numbers. The fluctuations in the proportions + during successive years are closely like those which occur with + mankind, when a small and thinly-populated area is considered: + thus in 1856 the male horses were as 107.1, and in 1867 as only + 92.6 to 100 females. In the tabulated returns the proportions + vary in cycles, for the males exceeded the females during six + successive years; and the females exceeded the males during two + periods each of four years: this, however, may be accidental; + at least I can detect nothing of the kind with man in the + decennial table in the Registrar’s Report for 1866. I may add + that certain, mares, and this holds good with certain cows and + with women, tend to produce more of one sex than of the other; + Mr. Wright of Yeldersley House, informs me that one of his Arab + mares, though put seven times to different horses, produced + seven fillies. + + + _Dogs._—During a period of twelve years, from 1857 to 1868, + the births of a large number of greyhounds, throughout England, + have been sent to the ‘Field’ newspaper; and I am again + indebted to Mr. Tegetmeier for carefully tabulating the + results. The recorded, births have been 6878, consisting of + 3605 males and 3273 females, that is, in the proportion of + 110.1 males to 100 females. The greatest fluctuations occurred + in 1864, when the proportion was as 95.3 males, and in 1867, as + 116.3 males to 100 females. The above average proportion of + 110.1 to 100 is probably nearly correct in the case of the + greyhound, but whether it would hold with other domesticated + breeds is in some degree doubtful. Mr. Cupples has enquired + from several great breeders of dogs, and finds that all without + exception believe that females are produced in excess; he + suggests that this belief may have arisen from females being + less valued and the consequent disappointment producing a + stronger impression on the mind. + + + _Sheep._—The sexes of sheep are not ascertained by + agriculturists until several months after birth, at the period + when the males are castrated; so that the following returns do + not give the proportions at birth. Moreover, I find that + several great breeders in Scotland, who annually raise some + thousand sheep, are firmly convinced that a larger proportion + of males than of females die during the first one or two years; + therefore the proportion of males would be somewhat greater at + birth than at the age of castration. This is a remarkable + coincidence with what occurs, as we have seen, with mankind, + and both cases probably depend on some common cause. I have + received returns from four gentlemen in England who have bred + lowland sheep, chiefly Leicesters, during the last ten or + sixteen years; they amount altogether to 8965 births, + consisting of 4407 males and 4558 females; that is in the + proportion of 96.7 males to 100 females. With respect to + Cheviot and black-faced sheep bred in Scotland, I have received + returns from six breeders, two of them on a large scale, + chiefly for the years 1867-1869, but some of the returns + extending back to 1862. The total number recorded amounts to + 50,685, consisting of 25,071 males and 25,614 females, or in + the proportion of 97.9 males to 100 females. If we take the + English and Scotch returns together, the total number amounts + to 59,650, consisting of 29,478 males and 30,172 females, or + as 97·7 to 100. So that with sheep at the age of castration the + females are certainly in excess of the males; but whether this + would hold good at birth is doubtful, owing to the greater + liability in the males to early death.[376] + + Of _Cattle_ I have received returns from nine gentlemen of 982 + births, too few to be trusted; these consisted of 477 + bull-calves and 505 cow-calves; _i.e._ in the proportion of + 94·4 males to 100 females. The Rev. W. D. Fox informs me that + in 1867 out of 34 calves born on a farm in Derbyshire only one + was a bull. Mr. Harrison Weir writes to me that he has enquired + from several breeders of _Pigs_, and most of them estimate the + male to the female births as about 7 to 6. This same gentleman + has bred _Rabbits_ for many years, and has noticed that a far + greater number of bucks are produced than does. + + Of mammalia in a state of nature I have been able to learn very + little. In regard to the common rat, I have received + conflicting statements. Mr. R. Elliot of Laighwood, informs me + that a rat-catcher assured him that he had always found the + males in great excess, even with the young in the nest. In + consequence of this, Mr. Elliot himself subsequently examined + some hundred old ones, and found the statement true. Mr. F. + Buckland has bred a large number of white rats, and he also + believes that the males greatly exceed the females. In regard + to Moles, it is said that “the males are much more numerous + than the females;”[377] and as the catching of these animals is + a special occupation, the statement may perhaps be trusted. Sir + A. Smith, in describing an antelope of S. Africa[378] (_Kobus + ellipsiprymnus_), remarks, that in the herds of this and other + species, the males are few in number compared with the females: + the natives believe that they are born in this proportion; + others believe that the younger males are expelled from the + herds, and Sir A. Smith says, that though he has himself never + seen herds consisting of young males alone, others affirm that + this does occur. It appears probable that the young males when + expelled from the herd, would be likely to fell a prey to the + many beasts of prey of the country. + + + BIRDS. + + With respect to the _Fowl_, I have received only one account, + namely, that out of 1001 chickens of a highly-bred stock of + Cochins, reared during eight years by Mr. Stretch, 487 proved + males and 514 females: _i.e._ as 94.7 to 100. In regard to + domestic pigeons there is good evidence that the males are + produced in excess, or that their lives are longer; for these + birds invariably pair, and single males, as Mr. Tegetmeier + informs me, can always be purchased cheaper than females. + Usually the two birds reared from the two eggs laid in the same + nest consist of a male and female; but Mr. Harrison Weir, who + has been so large a breeder, says that he has often bred two + cocks from the same nest, and seldom two hens; moreover the hen + is generally the weaker of the two, and more liable to perish. + + With respect to birds in a state of nature, Mr. Gould and + others[379] are convinced that the males are generally the more + numerous; and as the young males of many species resemble the + females, the latter would naturally appear to be the most + numerous. Large numbers of pheasants are reared by Mr. Baker of + Leadenhall from eggs laid by wild birds, and he informs Mr. + Jenner Weir that four or five males to one female are generally + produced. An experienced observer remarks[380] that in + Scandinavia the broods of the capercailzie and black-cock + contain more males than females; and that with the Dal-ripa (a + kind of ptarmigan) more males than females attend the _leks_ or + places of courtship; but this latter circumstance is accounted + for by some observers by a greater number of hen birds being + killed by vermin. From various facts given by White of + Selbourne,[381] it seems clear that the males of the partridge + must be in considerable excess in the south of England; and I + have been assured that this is the case in Scotland. Mr. Weir + on enquiring from the dealers who receive at certain seasons + large numbers of ruffs (_Machetes pugnax_) was told that the + males are much the most numerous. This same naturalist has also + enquired for me from the bird-catchers, who annually catch an + astonishing number of various small species alive for the + London market, and he was unhesitatingly answered by an old and + trustworthy man, that with the chaffinch the males are in large + excess; he thought as high as 2 males to 1 female, or at least + as high as 5 to 3.[382] The males of the blackbird, he likewise + maintained, were by far the most numerous, whether caught by + traps or by netting at night. These statements may apparently + be trusted, because the same man said that the sexes are about + equal with the lark, the twite (_Linaria montana_), and + goldfinch. On the other hand he is certain that with the common + linnet, the females preponderate greatly, but unequally during + different years; during some years he has found the females to + the males as four to one. It should, however, be borne in mind, + that the chief season for catching birds does not begin till + September, so that with some species partial migrations may + have begun, and the flocks at this period often consist of hens + alone. Mr. Salvin paid particular attention to the sexes of the + humming-birds in Central America, and he is convinced that with + most of the species the males are in excess; thus one year he + procured 204 specimens belonging to ten species, and these + consisted of 166 males and of 38 females. With two other + species the females were in excess: but the proportions + apparently vary either during different seasons or in different + localities; for on one occasion the males of _Campylopterus + hemileucurus_ were to the females as five to two, and on + another occasion[383] in exactly the reversed ratio. As bearing + on this latter point, I may add, that Mr. Powys found in Corfu + and Epirus the sexes of the chaffinch keeping apart, and “the + females by far the most numerous;” whilst in Palestine Mr. + Tristram found “the male flocks appearing greatly to exceed the + female in number.”[384] So again with the _Quiscalus major_, + Mr. G. Taylor[385] says, that in Florida there were “very few + females in proportion to the males,” whilst in Honduras the + proportion was the other way, the species there having the + character of a polygamist. + + + FISH. + + With Fish the proportional numbers of the sexes can be + ascertained only by catching them in the adult or nearly adult + state; and there are many difficulties in arriving at any just + conclusion.[386] Infertile females might readily be mistaken + for males, as Dr. Günther has remarked to me in regard to + trout. With some species the males are believed to die soon + after fertilising the ova. With many species the males are of + much smaller size than the females, so that a large number of + males would escape from the same net by which the females were + caught. M. Carbonnier,[387] who has especially attended to the + natural history of the pike (_Esox lucius_) states that many + males, owing to their small size, are devoured by the larger + females; and he believes that the males of almost all fish are + exposed from the same cause to greater danger than the females. + Nevertheless in the few cases in which the proportional numbers + have been actually observed, the males appear to be largely in + excess. Thus Mr. R. Buist, the superintendent of the + Stormontfield experiments, says that in 1865, out of 70 salmon + first landed for the purpose of obtaining the ova, upwards of + 60 were males. In 1867 he again “calls attention to the vast + disproportion of the males to the females. We had at the outset + at least ten males to one female.” Afterwards sufficient + females for obtaining ova were procured. He adds, “from the + great proportion of the males, they are constantly fighting and + tearing each other on the spawning-beds.”[388] This + disproportion, no doubt, can be accounted for in part, but + whether wholly is very doubtful, by the males ascending the + rivers before the females. Mr. F. Buckland remarks in regard to + trout, that “it is a curious fact that the males preponderate + very largely in number over the females. It _invariably_ + happens that when the first rush of fish is made to the net, + there will be at least seven or eight males to one female found + captive. I cannot quite account for this; either the males are + more numerous than the females, or the latter seek safety by + concealment rather than flight.” He then adds, that by + carefully searching the banks, sufficient females for obtaining + ova can be found.[389] Mr. H. Lee informs me that out of 212 + trout, taken for this purpose in Lord Portsmouth’s park, 150 + were males and 62 females. + + With the Cyprinidæ the males likewise seem to be in excess; but + several members of this Family, viz., the carp, tench, bream + and minnow, appear regularly to follow the practice, rare in + the animal kingdom, of polyandry; for the female whilst + spawning is always attended by two males, one on each side, and + in the case of the bream by three or four males. This fact is + so well known, that it is always recommended to stock a pond + with two male tenches to one female, or at least with three + males to two females. With the minnow, an excellent observer + states, that on the spawning-beds the males are ten times as + numerous as the females; when a female comes amongst the males, + “she is immediately pressed closely by a male on each side; and + when they have been in that situation for a time, are + superseded by other two males.”[390] + + + INSECTS. + + In this class, the Lepidoptera alone afford the means of + judging of the proportional numbers of the sexes; for they have + been collected with special care by many good observers, and + have been largely bred from the egg or caterpillar state. I had + hoped that some breeders of silk-moths might have kept an exact + record, but after writing to France and Italy, and consulting + various treatises, I cannot find that this has ever been done. + The general opinion appears to be that the sexes are nearly + equal, but in Italy as I hear from Professor Canestrini, many + breeders are convinced that the females are produced in excess. + The same naturalist, however, informs me, that in the two + yearly broods of the Ailanthus silk-moth (_Bombyx cynthia_), + the males greatly preponderate in the first, whilst in the + second the two sexes are nearly equal, or the females rather in + excess. + + In regard to Butterflies in a state of nature, several + observers have been much struck by the apparently enormous + preponderance of the males.[391] Thus Mr. Bates,[392] in + speaking of the species, no less than about a hundred in + number, which inhabit the Upper Amazons, says that the males + are much more numerous than the females, even in the proportion + of a hundred to one. In North America, Edwards, who had great + experience, estimates in the genus Papilio the males to the + females as four to one; and Mr. Walsh, who informed me of this + statement, says that with _P. turnus_ this is certainly the + case. In South Africa, Mr. R. Trimen found the males in excess + in 19 species;[393] and in one of these, which swarms in open + places, he estimated the number of males as fifty to one + female. With another species, in which the males are numerous + in certain localities, he collected during seven years only + five females. In the island of Bourbon, M. Maillard states that + the males of one species of Papilio are twenty times as + numerous as the females.[394] Mr. Trimen informs me that as far + as he has himself seen, or heard from others, it is rare for + the females of any butterfly to exceed in number the males; but + this is perhaps the case with three South African species. Mr. + Wallace[395] states that the females of _Ornithoptera + crœsus_, in the Malay archipelago, are more common and more + easily caught than the males; but this is a rare butterfly. I + may here add, that in Hyperythra, a genus of moths, Guenée + says, that from four to five females are sent in collections + from India for one male. + + When this subject of the proportional numbers of the sexes of + insects was brought before the Entomological Society,[396] it + was generally admitted that the males of most Lepidoptera, in + the adult or imago state, are caught in greater numbers than + the females; but this fact was attributed by various observers + to the more retiring habits of the females, and to the males + emerging earlier from the cocoon. This latter circumstance is + well known to occur with most Lepidoptera, as well as with + other insects. So that, as M. Personnat remarks, the males of + the domesticated _Bombyx yamamai_, are lost at the beginning of + the season, and the females at the end, from the want of + mates.[397] I cannot however persuade myself that these causes + suffice to explain the great excess of males in the cases, + above given, of butterflies which are extremely common in their + native countries. Mr. Stainton, who has paid such close + attention during many years to the smaller moths, informs me + that when he collected them in the imago state, he thought that + the males were ten times as numerous as the females, but that + since he has reared them on a large scale from the caterpillar + state, he is convinced that the females are the most numerous. + Several entomologists concur in this view. Mr. Doubleday, + however, and some others, take an opposite view, and are + convinced that they have reared from the egg and caterpillar + states a larger proportion of males than of females. + + Besides the more active habits of the males, their earlier + emergence from the cocoon, and their frequenting in some cases + more open stations, other causes may be assigned for an + apparent or real difference in the proportional numbers of the + sexes of Lepidoptera, when captured in the imago state, and + when reared from the egg or caterpillar state. It is believed + by many breeders in Italy, as I hear from Professor Canestrini, + that the female caterpillar of the silk-moth suffers more from + the recent disease than the male; and Dr. Staudinger informs me + that in rearing Lepidoptera more females die in the cocoon than + males. With many species the female caterpillar is larger than + the male, and a collector would naturally choose the finest + specimens, and thus unintentionally collect a larger number of + females. Three collectors have told me that this was their + practice; but Dr. Wallace is sure that most collectors take all + the specimens which they can find of the rarer kinds, which + alone are worth the trouble of rearing. Birds when surrounded + by caterpillars would probably devour the largest; and + Professor Canestrini informs me that in Italy some breeders + believe, though on insufficient evidence, that in the first + brood of the Ailanthus silk-moth, the wasps destroy a larger + number of the female than of the male caterpillars. Dr. Wallace + further remarks that female caterpillars, from being larger + than the males, require more time for their development and + consume more food and moisture; and thus they would be exposed + during a longer time to danger from ichneumons, birds, &c., and + in times of scarcity would perish in greater numbers. Hence it + appears quite possible that, in a state of nature, fewer female + Lepidoptera may reach maturity than males; and for our special + object we are concerned with the numbers at maturity, when the + sexes are ready to propagate their kind. + + The manner in which the males of certain moths congregate in + extraordinary numbers round a single female, apparently + indicates a great excess of males, though this fact may perhaps + be accounted for by the earlier emergence of the males from + their cocoons. Mr. Stainton informs me that from twelve to + twenty males may often be seen congregated round a female + _Elachista rufocinerea_. It is well known that if a virgin + _Lasiocampa quercus_ or _Saturnia carpini_ be exposed in a + cage, vast numbers of males collect round her, and if confined + in a room will even come down the chimney to her. Mr. + Doubleday believes that he has seen from fifty to a hundred + males of both these species attracted in the course of a single + day by a female under confinement. Mr. Trimen exposed in the + Isle of Wight a box in which a female of the Lasiocampa had + been confined on the previous day, and five males soon + endeavoured to gain admittance. M. Verreaux, in Australia, + having placed the female of a small Bombyx in a box in his + pocket, was followed by a crowd of males, so that about 200 + entered the house with him.[398] + + Mr. Doubleday has called my attention to Dr. Staudinger’s[399] + list of Lepidoptera, which gives the prices of the males and + females of 300 species or well-marked varieties of + (Rhopalocera) butterflies. The prices for both sexes of the + very common species are of course the same; but with 114 of the + rarer species they differ; the males being in all cases, + excepting one, the cheapest. On an average of the prices of the + 113 species, the price of the male to that of the female is as + 100 to 149; and this apparently indicates that inversely the + males exceed the females in number in the same proportion. + About 2000 species or varieties of moths (Heterocera) are + catalogued, those with wingless females being here excluded on + account of the difference in habits of the two sexes: of these + 2000 species, 141 differ in price according to sex, the males + of 130 being cheaper, and the males of only 11 being dearer + than the females. The average price of the males of the 130 + species, to that of the females, is as 100 to 143. With respect + to the butterflies in this priced list, Mr. Doubleday thinks + (and no man in England has had more experience), that there is + nothing in the habits of the species which can account for the + difference in the prices of the two sexes, and that it can be + accounted for only by an excess in the numbers of the males. + But I am bound to add that Dr. Staudinger himself, as he + informs me, is of a different opinion. He thinks that the less + active habits of the females and the earlier emergence of the + males will account for his collectors securing a larger number + of males than of females, and consequently for the lower prices + of the former With respect to specimens reared from the + caterpillar-state, Dr. Staudinger believes, as previously + stated, that a greater number of females than of males die + under confinement in the cocoons. He adds that with certain + species one sex seems to preponderate over the other during + certain years. + + Of direct observations on the sexes of Lepidoptera, reared + either from eggs or caterpillars, I have received only the few + following cases:— + + ——————————————————————————————————————————————————————————————————— + Males. | Females. + The Rev. J. Hellins[400] of Exeter reared, during | + 1868, imagos of 73 species, which consisted of 153 | 137 + | + Mr. Albert Jones of Eltham reared, during 1868, | + imagos of 9 species, which, consisted of 159 | 126 + | + During 1869 he reared imagos from 4 species, | + consisting of 114 | 112 + | + Mr. Buckler of Emsworth, Hants, during 1869, | + reared imagos from 74 species, consisting of 180 | 169 + | + Dr. Wallace of Colchester reared from one brood | + of Bombyx cynthia 52 | 48 + | + Dr. Wallace raised, from cocoons of Bombyx | + Pernyi sent from China, during 1869 224 | 123 + | + Dr. Wallace raised, during 1868 and 1869, | + from two lots of cocoons of Bombyx yamamai 52 | 46 + ———————+—————— + Total 934 | 761 + —————————————————————————————————————————————————————————+————————— + + So that in these eight lots of cocoons and eggs, males were + produced in excess. Taken together the proportion of males is + as 122.7 to 100 females. But the numbers are hardly large + enough to be trustworthy. + + On the whole, from the above various sources of evidence, all + pointing to the same direction, I infer that with most species + of Lepidoptera, the males in the imago state generally exceed + the females in number, whatever the proportions may be at their + first emergence from the egg. + + With reference to the other Orders of insects, I have been able + to collect very little reliable information. With the + stag-beetle (_Lucanus cervus_) “the males appear to be much + more numerous than the females;” but when, as Cornelius + remarked during 1867, an unusual number of these beetles + appeared in one part of Germany, the females appeared to exceed + the males as six so one. With one of the Elateridæ, the males + are said to be much more numerous than the females, and “two or + three are often found united with one female;”[401] so that here + polyandry seems to prevail. With Siagonium (Staphylinidæ), in + which the males are furnished with horns, “the females are far + more numerous than the opposite sex.” Mr. Janson stated at the + Entomological Society that the females of the bark-feeding + _Tomicus villosus_ are so common as to be a plague, whilst the + males are so rare as to be hardly known. In other Orders, from + unknown causes, but apparently in some instances owing to + parthenogenesis, the males of certain species have never been + discovered or are excessively rare, as with several of the + Cynipidæ.[402] In all the gall-making Cynipidæ known to Mr. + Walsh, the females are four or five times as numerous as the + males; and so it is, as he informs me, with the gall-making + Cecidomyiiæ (Diptera). With some common species of Saw-flies + (Tenthredinæ) Mr. F. Smith has reared hundreds of specimens + from larvæ of all sizes, but has never reared a single male: on + the other hand Curtis says,[403] that with certain species + (Athalia), bred by him, the males to the females were as six to + one; whilst exactly the reverse occurred with the mature + insects of the same species caught in the fields. With the + Neuroptera, Mr. Walsh states that in many, but by no means in + all, the species of the Odonatous groups (Ephemerina), there is + a great overplus of males: in the genus Hetærina, also, the + males are generally at least four times as numerous as the + females. In certain species in the genus Gomphus the males are + equally numerous, whilst in two other species, the females are + twice or thrice as numerous as the males. In some European + species of Psocus thousands of females may be collected without + a single male, whilst with other species of the same genus both + sexes are common.[404] In England, Mr. MacLachlan has captured + hundreds of the female _Apatania muliebris_, but has never seen + the male; and of _Boreus hyemalis_ only four or five males have + been here seen.[405] With most of these species (excepting, as + I have heard, with the Tenthredinæ) there is no reason to + suppose that the females are subject to parthenogenesis; and + thus we see how ignorant we are on the causes of the apparent + discrepancy in the proportional numbers of the two sexes. + + In the other Classes of the Articulata I have been able to + collect still less information. With Spiders, Mr. Blackwall, + who has carefully attended to this class during many years, + writes to me that the males from their more erratic habits are + more commonly seen, and therefore appear to be the more + numerous. This is actually the case with a few species; but he + mentions several species in six genera, in which the females + appear to be much more numerous than the males.[406] The small + size of the males in comparison with the females, which is + sometimes carried to an extreme degree, and their widely + different appearance, may account in some instances for their + rarity in collections.[407] + + Some of the lower Crustaceans are able to propagate their kind + asexually, and this will account for the extreme rarity of the + males. With some other forms (as with Tanais and Cypris) there + is reason to believe, as Fritz Müller informs me, that the male + is much shorter-lived than the female, which, supposing the two + sexes to be at first equal in number, would explain the + scarcity of the males. On the other hand this same naturalist + has invariably taken, on the shores of Brazil, far more males + than females of the Diastylidæ and of Cypridina; thus with a + species in the latter genus, 63 specimens caught the same day, + included 57 males; but he suggests that this preponderance may + be due to some unknown difference in the habits of the two + sexes. With one of the higher Brazilian crabs, namely a + Gelasimus, Fritz Müller found the males to be more numerous + than the females. The reverse seems to be the case, according + to the large experience of Mr. C. Spence Bate, with six common + British crabs, the names of which he has given me. + + +_On the Power of Natural Selection to regulate the proportional Numbers +of the Sexes, and General Fertility._—In some peculiar cases, an excess +in the number of one sex over the other might be a great advantage to a +species, as with the sterile females of social insects, or with those +animals in which more than one male is requisite to fertilise the +female, as with certain cirripedes and perhaps certain fishes. An +inequality between the sexes in these cases might have been acquired +through natural selection, but from their rarity they need not here be +further considered. In all ordinary cases an inequality would be no +advantage or disadvantage to certain individuals more than to others; +and therefore it could hardly have resulted from natural selection. We +must attribute the inequality to the direct action of those unknown +conditions, which with mankind lead to the males being born in a +somewhat larger excess in certain countries than in others, or which +cause the proportion between the sexes to differ slightly in legitimate +and illegitimate births. + +Let us now take the case of a species producing from the unknown causes +just alluded to, an excess of one sex—we will say of males—these being +superfluous and useless, or nearly useless. Could the sexes be equalised +through natural selection? We may feel sure, from all characters being +variable, that certain pairs would produce a somewhat less excess of +males over females than other pairs. The former, supposing the actual +number of the offspring to remain constant, would necessarily produce +more females, and would therefore be more productive. On the doctrine of +chances a greater number of the offspring of the more productive pairs +would survive; and these would inherit a tendency to procreate fewer +males and more females. Thus a tendency towards the equalisation of the +sexes would be brought about. But our supposed species would by this +process be rendered, as just remarked, more productive; and this would +in many cases be far from an advantage; for whenever the limit to the +numbers which exist, depends, not on destruction by enemies, but on the +amount of food, increased fertility will lead to severer competition and +to most of the survivors being badly fed. In this case, if the sexes +were equalised by an increase in the number of the females, a +simultaneous decrease in the total number of the offspring would be +beneficial, or even necessary, for the existence of the species; and +this, I believe, could be effected through natural selection in the +manner hereafter to be described. The same train of reasoning is +applicable in the above, as well as in the following case, if we assume +that females instead of males are produced in excess, for such females +from not uniting with males would be superfluous and useless. So it +would be with polygamous species, if we assume the excess of females to +be inordinately great. + +An excess of either sex, we will again say of the males, could, however, +apparently be eliminated through natural selection in another and +indirect manner, namely by an actual diminution of the males, without +any increase of the females, and consequently without any increase in +the productiveness of the species. From the variability of all +characters, we may feel assured that some pairs, inhabiting any +locality, would produce a rather smaller excess of superfluous males, +but an equal number of productive females. When the offspring from the +more and the less male-productive parents were all mingled together, +none would have any direct advantage over the others; but those that +produced few superfluous males would have one great indirect advantage, +namely that their ova or embryos would probably be larger and finer, or +their young better nurtured in the womb and afterwards. We see this +principle illustrated with plants; as those which bear a vast number of +seed produce small ones; whilst those which bear comparatively few +seeds, often produce large ones well-stocked with nutriment for the use +of the seedlings.[408] Hence the offspring of the parents which had +wasted least force in producing superfluous males would be the most +likely to survive, and would inherit the same tendency not to produce +superfluous males, whilst retaining their full fertility in the +production of females. So it would be with the converse case of the +female sex. Any slight excess, however, of either sex could hardly be +checked in so indirect a manner. Nor indeed has a considerable +inequality between the sexes been always prevented, as we have seen in +some of the cases given in the previous discussion. In these cases the +unknown causes which determine the sex of the embryo, and which under +certain conditions lead to the production of one sex in excess over the +other, have not been mastered by the survival of those varieties which +were subjected to the least waste of organised matter and force by the +production of superfluous individuals of either sex. Nevertheless we may +conclude that natural selection will always tend, though sometimes +inefficiently, to equalise the relative numbers of the two sexes. + +Having said this much on the equalisation of the sexes, it may be well +to add a few remarks on the regulation through natural selection of the +ordinary fertility of species. Mr. Herbert Spencer has shewn in an able +discussion[409] that with all organisms a ratio exists between what he +calls individuation and genesis; whence it follows that beings which +consume much matter or force in their growth, complicated structure or +activity, or which produce ova and embryos of large size, or which +expend much energy in nurturing their young, cannot be so productive as +beings of an opposite nature. Mr. Spencer further shews that minor +differences in fertility will be regulated through natural selection. +Thus the fertility of each species will tend to increase, from the more +fertile pairs producing a larger number of offspring, and these from +their mere number will have the best chance of surviving, and will +transmit their tendency to greater fertility. The only check to a +continued augmentation of fertility in each organism seems to be either +the expenditure of more power and the greater risks run by the parents +that produce a more numerous progeny, or the contingency of very +numerous eggs and young being produced of smaller size, or less +vigorous, or subsequently not so well nurtured. To strike a balance in +any case between the disadvantages which follow from the production of a +numerous progeny, and the advantages (such as the escape of at least +some individuals from various dangers) is quite beyond our power of +judgment. + +When an organism has once been rendered extremely fertile, how its +fertility can be reduced through natural selection is not so clear as +how this capacity was first acquired. Yet it is obvious that if +individuals of a species, from a decrease of their natural enemies, were +habitually reared in larger numbers than could be supported, all the +members would suffer. Nevertheless the offspring from the less fertile +parents would have no direct advantage over the offspring from the more +fertile parents, when all were mingled together in the same district. +All the individuals would mutually tend to starve each other. The +offspring indeed of the less fertile parents would lie under one great +disadvantage, for from the simple fact of being produced in smaller +numbers, they would be the most liable to extermination. Indirectly, +however, they would partake of one great advantage; for under the +supposed condition of severe competition, when all were pressed for +food, it is extremely probable that those individuals which from some +variation in their constitution produced fewer eggs or young, would +produce them of greater size or vigour; and the adults reared from such +eggs or young would manifestly have the best chance of surviving, and +would inherit a tendency towards lessened fertility. The parents, +moreover, which had to nourish or provide for fewer offspring would +themselves be exposed to a less severe strain in the struggle for +existence, and would have a better chance of surviving. By these steps, +and by no others as far as I can see, natural selection under the above +conditions of severe competition for food, would lead to the formation +of a new race less fertile, but better adapted for survival, than the +parent-race. + + + + +CHAPTER IX. + +SECONDARY SEXUAL CHARACTERS IN THE LOWER CLASSES OF THE ANIMAL KINGDOM. + + + These characters absent in the lowest classes—Brilliant + colours—Mollusca—Annelids—Crustacea, secondary sexual + characters strongly developed; dimorphism; colour; characters + not acquired before maturity—Spiders, sexual colours of; + stridulation by the males—Myriapoda. + + +In the lowest classes the two sexes are not rarely united in the same +individual, and therefore secondary sexual characters cannot be +developed. In many cases in which the two sexes are separate, both are +permanently attached to some support, and the one cannot search or +struggle for the other. Moreover it is almost certain that these animals +have too imperfect senses and much too low mental powers to feel mutual +rivalry, or to appreciate each other’s beauty or other attractions. + +Hence in these classes, such as the Protozoa, Cœlenterata, +Echinodermata, Scolecida, true secondary sexual characters do not occur; +and this fact agrees with the belief that such characters in the higher +classes have been acquired through sexual selection, which depends on +the will, desires, and choice of either sex. Nevertheless some few +apparent exceptions occur; thus, as I hear from Dr. Baird, the males of +certain Entozoa, or internal parasitic worms, differ slightly in colour +from the females; but we have no reason to suppose that such differences +have been augmented through sexual selection. + +Many of the lower animals, whether hermaphrodites or with the sexes +separate, are ornamented with the most brilliant tints, or are shaded +and striped in an elegant manner. This is the case with many corals and +sea-anemonies (Actiniæ), with some jelly-fish (Medusæ, Porpita, &c.), +with some Planariæ, Ascidians, numerous Star-fishes, Echini, &c.; but we +may conclude from the reasons already indicated, namely the union of the +two sexes in some of these animals, the permanently affixed condition of +others, and the low mental powers of all, that such colours do not serve +as a sexual attraction, and have not been acquired through sexual +selection. With the higher animals the case is very different; for with +them when one sex is much more brilliantly or conspicuously coloured +than the other, and there is no difference in the habits of the two +sexes which will account for this difference, we have reason to believe +in the influence of sexual selection; and this belief is strongly +confirmed when the more ornamented individuals, which are almost always +the males, display their attractions before the other sex. We may also +extend this conclusion to both sexes, when coloured alike, if their +colours are plainly analogous to those of one sex alone in certain other +species of the same group. + +How, then, are we to account for the beautiful or even gorgeous colours +of many animals in the lowest classes? It appears very doubtful whether +such colours usually serve as a protection; but we are extremely liable +to err in regard to characters of all kinds in relation to protection, +as will be admitted by every one who has read Mr. Wallace’s excellent +essay on this subject. It would not, for instance, at first occur to any +one that the perfect transparency of the Medusæ, or jelly-fishes, was of +the highest service to them as a protection; but when we are reminded +by Häckel that not only the medusæ but many floating mollusca, +crustaceans, and even small oceanic fishes partake of this same +glass-like structure, we can hardly doubt that they thus escape the +notice of pelagic birds and other enemies. + +Notwithstanding our ignorance how far colour in many cases serves as a +protection, the most probable view in regard to the splendid tints of +many of the lowest animals seems to be that their colours are the direct +result either of the chemical nature or the minute structure of their +tissues, independently of any benefit thus derived. Hardly any colour is +finer than that of arterial blood; but there is no reason to suppose +that the colour of the blood is in itself any advantage; and though it +adds to the beauty of the maiden’s cheek, no one will pretend that it +has been acquired for this purpose. So again with many animals, +especially the lower ones, the bile is richly coloured; thus the extreme +beauty of the Eolidæ (naked sea-slugs) is chiefly due, as I am informed +by Mr. Hancock, to the biliary glands seen through the translucent +integuments; this beauty being probably of no service to these animals. +The tints of the decaying leaves in an American forest are described by +every one as gorgeous; yet no one supposes that these tints are of the +least advantage to the trees. Bearing in mind how many substances +closely analogous to natural organic compounds have been recently formed +by chemists, and which exhibit the most splendid colours, it would have +been a strange fact if substances similarly coloured had not often +originated, independently of any useful end being thus gained, in the +complex laboratory of living organisms. + + +_The sub-kingdom of the Mollusca._—Throughout this great division +(taken in its largest acceptation) of the animal kingdom, secondary +sexual characters, such as we are here considering, never, as far as I +can discover, occur. Nor could they be expected in the three lowest +classes, namely in the Ascidians, Polyzoa, and Brachiopods (constituting +the Molluscoida of Huxley), for most of these animals are permanently +affixed to a support or have their sexes united in the same individual. +In the Lamellibranchiata, or bivalve shells, hermaphroditism is not +rare. In the next higher class of the Gasteropoda, or marine univalve +shells, the sexes are either united or separate. But in this latter case +the males never possess special organs for finding, securing, or +charming the females, or for fighting with other males. The sole +external difference between the sexes consists, as I am informed by Mr. +Gwyn Jeffreys, in the shell sometimes differing a little in form; for +instance, the shell of the male periwinkle (_Littorina littorea_) is +narrower and has a more elongated spire than that of the female. But +differences of this nature, it may be presumed, are directly connected +with the act of reproduction or with the development of the ova. + +The Gasteropoda, though capable of locomotion and furnished with +imperfect eyes, do not appear to be endowed with sufficient mental +powers for the members of the same sex to struggle together in rivalry, +and thus to acquire secondary sexual characters. Nevertheless with the +pulmoniferous gasteropods, or land-snails, the pairing is preceded by +courtship; for these animals, though hermaphrodites, are compelled by +their structure to pair together. Agassiz remarks,[410] “Quiconque a eu +l’occasion d’observer les amours des limaçons, ne saurait mettre en +doute la séduction déployée dans les mouvements et les allures qui +préparent et accomplissent le double embrassement de ces +hermaphrodites.” These animals appear also susceptible of some degree of +permanent attachment: an accurate observer, Mr. Lonsdale, informs me +that he placed a pair of land-shells (_Helix pomatia_), one of which was +weakly, into a small and ill-provided garden. After a short time the +strong and healthy individual disappeared, and was traced by its track +of slime over a wall into an adjoining well-stocked garden. Mr. Lonsdale +concluded that it had deserted its sickly mate; but after an absence of +twenty-four hours it returned, and apparently communicated the result of +its successful exploration, for both then started along the same track +and disappeared over the wall. + +Even in the highest class of the Mollusca, namely the Cephalopoda or +cuttle-fishes, in which the sexes are separate, secondary sexual +characters of the kind which we are here considering, do not, as far as +I can discover, occur. This is a surprising circumstance, as these +animals possess highly-developed sense-organs and have considerable +mental powers, as will be admitted by every one who has watched their +artful endeavours to escape from an enemy.[411] Certain Cephalopoda, +however, are characterised by one extraordinary sexual character, +namely, that the male element collects within one of the arms or +tentacles, which is then cast off, and, clinging by its sucking-discs to +the female, lives for a time an independent life. So completely does the +cast-off arm resemble a separate animal, that it was described by Cuvier +as a parasitic worm under the name of Hectocotyle. But this marvellous +structure may be classed as a primary rather than as a secondary sexual +character. + +Although with the Mollusca sexual selection does not seem to have come +into play; yet many univalve and bivalve shells, such as volutes, cones, +scallops, &c., are beautifully coloured and shaped. The colours do not +appear in most cases to be of any use as a protection; they are probably +the direct result, as in the lowest classes, of the nature of the +tissues; the patterns and the sculpture of the shell depending on its +manner of growth. The amount of light seems to a certain extent to be +influential; for although, as repeatedly stated by Mr. Gwyn Jeffreys, +the shells of some species living at a profound depth are brightly +coloured, yet we generally see the lower surfaces and the parts covered +by the mantle less highly coloured than the upper and exposed +surfaces.[412] In some cases, as with shells living amongst corals or +brightly-tinted sea-weeds, the bright colours may serve as a protection. +But many of the nudibranch mollusca, or sea-slugs, are as beautifully +coloured as any shells, as may be seen in Messrs. Alder and Hancock’s +magnificent work; and from information kindly given me by Mr. Hancock, +it is extremely doubtful whether these colours usually serve as a +protection. With some species this may be the case, as with one which +lives on the green leaves of algæ, and is itself bright-green. But many +brightly-coloured, white or otherwise conspicuous species, do not seek +concealment; whilst again some equally conspicuous species, as well as +other dull-coloured kinds, live under stones and in dark recesses. So +that with these nudibranch molluscs, colour apparently does not stand in +any close relation to the nature of the places which they inhabit. + +These naked sea-slugs are hermaphrodites, yet they pair together, as do +land-snails, many of which have extremely pretty shells. It is +conceivable that two hermaphrodites, attracted by each others’ greater +beauty, might unite and leave offspring which would inherit their +parents’ greater beauty. But with such lowly-organised creatures this is +extremely improbable. Nor is it at all obvious how the offspring from +the more beautiful pairs of hermaphrodites would have any advantage, so +as to increase in numbers, over the offspring of the less beautiful, +unless indeed vigour and beauty generally coincided. We have not here a +number of males becoming mature before the females, and the more +beautiful ones selected by the more vigorous females. If, indeed, +brilliant colours were beneficial to an hermaphrodite animal in relation +to its general habits of life, the more brightly-tinted individuals +would succeed best and would increase in number; but this would be a +case of natural and not of sexual selection. + + +_Sub-kingdom of the Vermes or Annulosa_: Class, _Annelida (or +Sea-worms)_.—In this class, although the sexes (when separate) +sometimes differ from each other in characters of such importance that +they have been placed under distinct genera or even families, yet the +differences do not seem of the kind which can be safely attributed to +sexual selection. These animals, like those in the preceding classes, +apparently stand too low in the scale, for the individuals of either sex +to exert any choice in selecting a partner, or for the individuals of +the same sex to struggle together in rivalry. + + +_Sub-kingdom of the Arthropoda_: Class, _Crustacea_.—In this great +class we first meet with undoubted secondary sexual characters, often +developed in a remarkable manner. Unfortunately the habits of +crustaceans are very imperfectly known, and we cannot explain the uses +of many structures peculiar to one sex. With the lower parasitic species +the males are of small size, and they alone are furnished with perfect +swimming-legs, antennæ and sense-organs; the females being destitute of +these organs, with their bodies often consisting of a mere distorted +mass. But these extraordinary differences between the two sexes are no +doubt related to their widely different habits of life, and consequently +do not concern us. In various crustaceans, belonging to distinct +families, the anterior antennæ are furnished with peculiar thread-like +bodies, which are believed to act as smelling-organs, and these are much +more numerous in the males than in the females. As the males, without +any unusual development of their olfactory organs, would almost +certainly be able sooner or later to find the females, the increased +number of the smelling-threads has probably been acquired through sexual +selection, by the better provided males having been the most successful +in finding partners and in leaving offspring. Fritz Müller has described +a remarkable dimorphic species of Tanais, in which the male is +represented by two distinct forms, never graduating into each other. In +the one form the male is furnished with more numerous smelling-threads, +and in the other form with more powerful and more elongated chelæ or +pincers which serve to hold the female. Fritz Müller suggests that these +differences between the two male forms of the same species must have +originated in certain individuals having varied in the number of the +smelling-threads, whilst other individuals varied in the shape and size +of their chelæ; so that of the former, those which were best able to +find the female, and of the latter, those which were best able to hold +her when found, have left the greater number of progeny to inherit their +respective advantages.[413] + +[Illustration: Fig. 3. Labidocera Darwinii, (from Lubbock). + +_a._ Part of right-hand anterior antenna of male, forming a prehensile +organ. + +_b._ Posterior pair of thoracic legs of male. + +_c._ Ditto of female.] + +In some of the lower crustaceans, the right-hand anterior antenna of the +male differs greatly in structure from the left-hand one, the latter +resembling in its simple tapering joints the antennæ of the female. In +the male the modified antenna is either swollen in the middle or +angularly bent, or converted (fig. 3) into an elegant, and sometimes +wonderfully complex, prehensile organ.[414] It serves, as I hear from +Sir J. Lubbock, to hold the female, and for this same purpose one of the +two posterior legs (_b_) on the same side of the body is converted into +a forceps. In another family the inferior or posterior antennæ are +“curiously zigzagged” in the males alone. + +[Illustration: Fig. 4. Anterior part of body of Callianassa (from +Milne-Edwards), showing the unequal and differently-constructed right +and left-hand chelæ of the male. + +N.B.—The artist by mistake has reversed the drawing, and made the +left-hand chela the largest.] + +[Illustration: Fig. 5. Second leg of male Orchestia Tucuratinga (from +Fritz Müller).] + +[Illustration: Fig. 6. Ditto of female.] + +In the higher crustaceans the anterior legs form a pair of chelæ or +pincers, and these are generally larger in the male than in the female. +In many species the chelæ on the opposite sides of the body are of +unequal size, the right-hand one being, as I am informed by Mr. C. +Spence Bate, generally, though not invariably, the largest. This +inequality is often much greater in the male than in the female. The two +chelæ also often differ in structure (figs. 4, 5, and 6), the smaller one +resembling those of the female. What advantage is gained by their +inequality in size on the opposite sides of the body, and by the +inequality being much greater in the male than in the female; and why, +when they are of equal size, both are often much larger in the male than +in the female, is not known. The chelæ are sometimes of such length and +size that they cannot possibly be used, as I hear from Mr. Spence Bate, +for carrying food to the mouth. In the males of certain freshwater +prawns (Palæmon) the right leg is actually longer than the whole +body.[415] It is probable that the great size of one leg with its chelæ +may aid the male in fighting with his rivals; but this use will not +account for their inequality in the female on the opposite sides +of the body. In Gelasimus, according to a statement quoted by +Milne-Edwards,[416] the male and female live in the same burrow, which +is worth notice, as shewing that they pair, and the male closes the +mouth of the burrow with one of its chelæ, which is enormously +developed; so that here it indirectly serves as a means of defence. +Their main use, however, probably is to seize and to secure the female, +and this in some instances, as with Gammarus, is known to be the case. +The sexes, however, of the common shore-crab (_Carcinus mænas_), as Mr. +Spence Bate informs me, unite directly after the female has moulted her +hard shell, and when she is so soft that she would be injured if seized +by the strong pincers of the male; but as she is caught and carried +about by the male previously to the act of moulting, she could then be +seized with impunity. + +Fritz Müller states that certain species of Melita are distinguished +from all other amphipods by the females having “the coxal lamellæ of the +penultimate pair of feet produced into hook-like processes, of which the +males lay hold with the hands of the first pair.” The development of +these hook-like processes probably resulted from those females which +were the most securely held during the act of reproduction, having left +the largest number of offspring. Another Brazilian amphipod (_Orchestia +Darwinii_, fig. 7) is described by Fritz Müller, as presenting a case of +dimorphism, like that of Tanais; for there are two male forms, which +differ in the structure of their chelæ.[417] As chelæ of either shape +would certainly have sufficed to hold the female, for both are now used +for this purpose, the two male forms probably originated, by some having +varied in one manner and some in another; both forms having derived +certain special, but nearly equal advantages, from their differently +shaped organs. + +It is not known that male crustaceans fight together for the possession +of the females, but this is probable; for with most animals when the +male is larger than the female, he seems to have acquired his greater +size by having conquered during many generations other males. Now Mr. +Spence Bate informs me that in most of the crustacean orders, especially +in the highest or the Brachyura, the male is larger than the female; the +parasitic genera, however, in which the sexes follow different habits of +life, and most of the Entomostraca must be excepted. The chelæ of many +crustaceans are weapons well adapted for fighting. Thus a Devil-crab +(_Portunus puber_) was seen by a son of Mr. Bate fighting with a +_Carcinus mænas_, and the latter was soon thrown on its back, and had +every limb torn from its body. When several males of a Brazilian +Gelasimus, a species furnished with immense pincers, were placed +together by Fritz Müller in a glass vessel, they mutilated and killed +each other. Mr. Bate put a large male _Carcinus mænas_ into a pan of +water, inhabited by a female paired with a smaller male; the latter was +soon dispossessed, but, as Mr. Bate adds, “if they fought, the victory +was a bloodless one, for I saw no wounds.” This same naturalist +separated a male sand-skipper (so common on our sea-shores), _Gammarus +marinus_, from its female, both of which were imprisoned in the same +vessel with many individuals of the same species. The female being thus +divorced joined her comrades. After an interval the male was again put +into the same vessel and he then, after swimming about for a time, +dashed into the crowd, and without any fighting at once took away his +wife. This fact shews that in the Amphipoda, an order low in the scale, +the males and females recognise each other, and are mutually attached. + +[Illustration: Fig. 7. Orchestia Darwinii (from Fritz Müller), showing +the differently-constructed chelæ of the two male forms.] + +The mental powers of the Crustacea are probably higher than might have +been expected. Any one who has tried to catch one of the shore-crabs, so +numerous on many tropical coasts, will have perceived how wary and alert +they are. There is a large crab (_Birgus latro_), found on coral +islands, which makes at the bottom of a deep burrow a thick bed of the +picked fibres of the cocoa-nut. It feeds on the fallen fruit of this +tree by tearing off the husk, fibre by fibre; and it always begins at +that end where the three eye-like depressions are situated. It then +breaks through one of these eyes by hammering with its heavy front +pincers, and turning round, extracts the albuminous core with its narrow +posterior pincers. But these actions are probably instinctive, so that +they would be performed as well by a young as by an old animal. The +following case, however, can hardly be so considered: a trustworthy +naturalist, Mr. Gardner,[418] whilst watching a shore-crab (Gelasimus) +making its burrow, threw some shells towards the hole. One rolled in, +and three other shells remained within a few inches of the mouth. In +about five minutes the crab brought out the shell which had fallen in, +and carried it away to the distance of a foot; it then saw the three +other shells lying near, and evidently thinking that they might likewise +roll in, carried them to the spot where it had laid the first. It would, +I think, be difficult to distinguish this act from one performed by man +by the aid of reason. + +With respect to colour which so often differs in the two sexes of +animals belonging to the higher classes, Mr. Spence Bate does not know +of any well-marked instances with our British crustaceans. In some +cases, however, the male and female differ slightly in tint, but Mr. +Bate thinks not more than may be accounted for by their different habits +of life, such as by the male wandering more about and being thus more +exposed to the light. In a curious Bornean crab, which inhabits sponges, +Mr. Bate could always distinguish the sexes by the male not having the +epidermis so much rubbed off. Dr. Power tried to distinguish by colour +the sexes of the species which inhabit the Mauritius, but always failed, +except with one species of Squilla, probably the _S. stylifera_, the +male of which is described as being “of a beautiful blueish-green,” with +some of the appendages cherry-red, whilst the female is clouded with +brown and grey, “with the red about her much less vivid than in the +male.”[419] In this case, we may suspect the agency of sexual selection. +With Saphirina (an oceanic genus of Entomostraca, and therefore low in +the scale) the males are furnished with minute shields or cell-like +bodies, which exhibit beautiful changing colours; these being absent in +the females, and in the case of one species in both sexes.[420] It +would, however, be extremely rash to conclude that these curious organs +serve merely to attract the females. In the female of a Brazilian +species of Gelasimus, the whole body, as I am informed by Fritz Müller, +is of a nearly uniform greyish-brown. In the male the posterior part of +the cephalo-thorax is pure white, with the anterior part of a rich +green, shading into dark brown; and it is remarkable that these colours +are liable to change in the course of a few minutes—the white becoming +dirty grey or even black, the green “losing much of its brilliancy.” The +males apparently are much more numerous than the females. It deserves +especial notice that they do not acquire their bright colours until they +become mature. They differ also from the females in the larger size of +their chelæ. In some species of the genus, probably in all, the sexes +pair and inhabit the same burrow. They are also, as we have seen, highly +intelligent animals. From these various considerations it seems highly +probable that the male in this species has become gaily ornamented in +order to attract or excite the female. + +It has just been stated that the male Gelasimus does not acquire his +conspicuous colours until mature and nearly ready to breed. This seems +the general rule in the whole class with the many remarkable differences +in structure between the two sexes. We shall hereafter find the same law +prevailing throughout the great sub-kingdom of the Vertebrata, and in +all cases it is eminently distinctive of characters which have been +acquired through sexual selection. Fritz Müller[421] gives some +striking instances of this law; thus the male sand-hopper (Orchestia) +does not acquire his large claspers, which are very differently +constructed from those of the female, until nearly full-grown; whilst +young his claspers resemble those of the female. Thus, again, the male +Brachyscelus possesses, like all other amphipods, a pair of posterior +antennæ; the female, and this is a most extraordinary circumstance, is +destitute of them, and so is the male as long as he remains immature. + + +Class, _Arachnida_ (Spiders).—The males are often darker, but sometimes +lighter than the females, as may be seen in Mr. Blackwall’s magnificent +work.[422] In some species the sexes differ conspicuously from each +other in colour; thus the female of _Sparassus smaragdulus_ is +dullish-green; whilst the adult male has the abdomen of a fine yellow, +with three longitudinal stripes of rich red. In some species of Thomisus +the two sexes closely resemble each other; in others they differ much; +thus in _T. citreus_ the legs and body of the female are pale-yellow or +green, whilst the front legs of the male are reddish-brown: in _T. +floricolens_, the legs of the female are pale-green, those of the male +being ringed in a conspicuous manner with various tints. Numerous +analogous cases could be given in the genera Epeira, Nephila, +Philodromus, Theridion, Linyphia, &c. It is often difficult to say which +of the two sexes departs most from the ordinary coloration of the genus +to which the species belong; but Mr. Blackwall thinks that, as a +general rule, it is the male. Both sexes whilst young, as I am informed +by the same author, usually resemble each other; and both often undergo +great changes in colour during their successive moults before arriving +at maturity. In other cases the male alone appears to change colour. +Thus the male of the above-mentioned brightly-coloured Sparassus at +first resembles the female and acquires his peculiar tints only when +nearly adult. Spiders are possessed of acute senses, and exhibit much +intelligence. The females often shew, as is well known, the strongest +affection for their eggs, which they carry about enveloped in a silken +web. On the whole it appears probable that well-marked differences in +colour between the sexes have generally resulted from sexual selection, +either on the male or female side. But doubts may be entertained on this +head from the extreme variability in colour of some species, for +instance of _Theridion lineatum_, the sexes of which differ when adult; +this great variability indicates that their colours have not been +subjected to any form of selection. + +Mr. Blackwall does not remember to have seen the males of any species +fighting together for the possession of the female. Nor, judging from +analogy, is this probable; for the males are generally much smaller than +the females, sometimes to an extraordinary degree.[423] Had the males +been in the habit of fighting together, they would, it is probable, have +gradually acquired greater size and strength. Mr. Blackwall has +sometimes seen two or more males on the same web with a single female; +but their courtship is too tedious and prolonged an affair to be easily +observed. The male is extremely cautious in making his advances, as the +female carries her coyness to a dangerous pitch. De Geer saw a male that +“in the midst of his preparatory caresses was seized by the object of +his attentions, enveloped by her in a web and then devoured, a sight +which, as he adds, filled him with horror and indignation.”[424] + +Westring has made the interesting discovery that the males of several +species of Theridion[425] have the power of making a stridulating sound +(like that made by many beetles and other insects, but feebler), whilst +the females are quite mute. The apparatus consists of a serrated ridge +at the base of the abdomen, against which the hard hinder part of the +thorax is rubbed; and of this structure not a trace could be detected in +the females. From the analogy of the Orthoptera and Homoptera, to be +described in the next chapter, we may feel almost sure that the +stridulation serves, as Westring remarks, either to call or to excite +the female; and this is the first case in the ascending scale of the +animal kingdom, known to me, of sounds emitted for this purpose. + + +Class, _Myriapoda_.—In neither of the two orders in this class, +including the millipedes and centipedes, can I find any well-marked +instances of sexual differences such as more particularly concern us. In +_Glomeris limbata_, however, and perhaps in some few other species, the +males differ slightly in colour from the females; but this Glomeris is a +highly variable species. In the males of the Diplopoda, the legs +belonging to one of the anterior segments of the body, or to the +posterior segment, are modified into prehensile hooks which serve to +secure the female. In some species of Iulus the tarsi of the male are +furnished with membranous suckers for the same purpose. It is a much +more unusual circumstance, as we shall see when we treat of Insects, +that it is the female in Lithobius which is furnished with prehensile +appendages at the extremity of the body for holding the male.[426] + + + + +CHAPTER X. + +SECONDARY SEXUAL CHARACTERS OF INSECTS. + + + Diversified structures possessed by the males for seizing the + females—Differences between the sexes, of which the meaning is + not understood—Difference in size between the + sexes—Thysanura—Diptera—Hemiptera—Homoptera, musical powers + possessed by the males alone—Orthoptera, musical instruments + of the males, much diversified in structure; pugnacity; + colours—Neuroptera, sexual differences in colour—Hymenoptera, + pugnacity and colours—Coleoptera, colours; furnished with + great horns, apparently as an ornament; battles; stridulating + organs generally common to both sexes. + + +In the immense class of insects the sexes sometimes differ in their +organs for locomotion, and often in their sense-organs, as in the +pectinated and beautifully plumose antennæ of the males of many species. +In one of the Ephemeræ, namely Chloëon, the male has great pillared +eyes, of which the female is entirely destitute.[427] The ocelli are +absent in the females of certain other insects, as in the Mutillidæ, +which are likewise destitute of wings. But we are chiefly concerned with +structures by which one male is enabled to conquer another, either in +battle or courtship, through his strength, pugnacity, ornaments, or +music. The innumerable contrivances, therefore, by which the male is +able to seize the female, may be briefly passed over. Besides the +complex structures at the apex of the abdomen, which ought perhaps to be +ranked as primary organs,[428] “it is astonishing,” as Mr. B. D. +Walsh[429] has remarked, “how many different organs are worked in by +nature, for the seemingly insignificant object of enabling the male to +grasp the female firmly.” The mandibles or jaws are sometimes used for +this purpose; thus the male _Corydalis cornutus_ (a neuropterous insect +in some degree allied to the Dragon-flies, &c.) has immense curved jaws, +many times longer than those of the female; and they are smooth instead +of being toothed, by which means he is enabled to seize her without +injury.[430] One of the stag-beetles of North America (_Lucanus +elaphus_) uses his jaws, which are much larger than those of the female, +for the same purpose, but probably likewise for fighting. In one of the +sand-wasps (_Ammophila_) the jaws in the two sexes are closely alike, +but are used for widely different purposes; the males, as Professor +Westwood observes, “are exceedingly ardent, seizing their partners round +the neck with their sickle-shaped jaws;”[431] whilst the females use +these organs for burrowing in sand-banks and making their nests. + +[Illustration: Fig. 8. Crabro cribrarius. Upper figure, male: lower +figure, female.] + +The tarsi of the front-legs are dilated in many male beetles, or are +furnished with broad cushions of hairs; and in many genera of +water-beetles they are armed with a round flat sucker, so that the male +may adhere to the slippery body of the female. It is a much more unusual +circumstance that the females of some water-beetles (Dytiscus) have +their elytra deeply grooved, and in _Acilius sulcatus_ thickly set with +hairs, as an aid to the male. The females of some other water-beetles +(Hydroporus) have their elytra punctured for the same object.[432] In +the male of _Crabro cribrarius_ (fig. 8.), it is the tibia which is +dilated into a broad horny plate, with minute membraneous dots, giving +to it a singular appearance like that of a riddle.[433] In the male of +Penthe (a genus of beetles) a few of the middle joints of the antennæ +are dilated and furnished on the inferior surface with cushions of +hair, exactly like those on the tarsi of the Carabidæ, “and obviously +for the same end.” In male dragon-flies, “the appendages at the tip of +the tail are modified in an almost infinite variety of curious patterns +to enable them to embrace the neck of the female.” Lastly in the males +of many insects, the legs are furnished with peculiar spines, knobs or +spurs; or the whole leg is bowed or thickened, but this is by no means +invariably a sexual character; or one pair, or all three pairs are +elongated, sometimes to an extravagant length.[434] + +[Illustration: Fig. 9. Taphroderes distortus (much enlarged). Upper +figure, male; lower figure, female.] + +In all the orders, the sexes of many species present differences, of +which the meaning is not understood. One curious case is that of a +beetle (fig. 9), the male of which has the left mandible much enlarged; +so that the mouth is greatly distorted. In another Carabidous beetle, +the Eurygnathus,[435] we have the unique case, as far as known to Mr. +Wollaston, of the head of the female being much broader and larger, +though in a variable degree, than that of the male. Any number of such +cases could be given. They abound in the Lepidoptera: one of the most +extraordinary is that certain male butterflies have their fore-legs more +or less atrophied, with the tibiæ and tarsi reduced to mere rudimentary +knobs. The wings, also, in the two sexes often differ in neuration,[436] +and sometimes considerably in outline, as in the _Aricoris epitus_, +which was shown to me in the British Museum by Mr. A. Butler. The males +of certain South American butterflies have tufts of hair on the margins +of the wings, and horny excrescences on the discs of the posterior +pair.[437] In several British butterflies, the males alone, as shewn by +Mr. Wonfor, are in parts clothed with peculiar scales. + +The purpose of the luminosity in the female glow-worm is likewise not +understood; for it is very doubtful whether the primary use of the light +is to guide the male to the female. It is no serious objection to this +latter belief that the males emit a feeble light; for secondary sexual +characters proper to one sex are often developed in a slight degree in +the other sex. It is a more valid objection that the larvæ shine, and in +some species brilliantly: Fritz Müller informs me that the most luminous +insect which he ever beheld in Brazil, was the larva of some beetle. +Both sexes of certain luminous species of Elater emit light. Kirby and +Spence suspect that the phosphorescence serves to frighten and drive +away enemies. + + +_Difference in Size between the Sexes._—With insects of all kinds the +males are commonly smaller than the females;[438] and this difference +can often be detected even in the larval state. So considerable is the +difference between the male and female cocoons of the silk-moth +(_Bombyx mori_), that in France they are separated by a particular mode +of weighing.[439] In the lower classes of the animal kingdom, the +greater size of the females seems generally to depend on their +developing an enormous number of ova; and this may to a certain extent +hold good with insects. But Dr. Wallace has suggested a much more +probable explanation. He finds, after carefully attending to the +development of the caterpillars of _Bombyx cynthia_ and _yamamai_, and +especially of some dwarfed caterpillars reared from a second brood on +unnatural food, “that in proportion as the individual moth is finer, so +is the time required for its metamorphosis longer; and for this reason +the female, which is the larger and heavier insect, from having to carry +her numerous eggs, will be preceded by the male, which is smaller and +has less to mature.”[440] Now as most insects are short-lived, and as +they are exposed to many dangers, it would manifestly be advantageous to +the female to be impregnated as soon as possible. This end would be +gained by the males being first matured in large numbers ready for the +advent of the females; and this again would naturally follow, as Mr. A. +E. Wallace has remarked,[441] through natural selection; for the smaller +males would be first matured, and thus would procreate a large number of +offspring which would inherit the reduced size of their male parents, +whilst the larger males from being matured later would leave fewer +offspring. + +There are, however, exceptions to the rule of male insects being smaller +than the females; and some of these exceptions are intelligible. Size +and strength would be an advantage to the males, which fight for the +possession of the female; and in these cases the males, as with the +stag-beetle (Lucanus), are larger than the females. There are, however, +other beetles which are not known to fight together, of which the males +exceed the females in size; and the meaning of this fact is not known; +but in some of these cases, as with the huge Dynastes and Megasoma, we +can at least see that there would be no necessity for the males to be +smaller than the females, in order to be matured before them, for these +beetles are not short-lived, and there would be ample time for the +pairing of the sexes. So, again, male dragon-flies (Libellulidæ) are +sometimes sensibly larger, and never smaller, than the females;[442] and +they do not, as Mr. MacLachlan believes, generally pair with the +females, until a week or fortnight has elapsed, and until they have +assumed their proper masculine colours. But the most curious case, +shewing on what complex and easily-overlooked relations, so trifling a +character as a difference in size between the sexes may depend, is that +of the aculeate Hymenoptera; for Mr. F. Smith informs me that throughout +nearly the whole of this large group the males, in accordance with the +general rule, are smaller than the females and emerge about a week +before them; but amongst the Bees, the males of _Apis mellifica_, +_Anthidium manicatum_ and _Anthophora acervorum_, and amongst the +Fossores, the males of the _Methoca ichneumonides_, are larger than the +females. The explanation of this anomaly is that a marriage-flight is +absolutely necessary with these species, and the males require great +strength and size in order to carry the females through the air. +Increased size has here been acquired in opposition to the usual +relation between size and the period of development, for the males, +though larger, emerge before the smaller females. + +We will now review the several Orders, selecting such facts as more +particularly concern us. The Lepidoptera (Butterflies and Moths) will be +retained for a separate chapter. + + +Order, _Thysanura_.—The members of this Order are lowly organised for +their class. They are wingless, dull-coloured, minute insects, with +ugly, almost misshapen heads and bodies. The sexes do not differ; but +they offer one interesting fact, by showing that the males pay sedulous +court to their females even low down in the animal scale. Sir J. +Lubbock[443] in describing the _Smynthurus luteus_, says: “it is very +amusing to see these little creatures coquetting together. The male, +which is much smaller than the female, runs round her, and they butt one +another, standing face to face, and moving backward and forward like two +playful lambs. Then the female pretends to run away and the male runs +after her with a queer appearance of anger, gets in front and stands +facing her again; then she turns coyly round, but he, quicker and more +active, scuttles round too, and seems to whip her with his antennæ; then +for a bit they stand face to face, play with their antennæ, and seem to +be all in all to one another.” + + +Order, _Diptera_ (Flies).—The sexes differ little in colour. The +greatest difference, known to Mr. F. Walker, is in the genus Bibio, in +which the males are blackish or quite black, and the females obscure +brownish-orange. The genus Elaphomyia, discovered by Mr. Wallace[444] in +New Guinea, is highly remarkable, as the males are furnished with horns, +of which the females are quite destitute. The horns spring from beneath +the eyes, and curiously resemble those of stags, being either branched +or palmated. They equal in length the whole of the body in one of the +species. They might be thought to serve for fighting, but as in one +species they are of a beautiful pink colour, edged with black, with a +pale central stripe, and as these insects have altogether a very elegant +appearance, it is perhaps more probable that the horns serve as +ornaments. That the males of some Diptera fight together is certain; for +Prof. Westwood[445] has several times seen this with some species of +Tipula or Harry-long-legs. Many observers believe that when gnats +(Culicidæ) dance in the air in a body, alternately rising and falling, +the males are courting the females. The mental faculties of the Diptera +are probably fairly well developed, for their nervous system is more +highly developed than in most other Orders of insects.[446] + + +Order, _Hemiptera_ (Field-Bugs).—Mr. J. W. Douglas, who has +particularly attended to the British species, has kindly given me an +account of their sexual differences. The males of some species are +furnished with wings, whilst the females are wingless; the sexes differ +in the form of the body and elytra; in the second joints of their +antennæ and in their tarsi; but as the signification of these +differences is quite unknown, they may be here passed over. The females +are generally larger and more robust than the males. With British, and, +as far as Mr. Douglas knows, with exotic species, the sexes do not +commonly differ much in colour; but in about six British species the +male is considerably darker than the female, and in about four other +species the female is darker than the male. Both sexes of some species +are beautifully marked with vermilion and black. It is doubtful whether +these colours serve as a protection. If in any species the males had +differed from the females in an analogous manner, we might have been +justified in attributing such conspicuous colours to sexual selection +with transference to both sexes. + +Some species of Reduvidæ make a stridulating noise; and, in the case of +_Pirates stridulus_, this is said[447] to be effected by the movement of +the neck within the pro-thoracic cavity. According to Westring, +_Reduvius personatus_ also stridulates. But I have not been able to +learn any particulars about these insects; nor have I any reason to +suppose that they differ sexually in this respect. + + +Order, _Homoptera._—Every one who has wandered in a tropical forest +must have been astonished at the din made by the male Cicadæ. The +females are mute; as the Grecian poet Xenarchus says, “Happy the Cicadas +live, since they all have voiceless wives.” The noise thus made could be +plainly heard on board the “Beagle,” when anchored at a quarter of a +mile from the shore of Brazil; and Captain Hancock says it can be heard +at the distance of a mile. The Greeks formerly kept, and the Chinese now +keep, these insects in cages for the sake of their song, so that it +must be pleasing to the ears of some men.[448] The Cicadidæ usually sing +during the day; whilst the Fulgoridæ appear to be night-songsters. The +sound, according to Landois,[449] who has recently studied the subject, +is produced by the vibration of the lips of the spiracles, which are set +into motion by a current of air emitted from the tracheæ. It is +increased by a wonderfully complex resounding apparatus, consisting of +two cavities covered by scales. Hence the sound may truly be called a +voice. In the female the musical apparatus is present, but very much +less developed than in the male, and is never used for producing sound. + +With respect to the object of the music, Dr. Hartman in speaking of the +_Cicada septemdecim_ of the United States, says,[450] “the drums are now +(June 6th and 7th, 1851) heard in all directions. This I believe to be +the marital summons from the males. Standing in thick chestnut sprouts +about as high as my head, where hundreds were around me, I observed the +females coming around the drumming males.” He adds, “this season (Aug. +1868) a dwarf pear-tree in my garden produced about fifty larvæ of _Cic. +pruinosa_; and I several times noticed the females to alight near a male +while he was uttering his clanging notes.” Fritz Müller writes to me +from S. Brazil that he has often listened to a musical contest between +two or three males of a Cicada, having a particularly loud voice, and +seated at a considerable distance from each other. As soon as the first +had finished his song, a second immediately began; and after he had +concluded, another began, and so on. As there is so much rivalry between +the males, it is probable that the females not only discover them by the +sounds emitted, but that, like female birds, they are excited or allured +by the male with the most attractive voice. + +I have not found any well-marked cases of ornamental differences between +the sexes of the Homoptera. Mr. Douglas informs me that there are three +British species, in which the male is black or marked with black bands, +whilst the females are pale-coloured or obscure. + + +Order, _Orthoptera_.—The males in the three saltatorial families +belonging to this Order are remarkable for their musical powers, namely +the Achetidæ or crickets, the Locustidæ for which there is no exact +equivalent name in English, and the Acridiidæ or grasshoppers. The +stridulation produced by some of the Locustidæ is so loud that it can be +heard during the night at the distance of a mile;[451] and that made by +certain species is not unmusical even to the human ear, so that the +Indians on the Amazons keep them in wicker cages. All observers agree +that the sounds serve either to call or excite the mute females. But it +has been noticed[452] that the male migratory locust of Russia (one of +the Acridiidæ) whilst coupled with the female, stridulates from anger or +jealousy when approached by another male. The house-cricket when +surprised at night uses its voice to warn its fellows.[453] In North +America the Katy-did (_Platyphyllum concavum_, one of the Locustidæ) is +described[454] as mounting on the upper branches of a tree, and in the +evening beginning “his noisy babble, while rival notes issue from the +neighbouring trees, and the groves resound with the call of +_Katy-did-she-did_, the live-long night.” Mr. Bates, in speaking of the +European field-cricket (one of the Achetidæ), says, “the male has been +observed to place itself in the evening at the entrance of its burrow, +and stridulate until a female approaches, when the louder notes are +succeeded by a more subdued tone, whilst the successful musician +caresses with his antennæ the mate he has won.”[455] Dr. Scudder was +able to excite one of these insects to answer him, by rubbing on a file +with a quill.[456] In both sexes a remarkable auditory apparatus has +been discovered by Von Siebold, situated in the front legs.[457] + +[Illustration: Fig. 10. Gryllus campestris (from Landois). + +Right-hand figure, under side of part of the wing-nervure, much +magnified, showing the teeth, _st_. + +Left-hand figure, upper surface of wing-cover, with the projecting, +smooth nervure, _r._, across which the teeth (_st_) are scraped.] + +In the three Families the sounds are differently produced. In the males +Of the Achetidæ both wing-covers have the same structure; and this in +the field-cricket (_Gryllus campestris_, fig. 10) consists, as +described by Landois,[458] of from 131 to 138 sharp, transverse ridges +or teeth (_st_) on the under side of one of the nervures of the +wing-cover. This toothed nervure is rapidly scraped across a projecting, +smooth, hard nervure (_r_) on the upper surface of the opposite wing. +First one wing is rubbed over the other, and then the movement is +reversed. Both wings are raised a little at the same time, so as to +increase the resonance. In some species the wing-covers of the males are +furnished at the base with a talc-like plate.[459] I have here given a +drawing (fig. 11) of the teeth on the under side of the nervure of +another species of Gryllus, viz. _G. domesticus_. + +[Illustration: Fig. 11. Teeth of Nervure of Gryllus domesticus (from +Landois).] + +In the Locustidæ the opposite wing-covers differ in structure (fig. 12), +and cannot, as in the last family, be indifferently used in a reversed +manner. The left wing, which acts as the bow of the fiddle, lies over +the right wing which serves as the fiddle itself. One of the nervures +(_a_) on the under surface of the former is finely serrated, and is +scraped across the prominent nervures on the upper surface of the +opposite or right wing. In our British _Phasgonura viridissima_ it +appeared to me that the serrated nervure is rubbed against the rounded +hind corner of the opposite wing, the edge of which is thickened, +coloured brown, and very sharp. In the right wing, but not in the left, +there is a little plate, as transparent as talc, surrounded by nervures, +and called the speculum. In _Ephippiger vitium_, a member of this same +family, we have a curious subordinate modification; for the wing-covers +are greatly reduced in size, but “the posterior part of the pro-thorax +is elevated into a kind of dome over the wing-covers, and which has +probably the effect of increasing the sound.”[460] + +[Illustration: Fig. 12. Chlorocœlus Tanana (from Bates). _a_, _b_. +Lobes of opposite wing-covers.] + +We thus see that the musical apparatus is more differentiated or +specialised in the Locustidæ, which includes I believe the most powerful +performers in the Order, than in the Achetidæ, in which both wing-covers +have the same structure and the same function.[461] Landois, however, +detected in one of the Locustidæ, namely in Decticus, a short and narrow +row of small teeth, mere rudiments, on the inferior surface of the +right wing-cover, which underlies the other and is never used as the +bow. I observed the same rudimentary structure on the under side of the +right wing-cover in _Phasgonura viridissima_. Hence we may with +confidence infer that the Locustidæ are descended from a form, in which, +as in the existing Achetidæ, both wing-covers had serrated nervures on +the under surface, and could be indifferently used as the bow; but that +in the Locustidæ the two wing-covers gradually became differentiated and +perfected, on the principle of the division of labour, the one to act +exclusively as the bow and the other as the fiddle. By what steps the +more simple apparatus in the Achetidæ originated, we do not know, but it +is probable that the basal portions of the wing-covers overlapped each +other formerly as at present, and that the friction of the nervures +produced a grating sound, as I find is now the case with the wing-covers +of the females.[462] A grating sound thus occasionally and accidentally +made by the males, if it served them ever so little as a love-call to +the females, might readily have been intensified through sexual +selection by fitting variations in the roughness of the nervures having +been continually preserved. + +[Illustration: Fig. 13, Hind-leg of Stenobothrus pratorum: _r_, the +stridulating ridge; lower figure, the teeth, forming the ridge, much +magnified (from Landois).] + +In the last and third Family, namely the Acridiidæ or grasshoppers, the +stridulation is produced in a very different manner, and is not so +shrill, according to Dr. Scudder, as in the preceding Families. The +inner surface of the femur (fig. 13, _r_) is furnished with a +longitudinal row of minute, elegant, lancet-shaped, elastic teeth, from +85 to 93 in number;[463] and these are scraped across the sharp, +projecting nervures on the wing-covers, which, are thus made to vibrate +and resound. Harris[464] says that when one of the males begins to play, +he first “bends the shank of the hind-leg beneath, the thigh, where it +is lodged in a furrow designed to receive it, and then draws the leg +briskly up and down. He does not play both fiddles together, but +alternately first upon one and then on the other.” In many species, the +base of the abdomen is hollowed out into a great cavity which is +believed to act as a resounding board. In Pneumora (fig. 14), a S. +African genus belonging to this same family, we meet with a new and +remarkable modification: in the males a small notched ridge projects +obliquely from each side of the abdomen, against which the hind femora +are rubbed.[465] As the male is furnished with wings, the female being +wingless, it is remarkable that the thighs are not rubbed in the usual +manner against the wing-covers; but this may perhaps be accounted for by +the unusually small size of the hind-legs. I have not been able to +examine the inner surface of the thighs, which, judging from analogy, +would be finely serrated. The species of Pneumora have been more +profoundly modified for the sake of stridulation than any other +orthopterous insect; for in the male the whole body has been converted +into a musical instrument, being distended with air, like a great +pellucid bladder, so as to increase the resonance. Mr. Trimen informs me +that at the Cape of Good Hope these insects make a wonderful noise +during the night There is one exception to the rule that the females in +these three Families are destitute of an efficient musical apparatus; +for both sexes of Ephippiger (Locustidæ) are said[466] to be thus +provided. This case may be compared with that of the reindeer, in which +species alone both sexes possess horns. Although the female orthoptera +are thus almost invariably mute, yet Landois[467] found rudiments of the +stridulating organs on the femora of the female Acridiidæ, and similar +rudiments on the under surface of the wing-covers of the female +Achetidæ; but he failed to find any rudiments in the females of +Decticus, one of the Locustidæ. In the Homoptera the mute females of +Cicada, have the proper musical apparatus in an undeveloped state; and +we shall hereafter meet in other divisions of the animal kingdom with +innumerable instances of structures proper to the male being present in +a rudimentary condition in the female. Such cases appear at first sight +to indicate that both sexes were primordially constructed in the same +manner, but that certain organs were subsequently lost by the females. +It is, however, a more probable view, as previously explained, that the +organs in question were acquired by the males and partially transferred +to the females. + +[Illustration: Fig. 14. Pneumora (from specimens in the British Museum). +Upper figure, male; lower figure, female.] + +Landois has observed another interesting fact, namely that in the +females of the Acridiidæ, the stridulating teeth on the femora remain +throughout life in the same condition in which they first appear in both +sexes during the larval state. In the males, on the other hand, they +become fully developed and acquire their perfect structure at the last +moult, when the insect is mature and ready to breed. + +From the facts now given, we see that the means by which the males +produce their sounds are extremely diversified in the Orthoptera, and +are altogether different from those employed by the Homoptera. But +throughout the animal kingdom we incessantly find the same object +gained by the most diversified means; this being due to the whole +organisation undergoing in the course of ages multifarious changes; and +as part after part varies, different variations are taken advantage of +for the same general purpose. The diversification of the means for +producing sound in the three families of the Orthoptera and in the +Homoptera, impresses the mind with the high importance of these +structures to the males, for the sake of calling or alluring the +females. We need feel no surprise at the amount of modification which +the Orthoptera have undergone in this respect, as we now know, from Dr. +Scudder’s remarkable discovery,[468] that there has been more than ample +time. This naturalist has lately found a fossil insect in the Devonian +formation of New Brunswick, which is furnished with “the well-known +tympanum or stridulating apparatus of the male Locustidæ.” This insect, +though in most respects related to the Neuroptera, appears to connect, +as is so often the case with very ancient forms, the two Orders of the +Neuroptera and Orthoptera which are now generally ranked as quite +distinct. + +I have but little more to say on the Orthoptera. Some of the species are +very pugnacious: when two male field-crickets (_Gryllus campestris_) are +confined together, they fight till one kills the other; and the species +of Mantis are described as manœuvring with their sword-like +front-limbs, like hussars with their sabres. The Chinese keep these +insects in little bamboo cages and match them like game-cocks.[469] With +respect to colour, some exotic locusts are beautifully ornamented; the +posterior wings being marked with red, blue, and black; but as +throughout the Order the two sexes rarely differ much in colour, it is +doubtful whether they owe these bright tints to sexual selection. +Conspicuous colours may be of use to these insects as a protection, on +the principle to be explained in the next chapter, by giving notice to +their enemies that they are unpalatable. Thus it has been observed[470] +that an Indian brightly-coloured locust was invariably rejected when +offered to birds and lizards. Some cases, however, of sexual differences +in colour in this Order are known. The male of an American cricket[471] +is described as being as white as ivory, whilst the female varies from +almost white to greenish-yellow or dusky. Mr. Walsh informs me that the +adult male of _Spectrum femoratum_ (one of the Phasmidæ) “is of a +shining brownish-yellow colour; the adult female being of a dull, +opaque, cinereous-brown; the young of both sexes being green.” Lastly, I +may mention that the male of one curious kind of cricket[472] is +furnished with “a long membranous appendage, which falls over the face +like a veil;” but whether this serves as an ornament is not known. + + +Order, _Neuroptera_.—Little need here be said, except in regard to +colour. In the Ephemeridæ the sexes often differ slightly in their +obscure tints;[473] but it is not probable that the males are thus +rendered attractive to the females. The Libellulidæ or dragon-flies are +ornamented with splendid green, blue, yellow, and vermilion metallic +tints; and the sexes often differ. Thus, the males of some of the +Agrionidæ, as Prof. Westwood remarks[474] “are of a rich blue with black +wings, whilst the females are fine green with colourless wings.” But in +_Agrion Ramburii_ these colours are exactly reversed in the two +sexes.[475] In the extensive N. American genus of Hetærina, the males +alone have a beautiful carmine spot at the base of each wing. In _Anax +junius_ the basal part of the abdomen in the male is a vivid +ultra-marine blue, and in the female grass-green. In the allied genus +Gomphus, on the other hand, and in some other genera, the sexes differ +but little in colour. Throughout the animal kingdom, similar cases of +the sexes of closely-allied forms either differing greatly, or very +little, or not at all, are of frequent occurrence. Although with many +Libellulidæ there is so wide a difference in colour between the sexes, +it is often difficult to say which is the most brilliant; and the +ordinary coloration of the two sexes is exactly reversed, as we have +just seen, in one species of Agrion. It is not probable that their +colours in any case have been gained as a protection. As Mr. MacLachlan, +who has closely attended to this family, writes to me, dragon-flies—the +tyrants of the insect-world—are the least liable of any insect to be +attacked by birds or other enemies. He believes that their bright +colours serve as a sexual attraction. It deserves notice, as bearing on +this subject, that certain dragon-flies appear to be attracted by +particular colours: Mr. Patterson observed[476] that the species of +Agrionidæ, of which the males are blue, settled in numbers on the blue +float of a fishing line; whilst two other species were attracted by +shining white colours. + +It is an interesting fact, first observed by Schelver, that the males, +in several genera belonging to two sub-families, when they first emerge +from the pupal state are coloured exactly like the females; but that +their bodies in a short time assume a conspicuous milky-blue tint, owing +to the exudation of a kind of oil, soluble in ether and alcohol. Mr. +MacLachlan believes that in the male of _Libellula depressa_ this change +of colour does not occur until nearly a fortnight after the +metamorphosis, when the sexes are ready to pair. + +Certain species of Neurothemis present, according to Brauer[477] a +curious case of dimorphism, some of the females having their wings +netted in the usual manner; whilst other females have them “very richly +netted as in the males of the same species.” Brauer “explains the +phenomenon on Darwinian principles by the supposition that the close +netting of the veins is a secondary sexual character in the males.” This +latter character is generally developed in the males alone, but being, +like every other masculine character, latent in the female, is +occasionally developed in them. We have here an illustration of the +manner in which the two sexes of many animals have probably come to +resemble each other, namely by variations first appearing in the males, +being preserved in them, and then transmitted to and developed in the +females; but in this particular genus a complete transference is +occasionally and abruptly effected. Mr. MacLachlan informs me of another +case of dimorphism occurring in several species of Agrion in which a +certain number of individuals are found of an orange colour, and these +are invariably females. This is probably a case of reversion, for in +the true Libellulæ, when the sexes differ in colour, the females are +always orange or yellow, so that supposing Agrion to be descended from +some primordial form having the characteristic sexual colours of the +typical Libellulæ, it would not be surprising that a tendency to vary in +this manner should occur in the females alone. + +Although many dragon-flies are such large, powerful, and fierce insects, +the males have not been observed by Mr. MacLachlan to fight together, +except, as he believes, in the case of some of the smaller species of +Agrion. In another very distinct group in this Order, namely in the +Termites or white ants, both sexes at the time of swarming may be seen +running about, “the male after the female, sometimes two chasing one +female, and contending with great eagerness who shall win the +prize.”[478] + + +Order, _Hymenoptera_.—That inimitable observer, M. Fabre,[479] in +describing the habits of Cerceris, a wasp-like insect, remarks that +“fights frequently ensue between the males for the possession of some +particular female, who sits an apparently unconcerned beholder of the +struggle for supremacy, and when the victory is decided, quietly flies +away in company with the conqueror.” Westwood[480] says that the males +of one of the saw-flies (Tenthredinæ) “have been found fighting +together, with their mandibles locked.” As M. Fabre speaks of the males +of Cerceris striving to obtain a particular female, it may be well to +bear in mind that insects belonging to this Order have the power of +recognising each other after long intervals of time, and are deeply +attached. For instance, Pierre Huber, whose accuracy no one doubts, +separated some ants, and when after an interval of four months they met +others which had formerly belonged to the same community, they mutually +recognised and caressed each other with their antennæ. Had they been +strangers they would have fought together. Again, when two communities +engage in a battle, the ants on the same side in the general confusion +sometimes attack each other, but they soon perceive their mistake, and +the one ant soothes the other.[481] + +In this Order slight differences in colour, according to sex, are +common, but conspicuous differences are rare except in the family of +Bees; yet both sexes of certain groups are so brilliantly coloured—for +instance in Chrysis, in which vermilion and metallic greens +prevail—that we are tempted to attribute the result to sexual +selection. In the Ichneumonidæ, according to Mr. Walsh,[482] the males +are almost universally lighter coloured than the females. On the other +hand, in the Tenthredinidæ the males are generally darker than the +females. In the Siricidæ the sexes frequently differ; thus the male of +_Sirex juvencus_ is banded with orange, whilst the female is dark +purple; but it is difficult to say which sex is the most ornamented. In +_Tremex columbæ_ the female is much brighter coloured than the male. +With ants, as I am informed by Mr. F. Smith, the males of several +species are black, the females being testaceous. In the family of Bees, +especially in the solitary species, as I hear from the same +distinguished entomologist, the sexes often differ in colour. The males +are generally the brightest, and in Bombus as well as in Apathus, much +more variable in colour than the females. In _Anthophora retusa_ the +male is of a rich fulvous-brown, whilst the female is quite black: so +are the females of several species of Xylocopa, the males being bright +yellow. In an Australian bee (_Lestis bombylans_), the female is of an +extremely brilliant steel-blue, sometimes tinted with vivid green; the +male being of a bright brassy colour clothed with rich fulvous +pubescence. As in this group the females are provided with excellent +defensive weapons in their stings, it is not probable that they have +come to differ in colour from the males for the sake of protection. + +_Mutilla Europæa_ emits a stridulating noise; and according to +Goureau[483] both sexes have this power. He attributes the sound to the +friction of the third and preceding abdominal segments; and I find that +these surfaces are marked with very fine concentric ridges, but so is +the projecting thoracic collar, on which the head articulates; and this +collar, when scratched with the point of a needle, emits the proper +sound. It is rather surprising that both sexes should have the power of +stridulating, as the male is winged and the female wingless. It is +notorious that Bees express certain emotions, as of anger, by the tone +of their humming, as do some dipterous insects; but I have not referred +to these sounds, as they are not known to be in any way connected with +the act of courtship. + + +Order, _Coleoptera_ (Beetles).—Many beetles are coloured so as to +resemble the surfaces which they habitually frequent. Other species are +ornamented with gorgeous metallic tints,—for instance, many Carabidæ, +which live on the ground and have the power of defending themselves by +an intensely acrid secretion,—the splendid diamond-beetles which are +protected by an extremely hard covering,—many species of Chrysomela, +such as _C. cerealis_, a large species beautifully striped with various +colours, and in Britain confined to the bare summit of Snowdon,—and a +host of other species. These splendid colours, which are often arranged +in stripes, spots, crosses and other elegant patterns, can hardly be +beneficial, as a protection, except in the case of some flower-feeding +species; and we cannot believe that they are purposeless. Hence the +suspicion arises, that they serve as a sexual attraction; but we have no +evidence on this head, for the sexes rarely differ in colour. Blind +beetles, which cannot of course behold each other’s beauty, never +exhibit, as I hear from Mr. Waterhouse, jun., bright colours, though +they often have polished coats: but the explanation of their obscurity +may be that blind insects inhabit caves and other obscure stations. + +Some Longicorns, however, especially certain Prionidæ, offer an +exception to the common rule that the sexes of beetles do not differ in +colour. Most of these insects are large and splendidly coloured. The +males in the genus Pyrodes,[484] as I saw in Mr. Bates’ collection, are +generally redder but rather duller than the females, the latter being +coloured of a more or less splendid golden green. On the other hand, in +one species the male is golden-green, the female being richly tinted +with red and purple. In the genus Esmeralda the sexes differ so greatly +in colour that they have been ranked as distinct species: in one species +both are of a beautiful shining green, but the male has a red thorax. On +the whole, as far as I could judge, the females of those Prionidæ, in +which the sexes differ, are coloured more richly than the males; and +this does not accord with the common rule in regard to colour when +acquired through sexual selection. + +[Illustration: Fig. 15. Chalcosoma atlas. Upper figure, male (reduced); +lower figure, female (nat. size).] + +[Illustration: Fig. 16. Copris isidis. (Left-hand figures, males.)] + +[Illustration: Fig. 17. Phanæus faunus.] + +[Illustration: Fig. 18. Dipelicus cantori.] + +[Illustration: Fig. 19. Onthophagus rangifer, enlarged.] + +A most remarkable distinction between the sexes of many beetles is +presented by the great horns which rise from the head, thorax, or +clypeus of the males; and in some few cases from the under surface of +the body. These horns, in the great family of the Lamellicorns, resemble +those of various quadrupeds, such as stags, rhinoceroses, &c., and are +wonderful both from their size and diversified shapes. Instead of +describing them, I have given figures of the males and females of some +of the more remarkable forms. (Figs. 15 to 19.) The females generally +exhibit rudiments of the horns in the form of small knobs or ridges; but +some are destitute of even a rudiment. On the other hand, the horns are +nearly as well developed in the female as in the male of _Phanæus +lancifer_; and only a little less well developed in the females of some +other species of the same genus and of Copris. In the several +subdivisions of the family, the differences in structure of the horns do +not run parallel, as I am informed by Mr. Bates, with their more +important and characteristic differences; thus within the same natural +section of the genus Onthophagus, there are species which have either a +single cephalic horn, or two distinct horns. + +In almost all cases, the horns are remarkable from their excessive +variability; so that a graduated series can be formed, from the most +highly developed males to others so degenerate that they can barely be +distinguished from the females. Mr. Walsh[485] found that in _Phanæus +carnifex_ the horns were thrice as long in some males as in others. Mr. +Bates, after examining above a hundred males of _Onthophagus rangifer_ +(fig. 19), thought that he had at last discovered a species in which +the horns did not vary; but further research proved the contrary. + +The extraordinary size of the horns, and their widely different +structure in closely-allied forms, indicate that they have been formed +for some important purpose; but their excessive variability in the males +of the same species leads to the inference that this purpose cannot be +of a definite nature. The horns do not show marks of friction, as if +used for any ordinary work. Some authors suppose[486] that as the males +wander much more than the females, they require horns as a defence +against their enemies; but in many cases the horns do not seem well +adapted for defence, as they are not sharp. The most obvious conjecture +is that they are used by the males for fighting together; but they have +never been observed to fight; nor could Mr. Bates, after a careful +examination of numerous species, find any sufficient evidence in their +mutilated or broken condition of their having been thus used. If the +males had been habitual fighters, their size would probably have been +increased through sexual selection, so as to have exceeded that of the +female; but Mr. Bates, after comparing the two sexes in above a hundred +species of the Copridæ, does not find in well-developed individuals any +marked difference in this respect. There is, moreover, one beetle, +belonging to the same great division of the Lamellicorns, namely +Lethrus, the males of which are known to fight, but they are not +provided with horns, though their mandibles are much larger than those +of the female. + +The conclusion, which best agrees with the fact of the horns having been +so immensely yet not fixedly developed,—as shewn by their extreme +variability in the same species and by their extreme diversity in +closely-allied species—is that they have been acquired as ornaments. +This view will at first appear extremely improbable; but we shall +hereafter find with many animals, standing much higher in the scale, +namely fishes, amphibians, reptiles and birds, that various kinds of +crests, knobs, horns and combs have been developed apparently for this +sole purpose. + +[Illustration: Fig. 20. Onitis furcifer, male, viewed from beneath.] + +[Illustration: Fig. 21. Left-hand figure, male of Onitis furcifer, +viewed laterally. Right-hand figure, female. _a._ Rudiment of cephalic +horn. _b._ Trace of thoracic horn or crest.] + +The males of _Onitis furcifer_ (fig. 20) are furnished with singular +projections on their anterior femora, and with a great fork or pair of +horns on the lower surface of the thorax. This situation seems extremely +ill adapted for the display of these projections, and they may be of +some real service; but no use can at present be assigned to them. It is +a highly remarkable fact, that although the males do not exhibit even a +trace of horns on the upper surface of the body, yet in the females a +rudiment of a single horn on the head (fig. 21, _a_), and of a crest +(_b_) on the thorax, are plainly visible. That the slight thoracic crest +in the female is a rudiment of a projection proper to the male, though +entirely absent in the male of this particular species, is clear: for +the female of _Bubas bison_ (a form which comes next to _Onitis_) has a +similar slight crest on the thorax, and the male has in the same +situation a great projection. So again there can be no doubt that the +little point (_a_) on the head of the female _Onitis furcifer_, as well +of the females of two or three allied species, is a rudimentary +representative of the cephalic horn, which is common to the males of so +many lamellicorn beetles, as in Phanæus, fig. 17. The males indeed of +some unnamed beetles in the British Museum, which are believed actually +to belong to the genus Onitis, are furnished with a similar horn. The +remarkable nature of this case will be best perceived by an +illustration: the Ruminant quadrupeds run parallel with the lamellicorn +beetles, in some females possessing horns as large as those of the male, +in others having them much smaller, or existing as mere rudiments +(though this is as rare with ruminants as it is common with +Lamellicorns), or in having none at all. Now if a new species of deer or +sheep were discovered with the female bearing distinct rudiments of +horns, whilst the head of the male was absolutely smooth, we should have +a case like that of _Onitis furcifer_. + +In this case the old belief of rudiments having been created to complete +the scheme of nature is so far from holding good, that all ordinary +rules are completely broken through. The view which seems the most +probable is that some early progenitor of Onitis acquired, like other +Lamellicorns, horns on the head and thorax, and then transferred them, +in a rudimentary condition, as with so many existing species, to the +female, by whom they have ever since been retained. The subsequent loss +of the horns by the male may have resulted through the principle of +compensation from the development of the projections on the lower +surface, whilst the female has not been thus affected, as she is not +furnished with these projections, and consequently has retained the +rudiments of the horns on the upper surface. Although this view is +supported by the case of Bledius immediately to be given, yet the +projections on the lower surface differ greatly in structure and +development in the males of the several species of Onitis, and are even +rudimentary in some; nevertheless the upper surface in all these species +is quite destitute of horns. As secondary sexual characters are so +eminently variable, it is possible that the projections on the lower +surface may have been first acquired by some progenitor of Onitis and +produced their effect through compensation, and then have been in +certain cases almost completely lost. + +[Illustration: Fig. 22. Bledius taurus, magnified. Left-hand figure, +male; right-hand figure, female.] + +All the cases hitherto given refer to the Lamellicorns, but the males of +some few other beetles, belonging to two widely distinct groups, namely, +the Curculionidæ and Staphylinidæ, are furnished with horns,—in the +former on the lower surface of the body,[487] in the latter on the upper +surface of the head and thorax. In the Staphylinidæ the horns of the +males in the same species are extraordinarily variable, just as we have +seen with the Lamellicorns. In Siagonium we have a case of dimorphism, +for the males can be divided into two sets, differing greatly in the +size of their bodies, and in the development of their horns, without any +intermediate gradations. In a species of Bledius (fig. 22), also +belonging to the Staphylinidæ, male specimens can be found in the same +locality, as Professor Westwood states, “in which the central horn of +the thorax is very large, but the horns of the head quite rudimental; +and others, in which the thoracic horn is much shorter, whilst the +protuberances on the head are long.”[488] Here, then, we apparently have +an instance of compensation of growth, which throws light on the curious +case just given of the loss of the upper horns by the males of _Onitis +furcifer_. + + +_Law of Battle._—Some male beetles, which seem ill fitted for fighting, +nevertheless engage in conflicts for the possession of the females. Mr. +Wallace[489] saw two males of _Leptorhynchus angustatus_, a linear +beetle with a much elongated rostrum, “fighting for a female, who stood +close by busy at her boring. They pushed at each other with their +rostra, and clawed and thumped, apparently in the greatest rage.” The +smaller male, however, “soon ran away, acknowledging himself +vanquished.” In some few cases the males are well adapted for fighting, +by possessing great toothed mandibles, much larger than those of the +females. This is the case with the common stag-beetle (_Lucanus +cervus_), the males of which emerge from the pupal state about a week +before the other sex, so that several may often be seen pursuing the +same female. At this period they engage in fierce conflicts. When Mr. A. +H. Davis[490] enclosed two males with one female in a box, the larger +male severely pinched the smaller one, until he resigned his +pretensions. A friend informs me that when a boy he often put the males +together to see them fight, and he noticed that they were much bolder +and fiercer than the females, as is well known to be the case with the +higher animals. The males would seize hold of his finger, if held in +front, but not so the females. With many of the Lucanidæ, as well as +with the above-mentioned Leptorhynchus, the males are larger and more +powerful insects than the females. The two sexes of _Lethrus cephalotes_ +(one of the Lamellicorns) inhabit the same burrow; and the male has +larger mandibles than the female. If, during the breeding-season, a +strange male attempts to enter the burrow, he is attacked; the female +does not remain passive, but closes the mouth of the burrow, and +encourages her mate by continually pushing him on from behind. The +action does not cease until the aggressor is killed or runs away.[491] +The two sexes of another lamellicorn beetle, the _Ateuchus cicatricosus_ +live in pairs, and seem much attached to each other; the male excites +the female to roll the balls of dung in which the ova are deposited; and +if she is removed, he becomes much agitated. If the male is removed, the +female ceases all work, and as M. Brulerie[492] believes, would remain +on the spot until she died. + +The great mandibles of the male Lucanidæ are extremely variable both in +size and structure, and in this respect resemble the horns on the head +and thorax of many male Lamellicorns and Staphylinidæ. A perfect series +can be formed from the best-provided to the worst-provided or degenerate +males. Although the mandibles of the common stag-beetle, and probably of +many other species, are used as efficient weapons for fighting, it is +doubtful whether their great size can thus be accounted for. We have +seen that with the _Lucanus elaphus_ of N. America they are used for +seizing the female. As they are so conspicuous and so elegantly +branched, the suspicion has sometimes crossed my mind that they may be +serviceable to the males as an ornament, in the same manner as the horns +on the head and thorax of the various above described species. The male +_Chiasognathus grantii_ of S. Chile—a splendid beetle belonging to the +same family—has enormously-developed mandibles (fig. 23); he is bold +and pugnacious; when threatened on any side he faces round, opening his +great jaws, and at the same time stridulating loudly; but the mandibles +were not strong enough to pinch my finger so as to cause actual pain. + +[Illustration: Fig. 23. Chiasognathus grantii, reduced. Upper figure, +male; lower figure, female.] + +Sexual selection, which implies the possession of considerable +perceptive powers and of strong passions, seems to have been more +effective with the Lamellicorns than with any other family of the +Coleoptera or beetles. With some species the males are provided with +weapons for fighting; some live in pairs and show mutual affection; +many have the power of stridulating when excited; many are furnished +with the most extraordinary horns, apparently for the sake of ornament; +some which are diurnal in their habits are gorgeously coloured; and, +lastly, several of the largest beetles in the world belong to this +family, which was placed by Linnæus and Fabricius at the head of the +Order of the Coleoptera.[493] + + +_Stridulating organs._—Beetles belonging to many and widely distinct +families possess these organs. The sound can sometimes be heard at the +distance of several feet or even yards,[494] but is not comparable with +that produced by the Orthoptera. The part which may be called the rasp +generally consists of a narrow slightly-raised surface, crossed by very +fine, parallel ribs, sometimes so fine as to cause iridescent colours, +and having a very elegant appearance under the microscope. In some +cases, for instance, with Typhæus, it could be plainly seen that +extremely minute, bristly, scale-like prominences, which cover the whole +surrounding surface in approximately parallel lines, give rise to the +ribs of the rasp by becoming confluent and straight, and at the same +time more prominent and smooth. A hard ridge on any adjoining part of +the body, which in some cases is specially modified for the purpose, +serves as the scraper for the rasp. The scraper is rapidly moved across +the rasp, or conversely the rasp across the scraper. + +[Illustration: Fig. 24. Necrophorus (from Landois). _r._ The two rasps. +Left-hand figure, part of the rasp highly magnified.] + +These organs are situated in widely different positions. In the +carrion-beetles (Necrophorus) two parallel rasps (_r_, fig. 24) stand on +the dorsal surface of the fifth abdominal segment, each rasp being +crossed, as described by Landois,[495] by from 126 to 140 fine ribs. +These ribs are scraped by the posterior margins of the elytra, a small +portion of which projects beyond the general outline. In many +Crioceridæ, and in _Clythra 4-punctata_ (one of the Chrysomelidæ), and +in some Tenebrionidæ, &c.,[496] the rasp is seated on the dorsal apex of +the abdomen, on the pygidium or pro-pygidium, and is scraped as above by +the elytra. In Heterocerus, which belongs to another family, the rasps +are placed on the sides of the first abdominal segment, and are scraped +by ridges on the femora.[497] In certain Curculionidæ and Carabidæ,[498] +the parts are completely reversed in position, for the rasps are seated +on the inferior surface of the elytra, near their apices, or along their +outer margins, and the edges of the abdominal segments serve as the +scrapers. In _Pelobius hermanni_ (one of Dytiscidæ or water-beetles) a +strong ridge runs parallel and near to the sutural margin of the elytra, +and is crossed by ribs, coarse in the middle part, but becoming +gradually finer at both ends, especially at the upper end; when this +insect is held under water or in the air, a stridulating noise is +produced by scraping the extreme horny margin of the abdomen against the +rasp. In a great number of long-horned beetles (Longicornia) the organs +are altogether differently situated, the rasp being on the meso-thorax, +which is rubbed against the pro-thorax; Landois counted 238 very fine +ribs on the rasp of _Cerambyx heros_. + +[Illustration: Fig. 25. Hind-leg of Geotrupes stercorarius (from +Landois). _r._ Rasp. _c._ Coxa. _f._ Femur. _t._ Tibia. _tr._ Tarsi.] + +Many Lamellicorns have the power of stridulating, and the organs differ +greatly in position. Some species stridulate very loudly, so that when +Mr. F. Smith caught a _Trox sabulosus_, a gamekeeper who stood by +thought that he had caught a mouse; but I failed to discover the proper +organs in this beetle. In Geotrupes and Typhæus a narrow ridge runs +obliquely across (_r_, fig. 25) the coxa of each hind-leg, having in +_G. stercorarius_ 84 ribs, which are scraped by a specially-projecting +part of one of the abdominal segments. In the nearly allied _Copris +lunaris_, an excessively narrow fine rasp runs along the sutural margin +of the elytra, with another short rasp near the basal outer margin; but +in some other Coprini the rasp is seated, according to Leconte,[499] on +the dorsal surface of the abdomen. In Oryctes it is seated on the +pro-pygidium, and in some other Dynastini, according to the same +entomologist, on the under surface of the elytra. Lastly, Westring +states that in _Omaloplia brunnea_ the rasp is placed on the +pro-sternum, and the scraper on the meta-sternum, the parts thus +occupying the under surface of the body, instead of the upper surface as +in the Longicorns. + +We thus see that the stridulating organs in the different coleopterous +families are wonderfully diversified in position, but not much in +structure. Within the same family some species are provided with these +organs, and some are quite destitute of them. This diversity is +intelligible, if we suppose that originally various species made a +shuffling or hissing noise by the rubbing together of the hard and rough +parts of their bodies which were in contact; and that from the noise +thus produced being in some way useful, the rough surfaces were +gradually developed into regular stridulating organs. Some beetles as +they move, now produce, either intentionally or unintentionally, a +shuffling noise, without possessing any proper organs for the purpose. +Mr. Wallace informs me that the _Euchirus longimanus_ (a Lamellicorn, +with the anterior legs wonderfully elongated in the male) “makes, whilst +moving, a low hissing sound by the protrusion and contraction of the +abdomen; and when seized it produces a grating sound by rubbing its +hind-legs against the edges of the elytra.” The hissing sound is clearly +due to a narrow rasp running along the sutural margin of each elytron; +and I could likewise make the grating sound by rubbing the shagreened +surface of the femur against the granulated margin of the corresponding +elytron; but I could not here detect any proper rasp; nor is it likely +that I could have overlooked it in so large an insect. After examining +Cychrus and reading what Westring has written in his two papers about +this beetle, it seems very doubtful whether it possesses any true rasp, +though it has the power of emitting a sound. + +From the analogy of the Orthoptera and Homoptera, I expected to find +that the stridulating organs in the Coleoptera differed according to +sex; but Landois, who has carefully examined several species, observed +no such difference; nor did Westring; nor did Mr. G. R. Crotch in +preparing the numerous specimens which he had the kindness to send me +for examination. Any slight sexual difference, however, would be +difficult to detect, on account of the great variability of these +organs. Thus in the first pair of the _Necrophorus humator_ and of the +_Pelobius_ which I examined, the rasp was considerably larger in the +male than in the female; but not so with succeeding specimens. In +_Geotrupes stercorarius_ the rasp appeared to me thicker, opaquer, and +more prominent in three males than in the same number of females; +consequently my son, Mr. F. Darwin, in order to discover whether the +sexes differed in their power of stridulating, collected 57 living +specimens, which he separated into two lots, according as they made, +when held in the same manner, a greater or lesser noise. He then +examined their sexes, but found that the males were very nearly in the +same proportion to the females in both lots. Mr. F. Smith has kept alive +numerous specimens of _Mononychus pseudacori_ (Curculionidæ), and is +satisfied that both sexes stridulate, and apparently in an equal degree. + +Nevertheless the power of stridulating is certainly a sexual character +in some few Coleoptera. Mr. Crotch has discovered that the males alone +of two species of Heliopathes (Tenebrionidæ) possess stridulating +organs. I examined five males of _H. gibbus_, and in all these there was +a well-developed rasp, partially divided into two, on the dorsal surface +of the terminal abdominal segment; whilst in the same number of females +there was not even a rudiment of the rasp, the membrane of this segment +being transparent and much thinner than in the male. In _H. +cribratostriatus_ the male has a similar rasp, excepting that it is not +partially divided into two portions, and the female is completely +destitute of this organ; but in addition the male has on the apical +margins of the elytra, on each side of the suture, three or four short +longitudinal ridges, which are crossed by extremely fine ribs, parallel +to and resembling those on the abdominal rasp; whether these ridges +serve as an independent rasp, or as a scraper for the abdominal rasp, I +could not decide: the female exhibits no trace of this latter structure. + +Again, in three species of the Lamellicorn genus Oryctes, we have a +nearly parallel case. In the females of _O. gryphus_ and _nasicornis_ +the ribs on the rasp of the pro-pygidium are less continuous and less +distinct than in the males; but the chief difference is that the whole +upper surface of this segment, when held in the proper light, is seen to +be clothed with hairs, which are absent or are represented by +excessively fine down in the males. It should be noticed that in all +Coleoptera the effective part of the rasp is destitute of hairs. In _O. +senegalensis_ the difference between the sexes is more strongly marked, +and this is best seen when the proper segment is cleaned and viewed as a +transparent object. In the female the whole surface is covered with +little separate crests, bearing spines; whilst in the male these crests +become, in proceeding towards the apex, more and more confluent, +regular, and naked; so that three-fourths of the segment is covered with +extremely fine parallel ribs, which are quite absent in the female. In +the females, however, of all three species of Oryctes, when the abdomen +of a softened specimen is pushed backwards and forwards, a slight +grating or stridulating sound can be produced. + +In the case of the Heliopathes and Oryctes there can hardly be a doubt +that the males stridulate in order to call or to excite the females; but +with most beetles the stridulation apparently serves both sexes as a +mutual call. This view is not rendered improbable from beetles +stridulating under various emotions; we know that birds use their voices +for many purposes besides singing to their mates. The great +Chiasognathus stridulates in anger or defiance; many species do the same +from distress or fear, when held so that they cannot escape; Messrs. +Wollaston and Crotch were able, by striking the hollow stems of trees in +the Canary Islands, to discover the presence of beetles belonging to the +genus Acalles by their stridulation. Lastly the male Ateuchus +stridulates to encourage the female in her work, and from distress when +she is removed.[500] Some naturalists believe that beetles make this +noise to frighten away their enemies; but I cannot think that the +quadrupeds and birds which are able to devour the larger beetles with +their extremely hard coats, would be frightened by so slight a grating +sound. The belief that the stridulation serves as a sexual call is +supported by the fact that death-ticks (_Anobium tessellatum_) are well +known to answer each other’s ticking, or, as I have myself observed, a +tapping noise artificially made; and Mr. Doubleday informs me that he +has twice or thrice observed a female ticking,[501] and in the course of +an hour or two has found her united with a male, and on one occasion +surrounded by several males. Finally, it seems probable that the two +sexes of many kinds of beetles were at first enabled to find each other +by the slight shuffling noise produced by the rubbing together of the +adjoining parts of their hard bodies; and that as the males or females +which made the greatest noise succeeded best in finding partners, the +rugosities on various parts of their bodies were gradually developed by +means of sexual selection into true stridulating organs. + + + + +CHAPTER XI. + +INSECTS, _continued_.—ORDER LEPIDOPTERA. + + + Courtship of butterflies—Battles—Ticking noise—Colours + common to both sexes, or more brilliant in the + males—Examples—Not due to the direct action of the conditions + of life—Colours adapted for protection—Colours of + moths—Display—Perceptive powers of the + Lepidoptera—Variability—Causes of the difference in colour + between the males and females—Mimickry, female butterflies + more brilliantly coloured than the males—Bright colours of + caterpillars—Summary and concluding remarks on the secondary + sexual characters of insects—Birds and insects compared. + + +In this great Order the most interesting point for us is the difference +in colour between the sexes of the same species, and between the +distinct species of the same genus. Nearly the whole of the following +chapter will be devoted to this subject; but I will first make a few +remarks on one or two other points. Several males may often be seen +pursuing and crowding round the same female. Their courtship appears to +be a prolonged affair, for I have frequently watched one or more males +pirouetting round a female until I became tired, without seeing the end +of the courtship. Although butterflies are such weak and fragile +creatures, they are pugnacious, and an Emperor butterfly[502] has been +captured with the tips of its wings broken from a conflict with another +male. Mr. Collingwood in speaking of the frequent battles between the +butterflies of Borneo says, “They whirl round each other with the +greatest rapidity, and appear to be incited by the greatest ferocity.” +One case is known of a butterfly, namely the _Ageronia feronia_, which +makes a noise like that produced by a toothed wheel passing under a +spring catch, and which could be heard at the distance of several yards. +At Rio de Janeiro this sound was noticed by me, only when two were +chasing each other in an irregular course, so that it is probably made +during the courtship of the sexes; but I neglected to attend to this +point.[503] + +Every one has admired the extreme beauty of many butterflies and of some +moths; and we are led to ask, how has this beauty been acquired? Have +their colours and diversified patterns simply resulted from the direct +action of the physical conditions to which these insects have been +exposed, without any benefit being thus derived? Or have successive +variations been accumulated and determined either as a protection or for +some unknown purpose, or that one sex might be rendered attractive to +the other? And, again, what is the meaning of the colours being widely +different in the males and females of certain species, and alike in the +two sexes of other species? Before attempting to answer these questions +a body of facts must be given. + +With most of our English butterflies, both those which are beautiful, +such as the admiral, peacock, and painted lady (Vanessæ), and those +which are plain-coloured, such as the meadow-browns (Hipparchiæ), the +sexes are alike. This is also the case with the magnificent Heliconidæ +and Danaidæ of the tropics. But in certain other tropical groups, and +with some of our English butterflies, as the purple emperor, orange-tip, +&c. (_Apatura Iris_ and _Anthocharis cardamines_), the sexes differ +either greatly or slightly in colour. No language suffices to describe +the splendour of the males of some tropical species. Even within the +same genus we often find species presenting an extraordinary difference +between the sexes, whilst others have their sexes closely alike. Thus in +the South American genus Epicalia, Mr. Bates, to whom I am much indebted +for most of the following facts and for looking over this whole +discussion, informs me that he knows twelve species, the two sexes of +which haunt the same stations (and this is not always the case with +butterflies), and therefore cannot have been differently affected by +external conditions[504]. In nine of these species the males rank +amongst the most brilliant of all butterflies, and differ so greatly +from the comparatively plain females that they were formerly placed in +distinct genera. The females of these nine species resemble each other +in their general type of coloration, and likewise resemble both sexes in +several allied genera, found in various parts of the world. Hence in +accordance with the descent-theory we may infer that these nine species, +and probably all the others of the genus, are descended from an +ancestral form which was coloured in nearly the same manner. In the +tenth species the female still retains the same general colouring, but +the male resembles her, so that he is coloured in a much less gaudy and +contrasted manner than the males of the previous species. In the +eleventh and twelfth species, the females depart from the type of +colouring which is usual with their sex in this genus, for they are +gaily decorated in nearly the same manner as the males, but in a +somewhat less degree. Hence in these two species the bright colours of +the males seem to have been transferred to the females; whilst the male +of the tenth species has either retained or recovered the plain colours +of the female as well as of the parent-form of the genus; the two sexes +being thus rendered in both cases, though in an opposite manner, nearly +alike. In the allied genus Eubagis, both sexes of some of the species +are plain-coloured and nearly alike; whilst with the greater number the +males are decorated with beautiful metallic tints, in a diversified +manner, and differ much from their females. The females throughout the +genus retain the same general style of colouring, so that they commonly +resemble each other much more closely than they resemble their own +proper males. + +In the genus Papilio, all the species of the Æneas group are remarkable +for their conspicuous and strongly contrasted colours, and they +illustrate the frequent tendency to gradation in the amount of +difference between the sexes. In a few species, for instance in _P. +ascanius_, the males and females are alike; in others the males are a +little or very much more superbly coloured than the females. The genus +Junonia allied to our Vanessæ offers a nearly parallel case, for +although the sexes of most of the species resemble each other and are +destitute of rich colours, yet in certain species, as in _J. œnone_, +the male is rather more brightly coloured than the female, and in a few +(for instance _J. andremiaja_) the male is so different from the female +that he might be mistaken for an entirely distinct species. + +Another striking case was pointed out to me in the British museum by Mr. +A. Butler, namely one of the Tropical American Theclæ, in which both +sexes are nearly alike and wonderfully splendid; in another, the male +is coloured in a similarly gorgeous manner, whilst the whole upper +surface of the female is of a dull uniform brown. Our common little +English blue butterflies of the genus Lycæna, illustrate the various +differences in colour between the sexes, almost as well, though not in +so striking a manner, as the above exotic genera. In _Lycæna agestis_ +both sexes have wings of a brown colour, bordered with small ocellated +orange spots, and are consequently alike. In _L. œgon_ the wings of +the male are of a fine blue, bordered with black; whilst the wings of +the female are brown, with a similar border, and closely resemble those +of _L. agestis_. Lastly, in _L. arion_ both sexes are of a blue colour +and nearly alike, though in the female the edges of the wings are rather +duskier, with the black spots plainer; and in a bright blue Indian +species both sexes are still more closely alike. + +I have given the foregoing cases in some detail in order to shew, in the +first place, that when the sexes of butterflies differ, the male as a +general rule is the most beautiful, and departs most from the usual type +of colouring of the group to which the species belongs. Hence in most +groups the females of the several species resemble each other much more +closely than do the males. In some exceptional cases, however, to which +I shall hereafter allude, the females are coloured more splendidly than +the males. In the second place these cases have been given to bring +clearly before the mind that within the same genus, the two sexes +frequently present every gradation from no difference in colour to so +great a difference that it was long before the two were placed by +entomologists in the same genus. In the third place, we have seen that +when the sexes nearly resemble each other, this apparently may be due +either to the male having transferred his colours to the female, or to +the male having retained, or perhaps recovered, the primordial colours +of the genus to which the species belongs. It also deserves notice that +in those groups in which the sexes present any difference of colour, the +females usually resemble the males to a certain extent, so that when the +males are beautiful to an extraordinary degree, the females almost +invariably exhibit some degree of beauty. From the numerous cases of +gradation in the amount of difference between the sexes, and from the +prevalence of the same general type of coloration throughout the whole +of the same group, we may conclude that the causes, whatever they may +be, which have determined the brilliant colouring of the males alone of +some species, and of both sexes in a more or less equal degree of other +species, have generally been the same. + +As so many gorgeous butterflies inhabit the tropics, it has often been +supposed that they owe their colours to the great heat and moisture of +these zones; but Mr. Bates[505] has shewn by the comparison of various +closely-allied groups of insects from the temperate and tropical +regions, that this view cannot be maintained; and the evidence becomes +conclusive when brilliantly-coloured males and plain-coloured females of +the same species inhabit the same district, feed on the same food, and +follow exactly the same habits of life. Even when the sexes resemble +each other, we can hardly believe that their brilliant and +beautifully-arranged colours are the purposeless result of the nature of +the tissues, and the action of the surrounding conditions. + +With animals of all kinds, whenever colour has been modified for some +special purpose, this has been, as far as we can judge, either for +protection or as an attraction between the sexes. With many species of +butterflies the upper surfaces of the wings are obscurely coloured, and +this in all probability leads to their escaping observation and danger. +But butterflies when at rest would be particularly liable to be attacked +by their enemies; and almost all the kinds when resting raise their +wings vertically over their backs, so that the lower sides alone are +exposed to view. Hence it is this side which in many cases is obviously +coloured so as to imitate the surfaces on which these insects commonly +rest. Dr. Rössler, I believe, first noticed the similarity of the closed +wings of certain Vanessæ and other butterflies to the bark of trees. +Many analogous and striking facts could be given. The most interesting +one is that recorded by Mr. Wallace[506] of a common Indian and Sumatran +butterfly (Kallima), which disappears like magic when it settles in a +bush; for it hides its head and antennæ between its closed wings, and +these in form, colour, and veining cannot be distinguished from a +withered leaf together with the footstalk. In some other cases the lower +surfaces of the wings are brilliantly coloured, and yet are protective; +thus in _Thecla rubi_ the wings when closed are of an emerald green and +resemble the young leaves of the bramble, on which this butterfly in the +spring may often be seen seated. + +Although the obscure tints of the upper or under surface of many +butterflies no doubt serve to conceal them, yet we cannot possibly +extend this view to the brilliant and conspicuous colours of many kinds, +such as our admiral and peacock Vanessæ, our white cabbage-butterflies +(Pieris), or the great swallow-tail Papilio which haunts the open +fens—for these butterflies are thus rendered visible to every living +creature. With these species both sexes are alike; but in the common +brimstone butterfly (_Gonepteryx rhamni_), the male is of an intense +yellow, whilst the female is much paler; and in the orange-tip +(_Anthocharis cardamines_) the males alone have the bright orange tips +to their wings. In these cases the males and females are equally +conspicuous, and it is not credible that their difference in colour +stands in any relation to ordinary protection. Nevertheless it is +possible that the conspicuous colours of many species may be in an +indirect manner beneficial, as will hereafter be explained, by leading +their enemies at once to recognise them as unpalatable. Even in this +case it does not certainly follow that their bright colours and +beautiful patterns were acquired for this special purpose. In some other +remarkable cases, beauty has been gained for the sake of protection, +through the imitation of other beautiful species, which inhabit the same +district and enjoy an immunity from attack by being in some way +offensive to their enemies. + +The female of our orange-tip butterfly, above referred to, and of an +American species (_Anth. genutia_) probably shew us, as Mr. Walsh has +remarked to me, the primordial colours of the parent-species of the +genus; for both sexes of four or five widely-distributed species are +coloured in nearly the same manner. We may infer here, as in several +previous cases, that it is the males of _Anth. cardamines_ and _genutia_ +which have departed from the usual type of colouring of their genus. In +the _Anth. sara_ from California, the orange-tips have become partially +developed in the female; for her wings are tipped with reddish-orange, +but paler than in the male, and slightly different in some other +respects. In an allied Indian form, the _Iphias glaucippe_, the +orange-tips are fully developed in both sexes. In this Iphias the under +surface of the wings marvellously resembles, as pointed out to me by Mr. +A. Butler, a pale-coloured leaf; and in our English orange-tip, the +under surface resembles the flower-head of the wild parsley, on which it +may be seen going to rest at night.[507] The same reasoning power which +compels us to believe that the lower surfaces have here been coloured +for the sake of protection, leads us to deny that the wings have been +tipped, especially when this character is confined to the males, with +bright orange for the same purpose. + + +Turning now to Moths: most of these rest motionless with their wings +depressed during the whole or greater part of the day; and the upper +surfaces of their wings are often shaded and coloured in an admirable +manner, as Mr. Wallace has remarked, for escaping detection. With most +of the Bombycidæ and Noctuidæ,[508] when at rest, the front-wings +overlap and conceal the hind-wings; so that the latter might be brightly +coloured without much risk; and they are thus coloured in many species +of both families. During the act of flight, moths would often be able to +escape from their enemies; nevertheless, as the hind-wings are then +fully exposed to view, their bright colours must generally have been +acquired at the cost of some little risk. But the following fact shews +us how cautious we ought to be in drawing conclusions on this head. The +common yellow under-wings (Triphaena) often fly about during the day or +early evening, and are then conspicuous from the colour of their +hind-wings. It would naturally be thought that this would be a source of +danger; but Mr. J. Jenner Weir believes that it actually serves them as +a means of escape, for birds strike at these brightly coloured and +fragile surfaces, instead of at the body. For instance, Mr. Weir turned +into his aviary a vigorous specimen of _Triphaena pronuba_, which was +instantly pursued by a robin; but the bird’s attention being caught by +the coloured wings, the moth was not captured until after about fifty +attempts, and small portions of the wings were repeatedly broken off. He +tried the same experiment, in the open air, with a _T. fimbria_ and +swallow; but the large size of this moth probably interfered with its +capture.[509] We are thus reminded of a statement made by Mr. +Wallace,[510] namely, that in the Brazilian forests and Malayan islands, +many common and highly-decorated butterflies are weak flyers, though +furnished with a broad expanse of wings; and they “are often captured +with pierced and broken wings, as if they had been seized by birds, from +which they had escaped: if the wings had been much smaller in proportion +to the body, it seems probable that the insect would more frequently +have been struck or pierced in a vital part, and thus the increased +expanse of the wings may have been indirectly beneficial.” + + +_Display._—The bright colours of butterflies and of some moths are +specially arranged for display, whether or not they serve in addition as +a protection. Bright colours would not be visible during the night; and +there can be no doubt that moths, taken as a body, are much less gaily +decorated than butterflies, all of which are diurnal in their habits. +But the moths in certain families, such as the Zygænidæ, various +Sphingidæ, Uraniidæ, some Arctiidæ and Saturniidæ, fly about during the +day or early evening, and many of these are extremely beautiful, being +far more brightly coloured than the strictly nocturnal kinds. A few +exceptional cases, however, of brightly-coloured nocturnal species have +been recorded.[511] + +There is evidence of another kind in regard to display. Butterflies, as +before remarked, elevate their wings when at rest, and whilst basking in +the sunshine often alternately raise and depress them, thus exposing to +full view both surfaces; and although the lower surface is often +coloured in an obscure manner as a protection, yet in many species it is +as highly coloured as the upper surface, and sometimes in a very +different manner. In some tropical species the lower surface is even +more brilliantly coloured than the upper.[512] In one English +fritillary, the _Argynnis aglaia_, the lower surface alone is ornamented +with shining silver discs. Nevertheless, as a general rule, the upper +surface, which is probably the most fully exposed, is coloured more +brightly and in a more diversified manner than the lower. Hence the +lower surface generally affords to entomologists the most useful +character for detecting the affinities of the various species. + +Now if we turn to the enormous group of moths, which do not habitually +expose to full view the under surface of their wings, this side is very +rarely, as I hear from Mr. Stainton, coloured more brightly than the +upper side, or even with equal brightness. Some exceptions to the rule, +either real or apparent, must be noticed, as that of Hypopyra, specified +by Mr. Wormald.[513] Mr. R. Trimen informs me that in Guenée’s great +work, three moths are figured, in which the under surface is much the +most brilliant. For instance, in the Australian Gastrophora the upper +surface of the fore-wing is pale greyish-ochreous, while the lower +surface is magnificently ornamented by an ocellus of cobalt-blue, placed +in the midst of a black mark, surrounded by orange-yellow, and this by +bluish-white. But the habits of these three moths are unknown; so that +no explanation can be given of their unusual style of colouring. Mr. +Trimen also informs me that the lower surface of the wings in certain +other Geometræ[514] and quadrifid Noctuæ are either more variegated or +more brightly-coloured than the upper surface; but some of these species +have the habit of “holding their wings quite erect over their backs, +retaining them in this position for a considerable time,” and thus +exposing to view the under surface. Other species when settled on the +ground or herbage have the habit of now and then suddenly and slightly +lifting up their wings. Hence the lower surface of the wings being more +brightly-coloured than the upper surface in certain moths is not so +anomalous a circumstance as it at first appears. The Saturniidæ include +some of the most beautiful of all moths, their wings being decorated, as +in our British Emperor moth, with fine ocelli; and Mr. T. W. Wood[515] +observes that they resemble butterflies in some of their movements; “for +instance, in the gentle waving up and down of the wings, as if for +display, which is more characteristic of diurnal than of nocturnal +Lepidoptera.” + +It is a singular fact that no British moths, nor as far as I can +discover hardly any foreign species, which are brilliantly coloured, +differ much in colour according to sex; though this is the case with +many brilliant butterflies. The male, however, of one American moth, the +_Saturnia Io_, is described as having its fore-wings deep yellow, +curiously marked with purplish-red spots; whilst the wings of the female +are purple-brown, marked with grey lines.[516] The British moths which +differ sexually in colour are all brown, or various tints of dull +yellow, or nearly white. In several species the males are much darker +than the females,[517] and these belong to groups which generally fly +about during the afternoon. On the other hand, in many genera, as Mr. +Stainton informs me, the males have the hind-wings whiter than those of +the female—of which, fact _Agrotis exclamationis_ offers a good +instance. The males are thus rendered more conspicuous than the females, +whilst flying about in the dusk. In the Ghost Moth (_Hepialus humuli_) +the difference is more strongly marked; the males being white and the +females yellow with darker markings. It is difficult to conjecture what +the meaning can be of these differences between the sexes in the shades +of darkness or lightness; but we can hardly suppose that they are the +result of mere variability with sexually-limited inheritance, +independently of any benefit thus derived. + +From the foregoing statements it is impossible to admit that the +brilliant colours of butterflies and of some few moths, have commonly +been acquired for the sake of protection. We have seen that their +colours and elegant patterns are arranged and exhibited as if for +display. Hence I am led to suppose that the females generally prefer, or +are most excited by the more brilliant males; for on any other +supposition the males would be ornamented, as far as we can see, for no +purpose. We know that ants and certain lamellicorn beetles are capable +of feeling an attachment for each other, and that ants recognise their +fellows after an interval of several months. Hence there is no abstract +improbability in the Lepidoptera, which probably stand nearly or quite +as high in the scale as these insects, having sufficient mental capacity +to admire bright colours. They certainly discover flowers by colour, +and, as I have elsewhere shewn, the plants which are fertilised +exclusively by the wind never have a conspicuously-coloured corolla. The +Humming-bird Sphinx may often be seen to swoop down from a distance on a +bunch of flowers in the midst of green foliage; and I have been assured +by a friend, that these moths repeatedly visited flowers painted on the +walls of a room in the South of France. The common white butterfly, as I +hear from Mr. Doubleday, often flies down to a bit of paper on the +ground, no doubt mistaking it for one of its own species. Mr. +Collingwood[518] in speaking of the difficulty of collecting certain +butterflies in the Malay Archipelago, states that “a dead specimen +pinned upon a conspicuous twig will often arrest an insect of the same +species in its headlong flight, and bring it down within easy reach of +the net, especially if it be of the opposite sex.” + +The courtship of butterflies is a prolonged affair. The males sometimes +fight together in rivalry; and many may be seen pursuing or crowding +round the same female. If, then, the females do not prefer one male to +another, the pairing must be left to mere chance, and this does not +appear to me a probable event. If, on the other hand, the females +habitually, or even occasionally, prefer the more beautiful males, the +colours of the latter will have been rendered brighter by degrees, and +will have been transmitted to both sexes or to one sex, according to +which law of inheritance prevailed. The process of sexual selection will +have been much facilitated, if the conclusions arrived at from various +kinds of evidence in the supplement to the ninth chapter can be trusted; +namely that the males of many Lepidoptera, at least in the imago state, +greatly exceed in number the females. + +Some facts, however, are opposed to the belief that female butterflies +prefer the more beautiful males; thus, as I have been assured by several +observers, fresh females may frequently be seen paired with battered, +faded or dingy males; but this is a circumstance which could hardly +fail often to follow from the males emerging from their cocoons earlier +than the females. With moths of the family of the Bombycidæ, the sexes +pair immediately after assuming the imago state; for they cannot feed, +owing to the rudimentary condition of their mouths. The females, as +several entomologists have remarked to me, lie in an almost torpid +state, and appear not to evince the least choice in regard to their +partners, This is the case with the common silk-moth (_B. mori_), as I +have been told by some continental and English breeders. Dr. Wallace, +who has had such immense experience in breeding _Bombyx cynthia_, is +convinced that the females evince no choice or preference. He has kept +above 300 of these moths living together, and has often found the most +vigorous females mated with stunted males. The reverse apparently seldom +occurs; for, as he believes, the more vigorous males pass over the +weakly females, being attracted by those endowed with most vitality. +Although we have been indirectly induced to believe that the females of +many species prefer the more beautiful males, I have no reason to +suspect, either with moths or butterflies, that the males are attracted +by the beauty of the females. If the more beautiful females had been +continually preferred, it is almost certain, from the colours of +butterflies being so frequently transmitted to one sex alone, that the +females would often have been rendered more beautiful than their male +partners. But this does not occur except in a few instances; and these +can be explained, as we shall presently see, on the principle of +mimickry and protection. + + +As sexual selection primarily depends on variability, a few words must +be added on this subject. In respect to colour there is no difficulty, +as any number of highly variable Lepidoptera could be named. One good +instance will suffice. Mr. Bates shewed me a whole series of specimens +of _Papilio sesostris_ and _childrenæ_; in the latter the males varied +much in the extent of the beautifully enamelled green patch on the +fore-wings, and in the size of the white mark, as well as of the +splendid crimson stripe on the hind-wings; so that there was a great +contrast between the most and least gaudy males. The male of _Papilio +sesostris_, though a beautiful insect, is much less so than _P. +childrenæ_. It likewise varies a little in the size of the green patch +on the fore-wings, and in the occasional appearance of a small crimson +stripe on the hind-wings, borrowed, as it would seem, from its own +female; for the females of this and of many other species in the Æneas +group possess this crimson stripe. Hence between the brightest specimens +of _P. sesostris_ and the least bright of _P. childrenæ_, there was but +a small interval; and it was evident that as far as mere variability is +concerned, there would be no difficulty in permanently increasing by +means of selection the beauty of either species. The variability is here +almost confined to the male sex; but Mr. Wallace and Mr. Bates have +shewn[519] that the females of some other species are extremely +variable, the males being nearly constant. As I have before mentioned +the Ghost Moth (_Hepialus humuli_) as one of the best instances in +Britain of a difference in colour between the sexes of moths, it may be +worth adding[520] that in the Shetland Islands, males are frequently +found which closely resemble the females. In a future chapter I shall +have occasion to shew that the beautiful eye-like spots or ocelli, so +common on the wings of many Lepidoptera, are eminently variable. + + +On the whole, although many serious objections may be urged, it seems +probable that most of the species of Lepidoptera which are brilliantly +coloured, owe their colours to sexual selection, excepting in certain +cases, presently to be mentioned, in which conspicuous colours are +beneficial as a protection. From the ardour of the male throughout the +animal kingdom, he is generally willing to accept any female; and it is +the female which usually exerts a choice. Hence if sexual selection has +here acted, the male, when the sexes differ, ought to be the most +brilliantly coloured; and this undoubtedly is the ordinary rule. When +the sexes are brilliantly coloured and resemble each other, the +characters acquired by the males appear to have been transmitted to both +sexes. But will this explanation of the similarity and dissimilarity in +colour between the sexes suffice? + +The males and females of the same species of butterfly are known[521] in +several cases to inhabit different stations, the former commonly basking +in the sunshine, the latter haunting gloomy forests. It is therefore +possible that different conditions of life may have acted directly on +the two sexes; but this is not probable,[522] as in the adult state they +are exposed during a very short period to different conditions; and the +larvæ of both are exposed to the same conditions. Mr. Wallace believes +that the less brilliant colours of the female have been specially +gained in all or almost all cases for the sake of protection. On the +contrary it seems to me more probable that the males alone, in the large +majority of cases, have acquired their bright colours through sexual +selection, the females having been but little modified. Consequently the +females of distinct but allied species ought to resemble each other much +more closely than do the males of the same species; and this is the +general rule. The females thus approximately show us the primordial +colouring of the parent-species of the group to which they belong. They +have, however, almost always been modified to a certain extent by some +of the successive steps of variation, through the accumulation of which +the males were rendered beautiful, having been transferred to them. The +males and females of allied though distinct species will also generally +have been exposed during their prolonged larval state to different +conditions, and may have been thus indirectly affected; though with the +males any slight change of colour thus caused will often have been +completely masked by the brilliant tints gained through sexual +selection. When we treat of Birds, I shall have to discuss the whole +question whether the differences in colour between the males and females +have been in part specially gained by the latter as a protection; so +that I will here only give unavoidable details. + +In all cases when the more common form of equal inheritance by both +sexes has prevailed, the selection of bright-coloured males would tend +to make the females bright-coloured; and the selection of dull-coloured +females would tend to make the males dull. If both processes were +carried on simultaneously, they would tend to neutralise each other. As +far as I can see, it would be extremely difficult to change through +selection the one form of inheritance into the other. But by the +selection of successive variations, which were from the first sexually +limited in their transmission, there would not be the slightest +difficulty in giving bright colours to the males alone, and at the same +time or subsequently, dull colours to the females alone. In this latter +manner female butterflies and moths may, as I fully admit, have been +rendered inconspicuous for the sake of protection, and widely different +from their males. + +Mr. Wallace[523] has argued with much force in favour of his view that +when the sexes differ, the female has been specially modified for the +sake of protection; and that this has been effected by one form of +inheritance, namely, the transmission of characters to both sexes, +having been changed through the agency of natural selection into the +other form, namely, transmission to one sex. I was at first strongly +inclined to accept this view; but the more I have studied the various +classes throughout the animal kingdom, the less probable it has +appeared. Mr. Wallace urges that both sexes of the _Heliconidæ_, +_Danaidæ_, _Acroeidæ_ are equally brilliant because both are protected +from the attacks of birds and other enemies, by their offensive odour; +but that in other groups, which do not possess this immunity, the +females have been rendered inconspicuous, from having more need of +protection than the males. This supposed difference in the “need of +protection by the two sexes” is rather deceptive, and requires some +discussion. It is obvious that brightly-coloured individuals, whether +males or females, would equally attract, and obscurely-coloured +individuals equally escape, the attention of their enemies. But we are +concerned with the effects of the destruction or preservation of certain +individuals of either sex, on the character of the race. With insects, +after the male has fertilised the female, and after the latter has laid +her eggs, the greater or less immunity from danger of either sex could +not possibly have any effect on the offspring. Before the sexes have +performed their proper functions, if they existed in equal numbers and +if they strictly paired (all other circumstances being the same), the +preservation of the males and females would be equally important for the +existence of the species and for the character of the offspring. But +with most animals, as is known to be the case with the domestic +silk-moth, the male can fertilise two or three females; so that the +destruction of the males would not be so injurious to the species as +that of the females. On the other hand, Dr. Wallace believes that with +moths the progeny from a second or third fertilisation is apt to be +weakly, and therefore would not have so good chance of surviving. When +the males exist in much greater numbers than the females, no doubt many +males might be destroyed with impunity to the species; but I cannot see +that the results of ordinary selection for the sake of protection would +be influenced by the sexes existing in unequal numbers; for the same +proportion of the more conspicuous individuals, whether males or +females, would probably be destroyed. If indeed the males presented a +greater range of variation in colour, the result would be different; but +we need not here follow out such complex details. On the whole I cannot +perceive that an inequality in the numbers of the two sexes would +influence in any marked manner the effects of ordinary selection on the +character of the offspring. + +Female Lepidoptera require, as Mr. Wallace insists, some days to +deposit their fertilised ova and to search for a proper place; during +this period (whilst the life of the male was of no importance) the +brighter-coloured females would be exposed to danger and would be liable +to be destroyed. The duller-coloured females on the other hand would +survive, and thus would influence, it might be thought, in a marked +manner the character of the species,—either of both sexes or of one +sex, according to which form of inheritance prevailed. But it must not +be forgotten that the males emerge from the cocoon-state some days +before the females, and during this period, whilst the unborn females +were safe, the brighter-coloured males would be exposed to danger; so +that ultimately both sexes would probably be exposed during a nearly +equal length of time to danger, and the elimination of conspicuous +colours would not be much more effective in the one than the other sex. + +It is a more important consideration that female Lepidoptera, as Mr. +Wallace remarks, and as is known to every collector, are generally +slower flyers than the males. Consequently the latter, if exposed to +greater danger from being conspicuously coloured, might be able to +escape from their enemies, whilst the similarly-coloured females would +be destroyed; and thus the females would have the most influence in +modifying the colour of their progeny. + +There is one other consideration: bright colours, as far as sexual +selection is concerned, are commonly of no service to the females; so +that if the latter varied in brightness, and the variations were +sexually limited in their transmission, it would depend on mere chance +whether the females had their bright colours increased; and this would +tend throughout the Order to diminish the number of species with +brightly-coloured females in comparison with the species having +brightly-coloured males. On the other hand, as bright colours are +supposed to be highly serviceable to the males in their love-struggles, +the brighter males (as we shall see in the chapter on Birds) although +exposed to rather greater danger, would on an average procreate a +greater number of offspring than the duller males. In this case, if the +variations were limited in their transmission to the male sex, the males +alone would be rendered more brilliantly coloured; but if the variations +were not thus limited, the preservation and augmentation of such +variations would depend on whether more evil was caused to the species +by the females being rendered conspicuous, than good to the males by +certain individuals being successful over their rivals. + +As there can hardly be a doubt that both sexes of many butterflies and +moths have been rendered dull-coloured for the sake of protection, so +it may have been with the females alone of some species in which +successive variations towards dullness first appeared in the female sex +and were from the first limited in their transmission to the same sex. +If not thus limited, both sexes would become dull-coloured. We shall +immediately see, when we treat of mimickry, that the females alone of +certain butterflies have been rendered extremely beautiful for the sake +of protection, without any of the successive protective variations +having been transferred to the male, to whom they could not possibly +have been in the least degree injurious, and therefore could not have +been eliminated through natural selection. Whether in each particular +species, in which the sexes differ in colour, it is the female which has +been specially modified for the sake of protection; or whether it is the +male which has been specially modified for the sake of sexual +attraction, the female having retained her primordial colouring only +slightly changed through the agencies before alluded to; or whether +again both sexes have been modified, the female for protection and the +male for sexual attraction, can only be definitely decided when we know +the life-history of each species. + +Without distinct evidence, I am unwilling to admit that a double process +of selection has long been going on with a multitude of species,—the +males having been rendered more brilliant by beating their rivals; and +the females more dull-coloured by having escaped from their enemies. We +may take as an instance the common brimstone butterfly (Gonepteryx), +which appears early in the spring before any other kind. The male of +this species is of a far more intense yellow than the female, though she +is almost equally conspicuous; and in this case it does not seem +probable that she specially acquired her pale tints as a protection, +though it is probable that the male acquired his bright colours as a +sexual attraction. The female of _Anthocharis cardamines_ does not +possess the beautiful orange tips to her wings with which the male is +ornamented; consequently she closely resembles the white butterflies +(Pieris) so common in our gardens; but we have no evidence that this +resemblance is beneficial. On the contrary, as she resembles both sexes +of several species of the same genus inhabiting various quarters of the +world, it is more probable that she has simply retained to a large +extent her primordial colours. + +Various facts support the conclusion that with the greater number of +brilliantly-coloured Lepidoptera, it is the male which has been +modified; the two sexes having come to differ from each other, or to +resemble each other, according to which form of inheritance has +prevailed. Inheritance is governed by so many unknown laws or +conditions, that they seem to us to be most capricious in their +action;[524] and we can so far understand how it is that with +closely-allied species the sexes of some differ to an astonishing +degree, whilst the sexes of others are identical in colour. As the +successive steps in the process of variation are necessarily all +transmitted through the female, a greater or less number of such steps +might readily become developed in her; and thus we can understand the +frequent gradations from an extreme difference to no difference at all +between the sexes of the species within the same group. These cases of +gradation are much too common to favour the supposition that we here see +females actually undergoing the process of transition and losing their +brightness for the sake of protection; for we have every reason to +conclude that at any one time the greater number of species are in a +fixed condition. With respect to the differences between the females of +the species in the same genus or family, we can perceive that they +depend, at least in part, on the females partaking of the colours of +their respective males. This is well illustrated in those groups in +which the males are ornamented to an extraordinary degree; for the +females in these groups generally partake to a certain extent of the +splendour of their male partners. Lastly, we continually find, as +already remarked, that the females of almost all the species in the same +genus, or even family, resemble each other much more closely in colour +than do the males; and this indicates that the males have undergone a +greater amount of modification than the females. + + +_Mimickry._—This principle was first made clear in an admirable paper +by Mr. Bates,[525] who thus threw a flood of light on many obscure +problems. It had previously been observed that certain butterflies in S. +America belonging to quite distinct families, resembled the Heliconidæ +so closely in every stripe and shade of colour that they could not be +distinguished except by an experienced entomologist. As the Heliconidæ +are coloured in their usual manner, whilst the others depart from the +usual colouring of the groups to which they belong, it is clear that the +latter are the imitators, and the Heliconidæ the imitated. Mr. Bates +further observed that the imitating species are comparatively rare, +whilst the imitated swarm in large numbers; the two sets living mingled +together. From the fact of the Heliconidæ being conspicuous and +beautiful insects, yet so numerous in individuals and species, he +concluded that they must be protected from the attacks of birds by some +secretion or odour; and this hypothesis has now been confirmed by a +considerable body of curious evidence.[526] From these considerations +Mr. Bates inferred that the butterflies which imitate the protected +species had acquired their present marvellously deceptive appearance, +through variation and natural selection, in order to be mistaken for the +protected kinds and thus to escape being devoured. No explanation is +here attempted of the brilliant colours of the imitated, but only of the +imitating butterflies. We must account for the colours of the former in +the same general manner, as in the cases previously discussed in this +chapter. Since the publication of Mr. Bates’ paper, similar and equally +striking facts have been observed by Mr. Wallace[527] in the Malayan +region, and by Mr. Trimen in South Africa. + +As some writers[528] have felt much difficulty in understanding how the +first steps in the process of mimickry could have been effected through +natural selection, it may be well to remark that the process probably +has never commenced with forms widely dissimilar in colour. But with two +species moderately like each other, the closest resemblance if +beneficial to either form could readily be thus gained; and if the +imitated form was subsequently and gradually modified through sexual +selection or any other means, the imitating form would be led along the +same track, and thus be modified to almost any extent, so that it might +ultimately assume an appearance or colouring wholly unlike that of the +other members of the group to which it belonged. As extremely slight +variations in colour would not in many cases suffice to render a species +so like another protected species as to lead to its preservation, it +should be remembered that many species of Lepidoptera are liable to +considerable and abrupt variations in colour. A few instances have been +given in this chapter; but under this point of view Mr. Bates’ original +paper on mimickry, as well as Mr. Wallace’s papers, should be consulted. + +In the foregoing cases both sexes of the imitating species +resemble the imitated; but occasionally the female alone mocks a +brilliantly-coloured and protected species inhabiting the same district. +Consequently the female differs in colour from her own male, and, which +is a rare and anomalous circumstance, is the more brightly-coloured of +the two. In all the few species of Pieridæ, in which the female is more +conspicuously coloured than the male, she imitates, as I am informed by +Mr. Wallace, some protected species inhabiting the same region. The +female of _Diadema anomala_ is rich purple-brown with almost the whole +surface glossed with satiny blue, and she closely imitates the +_Euplœa midamus_, “one of the commonest butterflies of the East;” +whilst the male is bronzy or olive-brown, with only a slight blue gloss +on the outer parts of the wings.[529] Both sexes of this Diadema and of +_D. bolina_ follow the same habits of life, so that the differences in +colour between the sexes cannot be accounted for by exposure to +different conditions;[530] even if this explanation were admissible in +other instances.[531] + +The above cases of female butterflies which are more brightly-coloured +than the males, shew us, firstly, that variations have arisen in a state +of nature in the female sex, and have been transmitted exclusively, or +almost exclusively, to the same sex; and, secondly, that this form of +inheritance has not been determined through natural selection. For if we +assume that the females, before they became brightly coloured in +imitation of some protected kind, were exposed during each season for a +longer period to danger than the males; or if we assume that they could +not escape so swiftly from their enemies, we can understand how they +alone might originally have acquired through natural selection and +sexually-limited inheritance their present protective colours. But +except on the principle of these variations having been transmitted +exclusively to the female offspring, we cannot understand why the males +should have remained dull-coloured; for it would surely not have been in +any way injurious to each individual male to have partaken by +inheritance of the protective colours of the female, and thus to have +had a better chance of escaping destruction. In a group in which +brilliant colours are so common as with butterflies, it cannot be +supposed that the males have been kept dull-coloured through sexual +selection by the females rejecting the individuals which were rendered +as beautiful as themselves. We may, therefore, conclude that in these +cases inheritance by one sex is not due to the modification through +natural selection of a tendency to equal inheritance by both sexes. + +It may be well here to give an analogous case in another Order, of +characters acquired only by the female, though not in the least +injurious, as far as we can judge, to the male. Amongst the Phasmidæ, or +spectre-insects, Mr. Wallace states that “it is often the females alone +that so strikingly resemble leaves, while the males show only a rude +approximation.” Now, whatever may be the habits of these insects, it is +highly improbable that it could be disadvantageous to the males to +escape detection by resembling leaves.[532] Hence we may conclude that +the females alone in this latter as in the previous cases originally +varied in certain characters; these characters having been preserved and +augmented through ordinary selection for the sake of protection and from +the first transmitted to the female offspring alone. + + +_Bright Colours of Caterpillars._—Whilst reflecting on the beauty of +many butterflies, it occurred to me that some caterpillars were +splendidly coloured, and as sexual selection could not possibly have +here acted, it appeared rash to attribute the beauty of the mature +insect to this agency, unless the bright colours of their larvæ could +be in some manner explained. In the first place it may be observed that +the colours of caterpillars do not stand in any close correlation with +those of the mature insect. Secondly, their bright colours do not serve +in any ordinary manner as a protection. As an instance of this, Mr. +Bates informs me that the most conspicuous caterpillar which he ever +beheld (that of a Sphinx) lived on the large green leaves of a tree on +the open llanos of South America; it was about four inches in length, +transversely banded with black and yellow, and with its head, legs, and +tail of a bright red. Hence it caught the eye of any man who passed by +at the distance of many yards, and no doubt of every passing bird. + +I then applied to Mr. Wallace, who has an innate genius for solving +difficulties. After some consideration he replied: “Most caterpillars +require protection, as may be inferred from some kinds being furnished +with spines or irritating hairs, and from many being coloured green like +the leaves on which they feed, or curiously like the twigs of the trees +on which they live.” I may add as another instance of protection, that +there is a caterpillar of a moth, as I am informed by Mr. J. Mansel +Weale, which lives on the mimosas in South Africa, and fabricates for +itself a case, quite undistinguishable from the surrounding thorns. From +such considerations Mr. Wallace thought it probable that +conspicuously-coloured caterpillars were protected by having a nauseous +taste; but as their skin is extremely tender, and as their intestines +readily protrude from a wound, a slight peck from the beak of a bird +would be as fatal to them as if they had been devoured. Hence, as Mr. +Wallace remarks, “distastefulness alone would be insufficient to protect +a caterpillar unless some outward sign indicated to its would-be +destroyer that its prey was a disgusting morsel.” Under these +circumstances it would be highly advantageous to a caterpillar to be +instantaneously and certainly recognised as unpalatable by all birds and +other animals. Thus the most gaudy colours would be serviceable, and +might have been gained by variation and the survival of the most +easily-recognised individuals. + +This hypothesis appears at first sight very bold; but when it was +brought before the Entomological Society[533] it was supported by +various statements; and Mr. J. Jenner Weir, who keeps a large number of +birds in an aviary, has made, as he informs me, numerous trials, and +finds no exception to the rule, that all caterpillars of nocturnal and +retiring habits with smooth skins, all of a green colour, and all which +imitate twigs, are greedily devoured by his birds. The hairy and spinose +kinds are invariably rejected, as were four conspicuously-coloured +species. When the birds rejected a caterpillar, they plainly shewed, by +shaking their heads and cleansing their beaks, that they were disgusted +by the taste.[534] Three conspicuous kinds of caterpillars and moths +were also given by Mr. A. Butler to some lizards and frogs, and were +rejected; though other kinds were eagerly eaten. Thus the probable truth +of Mr. Wallace’s view is confirmed, namely, that certain caterpillars +have been made conspicuous for their own good, so as to be easily +recognised by their enemies, on nearly the same principle that certain +poisons are coloured by druggists for the good of man. This view will, +it is probable, be hereafter extended to many animals, which are +coloured in a conspicuous manner. + + +_Summary and Concluding Remarks on Insects._—Looking back to the +several Orders, we have seen that the sexes often differ in various +characters, the meaning of which is not understood. The sexes, also, +often differ in their organs of sense or locomotion, so that the males +may quickly discover or reach the females, and still oftener in the +males possessing diversified contrivances for retaining the females when +found. But we are not here much concerned with sexual differences of +these kinds. + +In almost all the Orders, the males of some species, even of weak and +delicate kinds, are known to be highly pugnacious; and some few are +furnished with special weapons for fighting with their rivals. But the +law of battle does not prevail nearly so widely with insects as with the +higher animals. Hence probably it is that the males have not often been +rendered larger and stronger than the females. On the contrary they are +usually smaller, in order that they may be developed within a shorter +time, so as to be ready in large numbers for the emergence of the +females. + +In two families of the Homoptera the males alone possess, in an +efficient state, organs which may be called vocal; and in three families +of the Orthoptera the males alone possess stridulating organs. In both +cases these organs are incessantly used during the breeding-season, not +only for calling the females, but for charming or exciting them in +rivalry with other males. No one who admits the agency of natural +selection, will dispute that these musical instruments have been +acquired through sexual selection. In four other Orders the members of +one sex, or more commonly of both sexes, are provided with organs for +producing various sounds, which apparently serve merely as call-notes. +Even when both sexes are thus provided, the individuals which were able +to make the loudest or most continuous noise would gain partners before +those which were less noisy, so that their organs have probably been +gained through sexual selection. It is instructive to reflect on the +wonderful diversity of the means for producing sound, possessed by the +males alone or by both sexes in no less than six Orders, and which were +possessed by at least one insect at an extremely remote geological +epoch. We thus learn how effectual sexual selection has been in leading +to modifications of structure, which sometimes, as with the Homoptera, +are of an important nature. + +From the reasons assigned in the last chapter, it is probable that the +great horns of the males of many lamellicorn, and some other beetles, +have been acquired as ornaments. So perhaps it may be with certain other +peculiarities confined to the male sex. From the small size of insects, +we are apt to undervalue their appearance. If we could imagine a male +Chalcosoma (fig. 15) with its polished, bronzed coat of mail, and vast +complex horns, magnified to the size of a horse or even of a dog, it +would be one of the most imposing animals in the world. + +The colouring of insects is a complex and obscure subject. When the male +differs slightly from the female, and neither are brilliantly coloured, +it is probable that the two sexes have varied in a slightly different +manner, with the variations transmitted to the same sex, without any +benefit having been thus derived or evil suffered. When the male is +brilliantly coloured and differs conspicuously from the female, as with +some dragon-flies and many butterflies, it is probable that he alone has +been modified, and that he owes his colours to sexual selection; whilst +the female has retained a primordial or very ancient type of colouring, +slightly modified by the agencies before explained, and has therefore +not been rendered obscure, at least in most cases, for the sake of +protection. But the female alone has sometimes been coloured +brilliantly so as to imitate other protected species inhabiting the same +district. When the sexes resemble each other and both are obscurely +coloured, there is no doubt that they have been in a multitude of cases +coloured for the sake of protection. So it is in some instances when +both are brightly coloured, causing them to resemble surrounding objects +such as flowers, or other protected species, or indirectly by giving +notice to their enemies that they are of an unpalatable nature. In many +other cases in which the sexes resemble each other and are brilliantly +coloured, especially when the colours are arranged for display, we may +conclude that they have been gained by the male sex as an attraction, +and have been transferred to both sexes. We are more especially led to +this conclusion whenever the same type of coloration prevails throughout +a group, and we find that the males of some species differ widely in +colour from the females, whilst both sexes of other species are quite +alike, with intermediate gradations connecting these extreme states. + +In the same manner as bright colours have often been partially +transferred from the males to the females, so it has been with the +extraordinary horns of many lamellicorn and some other beetles. So, +again, the vocal or instrumental organs proper to the males of the +Homoptera and Orthoptera have generally been transferred in a +rudimentary, or even in a nearly perfect condition to the females; yet +not sufficiently perfect to be used for producing sound. It is also an +interesting fact, as bearing on sexual selection, that the stridulating +organs of certain male Orthoptera are not fully developed until the last +moult; and that the colours of certain male dragon-flies are not fully +developed until some little time after their emergence from the pupal +state, and when they are ready to breed. + +Sexual selection implies that the more attractive individuals are +preferred by the opposite sex; and as with insects, when the sexes +differ, it is the male which, with rare exceptions, is the most +ornamented and departs most from the type to which the species +belongs;—and as it is the male which searches eagerly for the female, +we must suppose that the females habitually or occasionally prefer the +more beautiful males, and that these have thus acquired their beauty. +That in most or all the orders the females have the power of rejecting +any particular male, we may safely infer from the many singular +contrivances possessed by the males, such as great jaws, adhesive +cushions, spines, elongated legs, &c., for seizing the female; for these +contrivances shew that there is some difficulty in the act. In the case +of unions between distinct species, of which many instances have been +recorded, the female must have been a consenting party. Judging from +what we know of the perceptive powers and affections of various insects, +there is no antecedent improbability in sexual selection having come +largely into action; but we have as yet no direct evidence on this head, +and some facts are opposed to the belief. Nevertheless, when we see many +males pursuing the same female, we can hardly believe that the pairing +is left to blind chance—that the female exerts no choice, and is not +influenced by the gorgeous colours or other ornaments, with which the +male alone is decorated. + +If we admit that the females of the Homoptera and Orthoptera appreciate +the musical tones emitted by their male partners, and that the various +instruments for this purpose have been perfected through sexual +selection, there is little improbability in the females of other insects +appreciating beauty in form or colour, and consequently in such +characters having been thus gained by the males. But from the +circumstance of colour being so variable, and from its having been so +often modified for the sake of protection, it is extremely difficult to +decide in how large a proportion of cases sexual selection has come into +play. This is more especially difficult in those Orders, such as the +Orthoptera, Hymenoptera, and Coleoptera, in which the two sexes rarely +differ much in colour; for we are thus cut off from our best evidence of +some relation between the reproduction of the species and colour. With +the Coleoptera, however, as before remarked, it is in the great +lamellicorn group, placed by some authors at the head of the Order, and +in which we sometimes see a mutual attachment between the sexes, that we +find the males of some species possessing weapons for sexual strife, +others furnished with wonderful horns, many with stridulating organs, +and others ornamented with splendid metallic tints. Hence it seems +probable that all these characters have been gained through the same +means, namely sexual selection. + +When we treat of Birds, we shall see that they present in their +secondary sexual characters the closest analogy with insects. Thus, many +male birds are highly pugnacious, and some are furnished with special +weapons for fighting with their rivals. They possess organs which are +used during the breeding-season for producing vocal and instrumental +music. They are frequently ornamented with combs, horns, wattles and +plumes of the most diversified kinds, and are decorated with beautiful +colours, all evidently for the sake of display. We shall find that, as +with insects, both sexes, in certain groups, are equally beautiful, and +are equally provided with ornaments which are usually confined to the +male sex. In other groups both sexes are equally plain-coloured and +unornamented. Lastly, in some few anomalous cases, the females are more +beautiful than the males. We shall often find, in the same group of +birds, every gradation from no difference between the sexes, to an +extreme difference. In the latter case we shall see that the females, +like female insects, often possess more or less plain traces of the +characters which properly belong to the males. The analogy, indeed, in +all these respects between birds and insects, is curiously close. +Whatever explanation applies to the one class probably applies to the +other; and this explanation, as we shall hereafter attempt to shew, is +almost certainly sexual selection. + + + FOOTNOTES: + + [1] As the works of the first-named authors are so well known, + I need not give the titles; but as those of the latter are less + well known in England, I will give them:—‘Sechs Vorlesungen + über die Darwin’sche Theorie:’ zweite Auflage, 1868, von Dr. L. + Büchner; translated into French under the title ‘Conférences + sur la Théorie Darwinienne,’ 1869. ‘Der Mensch, im Lichte der + Darwin’sche Lehre,’ 1865, von Dr. F. Rolle. I will not attempt + to give references to all the authors who have taken the same + side of the question. Thus G. Canestrini has published + (‘Annuario della Soc. d. Nat.,’ Modena, 1867, p. 81) a very + curious paper on rudimentary characters, as bearing on the + origin of man. Another work has (1869) been published by Dr. + Barrago Francesco, bearing in Italian the title of “Man, made + in the image of God, was also made in the image of the ape.” + + [2] Prof. Häckel is the sole author who, since the publication + of the ‘Origin,’ has discussed, in his various works, in a very + able manner, the subject of sexual selection, and has seen its + full importance. + + [3] ‘Grosshirnwindungen des Menschen,’ 1868, s. 96. + + [4] ‘Leç. sur la Phys.’ 1866, p. 890, as quoted by M. Dally, + ‘L’Ordre des Primates et le Transformisme,’ 1868, p. 29. + + [5] ‘Naturgeschichte der Säugethiere von Paraguay,’ 1830, s. + 50. + + [6] Brehm, ‘Thierleben,’ B. i. 1864, s. 75, 86. On the Ateles, + s. 105. For other analogous statements, see s. 25, 107. + + [7] With respect to insects see Dr. Laycock ‘On a General Law + of Vital Periodicity,’ British Association, 1842. Dr. + Macculloch, ‘Silliman’s North American Journal of Science,’ + vol. xvii. p. 305, has seen a dog suffering from tertian ague. + + [8] I have given the evidence on this head in my ‘Variation of + Animals and Plants under Domestication,’ vol. ii. p. 15. + + [9] “Mares e diversis generibus Quadrumanorum sine dubio + dignoscunt feminas humanas a maribus. Primum, credo, odoratu, + postea aspectu. Mr. Youatt, qui diu in Hortis Zoologicis + (Bestiariis) medicus animalium erat, vir in rebus observandis + cautus et sagax, hoc mihi certissime probavit, et curatores + ejusdem loci et alii e ministris confirmaverunt. Sir Andrew + Smith et Brehm notabant idem in Cynocephalo. Illustrissimus + Cuvier etiam narrat multa de hac re quâ ut opinor nihil turpius + potest indicari inter omnia hominibus et Quadrumanis communia. + Narrat enim Cynocephalum quendam in furorem incidere aspectu + feminarum aliquarum, sed nequaquam accendi tanto furore ab + omnibus. Semper eligebat juniores, et dignoscebat in turba, et + advocabat voce gestuque.” + + [10] This remark is made with respect to Cynocephalus and the + anthropomorphous apes by Geoffroy Saint-Hilaire and F. Cuvier, + ‘Hist. Nat. des Mammifères,’ tom. i. 1824. + + [11] Huxley, ‘Man’s Place in Nature,’ 1863, p. 34. + + [12] ‘Man’s Place in Nature,’ 1863, p. 67. + + [13] The human embryo (upper fig.) is from Ecker, ‘Icones + Phys.,’ 1851-1859, tab. xxx. fig. 2. This embryo was ten lines + in length, so that the drawing is much magnified. The embryo of + the dog is from Bischoff, ‘Entwicklungsgeschichte des + Hunde-Eies,’ 1845, tab. xi. fig. 42 B. This drawing is five + times magnified, the embryo being 25 days old. The internal + viscera have been omitted, and the uterine appendages in both + drawings removed. I was directed to these figures by Prof. + Huxley, from whose work, ‘Man’s Place in Nature.’ the idea of + giving them was taken. Häckel has also given analogous drawings + in his ‘Schöpfungsgeschichte.’ + + [14] Prof. Wyman in ‘Proc. of American Acad. of Sciences,’ vol. + iv. 1860, p. 17. + + [15] Owen, ‘Anatomy of Vertebrates,’ vol. i. p. 533. + + [16] ‘Die Grosshirnwindungen des Menschen,’ 1868, s. 95. + + [17] ‘Anatomy of Vertebrates,’ vol. ii. p. 553. + + [18] ‘Proc. Soc. Nat. Hist.’ Boston, 1863, vol. ix. p. 185. + + [19] ‘Man’s Place in Nature,’ p. 65. + + [20] I had written a rough copy of this chapter before reading + a valuable paper, “Caratteri rudimentali in ordine all’origine + del uomo” (‘Annuario della Soc. d. Nat.,’ Modena, 1867, p. 81), + by G. Canestrini, to which paper I am considerably indebted. + Häckel has given admirable discussions on this whole subject, + under the title of Dysteleology, in his ‘Generelle Morphologie’ + and ‘Schöpfungsgeschichte.’ + + [21] Some good criticisms on this subject have been given by + Messrs. Murie and Mivart, in ‘Transact. Zoolog. Soc.’ 1869, + vol. vii. p. 92. + + [22] ‘Variation of Animals and Plants under Domestication,’ + vol. ii. pp. 317 and 397. See also ‘Origin of Species,’ 5th + edit. p. 535. + + [23] For instance M. Richard (‘Annales des Sciences Nat.’ 3rd + series, Zoolog. 1852, tom. xviii. p. 13) describes and figures + rudiments of what he calls the “muscle pédieux de la main,” + which he says is sometimes “infiniment petit.” Another muscle, + called “le tibial postérieur,” is generally quite absent in the + hand, but appears from time to time in a more or less + rudimentary condition. + + [24] Prof. W. Turner, ‘Proc. Royal Soc. Edinburgh,’ 1866-67, p. + 65. + + [25] Canestrini quotes Hyrt. (‘Annuario della Soc. dei + Naturalisti,’ Modena, 1867, p. 97) to the same effect. + + [26] ‘The Diseases of the Ear,’ by J. Toynbee, F.R.S., 1860, p. + 12. + + [27] See also some remarks, and the drawings of the ears of the + Lemuroidea, in Messrs. Murie and Mivart’s excellent paper in + ‘Transact. Zoolog. Soc.’ vol. vii. 1869, pp. 6 and 90. + + [28] Müller’s ‘Elements of Physiology,’ Eng. translat., 1842, + vol. ii. p. 1117. Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. + 260; ibid. on the Walrus, ‘Proc. Zoolog. Soc.’ November 8th, + 1854. See also R. Knox, ‘Great Artists and Anatomists,’ p. 106. + This rudiment apparently is somewhat larger in Negroes and + Australians than in Europeans, see Carl Vogt, ‘Lectures on + Man,’ Eng. translat. p. 129. + + [29] ‘The Physiology and Pathology of Mind,’ 2nd edit. 1868, p. + 134. + + [30] Eschricht, Ueber die Richtung der Haare am menschlichen + Körper ‘Müllers Archiv für Anat. und Phys.’ 1837, s. 47. I + shall often have to refer to this very curious paper. + + [31] Paget, ‘Lectures on Surgical Pathology,’ 1853, vol. i. p. + 71. + + [32] Eschricht, ibid. s. 40, 47. + + [33] Dr. Webb, ‘Teeth in Man and the Anthropoid Apes,’ as + quoted by Dr. C. Carter Blake in ‘Anthropological Review,’ + July, 1867, p. 299. + + [34] Owen, ‘Anatomy of Vertebrates,’ vol. iii. pp. 320, 321, + and 325. + + [35] ‘On the Primitive Form of the Skull,’ Eng. translat. in + ‘Anthropological Review,’ Oct. 1868, p. 426. + + [36] Owen, ‘Anatomy of Vertebrates,’ vol. iii. pp. 416, 434, + 441. + + [37] ‘Annuario della Soc. d. Nat.’ Modena, 1867, p. 94. + + [38] M. C. Martins (“De l’Unité Organique,” in ‘Revue des Deux + Mondes,’ June 15, 1862, p. 16), and Häckel (‘Generelle + Morphologie,’ B. ii. s. 278), have both remarked on the + singular fact of this rudiment sometimes causing death. + + [39] ‘The Lancet,’ Jan. 24, 1863, p. 83. Dr. Knox, ‘Great + Artists and Anatomists,’ p. 63. See also an important memoir on + this process by Dr. Grube, in the ‘Bulletin de l’Acad. Imp. de + St. Pétersbourg,’ tom. xii. 1867, p. 448. + + [40] “On the Caves of Gibraltar,” ‘Transact. Internat. Congress + of Prehist. Arch.’ Third Session, 1869, p. 54. + + [41] Quatrefages has lately collected the evidence on this + subject. ‘Revue des Cours Scientifiques,’ 1867-1868, p. 625. + + [42] Owen, ‘On the Nature of Limbs,’ 1849, p. 114. + + [43] Leuckart, in Todd’s ‘Cyclop. of Anat.’ 1849-52, vol. iv. + p. 1415. In man this organ is only from three to six lines in + length, but, like so many other rudimentary parts, it is + variable in development as well as in other characters. + + [44] See, on this subject, Owen, ‘Anatomy of Vertebrates,’ vol. + iii. pp. 675, 676, 706. + + [45] See the evidence on these points, as given by Lubbock, + ‘Prehistoric Times,’ p. 354, &c. + + [46] ‘L’Instinct chez les Insectes.’ ‘Revue des Deux Mondes,’ + Feb. 1870, p. 690. + + [47] ‘The American Beaver and his Works,’ 1868. + + [48] ‘The Principles of Psychology,’ 2nd edit. 1870, pp. + 418-443. + + [49] ‘Contributions to the Theory of Natural Selection,’ 1870, + p. 212 + + [50] ‘Recherches sur les Mœurs des Fourmis,’ 1810, p. 173. + + [51] All the following statements, given on the authority of + these two naturalists, are taken from Rengger’s ‘Naturges. der + Säugethiere von Paraguay,’ 1830, s. 41-57, and from Brehm’s + ‘Thierleben,’ B. i. s. 10-87. + + [52] ‘Bridgewater Treatise,’ p. 263. + + [53] W. C. L. Martin, ‘Nat. Hist. of Mammalia,’ 1841, p. 405. + + [54] Quoted by Vogt, ‘Mémoire sur les Microcéphales,’ 1867, p. + 168. + + [55] ‘The Variation of Animals and Plants under Domestication,’ + vol. i. p. 27. + + [56] ‘Les Mœurs des Fourmis,’ 1810, p. 150. + + [57] Quoted in Dr. Maudsley’s ‘Physiology and Pathology of + Mind,’ 1868, pp. 19, 220. + + [58] Dr. Jerdon, ‘Birds of India,’ vol. i. 1862, p. xxi. + + [59] Mr. L. H. Morgan’s work on ‘The American Beaver,’ 1868, + offers a good illustration of this remark. I cannot, however, + avoid thinking that he goes too far in underrating the power of + Instinct. + + [60] ‘The Moor and the Loch,’ p. 45. Col. Hutchinson on ‘Dog + Breaking,’ 1850, p. 46. + + [61] ‘Personal Narrative,’ Eng. translat., vol. iii. p. 106. + + [62] Quoted by Sir C. Lyell, ‘Antiquity of Man,’ p. 497. + + [63] ‘Journal of Researches during the Voyage of the “Beagle,”’ + 1845, p. 398. ‘Origin of Species,’ 5th edit. p. 260. + + [64] ‘Lettres Phil. sur l’Intelligence des Animaux,’ nouvelle + edit. 1802, p. 86. + + [65] See the evidence on this head in chap. i. vol. i. ‘On the + Variation of Animals and Plants under Domestication.’ + + [66] ‘Proc. Zoolog. Soc.’ 1864, p. 186. + + [67] Savage and Wyman in ‘Boston Journal of Nat. Hist.’ vol. + iv. 1843-44, p. 383. + + [68] ‘Säugethiere von Paraguay,’ 1830, s. 51-56. + + [69] ‘Thierleben,’ B. i. s. 79, 82. + + [70] ‘The Malay Archipelago,’ vol. i. 1869, p. 87. + + [71] ‘Primeval Man,’ 1869, pp. 145, 147. + + [72] ‘Prehistoric Times,’ 1865, p. 473, &c. + + [73] Quoted in ‘Anthropological Review,’ 1864, p. 158. + + [74] Rengger, ibid. s. 45. + + [75] See my ‘Variation of Animals and Plants under + Domestication,’ vol. i. p. 27. + + [76] See a discussion on this subject in Mr. E. B. Tylor’s very + interesting work, ‘Researches into the Early History of + Mankind,’ 1865, chaps. ii. to iv. + + [77] Hon. Daines Barrington in ‘Philosoph. Transactions,’ 1773, + p. 262. See also Dureau de la Malle, in ‘Ann. des Sc. Nat.’ 3rd + series, Zoolog. tom. x. p. 119. + + [78] ‘On the Origin of Language,’ by H. Wedgwood, 1866. + ‘Chapters on Language,’ by the Rev. F. W. Farrar, 1865. These + works are most interesting. See also ‘De la Phys. et de + Parole,’ par Albert Lemoine, 1865, p. 190. The work on this + subject, by the late Prof. Aug. Schleicher, has been translated + by Dr. Bikkers into English, under the title of ‘Darwinism + tested by the Science of Language,’ 1869. + + [79] Vogt, ‘Mémoire sur les Microcéphales,’ 1867, p. 169. With + respect to savages, I have given some facts in my ‘Journal of + Researches,’ &c., 1845, p. 206. + + [80] See clear evidence on this head in the two works so often + quoted, by Brehm and Rengger. + + [81] See remarks on this head by Dr. Maudsley, ‘The Physiology + and Pathology of Mind,’ 2nd edit. 1868, p. 199. + + [82] Many curious cases have been recorded. See, for instance, + ‘Inquiries Concerning the Intellectual Powers,’ by Dr. + Abercrombie, 1838, p. 150. + + [83] ‘The Variation of Animals and Plants under Domestication,’ + vol. ii. p. 6. + + [84] See some good remarks to this effect by Dr. Maudsley, ‘The + Physiology and Pathology of Mind,’ 1808, p. 199. + + [85] Macgillivray, ‘Hist. of British Birds,’ vol. ii. 1839, p. + 29. An excellent observer, Mr. Blackwall, remarks that the + magpie learns to pronounce single words, and even short + sentences, more readily than almost any other British bird; + yet, as he adds, after long and closely investigating its + habits, he has never known it, in a state of nature, display + any unusual capacity for imitation. ‘Researches in Zoology,’ + 1834, p. 158. + + [86] See the very interesting parallelism between the + development of species and languages, given by Sir C. Lyell in + ‘The Geolog. Evidences of the Antiquity of Man,’ 1863, chap. + xxiii. + + [87] See remarks to this effect by the Rev. F. W. Farrar, in an + interesting article, entitled “Philology and Darwinism” in + ‘Nature,’ March 24th, 1870, p. 528. + + [88] ‘Nature,’ Jan. 6th, 1870, p. 257. + + [89] Quoted by C. S. Wake, ‘Chapters on Man,’ 1868, p. 101. + + [90] Buckland, ‘Bridgewater Treatise,’ p. 411. + + [91] See some good remarks on the simplification of languages, + by Sir J. Lubbock, ‘Origin of Civilisation,’ 1870, p. 278. + + [92] ‘Conférences sur la Théorie Darwinienne,’ French + translat., 1869, p. 132. + + [93] The Rev. Dr. J. M’Cann, ‘Anti-Darwinism,’ 1869, p. 13. + + [94] ‘The Spectator,’ Dec. 4th, 1869, p. 1430. + + [95] See an excellent article on this subject by the Rev. F. W. + Farrar, in the ‘Anthropological Review,’ Aug. 1864, p. ccxvii. + For further facts see Sir J. Lubbock, ‘Prehistoric Times,’ 2nd + edit. 1869. p. 564; and especially the chapters on Religion in + his ‘Origin of Civilisation,’ 1870. + + [96] The Worship of Animals and Plants, in the ‘Fortnightly + Review,’ Oct. 1, 1869, p. 422. + + [97] Tylor, ‘Early History of Mankind,’ 1865, p. 6. See also + the three striking chapters on the Development of Religion, in + Lubbock’s ‘Origin of Civilisation,’ 1870. In a like manner Mr. + Herbert Spencer, in his ingenious essay in the ‘Fortnightly + Review’ (May 1st, 1870, p. 535), accounts for the earliest + forms of religious belief throughout the world, by man being + led through dreams, shadows, and other causes, to look at + himself as a double essence, corporeal and spiritual. As the + spiritual being is supposed to exist after death and to be + powerful, it is propitiated by various gifts and ceremonies, + and its aid invoked. He then further shews that names or + nicknames given from some animal or other object to the early + progenitors or founders of a tribe, are supposed after a long + interval to represent the real progenitor of the tribe; and + such animal or object is then naturally believed still to exist + as a spirit, is held sacred, and worshipped as a god. + Nevertheless I cannot but suspect that there is a still earlier + and ruder stage, when anything which manifests power or + movement is thought to be endowed with some form of life, and + with mental faculties analogous to our own. + + [98] See an able article on the Psychical Elements of Religion, + by Mr. L. Owen Pike, in ‘Anthropolog. Review,’ April, 1870, p. + lxiii. + + [99] ‘Religion, Moral, &c., der Darwin’schen Art-Lehre,’ 1869, + s. 53. + + [100] ‘Prehistoric Times,’ 2nd edit. p. 571. In this work (at + p. 553) there will be found an excellent account of the many + strange and capricious customs of savages. + + [101] See, for instance, on this subject, Quatrefages, ‘Unité + de l’Espèce Humaine,’ 1861, p. 21, &c. + + [102] ‘Dissertation on Ethical Philosophy,’ 1837, p. 231, &c. + + [103] ‘Metaphysics of Ethics,’ translated by J. W. Semple, + Edinburgh, 1836, p. 136. + + [104] Mr. Bain gives a list (‘Mental and Moral Science,’ 1868, + p. 543-725) of twenty-six British authors who have written on + this subject, and whose names are familiar to every reader; to + these, Mr. Bain’s own name, and those of Mr. Lecky, Mr. + Shadworth Hodgson, and Sir J. Lubbock, as well as of others, + may be added. + + [105] Sir B. Brodie, after observing that man is a social + animal (‘Psychological Enquiries,’ 1854, p. 192), asks the + pregnant question, “ought not this to settle the disputed + question as to the existence of a moral sense?” Similar ideas + have probably occurred to many persons, as they did long ago to + Marcus Aurelius. Mr. J. S. Mill speaks, in his celebrated work, + ‘Utilitarianism,’ (1864, p. 46), of the social feelings as a + “powerful natural sentiment,” and as “the natural basis of + sentiment for utilitarian morality;” but on the previous page + he says, “if, as is my own belief, the moral feelings are not + innate, but acquired, they are not for that reason less + natural.” It is with hesitation that I venture to differ from + so profound a thinker, but it can hardly be disputed that the + social feelings are instinctive or innate in the lower animals; + and why should they not be so in man? Mr. Bain (see, for + instance, ‘The Emotions and the Will,’ 1865, p. 481) and others + believe that the moral sense is acquired by each individual + during his lifetime. On the general theory of evolution this is + at least extremely improbable. + + [106] ‘Die Darwin’sche Theorie,’ s. 101. + + [107] Mr. R. Brown in ‘Proc. Zoolog. Soc.’ 1868, p. 409. + + [108] Brehm, ‘Thierleben,’ B. i. 1864, s. 52, 79. For the case + of the monkeys extracting thorns from each other, see s. 54. + With respect to the Hamadryas turning over stones, the fact is + given (s. 76) on the evidence of Alvarez, whose observations + Brehm thinks quite trustworthy. For the cases of the old male + baboons attacking the dogs, see s. 79; and with respect to the + eagle, s. 56. + + [109] ‘Annals and Mag. of Nat. Hist.’ November, 1868, p. 382. + + [110] Sir J. Lubbock, ‘Prehistoric Times,’ 2nd edit. p. 446. + + [111] As quoted by Mr. L. H. Morgan, ‘The American Beaver,’ + 1868, p. 272. Capt. Stansbury also gives an interesting account + of the manner in which a very young pelican, carried away by a + strong stream, was guided and encouraged in its attempts to + reach the shore by half a dozen old birds. + + [112] As Mr. Bain states, “effective aid to a sufferer springs + from sympathy proper:” ‘Mental and Moral Science,’ 1868, p. + 245. + + [113] ‘Thierleben,’ B. i. s. 85. + + [114] ‘De l’Espèce et de la Class.’ 1869, p. 97. + + [115] ‘Der Darwin’schen Art-Lehre,’ 1869, s. 54. + + [116] Brehm, ‘Thierleben,’ B. i. s. 76. + + [117] See the first and striking chapter in Adam Smith’s + ‘Theory of Moral Sentiments.’ Also Mr. Bain’s ‘Mental and Moral + Science,’ 1868, p. 244, and 275-282. Mr. Bain states, that + “sympathy is, indirectly, a source of pleasure to the + sympathiser;” and he accounts for this through reciprocity. He + remarks that “the person benefited, or others in his stead, may + make up, by sympathy and good offices returned, for all the + sacrifice.” But if, as appears to be the case, sympathy is + strictly an instinct, its exercise would give direct pleasure, + in the same manner as the exercise, as before remarked, of + almost every other instinct. + + [118] This fact, the Rev. L. Jenyns states (see his edition of + ‘White’s Nat. Hist. of Selborne,’ 1853, p. 204) was first + recorded by the illustrious Jenner, in ‘Phil. Transact.’ 1824, + and has since been confirmed by several observers, especially + by Mr. Blackwall. This latter careful observer examined, late + in the autumn, during two years, thirty-six nests; he found + that twelve contained young dead birds, five contained eggs on + the point of being hatched, and three eggs not nearly hatched. + Many birds not yet old enough for a prolonged flight are + likewise deserted and left behind. See Blackwall, ‘Researches + in Zoology,’ 1834, pp. 108, 118. For some additional evidence, + although this is not wanted, see Leroy, ‘Lettres Phil.’ 1802, + p. 217. + + [119] Hume remarks (‘An Enquiry Concerning the Principles of + Morals,’ edit. of 1751, p. 132), “there seems a necessity for + confessing that the happiness and misery of others are not + spectacles altogether indifferent to us, but that the view of + the former ... communicates a secret joy; the appearance of the + latter ... throws a melancholy damp over the imagination.” + + [120] ‘Mental and Moral Science,’ 1868, p. 254. + + [121] I have given one such case, namely of three Patagonian + Indians who preferred being shot, one after the other, to + betraying the plans of their companions in war (‘Journal of + Researches,’ 1845, p. 103). + + [122] Dr. Prosper Despine, in his ‘Psychologie Naturelle,’ 1868 + (tom. i. p. 243; tom ii. p. 169) gives many curious cases of + the worst criminals, who apparently have been entirely + destitute of conscience. + + [123] See an able article in the ‘North British Review,’ 1867, + p. 395. See also Mr. W. Bagehot’s articles on the Importance of + Obedience and Coherence to Primitive Man, in the ‘Fortnightly + Review,’ 1867, p. 529, and 1868, p. 457, &c. + + [124] The fullest account which I have met with is by Dr. + Gerland, in his ‘Ueber das Aussterben der Naturvölker,’ 1868; + but I shall have to recur to the subject of infanticide in a + future chapter. + + [125] See the very interesting discussion on Suicide in Lecky’s + ‘History of European Morals,’ vol. i. 1869, p. 223. + + [126] See, for instance, Mr. Hamilton’s account of the Kaffirs, + ‘Anthropological Review,’ 1870, p. xv. + + [127] Mr. M’Lennan has given ‘Primitive Marriage,’ 1865, p. + 176, a good collection of facts on this head. + + [128] Lecky, ‘History of European Morals,’ vol. i. 1869, p. + 109. + + [129] ‘Embassy to China,’ vol. ii. p. 348. + + [130] See on this subject copious evidence in Chap. vii. of Sir + J. Lubbock, ‘Origin of Civilisation,’ 1870. + + [131] For instance Lecky, ‘Hist. European Morals,’ vol. i. p. + 124. + + [132] This term is used in an able article in the ‘Westminster + Review,’ Oct. 1869, p. 498. For the Greatest Happiness + principle, see J. S. Mill, ‘Utilitarianism,’ p. 17. + + [133] Good instances are given by Mr. Wallace in ‘Scientific + Opinion,’ Sept. 15, 1869; and more fully in his ‘Contributions + to the Theory of Natural Selection,’ 1870, p. 353. + + [134] Tennyson, ‘Idylls of the King,’ p. 244. + + [135] ‘The Thoughts of the Emperor M. Aurelius Antoninus,’ Eng. + translat., 2nd edit., 1869, p. 112. Marcus Aurelius was born + A.D. 121. + + [136] Letter to Mr. Mill in Bain’s ‘Mental and Moral Science,’ + 1868, p. 722. + + [137] A writer in the ‘North British Review’ (July, 1869, p. + 531), well capable of forming a sound judgment, expresses + himself strongly to this effect. Mr. Lecky (‘Hist. of Morals,’ + vol. i. p. 143) seems to a certain extent to coincide. + + [138] See his remarkable work on ‘Hereditary Genius,’ 1869, p. + 349. The Duke of Argyll (‘Primeval Man,’ 1869, p. 188) has some + good remarks on the contest in man’s nature between right and + wrong. + + [139] ‘The Thoughts of Marcus Aurelius,’ &c., p. 139. + + [140] ‘Investigations in Military and Anthropolog. Statistics + of American Soldiers,’ by B. A. Gould, 1869, p. 256. + + [141] With respect to the “Cranial forms of the American + aborigines,” see Dr. Aitken Meigs in ‘Proc. Acad. Nat. Sci.’ + Philadelphia, May, 1866. On the Australians, see Huxley, in + Lyell’s ‘Antiquity of Man,’ 1863, p. 87. On the Sandwich + Islanders, Prof. J. Wyman, ‘Observations on Crania,’ Boston, + 1868, p. 18. + + [142] ‘Anatomy of the Arteries,’ by R. Quain. + + [143] ‘Transact. Royal Soc.’ Edinburgh, vol. xxiv. p. 175, 189. + + [144] ‘Proc. Royal Soc.’ 1867, p. 544; also 1868, p. 483, 524. + There is a previous paper, 1866, p. 229. + + [145] ‘Proc. R. Irish Academy,’ vol. x. 1868, p. 141. + + [146] ‘Act. Acad.,’ St. Petersburg, 1778, part ii. p. 217. + + [147] Brehm, ‘Thierleben,’ B. i. s. 58, 87. Rengger, + ‘Säugethiere von Paraguay,’ s. 57. + + [148] ‘Variation of Animals and Plants under Domestication,’ + vol. ii. chap. xii. + + [149] ‘Hereditary Genius: an Inquiry into its Laws and + Consequences,’ 1869. + + [150] Mr. Bates remarks (‘The Naturalist on the Amazons,’ 1863, + vol. ii. p. 159), with respect to the Indians of the same S. + American tribe, “no two of them were at all similar in the + shape of the head; one man had an oval visage with fine + features, and another was quite Mongolian in breadth and + prominence of cheek, spread of nostrils, and obliquity of + eyes.” + + [151] Blumenbach, ‘Treatises on Anthropolog.’ Eng. translat., + 1865, p. 205. + + [152] Godron, ‘De l’Espèce,’ 1859, tom. ii. livre 3. + Quatrefages, ‘Unité de l’Espèce Humaine,’ 1861. Also Lectures + on Anthropology, given in the ‘Revue des Cours Scientifiques,’ + 1866-1868. + + [153] ‘Hist. Gen. et Part. des Anomalies de l’Organisation,’ in + three volumes, tom. i. 1832. + + [154] I have fully discussed these laws in my ‘Variation of + Animals and Plants under Domestication,’ vol. ii. chap. xxii. + and xxiii. M. J. P. Durand has lately 1868; published a + valuable essay ‘De l’Influence des Milieux, &c.’ He lays much + stress on the nature of the soil. + + [155] ‘Investigations in Military and Anthrop. Statistics,’ &c. + 1869, by B. A. Gould, p. 93, 107, 126, 131, 134. + + [156] For the Polynesians, see Prichard’s ‘Physical Hist. of + Mankind,’ vol. v. 1847, p. 145, 283. Also Godron, ‘De + l’Espèce,’ tom. ii. p. 289. There is also a remarkable + difference in appearance between the closely-allied Hindoos + inhabiting the Upper Ganges and Bengal; see Elphinstone’s + ‘History of India,’ vol. i. p. 324. + + [157] ‘Memoirs, Anthropolog. Soc.’ vol. iii. 1867-69, p. 561, + 565, 567. + + [158] Dr. Brakenridge, ‘Theory of Diathesis,’ ‘Medical Times,’ + June 19 and July 17, 1869. + + [159] I have given authorities for these several statements in + my ‘Variation of Animals under Domestication,’ vol. ii. p. + 297-300. Dr. Jaeger, “Ueber das Längenwachsthum der Knochen,” + ‘Jenaischen Zeitschrift,’ B. v. Heft i. + + [160] ‘Investigations,’ &c. By B. A. Gould, 1869, p. 288. + + [161] ‘Säugethiere von Paraguay,’ 1830, s. 4. + + [162] ‘History of Greenland,’ Eng. translat. 1767, vol. i. p. + 230. + + [163] ‘Intermarriage.’ By Alex. Walker, 1838. p. 377. + + [164] ‘The Variation of Animals under Domestication,’ vol. i. + p. 173. + + [165] ‘Principles of Biology,’ vol. i. p. 455. + + [166] Paget, ‘Lectures on Surgical Pathology,’ vol. i. 1853, p. + 209. + + [167] ‘The Variation of Animals under Domestication,’ vol. ii. + p. 8. + + [168] ‘Säugethiere von Paraguay,’ s. 8, 10. I have had good + opportunities for observing the extraordinary power of eyesight + in the Fuegians.’ See also Lawrence (‘Lectures on Physiology,’ + &c., 1822, p. 404) on this same subject. M. Giraud-Teulon has + recently collected (‘Revue des Cours Scientifiques,’ 1870, p. + 625) a large and valuable body of evidence proving that the + cause of short-sight, “_C’est le travail assidu, de près._” + + [169] Prichard, ‘Phys. Hist. of Mankind,’ on the authority of + Blumenbach, vol. i. 1851, p. 311; for the statement by Pallas, + vol. iv. 1844, p. 407. + + [170] Quoted by Prichard, ‘Researches into the Phys. Hist. of + Mankind,’ vol. v. p. 463. + + [171] Mr. Forbes’ valuable paper is now published in the + ‘Journal of the Ethnological Soc. of London,’ new series, vol. + ii. 1870, p. 193. + + [172] Dr. Wilckens (‘Landwirthschaft. Wochenblatt,’ No. 10, + 1869) has lately published an interesting essay shewing how + domestic animals, which live in mountainous regions, have their + frames modified. + + [173] ‘Mémoire sur les Microcéphales,’ 1867, p. 50, 125, 169, + 171, 184-198. + + [174] See Dr. A. Farre’s well-known article in the ‘Cyclop. of + Anat. and Phys.’ vol. v. 1859, p. 642. Owen ‘Anatomy of + Vertebrates,’ vol. iii. 1868, p. 687. Prof. Turner in + ‘Edinburgh Medical Journal,’ Feb. 1865. + + [175] ‘Annuario della Soc. dei Naturalisti in Modena,’ 1867, p. + 83. Prof. Canestrini gives extracts on this subject from + various authorities. Laurillard remarks, that as he has found a + complete similarity in the form, proportions, and connexion of + the two malar bones in several human subjects and in certain + apes, he cannot consider this disposition of the parts as + simply accidental. + + [176] A whole series of cases is given by Isid. Geoffroy + St.-Hilaire, ‘Hist. des Anomalies,’ tom. iii. p. 437. + + [177] In my ‘Variation of Animals under Domestication’ (vol. + ii. p. 57) I attributed the not very rare cases of + supernumerary mammæ in women to reversion. I was led to this as + a _probable_ conclusion, by the additional mammæ being + generally placed symmetrically on the breast, and more + especially from one case, in which a single efficient mamma + occurred in the inguinal region of a woman, the daughter of + another woman with supernumerary mammæ. But Prof. Preyer (‘Der + Kampf um das Dasein,’ 1869, s. 45) states that _mammæ erraticæ_ + have been known to occur in other situations, even on the back; + so that the force of my argument is greatly weakened or perhaps + quite destroyed. + + With much hesitation I, in the same work (vol. ii. p. 12), + attributed the frequent cases of polydactylism in men to + reversion. I was partly led to this through Prof. Owen’s + statement, that some of the Ichthyopterygia possess more than + five digits, and therefore, as I supposed, had retained a + primordial condition; but after reading Prof. Gegenbaur’s paper + (‘Jenaischen Zeitschrift,’ B. v. Heft 3, s. 341), who is the + highest authority in Europe on such a point, and who disputes + Owen’s conclusion, I see that it is extremely doubtful whether + supernumerary digits can thus be accounted for. It was the fact + that such digits not only frequently occur and are strongly + inherited, but have the power of regrowth after amputation, + like the normal digits of the lower vertebrata, that chiefly + led me to the above conclusion. This extraordinary fact of + their regrowth remains inexplicable, if the belief in reversion + to some extremely remote progenitor must be rejected. I cannot, + however, follow Prof. Gegenbaur in supposing that additional + digits could not reappear through reversion, without at the + same time other parts of the skeleton being simultaneously and + similarly modified; for single characters often reappear + through reversion. + + [178] ‘Anatomy of Vertebrates,’ vol. iii. 1868, p. 323. + + [179] ‘Generelle Morphologie,’ 1866, B. ii. s. clv. + + [180] Carl Vogt’s ‘Lectures on Man,’ Eng. translat. 1864, p. + 151. + + [181] C. Carter Blake, on a jaw from La Naulette, ‘Anthropolog. + Review,’ 1867, p. 295. Schaaffhausen, ibid. 1868, p. 426. + + [182] ‘The Anatomy of Expression,’ 1844, p. 110, 131. + + [183] Quoted by Prof. Canestrini in the ‘Annuario,’ &c., 1867, + p. 90. + + [184] These papers deserve careful study by any one who desires + to learn how frequently our muscles vary, and in varying come + to resemble those of the Quadrumana. The following references + relate to the few points touched on in my text: Proc. Royal + Soc. vol. xiv. 1865, p. 379-384; vol. xv. 1866, p. 241, 242; + vol. xv. 1867, p. 544; vol. xvi. 1868, p. 524. I may here add + that Dr. Murie and Mr. St. George Mivart have shewn in their + Memoir on the Lemuroidea (‘Transact. Zoolog. Soc.’ vol. vii. + 1869, p. 96), how extraordinarily variable some of the muscles + are in these animals, the lowest members of the Primates. + Gradations, also, in the muscles leading to structures found in + animals still lower in the scale, are numerous in the + Lemuroidea. + + [185] Prof. Macalister in ‘Proc. R. Irish Academy,’ vol. x. + 1868, p. 124. + + [186] Prof. Macalister (ibid. p. 121) has tabulated his + observations, and finds that muscular abnormalities are most + frequent in the fore-arms, secondly in the face, thirdly in the + foot, &c. + + [187] The Rev. Dr. Haughton, after giving (‘Proc. R. Irish + Academy,’ June 27, 1864, p. 715) a remarkable case of variation + in the human _flexor pollicis longus_, adds, “This remarkable + example shews that man may sometimes possess the arrangement of + tendons of thumb and fingers characteristic of the macaque; but + whether such a case should be regarded as a macaque passing + upwards into a man, or a man passing downwards into a macaque, + or as a congenital freak of nature, I cannot undertake to say.” + It is satisfactory to hear so capable an anatomist, and so + embittered an opponent of evolutionism, admitting even the + possibility of either of his first propositions. Prof. + Macalister has also described (‘Proc. R. Irish Acad.’ vol. x. + 1864, p. 138) variations in the _flexor pollicis longus_, + remarkable from their relations to the same muscle in the + Quadrumana. + + [188] The authorities for these several statements are given in + my ‘Variation of Animals under Domestication,’ vol. ii. p. + 320-335. + + [189] This whole subject has been discussed in chap. xxiii. + vol. ii. of my ‘Variation of Animals and Plants under + Domestication.’ + + [190] See the ever memorable ‘Essay on the Principle of + Population,’ by the Rev. T. Malthus, vol. i. 1826, p. 6, 517. + + [191] ‘Variation of Animals and Plants under Domestication,’ + vol. ii. p. 111-113, 163. + + [192] Mr. Sedgwick, ‘British and Foreign Medico-Chirurg. + Review,’ July, 1863, p. 170. + + [193] ‘The Annals of Rural Bengal,’ by W. W. Hunter, 1868, p. + 259. + + [194] ‘Primitive Marriage,’ 1865. + + [195] See some good remarks to this effect by W. Stanley + Jevons, “A Deduction from Darwin’s Theory,” ‘Nature,’ 1869, p. + 231. + + [196] Latham, ‘Man and his Migrations,’ 1851, p. 135. + + [197] Messrs. Murie and Mivart in their “Anatomy of the + Lemuroidea” (‘Transact. Zoolog. Soc.’ vol. vii. 1869, p. 96-98) + say, “some muscles are so irregular in their distribution that + they cannot be well classed in any of the above groups.” These + muscles differ even on the opposite sides of the same + individual. + + [198] ‘Quarterly Review,’ April, 1869, p. 392. This subject is + more fully discussed in Mr. Wallace’s ‘Contributions to the + Theory of Natural Selection,’ 1870, in which all the essays + referred to in this work are republished. The ‘Essay on Man’ + has been ably criticised by Prof. Claparède, one of the most + distinguished zoologists in Europe, in an article published in + the ‘Bibliothèque Universelle,’ June, 1870. The remark quoted + in my text will surprise every one who has read Mr. Wallace’s + celebrated paper on ‘The Origin of Human Races deduced from the + Theory of Natural Selection,’ originally published in the + ‘Anthropological Review,’ May, 1864, p. clviii. I cannot here + resist quoting a most just remark by Sir J. Lubbock + (‘Prehistoric Times,’ 1865, p. 479) in reference to this paper, + namely, that Mr. Wallace, “with characteristic unselfishness, + ascribes it (_i.e._ the idea of natural selection) unreservedly + to Mr. Darwin, although, as is well known, he struck out the + idea independently, and published it, though not with the same + elaboration, at the same time.” + + [199] Quoted by Mr. Lawson Tait in his “Law of Natural + Selection,”—‘Dublin Quarterly Journal of Medical Science,’ + Feb. 1869. Dr. Keller is likewise quoted to the same effect. + + [200] Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 71. + + [201] ‘Quarterly Review,’ April, 1869, p. 392. + + [202] In _Hylobates syndactylus_, as the name expresses, two of + the digits regularly cohere; and this, as Mr. Blyth informs me, + is occasionally the case with the digits of _H. agilis_, _lar_, + and _leuciscus_. + + [203] Brehm, ‘Thierleben,’ B. i. s. 80. + + [204] “The Hand, its mechanism,” &c. ‘Bridgewater Treatise,’ + 1833, p. 38. + + [205] Häckel has an excellent discussion on the steps by which + man became a biped: ‘Natürliche Schöpfungsgeschichte,’ 1868, s. + 507. Dr. Büchner (‘Conférences sur la Théorie Darwinienne,’ + 1869, p. 135) has given good cases of the use of the foot as a + prehensile organ by man; also on the manner of progression of + the higher apes to which I allude in the following paragraph: + see also Owen (‘Anatomy of Vertebrates,’ vol. iii. p. 71) on + this latter subject. + + [206] “On the Primitive Form of the Skull,” translated in + ‘Anthropological Review,’ Oct. 1868, p. 428. Owen (‘Anatomy of + Vertebrates,’ vol. ii. 1866, p. 551) on the mastoid processes + in the higher apes. + + [207] ‘Die Grenzen der Thierwelt, eine Betrachtung zu Darwin’s + Lehre,’ 1868, s. 51. + + [208] Dujardin, ‘Annales des Sc. Nat.’ 3rd series, Zoolog. tom. + xiv. 1850, p. 203. See also Mr. Lowne, ‘Anatomy and Phys. of + the _Musca vomitoria_,’ 1870, p. 14. My son, Mr. F. Darwin, + dissected for me the cerebral ganglia of the _Formica rufa_. + + [209] ‘Philosophical Transactions,’ 1869, p. 513. + + [210] Quoted in C. Vogt’s ‘Lectures on Man,’ Eng. translat. + 1864, p. 88, 90. Prichard, ‘Phys. Hist. of Mankind,’ vol. i. + 1838, p. 305. + + [211] ‘Comptes Rendus des Séances,’ &c. June 1, 1868. + + [212] ‘The Variation of Animals and Plants under + Domestication,’ vol. ii. p. 124-129. + + [213] Schaaffhausen gives from Blumenbach and Busch, the cases + of the spasms and cicatrix, in ‘Anthropolog. Review,’ Oct. + 1868, p. 420. Dr. Jarrold (‘Anthropologia,’ 1808, p. 115, 116) + adduces from Camper and from his own observations, cases of the + modification of the skull from the head being fixed in an + unnatural position. He believes that certain trades, such as + that of a shoemaker, by causing the head to be habitually held + forward, makes the forehead more rounded and prominent. + + [214] ‘Variation of Animals,’ &c., vol. i. p. 117 on the + elongation of the skull; p. 119, on the effect of the lopping + of one ear. + + [215] Quoted by Schaaffhausen, in ‘Anthropolog. Review,’ Oct. + 1868, p. 419. + + [216] Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 619. + + [217] Isidore Geoffroy St.-Hilaire remarks (‘Hist. Nat. + Générale,’ tom. ii. 1859, p. 215-217) on the head of man being + covered with long hair; also on the upper surfaces of monkeys + and of other mammals being more thickly clothed than the lower + surfaces. This has likewise been observed by various authors. + Prof. P. Gervais (‘Hist. Nat. des Mammifères,’ tom. i. 1854, p. + 28), however, states that in the Gorilla the hair is thinner on + the back, where it is partly rubbed off, than on the lower + surface. + + [218] Mr. St. George Mivart, ‘Proc. Zoolog. Soc.’ 1865, p. 562, + 583. Dr. J. E. Gray, ‘Cat. Brit. Mus.: Skeletons.’ Owen, + ‘Anatomy of Vertebrates,’ vol. ii. p. 517. Isidore Geoffroy, + ‘Hist. Nat. Gén.’ tom. ii. p. 244. + + [219] ‘The Variation of Animals and Plants under + Domestication,’ vol. ii. p. 280, 282. + + [220] ‘Primeval Man,’ 1869, p. 66. + + [221] ‘Anthropological Review,’ May, 1864, p. clviii. + + [222] After a time the members or tribes which are absorbed + into another tribe assume, as Mr. Maine remarks (‘Ancient Law,’ + 1861, p. 131), that they are the co-descendants of the same + ancestors. + + [223] Morlot, ‘Soc. Vaud. Sc. Nat.’ 1860, p. 294. + + [224] I have given instances in my ‘Variation of Animals under + Domestication,’ vol. ii. p. 196. + + [225] See a remarkable series of articles on Physics and + Politics in the ‘Fortnightly Review,’ Nov. 1867; April 1, 1868; + July 1, 1869. + + [226] ‘Origin of Civilisation,’ 1870, p. 265. + + [227] Mr. Wallace gives cases in his ‘Contributions to the + Theory of Natural Selection,’ 1870, p. 354. + + [228] ‘Ancient Law,’ 1861, p. 22. For Mr. Bagehot’s remarks, + ‘Fortnightly Review,’ April 1, 1868, p. 452. + + [229] ‘The Variation of Animals and Plants under + Domestication,’ vol. i. p. 309. + + [230] ‘Fraser’s Magazine,’ Sept. 1868, p. 353. This article + seems to have struck many persons, and has given rise to two + remarkable essays and a rejoinder in the ‘Spectator,’ Oct. 3rd + and 17th 1868. It has also been discussed in the ‘Q. Journal of + Science,’ 1869, p. 152, and by Mr. Lawson Tait in the ‘Dublin + Q. Journal of Medical Science,’ Feb. 1869, and by Mr. E. Ray + Lankester in his ‘Comparative Longevity,’ 1870, p. 128. Similar + views appeared previously in the ‘Australasian,’ July 13, 1867. + I have borrowed ideas from several of these writers. + + [231] For Mr. Wallace, see ‘Anthropolog. Review,’ as before + cited. Mr. Galton in ‘Macmillan’s Magazine,’ Aug. 1865, p. 318; + also his great work, ‘Hereditary Genius,’ 1870. + + [232] ‘Hereditary Genius,’ 1870, p. 132-140. + + [233] See the fifth and sixth columns, compiled from good + authorities, in the table given in Mr. E. R. Lankester’s + ‘Comparative Longevity,’ 1870, p. 115. + + [234] ‘Hereditary Genius,’ 1870, p. 330. + + [235] ‘Origin of Species’ (fifth edition, 1869), p. 104. + + [236] ‘Hereditary Genius,’ 1870, p. 347. + + [237] E. Ray Lankester, ‘Comparative Longevity,’ 1870, p. 115. + The table of the intemperate is from Nelson’s ‘Vital + Statistics.’ In regard to profligacy, see Dr. Farr, “Influence + of Marriage on Mortality,” ‘Nat. Assoc. for the Promotion of + Social Science,’ 1858. + + [238] ‘Fraser’s Magazine,’ Sept. 1868, p. 353. ‘Macmillan’s + Magazine,’ Aug. 1865, p. 318. The Rev. F. W. Farrar (‘Fraser’s + Mag.,’ Aug. 1870, p. 264) takes a different view. + + [239] “On the Laws of the Fertility of Women,” in ‘Transact. + Royal Soc.’ Edinburgh, vol. xxiv. p. 287. See, also, Mr. + Galton, ‘Hereditary Genius,’ p. 352-357, for observations to + the above effect. + + [240] ‘Tenth Annual Report of Births, Deaths, &c., in + Scotland,’ 1867, p. xxix. + + [241] These quotations are taken from our highest authority on + such questions, namely, Dr. Farr, in his paper “On the + Influence of Marriage on the Mortality of the French People,” + read before the Nat. Assoc. for the Promotion of Social + Science, 1858. + + [242] Dr. Farr, ibid. The quotations given below are extracted + from the same striking paper. + + [243] I have taken the mean of the quinquennial means, given in + ‘The Tenth Annual Report of Births, Deaths, &c., in Scotland,’ + 1867. The quotation from Dr. Stark is copied from an article in + the ‘Daily News,’ Oct. 17th, 1868, which Dr. Farr considers + very carefully written. + + [244] See the ingenious and original argument on this subject + by Mr. Galton, ‘Hereditary Genius,’ p. 340-342. + + [245] Mr. Greg, ‘Fraser’s Magazine,’ Sept. 1868, p. 357. + + [246] ‘Hereditary Genius,’ 1870, p. 357-359. The Rev. F. H. + Farrar (‘Fraser’s Mag.’ , Aug. 1870, p. 257) advances arguments + on the other side. Sir C. Lyell had already (‘Principles of + Geology,’ vol. ii. 1868, p. 489) called attention, in a + striking passage, to the evil influence of the Holy Inquisition + in having lowered, through selection, the general standard of + intelligence in Europe. + + [247] Mr. Galton, ‘Macmillan’s Magazine,’ August, 1865, p. 325. + See, also, ‘Nature,’ “On Darwinism and National Life,” Dec. + 1869, p. 184. + + [248] ‘Last Winter in the United States,’ 1868, p. 29. + + [249] ‘On the Origin of Civilisation,’ ‘Proc. Ethnological + Soc.’ Nov. 26, 1867. + + [250] ‘Primeval Man,’ 1869. + + [251] ‘Royal Institution of Great Britain,’ March 15, 1867. + Also, ‘Researches into the Early History of Mankind,’ 1865. + + [252] ‘Primitive Marriage,’ 1865. See, likewise, an excellent + article, evidently by the same author, in the ‘North British + Review,’ July, 1869. Also, Mr. L. H. Morgan, “A Conjectural + Solution of the Origin of the Class. System of Relationship,” + in ‘Proc. American Acad. of Sciences,’ vol. vii. Feb. 1868. + Prof. Schaaffhausen (‘Anthropolog. Review,’ Oct. 1869, p. 373) + remarks on “the vestiges of human sacrifices found both in + Homer and the Old Testament.” + + [253] Sir J. Lubbock, ‘Prehistoric Times,’ 2nd edit. 1869, + chap. xv. and xvi. _et passim_. + + [254] Dr. F. Müller has made some good remarks to this effect + in the ‘Reise der Novara: Anthropolog. Theil,’ Abtheil. iii. + 1868, s. 127. + + [255] Isidore Geoffroy St.-Hilaire gives a detailed account of + the position assigned to man by various naturalists in their + classifications: ‘Hist. Nat. Gén.’ tom. ii. 1859, p. 170-189. + + [256] See the very interesting article, “L’Instinct chez les + Insectes,” by M. George Pouchet, ‘Revue des Deux Mondes,’ Feb. + 1870, p. 682. + + [257] Westwood, ‘Modern Class. of Insects,’ vol. ii. 1840, p. + 87. + + [258] ‘Proc. Zoolog. Soc.’ 1869, p. 4. + + [259] ‘Evidence as to Man’s Place in Nature,’ 1863, p. 70, _et + passim_. + + [260] Isid. Geoffroy, ‘Hist. Nat. Gén.’ tom. ii. 1859, p. 217. + + [261] “Ueber die Richtung der Haare,” &c., Müller’s ‘Archiv für + Anat. und Phys.’ 1837, s. 51. + + [262] On the hair in Hylobates, see ‘Nat. Hist. of Mammals,’ by + C. L. Martin, 1841, p. 415. Also, Isid. Geoffroy on the + American monkeys and other kinds, ‘Hist. Nat. Gén.’ vol. ii. + 1859, p. 216, 243. Eschricht, ibid., s. 46, 55, 61. Owen, ‘Anat. + of Vertebrates,’ vol. iii. p. 619. Wallace, ‘Contributions to + the Theory of Natural Selection,’ 1870. p. 344. + + [263] ‘Origin of Species,’ 5th edit. 1869, p. 194. ‘The + Variation of Animals and Plants under Domestication,’ vol. ii. + 1868, p. 348. + + [264] ‘An Introduction to the Classification of Animals,’ 1869, + p. 99. + + [265] This is nearly the same classification as that + provisionally adopted by Mr. St. George Mivart (‘Transact. + Philosoph. Soc.’ 1867, p. 300), who, after separating the + Lemuridæ, divides the remainder of the Primates into the + Hominidæ, the Simiadæ answering to the Catarhines, the Cebidæ, + and the Hapalidæ,—these two latter groups answering to the + Platyrhines. + + [266] ‘Transact. Zoolog. Soc.’ vol. vi. 1867, p. 214. + + [267] Mr. St. G. Mivart, ‘Transact. Phil. Soc.’ 1867, p. 410. + + [268] Messrs. Murie and Mivart on the Lemuroidea. ‘Transact. + Zoolog. Soc.’ vol. vii. 1869, p. 5. + + [269] Häckel has come to this same conclusion. See ‘Ueber die + Entstehung des Menschengeschlechts,’ in Virchow’s ‘Sammlung. + gemein. wissen. Vorträge,’ 1868, s. 61. Also his ‘Natürliche + Schöpfungsgeschichte,’ 1868, in which he gives in detail his + views on the genealogy of man. + + [270] ‘Anthropological Review,’ April, 1867, p. 236. + + [271] ‘Elements of Geology,’ 1865, p. 583-585. ‘Antiquity of + Man’, 1863; p. 145. + + [272] ‘Man’s Place in Nature,’ p. 105. + + [273] Elaborate tables are given in his ‘Generelle Morphologie’ + (B. ii. s. cliii. and s. 425); and with more especial reference + to man in his ‘Natürliche Schöpfungsgeschichte,’ 1868. Prof. + Huxley, in reviewing this latter work (‘The Academy,’ 1869, p. + 42) says, that he considers the phylum or lines of descent of + the Vertebrata to be admirably discussed by Häckel, although he + differs on some points. He expresses, also, his high estimate + of the value of the general tenor and spirit of the whole work. + + [274] ‘Palæontology,’ 1860, p. 199. + + [275] I had the satisfaction of seeing, at the Falkland + Islands, in April, 1833, and therefore some years before any + other naturalist, the locomotive larvæ of a compound Ascidian, + closely allied to, but apparently generically distinct from, + Synoicum. The tail was about five times as long as the oblong + head, and terminated in a very fine filament. It was plainly + divided, as sketched by me under a simple microscope, by + transverse opaque partitions, which I presume represent the + great cells figured by Kowalevsky. At an early stage of + development the tail was closely coiled round the head of the + larva. + + [276] ‘Mémoires de l’Acad. des Sciences de St. Pétersbourg,’ + tom. x. No. 15, 1866. + + [277] This is the conclusion of one of the highest authorities + in comparative anatomy, namely, Prof. Gegenbaur: ‘Grundzüge der + vergleich. Anat.’ 1870, s. 876. The result has been arrived at + chiefly from the study of the Amphibia; but it appears from the + researches of Waldeyer (as quoted in Humphry’s ‘Journal of + Anat. and Phys.’ 1869, p. 161), that the sexual organs of even + “the higher vertebrata are, in their early condition, + hermaphrodite.” Similar views have long been held by some + authors, though until recently not well based. + + [278] The male Thylacinus offers the best instance. Owen, + ‘Anatomy of Vertebrates,’ vol. iii. p. 771. + + [279] Serranus is well known often to be in an hermaphrodite + condition; but Dr. Günther informs me that he is convinced that + this is not its normal state. Descent from an ancient + androgynous prototype would, however, naturally favour and + explain, to a certain extent, the recurrence of this condition + in these fishes. + + [280] Mr. Lockwood believes (as quoted in ‘Quart. Journal of + Science,’ April, 1868, p. 269), from what he has observed of + the development of Hippocampus, that the walls of the abdominal + pouch of the male in some way afford nourishment. On male + fishes hatching the ova in their mouths, see a very interesting + paper by Prof. Wyman, in ‘Proc. Boston Soc. of Nat. Hist.’ + Sept. 15, 1857; also Prof. Turner, in ‘Journal of Anat. and + Phys.’ Nov. 1, 1866, p. 78. Dr. Günther has likewise described + similar cases. + + [281] All vital functions tend to run their course in fixed and + recurrent periods, and with tidal animals the periods would + probably be lunar; for such animals must have been left dry or + covered deep with water,—supplied with copious food or + stinted,—during endless generations, at regular lunar + intervals. If then the Vertebrata are descended from an animal + allied to the existing tidal Ascidians, the mysterious fact, + that with the higher and now terrestrial Vertebrata, not to + mention other classes, many normal and abnormal vital processes + run their course according to lunar periods, is rendered + intelligible. A recurrent period, if approximately of the right + duration, when once gained, would not, as far as we can judge, + be liable to be changed; consequently it might be thus + transmitted during almost any number of generations. This + conclusion, if it could be proved sound, would be curious; for + we should then see that the period of gestation in each mammal, + and the hatching of each bird’s eggs, and many other vital + processes, still betrayed the primordial birthplace of these + animals. + + [282] ‘History of India,’ 1841, vol. i. p. 323. Father Ripa + makes exactly the same remark with respect to the Chinese. + + [283] A vast number of measurements of Whites, Blacks, and + Indians, are given in the ‘Investigations in the Military and + Anthropolog. Statistics of American Soldiers,’ by B. A. Gould, + 1869, p. 298-358; on the capacity of the lungs, p. 471. See + also the numerous and valuable tables, by Dr. Weisbach, from + the observations of Dr. Scherzer and Dr. Schwarz, in the ‘Reise + der Novara: Anthropolog. Theil,’ 1867. + + [284] See, for instance, Mr. Marshall’s account of the brain of + a Bush-woman, in ‘Phil. Transact.’ 1864, p. 519. + + [285] Wallace, ‘The Malay Archipelago,’ vol. ii. 1869, p. 178. + + [286] With respect to the figures in the famous Egyptian caves + of Abou-Simbel, M. Pouchet says (‘The Plurality of the Human + Races,’ Eng. translat. 1864, p. 50), that he was far from + finding recognisable representations of the dozen or more + nations which some authors believe that they can recognise. + Even some of the most strongly-marked races cannot be + identified with that degree of unanimity which might have been + expected from what has been written on the subject. Thus + Messrs. Nott and Gliddon (‘Types of Mankind,’ p. 148) state + that Rameses II., or the Great, has features superbly European; + whereas Knox, another firm believer in the specific distinction + of the races of man (‘Races of Man,’ 1850, p. 201), speaking of + young Memnon (the same person with Rameses II., as I am + informed by Mr. Birch) insists in the strongest manner that he + is identical in character with the Jews of Antwerp. Again, + whilst looking in the British Museum with two competent judges, + officers of the establishment, at the statue of Amunoph III., + we agreed that he had a strongly negro cast of features; but + Messrs. Nott and Gliddon (ibid. p. 146, fig. 53) describe him + as “a hybrid, but not of negro intermixture.” + + [287] As quoted by Nott and Gliddon, ‘Types of Mankind,’ 1854, + p. 439. They give also corroborative evidence; but C. Vogt + thinks that the subject requires further investigation. + + [288] “Diversity of Origin of the Human Races,” in the + ‘Christian Examiner,’ July, 1850. + + [289] ‘Transact. B. Soc. of Edinburgh,’ vol. xxii. 1861, p. + 567. + + [290] ‘On the Phenomena of Hybridity in the Genus Homo,’ Eng. + translat. 1864. + + [291] See the interesting letter by Mr. T. A. Murray, in the + ‘Anthropolog. Review,’ April, 1868, p. liii. In this letter + Count Strzelecki’s statement, that Australian women who have + borne children to a white man are afterwards sterile with their + own race, is disproved. M. A. de Quatrefages has also collected + (‘Revue des Cours Scientifiques,’ March, 1869, p. 239) much + evidence that Australians and Europeans are not sterile when + crossed. + + [292] ‘An Examination of Prof. Agassiz’s Sketch of the Nat. + Provinces of the Animal World,’ Charleston, 1855, p. 44. + + [293] ‘Military and Anthropolog. Statistics of American + Soldiers,’ by B. A. Gould, 1869, p. 319. + + [294] ‘The Variation of Animals and Plants under + Domestication,’ vol. ii. p. 109. I may here remind the reader + that the sterility of species when crossed is not a + specially-acquired quality; but, like the incapacity of certain + trees to be grafted together, is incidental on other acquired + differences. The nature of these differences is unknown, but + they relate more especially to the reproductive system, and + much less to external structure or to ordinary differences in + constitution. One important element in the sterility of crossed + species apparently lies in one or both having been long + habituated to fixed conditions; for we know that changed + conditions have a special influence on the reproductive system, + and we have good reason to believe (as before remarked) that + the fluctuating conditions of domestication tend to eliminate + that sterility which is so general with species in a natural + state when crossed. It has elsewhere been shewn by me (ibid. + vol. ii. p. 185, and ‘Origin of Species,’ 5th edit. p. 317) + that the sterility of crossed species has not been acquired + through natural selection: we can see that when two forms have + already been rendered very sterile, it is scarcely possible + that their sterility should be augmented by the preservation or + survival of the more and more sterile individuals; for as the + sterility increases fewer and fewer offspring will be produced + from which to breed, and at last only single individuals will + be produced, at the rarest intervals. But there is even a + higher grade of sterility than this. Both Gärtner and Kölreuter + have proved that in genera of plants including numerous + species, a series can be formed from species which when crossed + yield fewer and fewer seeds, to species which never produce a + single seed, but yet are affected by the pollen of the other + species, for the germen swells. It is here manifestly + impossible to select the more sterile individuals, which have + already ceased to yield seeds; so that the acme of sterility, + when the germen alone is affected, cannot be gained through + selection. This acme, and no doubt the other grades of + sterility, are the incidental results of certain unknown + differences in the constitution of the reproductive system of + the species which are crossed. + + [295] ‘The Variation of Animals,’ &c., vol. ii. p. 92. + + [296] M. de Quatrefages has given (‘Anthropolog. Review,’ Jan. + 1869, p. 22) an interesting account of the success and energy + of the Paulistas in Brazil, who are a much crossed race of + Portuguese and Indians, with a mixture of the blood of other + races. + + [297] For instance with the aborigines of America and + Australia. Prof. Huxley says (‘Transact. Internat. Congress of + Prehist. Arch.’ 1868. p. 105) that the skulls of many South + Germans and Swiss are “as short and as broad as those of the + Tartars,” &c. + + [298] See a good discussion on this subject in Waitz, + ‘Introduct. to Anthropology,’ Eng. translat. 1863, p. 198-208, + 227. I have taken some of the above statements from H. Tuttle’s + ‘Origin and Antiquity of Physical Man,’ Boston, 1866, p. 35. + + [299] Prof. Nägeli has carefully described several striking + cases in his ‘Botanische Mittheilungen,’ B. ii. 1866, s. + 294-369. Prof. Asa Gray has made analogous remarks on some + intermediate forms in the Compositæ of N. America. + + [300] ‘Origin of Species,’ 5th edit. p. 68. + + [301] See Prof. Huxley to this effect in the ‘Fortnightly + Review,’ 1865, p. 275. + + [302] ‘Lectures on Man,’ Eng. translat. 1864, p. 468. + + [303] ‘Die Racen des Schweines,’ 1860, s. 46. ‘Vorstudien für + Geschichte, &c., Schweineschädel,’ 1864, s. 104. With respect + to cattle, see M. de Quatrefages, ‘Unité de l’Espèce Humaine,’ + 1861, p. 119. + + [304] Tylor’s ‘Early History of Mankind,’ 1865; for the + evidence with respect to gesture-language, see p. 54. Lubbock’s + ‘Prehistoric Times,’ 2nd edit. 1869. + + [305] ‘The Primitive Inhabitants of Scandinavia,’ Eng. + translat. edited by Sir J. Lubbock, 1868, p. 104. + + [306] Hodder M. Westropp, on Cromlechs, &c., ‘Journal of + Ethnological Soc.’ as given in ‘Scientific Opinion,’ June 2nd. + 1869, p. 3. + + [307] ‘Journal of Researches: Voyage of the “Beagle,”’ p. 46. + + [308] ‘Prehistoric Times,’ 1869, p. 574. + + [309] Translation in ‘Anthropological Review,’ Oct. 1868, p. + 431. + + [310] ‘Transact. Internat. Congress of Prehistoric Arch.’ 1868, + p. 172-175. See also Broca (translation) in ‘Anthropological + Review,’ Oct. 1868, p. 410. + + [311] Dr. Gerland, ‘Ueber das Aussterben der Naturvölker,’ + 1868, s. 82. + + [312] Gerland (ibid. s. 12) gives facts in support of this + statement. + + [313] See remarks to this effect in Sir H. Holland’s ‘Medical + Notes and Reflections,’ 1839, p. 390. + + [314] I have collected (‘Journal of Researches, Voyage of the + “Beagle,”’ p. 435) a good many cases bearing on this subject: + see also Gerland, ibid. s. 8. Poeppig speaks of the “breath of + civilisation as poisonous to savages.” + + [315] Sproat, ‘Scenes and Studies of Savage Life,’ 1868, p. + 284. + + [316] Bagehot, “Physics and Politics,” ‘Fortnightly Review,’ + April 1, 1868, p. 455. + + [317] “On Anthropology,” translation, ‘Anthropolog. Review,’ + Jan. 1868, p. 38. + + [318] ‘The Annals of Rural Bengal,’ 1868, p. 134. + + [319] ‘The Variation of Animals and Plants under + Domestication,’ vol. ii. p. 95. + + [320] Pallas, ‘Act. Acad. St. Petersburgh,’ 1780, part ii. p. + 69. He was followed by Rudolphi, in his ‘Beyträge zur + Anthropologie,’ 1812. An excellent summary of the evidence is + given by Godron, ‘De l’Espèce,’ 1859, vol. ii. p. 246, &c. + + [321] Sir Andrew Smith, as quoted by Knox, ‘Races of Man,’ + 1850, p. 473. + + [322] See De Quatrefages on this head, ‘Revue des Cours + Scientifiques,’ Oct. 17, 1868, p. 731. + + [323] Livingstone’s ‘Travels and Researches in S. Africa,’ + 1857, p. 338, 329. D’Orbigny, as quoted by Godron, ‘De + l’Espèce,’ vol. ii. p. 266. + + [324] See a paper read before the Royal Soc. in 1813, and + published in his Essays in 1818. I have given an account of Dr. + Wells’ views in the Historical Sketch (p. xvi) to my ‘Origin of + Species.’ Various cases of colour correlated with + constitutional peculiarities are given in my ‘Variation of + Animals under Domestication,’ vol. ii. p. 227, 335. + + [325] See, for instance, Nott and Gliddon, ‘Types of Mankind,’ + p. 68. + + [326] Major Tulloch, in a paper read before the Statistical + Society, April 20th, 1840, and given in the ‘Athenæum,’ 1840, + p. 353. + + [327] ‘The Plurality of the Human Race’ (translat.), 1864, p. + 60. + + [328] Quatrefages, ‘Unité de l’Espèce Humaine,’ 1861, p. 205. + Waitz, ‘Introduct. to Anthropology,’ translat. vol. i. 1863, p. + 124. Livingstone gives analogous cases in his ‘Travels.’ + + [329] In the spring of 1862 I obtained permission from the + Director-General of the Medical department of the Army, to + transmit to the surgeons of the various regiments on foreign + service a blank table, with the following appended remarks, but + I have received no returns. “As several well-marked cases have + been recorded with our domestic animals of a relation between + the colour of the dermal appendages and the constitution; and + it being notorious that there is some limited degree of + relation between the colour of the races of man and the climate + inhabited by them; the following investigation seems worth + consideration. Namely, whether there is any relation in + Europeans between the colour of their hair, and their liability + to the diseases of tropical countries. If the surgeons of the + several regiments, when stationed in unhealthy tropical + districts, would be so good as first to count, as a standard of + comparison, how many men, in the force whence the sick are + drawn, have dark and light-coloured hair, and hair of + intermediate or doubtful tints; and if a similar account were + kept by the same medical gentlemen, of all the men who suffered + from malarious and yellow fevers, or from dysentery, it would + soon be apparent, after some thousand cases had been tabulated, + whether there exists any relation between the colour of the + hair and constitutional liability to tropical diseases. Perhaps + no such relation would be discovered, but the investigation is + well worth making. In case any positive result were obtained, + it might be of some practical use in selecting men for any + particular service. Theoretically the result would be of high + interest, as indicating one means by which a race of men + inhabiting from a remote period an unhealthy tropical climate, + might have become dark-coloured by the better preservation of + dark-haired or dark-complexioned individuals during a long + succession of generations.” + + [330] ‘Anthropological Review,’ Jan. 1866, p. xxi. + + [331] See, for instance, Quatrefages (‘Revue des Cours + Scientifiques,’ Oct. 10, 1868, p. 724) on the effects of + residence in Abyssinia and Arabia, and other analogous cases. + Dr. Rolle (‘Der Mensch, seine Abstammung,’ &c., 1865, s. 99) + states, on the authority of Khanikof, that the greater number + of German families settled in Georgia, have acquired in the + course of two generations dark hair and eyes. Mr. D. Forbes + informs me that the Quichuas in the Andes vary greatly in + colour, according to the position of the valleys inhabited by + them. + + [332] Harlan, ‘Medical Researches,’ p. 532. Quatrefages (‘Unité + de l’Espèce Humaine,’ 1861, p. 128) has collected much evidence + on this head. + + [333] See Prof. Schaaffhausen, translat. in ‘Anthropological + Review,’ Oct. 1868, p. 429. + + [334] Mr. Catlin states (‘N. American Indians,’ 3rd edit. 1842, + vol. i. p. 49) that in the whole tribe of the Mandans, about + one in ten or twelve of the members of all ages and both sexes + have bright silvery grey hair, which is hereditary. Now this + hair is as coarse and harsh as that of a horse’s mane, whilst + the hair of other colours is fine and soft. + + [335] On the odour of the skin, Godron, ‘Sur l’Espèce,’ tom. + ii. p. 217. On the pores in the skin, Dr. Wilckens, ‘Die + Aufgaben der landwirth. Zootechnik,’ 1869, s. 7. + + [336] Westwood, ‘Modern Class. of Insects,’ vol. ii. 1810, p. + 541. In regard to the statement about Tanais, mentioned below, + I am indebted to Fritz Müller. + + [337] Kirby and Spence, ‘Introduction to Entomology,’ vol. iii. + 1826, p. 309. + + [338] Even with those of plants in which the sexes are + separate, the male flowers are generally mature before the + female. Many hermaphrodite plants are, as first shewn by C. K. + Sprengel, dichogamous; that is, their male and female organs + are not ready at the same time, so that they cannot be + self-fertilised. Now with such plants the pollen is generally + mature in the same flower before the stigma, though there are + some exceptional species in which the female organs are mature + before the male. + + [339] I have received information, hereafter to be given, to + this effect with respect to poultry. Even with birds, such as + pigeons, which pair for life, the female, as I hear from Mr. + Jenner Weir, will desert her mate if he is injured or grows + weak. + + [340] On the Gorilla, Savage and Wyman, ‘Boston Journal of Nat. + Hist.’ vol. v. 1845-47, p. 423. On Cynocephalus, Brehm, + ‘Illust. Thierleben,’ B. i. 1864, s. 77. On Mycetes, Rengger, + ‘Naturgesch.: Säugethiere von Paraguay,’ 1830, s. 14, 20. On + Cebus, Brehm, ibid. s. 108. + + [341] Pallas, ‘Spicilegia Zoolog.’ Fasc. xii. 1777, p. 29. Sir + Andrew Smith, ‘Illustrations of the Zoology of S. Africa,’ + 1849, pl. 29, on the Kobus. Owen, in his ‘Anatomy of + Vertebrates’ (vol. iii. 1868, p. 633) gives a table + incidentally showing which species of Antelopes pair and which + are gregarious. + + [342] Dr. Campbell, in ‘Proc. Zoolog. Soc.’ 1869, p. 138. See + also an interesting paper, by Lieut. Johnstone, in ‘Proc. + Asiatic Soc. of Bengal,’ May, 1868. + + [343] ‘The Ibis,’ vol. iii. 1861, p. 133, on the Progne + Widow-bird. See also on the Vidua axillaris, ibid. vol. ii. + 1860, p. 211. On the polygamy of the Capercailzie and Great + Bustard, see L. Lloyd, ‘Game Birds of Sweden,’ 1867, p. 19, and + 182. Montagu and Selby speak of the Black Grouse as polygamous + and of the Red Grouse as monogamous. + + [344] The Rev. E. S. Dixon, however, speaks positively + (‘Ornamental Poultry,’ 1848, p. 76) about the eggs of the + guinea-fowl being infertile when more than one female is kept + with the same male. + + [345] Noel Humphreys, ‘River Gardens,’ 1857. + + [346] Kirby and Spence, ‘Introduction to Entomology,’ vol. iii. + 1826, p. 342. + + [347] One parasitic Hymenopterous insect (Westwood, ‘Modern + Class. of Insects,’ vol. ii, p. 160) forms an exception to the + rule, as the male has rudimentary wings, and never quits the + cell in which it is born, whilst the female has well-developed + wings. Audouin believes that the females are impregnated by the + males which are born in the same cells with them; but it is + much more probable that the females visit other cells, and thus + avoid close inter-breeding. We shall hereafter meet with a few + exceptional cases, in various classes, in which the female, + instead of the male, is the seeker and wooer. + + [348] ‘Essays and Observations,’ edited by Owen, vol. i. 1861, + p. 194. + + [349] Prof. Sachs (‘Lehrbuch der Botanik,’ 1870, s. 633) in + speaking of the male and female reproductive cells, remarks, + “verhält sich die eine bei der Vereinigung activ, ... die + andere erscheint bei der Vereinigung passiv.” + + [350] ‘Reise der Novara: Anthropolog. Theil,’ 1867, s. 216-269. + The results were calculated by Dr. Weisbach from measurements + made by Drs. K. Scherzer and Schwarz. On the greater + variability of the males of domesticated animals, see my + ‘Variation of Animals and Plants under Domestication,’ vol. ii. + 1868, p. 75. + + [351] ‘Proceedings Royal Soc.’ vol. xvi. July, 1868, p. 519 and + 524. + + [352] ‘Proc. Royal Irish Academy,’ vol. x. 1868, p. 123. + + [353] ‘Massachusetts Medical Soc.’ vol. ii. No. 3, 1808, p. 9. + + [354] ‘The Variation of Animals and Plants under + Domestication,’ vol. ii. 1868, p. 75. In the last chapter but + one, the provisional hypothesis of pangenesis, above alluded + to, is fully explained. + + [355] These facts are given on the high authority of a great + breeder, Mr. Teebay, in Tegetmeier’s ‘Poultry Book,’ 1868, p. + 158. On the characters of chickens of different breeds, and on + the breeds of the pigeon, alluded to in the above paragraph, + see ‘Variation of Animals,’ &c., vol. i. p. 160, 249; vol. ii. + p. 77. + + [356] ‘Novæ species Quadrupedum e Glirium ordine,’ 1778, p. 7. + On the transmission of colour by the horse, see ‘Variation of + Animals, &c. under Domestication,’ vol. i. p. 21. Also vol. ii. + p. 71, for a general discussion on Inheritance as limited by + Sex. + + [357] Dr. Chapuis, ‘Le Pigeon Voyageur Belge,’ 1865, p. 87. + Boitard et Corbié, ‘Les Pigeons de Volière,’ &c., 1824, p. 173. + + [358] References are given in my ‘Variation of Animals under + Domestication,’ vol. ii. p. 72. + + [359] I am much obliged to Mr. Cupples for having made + enquiries for me in regard to the Roebuck and Red Deer of + Scotland from Mr. Robertson, the experienced head-forester to + the Marquis of Breadalbane. In regard to Fallow-deer, I am + obliged to Mr. Eyton and others for information. For the + _Cervus alces_ of N. America, see ‘Land and Water,’ 1868, p. + 221 and 254; and for the _C. Virginianus_ and _strongyloceros_ + of the same continent, see J. D. Caton, in ‘Ottawa Acad. of + Nat. Sc.’ 1868, p. 13. For _Cervus Eldi_ of Pegu, see Lieut. + Beavan, ‘Proc. Zoolog. Soc.’ 1867, p. 762. + + [360] _Antilocapra Americana._ Owen, ‘Anatomy of Vertebrates,’ + vol. iii. p. 627. + + [361] I have been assured that the horns of the sheep in North + Wales can always be felt, and are sometimes even an inch in + length, at birth. With cattle Youatt says (‘Cattle,’ 1834, p. + 277) that the prominence of the frontal bone penetrates the + cutis at birth, and that the horny matter is soon formed over + it. + + [362] I am greatly indebted to Prof. Victor Carus for having + made inquiries for me, from the highest authorities, with + respect to the merino sheep of Saxony. On the Guinea coast of + Africa there is a breed of sheep in which, as with merinos, the + rams alone bear horns; and Mr. Winwood Reade informs me that in + the one case observed, a young ram born on Feb. 10th first + showed horns on March 6th, so that in this instance the + development of the horns occurred at a later period of life, + conformably with our rule, than in the Welsh sheep, in which + both sexes are horned. + + [363] In the common peacock (_Pavo cristatus_) the male alone + possesses spurs, whilst both sexes of the Java peacock (_P. + muticus_) offer the unusual case of being furnished with spurs. + Hence I fully expected that in the latter species they would + have been developed earlier in life than in the common peacock; + but M. Hegt of Amsterdam informs me, that with young birds of + the previous year, belonging to both species, compared on April + 23rd, 1869, there was no difference in the development of the + spurs. The spurs, however, were as yet represented merely by + slight knobs or elevations. I presume that I should have been + informed if any difference in the rate of development had + subsequently been observed. + + [364] In some other species of the Duck Family the speculum in + the two sexes differs in a greater degree; but I have not been + able to discover whether its full development occurs later in + life in the males of such species, than in the male of the + common duck, as ought to be the case according to our rule. + With the allied _Mergus cucullatus_ we have, however, a case of + this kind: the two sexes differ conspicuously in general + plumage, and to a considerable degree in the speculum, which is + pure white in the male and greyish-white in the female. Now the + young males at first resemble, in all respects, the female, and + have a greyish-white speculum, but this becomes pure white at + an earlier age than that at which the adult male acquires his + other more strongly-marked sexual differences in plumage: see + Audubon, ‘Ornithological Biography,’ vol. iii. 1835, p. + 249-250. + + [365] ‘Das Ganze der Taubenzucht,’ 1837, s. 21, 24. For the + case of the streaked pigeons, see Dr. Chapuis, ‘Le Pigeon + Voyageur Belge.’ 1865, p. 87. + + [366] For full particulars and references on all these points + respecting the several breeds of the Fowl, see ‘Variation of + Animals and Plants under Domestication,’ vol. i. p. 250, 256. + In regard to the higher animals, the sexual differences which + have arisen under domestication are described in the same work + under the head of each species. + + [367] ‘Twenty-ninth Annual Report of the Registrar-General for + 1866.’ In this report (p. xii) a special decennial table is + given. + + [368] For Norway and Russia, see abstract of Prof. Faye’s + researches, in ‘British and Foreign Medico-Chirurg. Review,’ + April, 1867, p. 343, 345. For France, the ‘Annuaire pour l’An + 1867.’ p. 213. + + [369] In regard to the Jews, see M. Thury, ‘La Loi de + Production des Sexes,’ 1863, p. 25. + + [370] Babbage, ‘Edinburgh Journal of Science,’ 1829, vol. i. p. + 88; also p. 90, on still-born children. On illegitimate + children in England, see ‘Report of Registrar-General for + 1866,’ p. xv. + + [371] ‘British and Foreign Medico-Chirurg. Review,’ April, + 1867, p. 343. Dr. Stark also remarks (‘Tenth Annual Report of + Births, Deaths, &c., in Scotland,’ 1867, p. xxviii) that “These + examples may suffice to shew that, at almost every stage of + life, the males in Scotland have a greater liability to death + and a higher death-rate than the females. The fact, however, of + this peculiarity being most strongly developed at that + infantile period of life when the dress, food, and general + treatment of both sexes are alike, seems to prove that the + higher male death-rate is an impressed, natural, and + constitutional peculiarity due to sex alone.” + + [372] With the savage Guaranys of Paraguay, according to the + accurate Azara (‘Voyages dans l’Amérique mérid.’ tom. ii. 1809, + p. 60, 179), the women in proportion to the men are as 14 to + 13. + + [373] Leuckart in Wagner, ‘Handwörterbuch der Phys.’ B. iv. + 1853, s. 774. + + [374] Anthropological Review, April, 1870, p. cviii. + + [375] During the last eleven years a record has been kept of + the number of mares which have proved barren or prematurely + slipped their foals; and it deserves notice, as shewing how + infertile these highly-nurtured and rather closely-interbred + animals have become, that not far from one-third of the mares + failed to produce living foals. Thus during 1866, 809 male + colts and 816 female colts were born, and 743 mares failed to + produce offspring. During 1867, 836 males and 902 females were + born, and 794 mares failed. + + [376] I am much indebted to Mr. Cupples for having procured for + me the above returns from Scotland, as well as some of the + following returns on cattle. Mr. R. Elliot, of Laighwood, first + called my attention to the premature deaths of the males,—a + statement subsequently confirmed by Mr. Aitchison and others. + To this latter gentleman, and to Mr. Payan, I owe my thanks for + the larger returns on sheep. + + [377] Bell, ‘History of British Quadrupeds,’ p. 100. + + [378] ‘Illustrations of the Zoology of S. Africa,’ 1849, pl. + 29. + + [379] Brehm (‘Illust. Thierleben,’ B. iv. s. 990) comes to the + same conclusion. + + [380] On the authority of L. Lloyd, ‘Game Birds of Sweden,’ + 1867, p. 12, 132. + + [381] ‘Nat. Hist. of Selbourne,’ letter xxix. edit. of 1825, + vol. i. p. 139. + + [382] Mr. Jenner Weir received similar information, on making + enquiries during the following year. To shew the number of + chaffinches caught, I may mention that in 1869 there was a + match between two experts; and one man caught in a day 62, and + another 40, male chaffinches. The greatest number ever caught + by one man in a single day was 70. + + [383] ‘Ibis,’ vol. ii. p. 260, as quoted in Gould’s + ‘Trochilidæ,’ 1861, p. 52. For the foregoing proportions, I am + indebted to Mr. Salvin for a table of his results. + + [384] ‘Ibis,’ 1860, p. 137; and 1867, p. 369. + + [385] ‘Ibis,’ 1862, p. 137. + + [386] Leuckart quotes Bloch (Wagner, ‘Handwörterbuch der Phys.’ + B. iv. 1853, s. 775), that with fish there are twice as many + males as females. + + [387] Quoted in the ‘Farmer,’ March 18, 1869, p. 369. + + [388] ‘The Stormontfield Piscicultural Experiments,’ 1866, p. + 23. The ‘Field’ newspaper, June 29th, 1867. + + [389] ‘Land and Water,’ 1868, p. 41. + + [390] Yarrell, ‘Hist. British Fishes,’ vol. i. 1836, p. 307; on + the _Cyprinus carpio_, p. 331; on the _Tinca vulgaris_, p. 331; + on the _Abramis brama_, p. 336. See, for the minnow (_Leuciscus + phoxinus_), ‘Loudon’s Mag. of Nat. Hist.’ vol. v. 1832, p. 682. + + [391] Leuckart quotes Meinecke (Wagner, ‘Handwörterbuch der + Phys.’ B. iv. 1853, s. 775) that with Butterflies the males are + three or four times as numerous as the females. + + [392] ‘The Naturalist on the Amazons,’ vol. ii. 1863, p. 228, + 347. + + [393] Four of these cases are given by Mr. Trimen in his + ‘Rhopalocera Africæ Australis.’ + + [394] Quoted by Trimen, ‘Transact. Ent. Soc.’ vol. v. part iv. + 1866, p. 330. + + [395] ‘Transact. Linn. Soc.’ vol. xxv. p. 37. + + [396] ‘Proc. Entomolog. Soc.’ Feb. 17th, 1868. + + [397] Quoted by Dr. Wallace in ‘Proc. Ent. Soc.’ 3rd series, + vol. v. 1867, p. 487. + + [398] Blanchard, ‘Metamorphoses, Mœurs des Insectes,’ 1868, p. + 225-226. + + [399] ‘Lepidopteren-Doubblettren Liste,’ Berlin, No. x. 1866. + + [400] This naturalist has been so kind as to send me some + results from former years, in which the females seemed to + preponderate; but so many of the figures were estimates, that I + found it impossible to tabulate them. + + [401] Günther’s ‘Record of Zoological Literature,’ 1867, p. + 260. On the excess of female Lucanus, ibid. p. 250. On the + males of Lucanus in England, Westwood, ‘Modern Class. of + Insects,’ vol. i. p. 187. On the Siagonium, ibid. p. 172. + + [402] Walsh, in ‘The American Entomologist,’ vol. i. 1869, p. + 103. F. Smith, ‘Record of Zoological Literature,’ 1867, p. 328. + + [403] ‘Farm Insects,’ p. 45-46. + + [404] ‘Observations on N. American Neuroptera,’ by H. Hagen and + B. D. Walsh, ‘Proc. Ent. Soc. Philadelphia,’ Oct. 1863, p. 168, + 223, 239. + + [405] ‘Proc. Ent. Soc. London,’ Feb. 17, 1868. + + [406] Another great authority in this class, Prof. Thorell of + Upsala (‘On European Spiders,’ 1869-70, part i. p. 205) speaks + as if female spiders were generally commoner than the males. + + [407] See, on this subject, Mr. Pickard-Cambridge, as quoted in + ‘Quarterly Journal of Science,’ 1868, p. 429. + + [408] I have often been struck with the fact, that in several + species of Primula the seeds in the capsules which contained + only a few were very much larger than the numerous seeds in the + more productive capsules. + + [409] ‘Principles of Biology,’ vol. ii. 1867, chaps. ii.-xi. + + [410] ‘De l’Espèce et de la Class.’ &c., 1869, p. 106. + + [411] See, for instance, the account which I have given in my + ‘Journal of Researches,’ 1845, p. 7. + + [412] I have given (‘Geolog. Observations on Volcanic Islands,’ + 1844, p. 53) a curious instance of the influence of light on + the colours of a frondescent incrustation, deposited by the + surf on the coast-rocks of Ascension, and formed by the + solution of triturated sea-shells. + + [413] ‘Facts and Arguments for Darwin,’ English translat. 1869, + p. 20. See the previous discussion on the olfactory threads. + Sars has described a somewhat analogous case (as quoted in + ‘Nature,’ 1870, p. 455) in a Norwegian crustacean, the + _Pontoporeia affinis_. + + [414] See Sir J. Lubbock in ‘Annals. and Mag. of Nat. Hist.’ + vol. xi. 1853, pl. i. and x.; and vol. xii. (1853) pl. vii. See + also Lubbock in ‘Transact. Ent. Soc.’ vol. iv. new series, + 1856-1858, p. 8. With respect to the zigzagged antennæ + mentioned below, see Fritz Müller, ‘Facts and Arguments for + Darwin’ 1869, p. 40, foot-note. + + [415] See a paper by Mr. C. Spence Bate, with figures, in + ‘Proc. Zoolog. Soc.’ 1868, p. 363; and on the nomenclature of + the genus, ibid. p. 585. I am greatly indebted to Mr. Spence + Bate for nearly all the above statements with respect to the + chelæ of the higher crustaceans. + + [416] ‘Hist. Nat. des Crust.’ tom. ii. 1837, p. 50. + + [417] Fritz Müller, ‘Facts and Arguments for Darwin,’ 1869, p. + 25-28. + + [418] ‘Travels in the Interior of Brazil,’ 1846, p. 111. I have + given, in my ‘Journal of Researches,’ p. 463, an account of the + habits of the Birgos. + + [419] Mr. Ch. Fraser, in ‘Proc. Zoolog. Soc.’ 1869, p. 3. I am + indebted to Mr. Bate for the statement from Dr. Power. + + [420] Claus, ‘Die freilebenden Copepoden,’ 1863, s. 35. + + [421] ‘Facts and Arguments,’ &c., p. 79. + + [422] ‘A History of the Spiders of Great Britain,’ 1861-64. For + the following facts, see p. 102, 77, 88. + + [423] Aug. Vinson (‘Aranéides des Iles de la Réunion,’ pl. vi. + figs. 1 and 2) gives a good instance of the small size of the + male in _Epeira nigra_. In this species, as I may add, the male + is testaceous and the female black with legs banded with red. + Other even more striking cases of inequality in size between + the sexes have been recorded (‘Quarterly Journal of Science,’ + 1868, July, p. 429); but I have not seen the original accounts. + + [424] Kirby and Spence, ‘Introduction to Entomology,’ vol. i. + 1818, p. 280. + + [425] Theridion (Asagena, Sund.) serratipes, 4-punctatum et + guttatum; see Westring, in Kroyer, ‘Naturhist. Tidskrift,’ vol. + iv. 1842-1843, p. 349; and vol. ii. 1846-1849, p. 342. See, + also, for other species, ‘Araneæ Svecicæ,’ p. 184. + + [426] Walckenaer et P. Gervais, ‘Hist. Nat. des Insectes: + Aptères,’ tom. iv. 1847, p. 17, 19, 68. + + [427] Sir J. Lubbock, ‘Transact. Linnean Soc.’ vol. xxv. 1866, + p. 484. With respect to the Mutillidæ see Westwood, ‘Modern + Class. of Insects,’ vol. ii. p. 213. + + [428] These organs in the male often differ in closely-allied + species, and afford excellent specific characters. But their + importance, under a functional point of view, as Mr. E. + MacLachlan has remarked to me, has probably been overrated. It + has been suggested, that slight differences in these organs + would suffice to prevent the intercrossing of well-marked + varieties or incipient species, and would thus aid in their + development. That this can hardly be the case, we may infer + from the many recorded cases (see for instance, Bronn, + ‘Geschichte der Natur,’ B. ii. 1843, s. 164; and Westwood, + ‘Transact. Ent. Soc.’ vol. iii. 1842, p. 195) of distinct + species having been observed in union. Mr. MacLachlan informs + me (vide ‘Stett. Ent. Zeitung,’ 1867, s. 155) that when several + species of Phryganidæ, which present strongly-pronounced + differences of this kind, were confined together by Dr. Aug. + Meyer, _they coupled_, and one pair produced fertile ova. + + [429] ‘The Practical Entomologist,’ Philadelphia, vol. ii. May, + 1867, p. 88. + + [430] Mr. Walsh, ibid. p. 107. + + [431] ‘Modern Classification of Insects,’ vol. ii. 1840, p. + 206, 205. Mr. Walsh, who called my attention to this double use + of the jaws, says that he has repeatedly observed this fact. + + [432] We have here a curious and inexplicable case of + dimorphism, for some of the females of four European species of + Dytiscus, and of certain species of Hydroporus, have their + elytra smooth; and no intermediate gradations between sulcated + or punctured and quite smooth elytra have been observed. See + Dr. H. Schaum, as quoted in the ‘Zoologist,’ vol. v.-vi. + 1847-48, p. 1896. Also Kirby and Spence, ‘Introduction to + Entomology,’ vol. iii. 1826, p. 305. + + [433] Westwood, ‘Modern Class.’ vol. ii. p. 193. The following + statement about Penthe, and others in inverted commas, are + taken from Mr. Walsh, ‘Practical Entomologist,’ Philadelphia, + vol. ii. p. 88. + + [434] Kirby and Spence, ‘Introduct.’ &c., vol. iii. p. 332-336. + + [435] ‘Insecta Maderensia,’ 1854, p. 20. + + [436] E. Doubleday, ‘Annals and Mag. of Nat. Hist.’ vol. i. + 1848, p. 379. I may add that the wings in certain Hymenoptera + (see Shuckard, ‘Fossorial Hymenop.’ 1837, p. 39-43) differ in + neuration according to sex. + + [437] H. W. Bates, in ‘Journal of Proc. Linn. Soc.’ vol. vi. + 1862, p. 74. Mr. Wonfor’s observations are quoted in ‘Popular + Science Review,’ 1868, p. 343. + + [438] Kirby and Spence, ‘Introduction to Entomology,’ vol. iii. + p. 299. + + [439] Robinet, ‘Vers à Soie,’ 1848, p. 207. + + [440] ‘Transact. Ent. Soc.’ 3rd series, vol. v. p. 486. + + [441] ‘Journal of Proc. Ent. Soc.’ Feb. 4th, 1867, p. lxxi. + + [442] For this and other statements on the size of the sexes, + see Kirby and Spence, ibid. vol. iii. p. 300; on the duration + of life in insects, see p. 344. + + [443] ‘Transact. Linnean Soc.’ vol. xxvi. 1868, p. 296. + + [444] ‘The Malay Archipelago,’ vol. ii. 1869, p. 313. + + [445] ‘Modern Classification of Insects,’ vol. ii. 1840, p. + 526. + + [446] See Mr. B. T. Lowne’s very interesting work, ‘On the + Anatomy of the Blow-Fly, Musca vomitoria,’ 1870, p. 14. + + [447] Westwood, ‘Modern Class. of Insects,’ vol. ii. p. 473. + + [448] These particulars are taken from Westwood’s ‘Modern + Class. of Insects,’ vol. ii. 1840, p. 422. See, also, on the + Fulgoridæ, Kirby and Spence, ‘Introduct.’ vol. ii. p. 401. + + [449] ‘Zeitschrift für wissenschaft. Zoolog.’ B. xvii. 1867, s. + 152-158. + + [450] I am indebted to Mr. Walsh for having sent me this + extract from a ‘Journal of the Doings of Cicada septemdecim,’ + by Dr. Hartman. + + [451] L. Guilding, ‘Transact. Linn. Soc.’ vol. xv. p. 154. + + [452] Köppen, as quoted in the ‘Zoological Record,’ for 1867, + p. 460. + + [453] Gilbert White, ‘Nat. Hist. of Selborne,’ vol. ii. 1825, + p. 262. + + [454] Harris, ‘Insects of New England,’ 1842, p. 128. + + [455] ‘The Naturalist on the Amazons,’ vol. i. 1863, p. 252. + Mr. Bates gives a very interesting discussion on the gradations + in the musical apparatus of the three families. See also + Westwood, ‘Modern Class.’ vol. ii. p. 445 and 453. + + [456] ‘Proc. Boston Soc. of Nat. Hist.’ vol. xi. April, 1868. + + [457] ‘Nouveau Manuel d’Anat. Comp.’ (French translat.), tom. + i. 1850 p. 567. + + [458] ‘Zeitschrift für wissenschaft. Zoolog.’ B. xvii. 1867, s. + 117. + + [459] Westwood, ‘Modern Class. of Insects,’ vol. i. p. 440. + + [460] Westwood, ‘Modern Class. of Insects,’ vol. i. p. 453. + + [461] Landois, ibid. s. 121, 122. + + [462] Mr. Walsh also informs me that he has noticed that the + female of the _Platyphyllum concavum_, “when captured makes a + feeble grating noise by shuffling her wing-covers together.” + + [463] Landois, ibid. s. 113. + + [464] ‘Insects of New England,’ 1842, p. 133. + + [465] Westwood, ‘Modern Classification,’ vol. i. p. 462. + + [466] Westwood, ibid. vol. i. p. 453. + + [467] Landois, ibid. s. 115, 116, 120, 122. + + [468] ‘Transact. Ent. Soc.’ 3rd series, vol. ii. (‘Journal of + Proceedings, p. 117.) + + [469] Westwood, ‘Modern Class. of Insects,’ vol. i. p. 427; for + crickets, p. 445. + + [470] Mr. Ch. Horne, in ‘Proc. Ent. Soc.’ May 3, 1869, p. xii. + + [471] The _Oecanthus nivalis_, Harris, ‘Insects of New + England,’ 1842, p. 124. + + [472] Platyblemnus: Westwood, ‘Modern. Class.’ vol. i. p. 447. + + [473] B. D. Walsh, the Pseudo-neuroptera of Illinois, in ‘Proc. + Ent. Soc. of Philadelphia,’ 1862, p. 361. + + [474] ‘Modern Class.’ vol. ii. p. 37. + + [475] Walsh, ibid. p. 381. I am indebted to this naturalist for + the following facts on Hetærina, Anax, and Gomphus. + + [476] ‘Transact. Ent. Soc’ vol. i. 1836, p. lxxxi. + + [477] See abstract in the ‘Zoological Record’ for 1867, p. 450. + + [478] Kirby and Spence, ‘Introduct. to Entomology,’ vol. ii. + 1818, p. 35. + + [479] See an interesting article, “The Writings of Fabre,” in + ‘Nat. Hist. Review,’ April, 1862, p. 122. + + [480] ‘Journal of Proc. of Entomolog. Soc.’ Sept. 7th, 1863, + p. 169. + + [481] P. Huber, ‘Recherches sur les Mœurs des Fourmis,’ 1810, + p. 150, 165. + + [482] ‘Proc. Entomolog. Soc. of Philadelphia,’ 1866, p. + 238-239. + + [483] Quoted by Westwood, ‘Modern Class. of Insects,’ vol. ii. + p. 214. + + [484] Pyrodes pulcherrimus, in which the sexes differ + conspicuously, has been described by Mr. Bates in ‘Transact. + Ent. Soc.’ 1869, p. 50. I will specify the few other cases in + which I have heard of a difference in colour between the sexes + of beetles. Kirby and Spence (‘Introduct. to Entomology,’ vol. + iii. p. 301) mention a Cantharis, Meloe, Rhagium, and the + _Leptura testacea_; the male of the latter being testaceous, + with a black thorax, and the female of a dull red all over. + These two latter beetles belong to the Order of Longicorns. + Messrs. R. Trimen and Waterhouse, junr., inform me of two + Lamellicorns, viz., a Peritrichia and Trichius, the male of the + latter being more obscurely coloured than the female. In + _Tillus elongatus_ the male is black, and the female always, as + it is believed, of a dark blue colour with a red thorax. The + male, also, of _Orsodacna atra_, as I hear from Mr. Walsh, is + black, the female (the so-called _O. ruficollis_) having a + rufous thorax. + + [485] ‘Proc. Entomolog. Soc. of Philadelphia,’ 1864, p. 228. + + [486] Kirby and Spence, ‘Introduct. Entomolog.’ vol. iii. p. + 300. + + [487] Kirby and Spence, ibid. vol. iii. p. 329. + + [488] ‘Modern Classification of Insects,’ vol. i. p. 172. On + the same page there is an account of Siagonium. In the British + Museum I noticed one male specimen of Siagonium in an + intermediate condition, so that the dimorphism is not strict. + + [489] ‘The Malay Archipelago,’ vol. ii. 1869, p. 276. + + [490] ‘Entomological Magazine,’ vol. i. 1833, p. 82. See also + on the conflicts of this species, Kirby and Spence, ibid. vol. + iii. p. 314; and Westwood, ibid. vol. i. p. 187. + + [491] Quoted from Fischer, in ‘Dict. Class. d’Hist. Nat.’ tom. + x. p. 324. + + [492] ‘Ann. Soc. Entomolog. France,’ 1866, as quoted in + ‘Journal of Travel,’ by A. Murray, 1868, p. 135. + + [493] Westwood, ‘Modern Class.’ vol. i. p. 184. + + [494] Wollaston, On certain musical Curculionidæ, ‘Annals and + Mag. of Nat. Hist.’ vol. vi. 1860, p. 14. + + [495] ‘Zeitschrift für wiss. Zoolog.’ B. xvii. 1867, s. 127. + + [496] I am greatly indebted to Mr. G. R. Crotch for having sent + me numerous prepared specimens of various beetles belonging to + these three families and others, as well as for valuable + information of all kinds. He believes that the power of + stridulation in the Clythra has not been previously observed. I + am also much indebted to Mr. E. W. Janson, for information and + specimens. I may add that my son, Mr. F. Darwin, finds that + _Dermestes murinus_ stridulates, but he searched in vain for + the apparatus. Scolytus has lately been described by Mr. Algen + as a stridulator, in the ‘Edinburgh Monthly Magazine,’ 1869, + Nov., p. 130. + + [497] Schiödte, translated in ‘Annals and Mag. of Nat. Hist.’ + vol. xx. 1867, p. 37. + + [498] Westring has described (Kroyer, ‘Naturhist. Tidskrift,’ + B. ii. 1848-49, p. 334) the stridulating organs in these two, + as well as in other families. In the Carabidæ I have examined + _Elaphrus uliginosus_ and _Blethisa multipunctata_, sent to me + by Mr. Crotch. In Blethisa the transverse ridges on the + furrowed border of the abdominal segment do not come into play, + as far as I could judge, in scraping the rasps on the elytra. + + [499] I am indebted to Mr. Walsh, of Illinois, for having sent + me extracts from Leconte’s ‘Introduction to Entomology,’ p. + 101, 143. + + [500] M. P. de la Brulerie, as quoted in ‘Journal of Travel,’ + A. Murray, vol. i. 1868, p. 135. + + [501] Mr. Doubleday informs me that “the noise is produced by + the insect raising itself on its legs as high as it can, and + then sinking its thorax five or six times, in rapid succession, + against the substance upon which it is sitting.” For references + on this subject see Landois, ‘Zeitschrift für wissen. Zoolog.’ + B. xvii. s. 131. Olivier says (as quoted by Kirby and Spence, + ‘Introduct.’ vol. ii. p. 395) that the female of _Pimelia + striata_ produces a rather loud sound by striking her abdomen + against any hard substance, “and that the male, obedient to + this call, soon attends her and they pair.” + + [502] Apatura Iris: ‘The Entomologist’s Weekly Intelligencer,’ + 1859, p. 139. For the Bornean Butterflies see C. Collingwood, + ‘Rambles of a Naturalist,’ 1868, p. 183. + + [503] See my ‘Journal of Researches,’ 1845, p. 33. Mr. + Doubleday has detected (‘Proc. Ent. Soc.’ March 3rd, 1845, p. + 123) a peculiar membranous sac at the base of the front wings, + which is probably connected with the production of the sound. + + [504] See also Mr. Bates’ paper in ‘Proc. Ent. Soc. of + Philadelphia,’ 1865, p. 206. Also Mr. Wallace on the same + subject, in regard to Diadema, in ‘Transact. Entomolog. Soc. of + London,’ 1869, p. 278. + + [505] ‘The Naturalist on the Amazons,’ vol. i. 1863, p. 19. + + [506] See the interesting article in the ‘Westminster Review,’ + July, 1867, p. 10. A woodcut of the Kallima is given by Mr. + Wallace in Hardwicke’s ‘Science Gossip,’ Sept. 1867, p. 196. + + [507] See the interesting observations by Mr. T. W. Wood, ‘The + Student,’ Sept. 1868, p. 81. + + [508] Mr. Wallace in ‘Hardwicke’s Science Gossip,’ Sept. 1867, + p. 193. + + [509] See also, on this subject, Mr. Weir’s paper in ‘Transact. + Ent. Soc.’ 1869, p. 23. + + [510] ‘Westminster Review,’ July, 1867, p. 16. + + [511] For instance, Lithosia; but Prof. Westwood (‘Modern + Class. of Insects,’ vol. ii. p. 390) seems surprised at this + case. On the relative colours of diurnal and nocturnal + Lepidoptera, see ibid. p. 333 and 392; also Harris, ‘Treatise + on the Insects of New England,’ 1842, p. 315. + + [512] Such differences between the upper and lower surfaces of + the wings of several species of Papilio, may be seen in the + beautiful plates to Mr. Wallace’s Memoir on the Papilionidæ of + the Malayan Region, in ‘Transact. Linn. Soc.’ vol. xxv. part i. + 1865. + + [513] ‘Proc. Ent. Soc.’ March 2nd, 1868. + + [514] See also an account of the S. American genus Erateina + (one of the Geometræ) in ‘Transact. Ent. Soc.’ new series, vol. + v. pl. xv. and xvi. + + [515] ‘Proc. Ent. Soc. of London,’ July 6, 1868, p. xxvii. + + [516] Harris, ‘Treatise,’ &c., edited by Flint, 1862, p. 395. + + [517] For instance, I observe in my son’s cabinet that the + males are darker than the females in the _Lasiocampa quercus_, + _Odonestis potatoria_, _Hypogymna dispar_, _Dasychira + pudibunda_, and _Cycnia mendica_. In this latter species the + difference in colour between the two sexes is strongly marked; + and Mr. Wallace informs me that we here have, as he believes, + an instance of protective mimickry confined to one sex, as will + hereafter be more fully explained. The white female of the + Cycnia resembles the very common _Spilosoma menthrasti_, both + sexes of which are white; and Mr. Stainton observed that this + latter moth was rejected with utter disgust by a whole brood of + young turkeys, which were fond of eating other moths; so that + if the Cycnia was commonly mistaken by British birds for the + Spilosoma, it would escape being devoured, and its white + deceptive colour would thus be highly beneficial. + + [518] ‘Rambles of a Naturalist in the Chinese Seas,’ 1868, p. + 182. + + [519] Wallace on the Papilionidæ of the Malayan Region, in + ‘Transact. Linn. Soc.’ vol. xxv. 1865, p. 8, 36. A striking case + of a rare variety, strictly intermediate between two other + well-marked female varieties, is given by Mr. Wallace. See also + Mr. Bates, in ‘Proc. Entomolog. Soc.’ Nov. 19th, 1866, p. xl. + + [520] Mr. R. MacLachlan, ‘Transact. Ent. Soc.’ vol. ii. part + 6th, 3rd series, 1866, p. 459. + + [521] H. W. Bates, ‘The Naturalist on the Amazons,’ vol. ii. + 1863, p. 228. A. R. Wallace, in ‘Transact. Linn. Soc.’ vol. xxv. + 1865, p. 10. + + [522] On this whole subject see ‘The Variation of Animals and + Plants under Domestication,’ vol. ii. 1868, chap. xxiii. + + [523] A. R. Wallace, in ‘The Journal of Travel,’ vol. i. 1868, + p. 88. ‘Westminster Review,’ July, 1857, p. 37. See also + Messrs. Wallace and Bates in ‘Proc. Ent. Soc.’ Nov. 19th, 1866, + p. xxxix. + + [524] ‘The Variation of Animals and Plants under + Domestication,’ vol. ii. chap. xii. p. 17. + + [525] ‘Transact. Linn. Soc.’ vol. xxiii. 1862, p. 495. + + [526] ‘Proc. Ent. Soc.’ Dec. 3rd, 1866, p. xlv. + + [527] ‘Transact. Linn. Soc.’ vol. xxv. 1865, p. 1; also + ‘Transact. Ent. Soc.’ vol. iv. (3rd series), 1867, p. 301. + + [528] See an ingenious article entitled, “Difficulties of the + Theory of Natural Selection,” in the ‘Month,’ 1869. The writer + strangely supposes that I attribute the variations in colour of + the Lepidoptera, by which certain species belonging to distinct + families have come to resemble others, to reversion to a common + progenitor; but there is no more reason to attribute these + variations to reversion than in the case of any ordinary + variation. + + [529] Wallace, “Notes on Eastern Butterflies,” ‘Transact. Ent. + Soc.’ 1869, p. 287. + + [530] Wallace, in ‘Westminster Review,’ July, 1867, p. 37; and + in ‘Journal of Travel and Nat. Hist.’ vol. i. 1868, p. 88. + + [531] See remarks by Messrs. Bates and Wallace, in ‘Proc. Ent. + Soc.’ Nov. 19, 1866, p. xxxix. + + [532] See Mr. Wallace in ‘Westminster Review,’ July, 1867, p. + 11 and 37. The male of no butterfly, as Mr. Wallace informs me, + is known to differ in colour, as a protection, from the female; + and he asks me how I can explain this fact on the principle + that one sex alone has varied and has transmitted its + variations exclusively to the same sex, without the aid of + selection to check the variations being inherited by the other + sex. No doubt if it could be shewn that the females of very + many species had been rendered beautiful through protective + mimickry, but that this has never occurred with the males, it + would be a serious difficulty. But the number of cases as yet + known hardly suffices for a fair judgment. We can see that the + males, from having the power of flying more swiftly, and thus + escaping danger, would not be so likely as the females to have + had their colours modified for the sake of protection; but this + would not in the least have interfered with their receiving + protective colours through inheritance from the females. In the + second place, it is probable that sexual selection would + actually tend to prevent a beautiful male from becoming + obscure, for the less brilliant individuals would be less + attractive to the females. Supposing that the beauty of the + male of any species had been mainly acquired through sexual + selection, yet if this beauty likewise served as a protection, + the acquisition would have been aided by natural selection. But + it would be quite beyond our power to distinguish between the + two processes of sexual and ordinary selection. Hence it is not + likely that we should be able to adduce cases of the males + having been rendered brilliant exclusively through protective + mimickry, though this is comparatively easy with the females, + which have rarely or never been rendered beautiful, as far as + we can judge, for the sake of sexual attraction, although they + have often received beauty through inheritance from their male + parents. + + [533] ‘Proc. Entomolog. Soc.’ Dec. 3rd, 1866, p. xlv., and + March 4th, 1867, p. lxxx. + + [534] See Mr. J. Jenner Weir’s paper on insects and + insectivorous birds, in ‘Transact. Ent. Soc.’ 1869, p. 21; also + Mr. Butler’s paper, ibid. p. 27. + + +END OF VOL. I. + +LONDON: + +PRINTED BY WILLIAM CLOWES AND SONS, DUKE STREET, STAMFORD STREET, AND +CHARING CROSS. + + +50A, ALBEMARLE STREET, LONDON, +_January, 1871_. + + +MR. MURRAY’S + +LIST OF POPULAR WORKS. + + +_One Shilling._ + +THE PRINCIPAL SPEECHES AND ADDRESSES OF H.R.H. THE PRINCE CONSORT, with +an Introduction giving some Outlines of his Character. With Portrait. + +MUSIC AND THE ART OF DRESS. Two Essays. By A LADY. + +THE FALL OF JERUSALEM. A Dramatic Poem. By DEAN MILMAN. + +THEODORE HOOK. A Sketch. By J. G. LOCKHART. + +HISTORY OF THE GUILLOTINE. By RT. HON. J. W. CROKER. Woodcuts. + +THE CHACE. A Descriptive Essay. By C. J. APPERLEY (NIMROD). With +Woodcuts. + +REJECTED ADDRESSES; or, The New Theatrum Poetarum. By HORACE and JAMES +SMITH. 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By HENRY TAYLOR. + +THE SIEGE OF GIBRALTAR, 1779-83. With a Description and Account of that +Garrison from, the Earliest Periods. By JOHN DRINKWATER. + +SIR ROBERT SALE’S BRIGADE IN AFFGHANISTAN AND THE DEFENCE OF JELLALABAD. +By REV. G. R. GLEIG. + +THE TWO SIEGES OF VIENNA BY THE TURKS. Translated from the German. By +LORD ELLESMERE. + +THE WAYSIDE CROSS; or, The Raid of Gomez: a Tale of the Carlist War. By +CAPT. MILMAN. + +ADVENTURES ON THE ROAD TO PARIS DURING THE CAMPAIGNS OF 1813-14. From +the Autobiography of HENRY STEFFENS. + +STOKERS AND POKERS; or, The North-Western Railway.—The Electric +Telegraph—and Railway Clearing House. By SIR FRANCIS HEAD. + +TRAVELS IN EGYPT, NUBIA, SYRIA, AND THE HOLY LAND, with a Journey round +the Dead Sea, and through the Country East of the Jordan. By IRBY and +MANGLES. + +WESTERN BARBARY; An Account of the Wild Tribes and Savage Animals. By +JOHN H. DRUMMOND HAY. + +LETTERS FROM THE SHORES OF THE BALTIC. By A LADY. + +NOTES AND SKETCHES OF NEW SOUTH WALES DURING A RESIDENCE OF MANY YEARS +IN THAT COLONY. By MRS. CHARLES MEREDITH. + +RECOLLECTIONS OF BUSH LIFE IN AUSTRALIA, during a Residence of Eight +Years in the Interior. By REV. H. W. HAYGARTH. + +JOURNAL OF A RESIDENCE AMONG THE NEGROES IN THE WEST INDIES. By M. G. +LEWIS. + +MEMOIRS OF FATHER RIPA DURING THIRTEEN YEARS’ RESIDENCE AT THE COURT OF +PEKIN, in the Service of the Emperor of China, with an Account of the +Foundation of the College for the Education of Young Chinese in Naples. +From the Italian. By FORTUNATO PRANDI. + +PHILIP MUSGRAVE; or, Memoirs of a Church of England Missionary in the +North American Colonies. By REV. J. ABBOTT. + +A MONTH IN NORWAY. By JOHN G. HOLLWAY. + +LETTERS FROM MADRAS; or, First Impressions of Life and Manners in India. +By A LADY. + +ROUGH NOTES TAKEN DURING SOME RAPID RIDES ACROSS THE PAMPAS AND AMONG +THE ANDES. By SIR FRANCIS HEAD. + +A VOYAGE UP THE RIVER AMAZON, including a Visit to PARA. By WILLIAM H. +EDWARDS. + +A POPULAR ACCOUNT OF THE MANNERS AND CUSTOMS OF INDIA. By REV. CHARLES +ACLAND. + +ADVENTURES IN THE LIBYAN DESERT AND THE OASIS OF JUPITER AMMON. By BAYLE +ST. JOHN. + +LIFE OF SIR FOWELL BUXTON. By his Son, CHARLES BUXTON. With an Inquiry +into the Results of Emancipation. + + +_Two Shillings and Sixpence._ + +THE CONTINUITY OF SCRIPTURE, as declared by the Testimony of our Lord +and of the Evangelists and Apostles. By LORD HATHERLEY. + +LIFE OF LORD BACON. By LORD CAMPBELL. + +CHILDE HAROLD’S PILGRIMAGE. A Romaunt. By LORD BYRON. + +TALES AND POEMS. By LORD BYRON. + +LITTLE ARTHUR’S HISTORY OF ENGLAND. By LADY CALLCOTT. With 28 Woodcuts. + +THE EMIGRANT. By SIR FRANCIS HEAD. + +THE FABLES OF ÆSOP; A NEW VERSION, chiefly from ORIGINAL SOURCES. By +REV. THOMAS JAMES. With 100 Woodcuts. + +STORIES FOR CHILDREN SELECTED FROM THE HISTORY OF ENGLAND. By RT. HON. +J. W. CROKER. With 24 Woodcuts. + +THE COMPLETE POETICAL WORKS OF LORD BYRON. + +THE PHILOSOPHY OF THE MORAL FEELINGS. By JOHN ABERCROMBIE. + +POEMS AND SONGS. By ALLAN CUNNINGHAM. With Biographical Notice. + +THE FRANCHISE; a Privilege and not a Right. Proved by the Political +Experience of the Ancients. By H. S. TREMENHEERE. + + +_Three Shillings._ + +“THE FORTY-FIVE;” A NARRATIVE OF THE INSURRECTION OF 1745 IN SCOTLAND. +To which are added Letters of PRINCE CHARLES STUART. By LORD MAHON. + + +_Three Shillings and Sixpence._ + +THE BIBLE IN SPAIN; or, The Adventures and Imprisonment of an Englishman +in attempting to Circulate the Scriptures in the Peninsula. By GEORGE +BORROW. + +THE GYPSIES OF SPAIN; their Manners, Customs, Religion, and Language. By +GEORGE BORROW. + +THE BEAUTIES OF LORD BYRON’S WRITINGS; POETRY AND PROSE. With Portrait. + +LIFE OF LOUIS PRINCE OF CONDÉ, SURNAMED THE GREAT. By LORD MAHON. + +SKETCHES OF GERMAN LIFE AND SCENES FROM THE WAR OF LIBERATION IN +GERMANY. By VARNHAGEN VON ERESE. Translated by SIR ALEXANDER DUFF +GORDON. + +THE TRUE STORY OF THE BATTLE OF WATERLOO. By REV. G. R. GLEIG. + +DEEDS OF NAVAL DARING; or, ANECDOTES OF THE BRITISH NAVY. By EDWARD +GIFFARD. + +TYPEE; or, The Marquesas Islanders. By HERMAN MELVILLE. + +OMOO: A Narrative of Adventures in the South Seas; a Sequel to ‘Typee.’ +By HERMAN MELVILLE. + +THE BIBLE IN THE HOLY LAND; being Extracts from Dean Stanley’s ‘Sinai +and Palestine.’ With Woodcuts. + +AN ESSAY ON ENGLISH POETRY. With Short Notices of the British Poets. By +THOMAS CAMPBELL. + +HISTORICAL AND CRITICAL ESSAYS. By LORD MAHON. + +LIFE OF LORD CLIVE. By REV. G. R. GLEIG. + +LIFE OF SIR THOMAS MUNRO. With Selections from his Correspondence. By +REV. G. R. GLEIG. + +THE RISE OF OUR INDIAN EMPIRE: being a History of British India from its +Origin till the Peace of 1783. By LORD MAHON. + +A NARRATIVE OF THE SIEGE OF KARS, and of the Six Months’ Resistance by +the Turkish Garrison under General Williams. With Travels and Adventures +in Armenia, &c. By HUMPHREY SANDWITH, M.D. + +SKETCHES OF PERSIA. By SIR JOHN MALCOLM. + +THE WILD SPORTS AND NATURAL HISTORY OF THE HIGHLANDS. By CHARLES ST. +JOHN. + +GATHERINGS FROM SPAIN. By RICHARD FORD. + +TRAVELS IN MEXICO AND THE ROCKY MOUNTAINS. By G. F. RUXTON. + +PORTUGAL AND GALLICIA; WITH AN ACCOUNT OF THE SOCIAL AND POLITICAL STATE +OF THE BASQUE PROVINCES. By LORD CARNARVON. + +A RESIDENCE AT SIERRA LEONE. Described from a Journal kept on the Spot, +and Letters written to Friends at Home. By A LADY. + +THE REMAINS IN VERSE AND PROSE OF ARTHUR HENRY HALLAM. With Preface, +Memoir, and Portrait. + +THE POETICAL WORKS OF BISHOP HEBER; containing Palestine, Europe, the +Red Sea, Hymns, &c. With Portrait. + +GLEANINGS IN NATURAL HISTORY. By EDWARD JESSE. With Woodcuts. + +THE REJECTED ADDRESSES: or, The New Theatrum Poetarum. By HORACE and +JAMES SMITH. With Portrait and Woodcuts. + +CONSOLATIONS IN TRAVEL; or, The Last Days of a Philosopher. 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SMITH. + +SPECIMENS OF THE CHIEF WRITERS IN ENGLISH LITERATURE. Chronologically +arranged. Edited by DR. WM. SMITH. + +A SMALLER SCRIPTURE HISTORY.—I. Old Testament History; II. Connection +of Old and New Testaments; III. New Testament History to A.D. 70. Edited +by DR. WM. SMITH. With Woodcuts. + + +_Four Shillings._ + +HISTORY OF ENGLAND, from the FIRST INVASION by the ROMANS, continued +down to 1865. With CONVERSATIONS at the end of each CHAPTER. By MRS. +MARKHAM. With 100 Woodcuts. + +HISTORY OF FRANCE, from the CONQUEST by the GAULS, continued down to +1867. With CONVERSATIONS at the end of each CHAPTER. By MRS. MARKHAM. +With 70 Woodcuts. + +HISTORY OF GERMANY, from the INVASION of the KINGDOM by the ROMANS under +MARIUS, continued down to 1867. On the Plan of MRS. MARKHAM. With 50 +Woodcuts. + +SHALL AND WILL; or, the Future Auxiliary Verb. By SIR EDMUND HEAD. + + +_Four Shillings and Sixpence._ + +CHILDREN OF THE LAKE. A Poem. 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With Woodcuts. + +HANDBOOK OF FAMILIAR QUOTATIONS, chiefly from English Authors. + +THE CHACE—THE TURF—AND THE ROAD. A Series of Popular Essays. By C. J. +APPERLEY (NIMROD). With Portrait and Illustrations. + +AUNT IDA’S WALKS AND TALKS. A Story Book for Children. By A LADY. + +STORIES FOR DARLINGS. A Book for Boys and Girls. With Illustrations. + +THE CHARMED ROE. A Story Book for Young People. Illustrated by OTTO +SPECKTER. + +DON JUAN AND BEPPO. By LORD BYRON. 2 vols. + +LIFE IN THE LIGHT OF GOD’S WORD. By ARCHBISHOP THOMSON, D.D. + +ATHENS AND ATTICA; Notes of a Tour. By BISHOP WORDSWORTH, D.D. With +Illustrations. + +ANNALS OF THE WARS—XVIIITH CENTURY, 1700-1799. Compiled from the most +Authentic Sources. By SIR EDWARD CUST, D.C.L. With Maps. 5 vols. Post +8vo. 5s. each. + +ANNALS OF THE WARS—XIXTH CENTURY, 1800-15. Compiled from the most +Authentic Sources. By SIR EDWARD CUST. 4 vols. Fcap. 8vo. 5s. each. + + +_Six Shillings._ + +BENEDICITE; or, THE SONG OF THE THREE CHILDREN. Being ILLUSTRATIONS of +the POWER, BENEFICENCE, and DESIGN manifested by the CREATOR in HIS +WORKS. By DR. CHAPLIN CHILD. + +OLD DECCAN DAYS; or, HINDOO FAIRY LEGENDS current in Southern India. By +M. FRERE. With Introduction by SIR BARTLE FRERE. With Illustrations. + +THE WILD GARDEN; or, OUR GROVES AND SHRUBBERIES MADE BEAUTIFUL BY THE +NATURALIZATION OF HARDY EXOTIC PLANTS. By WILLIAM ROBINSON. With +Frontispiece. + +MISSIONARY TRAVELS AND RESEARCHES IN SOUTH AFRICA. By DAVID LIVINGSTONE, +M.D. With Map and Illustrations. + +FIVE YEARS OF A HUNTER’S LIFE IN SOUTH AFRICA; By GORDON CUMMING. With +Illustrations. + +THOUGHTS ON ANIMALCULES; or, The Invisible World, as revealed by the +Microscope. By GIDEON A. MANTELL. With Plates. + +LIVES OF BRINDLEY AND THE EARLY ENGINEERS. By SAMUEL SMILES. With +Woodcuts. + +LIFE OF TELFORD. With a History of Roads and Travelling in England. By +SAMUEL SMILES. With Woodcuts. + +LIVES OF GEORGE AND ROBERT STEPHENSON. By SAMUEL SMILES. 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You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: The Descent of Man and Selection in Relation to Sex, Vol. I (1st edition) + +Author: Charles Darwin + +Release Date: January 15, 2011 [EBook #34967] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK THE DESCENT OF MAN AND *** + + + + +Produced by StevenGibbs, Turgut Dincer and the Online +Distributed Proofreading Team at http://www.pgdp.net + + + + + + + + + +THE DESCENT OF MAN, + +AND + +SELECTION IN RELATION TO SEX. + + + + +BY CHARLES DARWIN, M.A., F.R.S., &c. + +IN TWO VOLUMES.--VOL. I. + +WITH ILLUSTRATIONS. + +LONDON: + +JOHN MURRAY, ALBEMARLE STREET. + +1871. + +[_The right of Translation is reserved._] + +BY THE SAME AUTHOR. + + + ON THE ORIGIN OF SPECIES BY MEANS OF NATURAL SELECTION; or, THE + PRESERVATION OF FAVOURED RACES IN THE STRUGGLE FOR LIFE; Fifth + Edition (_Tenth Thousand_), with Additions and Corrections. + 1869. MURRAY. + + THE VARIATION OF ANIMALS AND PLANTS UNDER DOMESTICATION. In two vols. + With Illustrations. 1868. MURRAY. + + ON THE VARIOUS CONTRIVANCES BY WHICH BRITISH AND FOREIGN ORCHIDS ARE + FERTILISED BY INSECTS; and on the GOOD EFFECTS OF CROSSING. With + numerous Woodcuts. MURRAY. + + A NATURALIST'S VOYAGE ROUND THE WORLD; or, A JOURNAL OF RESEARCHES INTO + THE NATURAL HISTORY AND GEOLOGY OF THE COUNTRIES visited during the + voyage of H.M.S. 'Beagle,' under the command of Captain FITZROY, R.N. + _Eleventh Thousand_. MURRAY. + + ON THE STRUCTURE AND DISTRIBUTION OF CORAL REEFS. SMITH, ELDER, & CO. + + GEOLOGICAL OBSERVATIONS ON VOLCANIC ISLANDS. SMITH, ELDER, & CO. + + GEOLOGICAL OBSERVATIONS ON SOUTH AMERICA. SMITH, ELDER & CO. + + A MONOGRAPH OF THE CIRRIPEDIA. With numerous Illustrations. 2 vols. + 8vo. HARDWICKE. + + ON THE MOVEMENTS AND HABITS OF CLIMBING PLANTS. With Woodcuts. WILLIAMS + & NORGATE. + + +LONDON: PRINTED BY WILLIAM CLOWES AND SONS, STAMFORD STREET, AND CHARING +CROSS. + + + + + CONTENTS. + + + INTRODUCTION Page 1-5 + + + PART I. + + ON THE DESCENT OF MAN. + + + CHAPTER I. + + THE EVIDENCE OF THE DESCENT OF MAN FROM SOME LOWER FORM. + + Nature of the evidence bearing on the origin of man--Homologous + structures in man and the lower animals--Miscellaneous points + of correspondence--Development--Rudimentary structures, + muscles, sense-organs, hair, bones, reproductive organs, + &c.--The bearing of these three great classes of facts on the + origin of man 9-33 + + + CHAPTER II. + + COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER ANIMALS. + + The difference in mental power between the highest ape and the + lowest savage, immense--Certain instincts in common--The + emotions--Curiosity--Imitation--Attention--Memory--Imagination-- + Reason--Progressive improvement--Tools and weapons used by + animals--Language--Self-consciousness--Sense of beauty--Belief + in God, spiritual agencies, superstitions 34-69 + + + CHAPTER III. + + COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER + ANIMALS--_continued_. + + The moral sense--Fundamental proposition--The qualities of + social animals--Origin of sociability--Struggle between opposed + instincts--Man and social animal--The more enduring social + instincts conquer other less persistent instincts--The social + virtues alone regarded by savages--The self-regarding virtues + acquired at a later stage of development--The importance of the + judgment of the members of the same community on + conduct--Transmission of moral tendencies--Summary 70-106 + + + CHAPTER IV. + + ON THE MANNER OF DEVELOPMENT OF MAN FROM SOME LOWER FORM. + + Variability of body and mind in man--Inheritance--Causes of + variability--Laws of variation the same in man as in the lower + animals--Direct action of the conditions of life--Effects of + the increased use and disuse of parts--Arrested + development--Reversion--Correlated variation--Rate of + increase--Checks to increase--Natural selection--Man the most + dominant animal in the world--Importance of his corporeal + structure--The causes which have led to his becoming + erect--Consequent changes of structure--Decrease in size of the + canine teeth--Increased size and altered shape of the + skull--Nakedness--Absence of a tail--Defenceless condition of + man 107-157 + + + CHAPTER V. + + ON THE DEVELOPMENT OF THE INTELLECTUAL AND MORAL FACULTIES + DURING PRIMEVAL AND CIVILISED TIMES. + + The advancement of the intellectual powers through natural + selection--Importance of imitation--Social and moral + faculties--Their development within the limits of the same + tribe--Natural selection as affecting civilised + nations--Evidence that civilised nations were once barbarous 158-184 + + + CHAPTER VI. + + ON THE AFFINITIES AND GENEALOGY OF MAN. + + Position of man in the animal series--The natural system + genealogical--Adaptive characters of slight value--Various + small points of resemblance between man and the + Quadrumana--Rank of man in the natural system--Birthplace and + antiquity of man--Absence of fossil connecting-links--Lower + stages in the genealogy of man, as inferred, firstly from his + affinities and secondly from his structure--Early androgynous + condition of the Vertebrata--Conclusion 185-213 + + + CHAPTER VII. + + ON THE RACES OF MAN. + + The nature and value of specific characters--Application to the + races of man--Arguments in favour of, and opposed to, ranking + the so-called races of man as distinct species--Sub-species-- + Monogenists and polygenists--Convergence of character--Numerous + points of resemblance in body and mind between the most + distinct races of man--The state of man when he first spread + over the earth--Each race not descended from a single + pair--The extinction of races--The formation of races-- + The effects of crossing--Slight influence of the direct + action of the conditions of life--Slight or no influence of + natural selection--Sexual selection. 214-250 + + + PART II. SEXUAL SELECTION. + + CHAPTER VIII. + + PRINCIPLES OF SEXUAL SELECTION. + + Secondary sexual characters--Sexual selection--Manner of + action--Excess of males--Polygamy--The male alone generally + modified through sexual selection--Eagerness of the + male--Variability of the male--Choice exerted by the + female--Sexual compared with natural selection--Inheritance at + corresponding periods of life, at corresponding seasons of the + year, and as limited by sex--Relations between the several + forms of inheritance--Causes why one sex and the young are not + modified through sexual selection--Supplement on the + proportional numbers of the two sexes throughout the animal + kingdom--On the limitation of the numbers of the two sexes + through natural selection 253-320 + + + CHAPTER IX. + + SECONDARY SEXUAL CHARACTERS IN THE LOWER CLASSES OF THE ANIMAL + KINGDOM. + + These characters absent in the lowest classes--Brilliant + colours--Mollusca--Annelids--Crustacea, secondary sexual + characters strongly developed; dimorphism; colour; characters + not acquired before maturity--Spiders, sexual colours of; + stridulation by the males--Myriapoda 321-340 + + + CHAPTER X. + + SECONDARY SEXUAL CHARACTERS OF INSECTS. + + Diversified structures possessed by the males for seizing the + females--Differences between the sexes, of which the meaning is + not understood--Difference in size between the + sexes--Thysanura--Diptera--Hemiptera--Homoptera, musical powers + possessed by the males alone--Orthoptera, musical instruments + of the males, much diversified in structure; pugnacity; + colours--Neuroptera, sexual differences in colour--Hymenoptera, + pugnacity and colours--Coleoptera, colours; furnished with + great horns, apparently as an ornament; battles; stridulating + organs generally common to both sexes 341-385 + + + CHAPTER XI. + + INSECTS, _continued_.--ORDER LEPIDOPTERA. + + Courtship of butterflies--Battles--Ticking noise--Colours + common to both sexes, or more brilliant in the + males--Examples--Not due to the direct action of the conditions + of life--Colours adapted for protection--Colours of + moths--Display--Perceptive powers of the + Lepidoptera--Variability--Causes of the difference in colour + between the males and females--Mimickry, female butterflies + more brilliantly coloured than the males--Bright colours of + caterpillars--Summary and concluding remarks on the secondary + sexual characters of insects--Birds and insects compared 386-423 + + + + +THE DESCENT OF MAN; + +AND ON + +SELECTION IN RELATION TO SEX. + + +INTRODUCTION. + +The nature of the following work will be best understood by a brief +account of how it came to be written. During many years I collected +notes on the origin or descent of man, without any intention of +publishing on the subject, but rather with the determination not to +publish, as I thought that I should thus only add to the prejudices +against my views. It seemed to me sufficient to indicate, in the first +edition of my 'Origin of Species,' that by this work "light would be +thrown on the origin of man and his history;" and this implies that man +must be included with other organic beings in any general conclusion +respecting his manner of appearance on this earth. Now the case wears a +wholly different aspect. When a naturalist like Carl Vogt ventures to +say in his address as President of the National Institution of Geneva +(1869), "personne, en Europe au moins, n'ose plus soutenir la cration +indpendante et de toutes pices, des espces," it is manifest that at +least a large number of naturalists must admit that species are the +modified descendants of other species; and this especially holds good +with the younger and rising naturalists. The greater number accept the +agency of natural selection; though some urge, whether with justice the +future must decide, that I have greatly overrated its importance. Of the +older and honoured chiefs in natural science, many unfortunately are +still opposed to evolution in every form. + +In consequence of the views now adopted by most naturalists, and which +will ultimately, as in every other case, be followed by other men, I +have been led to put together my notes, so as to see how far the general +conclusions arrived at in my former works were applicable to man. This +seemed all the more desirable as I had never deliberately applied these +views to a species taken singly. When we confine our attention to any +one form, we are deprived of the weighty arguments derived from the +nature of the affinities which connect together whole groups of +organisms--their geographical distribution in past and present times, +and their geological succession. The homological structure, +embryological development, and rudimentary organs of a species, whether +it be man or any other animal, to which our attention may be directed, +remain to be considered; but these great classes of facts afford, as it +appears to me, ample and conclusive evidence in favour of the principle +of gradual evolution. The strong support derived from the other +arguments should, however, always be kept before the mind. + +The sole object of this work is to consider, firstly, whether man, like +every other species, is descended from some pre-existing form; secondly, +the manner of his development; and thirdly, the value of the +differences between the so-called races of man. As I shall confine +myself to these points, it will not be necessary to describe in detail +the differences between the several races--an enormous subject which has +been fully discussed in many valuable works. The high antiquity of man +has recently been demonstrated by the labours of a host of eminent men, +beginning with M. Boucher de Perthes; and this is the indispensable +basis for understanding his origin. I shall, therefore, take this +conclusion for granted, and may refer my readers to the admirable +treatises of Sir Charles Lyell, Sir John Lubbock, and others. Nor shall +I have occasion to do more than to allude to the amount of difference +between man and the anthropomorphous apes; for Prof. Huxley, in the +opinion of most competent judges, has conclusively shewn that in every +single visible character man differs less from the higher apes than +these do from the lower members of the same order of Primates. + +This work contains hardly any original facts in regard to man; but as +the conclusions at which I arrived, after drawing up a rough draft, +appeared to me interesting, I thought that they might interest others. +It has often and confidently been asserted, that man's origin can never +be known: but ignorance more frequently begets confidence than does +knowledge: it is those who know little, and not those who know much, who +so positively assert that this or that problem will never be solved by +science. The conclusion that man is the co-descendant with other species +of some ancient, lower, and extinct form, is not in any degree new. +Lamarck long ago came to this conclusion, which has lately been +maintained by several eminent naturalists and philosophers; for instance +by Wallace, Huxley, Lyell, Vogt, Lubbock, Bchner, Rolle, &c.,[1] and +especially by Hckel. This last naturalist, besides his great work, +'Generelle Morphologie' (1866), has recently (1868, with a second edit. +in 1870), published his 'Natrliche Schpfungsgeschichte,' in which he +fully discusses the genealogy of man. If this work had appeared before +my essay had been written, I should probably never have completed it. +Almost all the conclusions at which I have arrived I find confirmed by +this naturalist, whose knowledge on many points is much fuller than +mine. Wherever I have added any fact or view from Prof. Hckel's +writings, I give his authority in the text, other statements I leave as +they originally stood in my manuscript, occasionally giving in the +foot-notes references to his works, as a confirmation of the more +doubtful or interesting points. + +During many years it has seemed to me highly probable that sexual +selection has played an important part in differentiating the races of +man; but in my 'Origin of Species' (first edition, p. 199) I contented +myself by merely alluding to this belief. When I came to apply this view +to man, I found it indispensable to treat the whole subject in full +detail.[2] Consequently the second part of the present work, treating of +sexual selection, has extended to an inordinate length, compared with +the first part; but this could not be avoided. + +I had intended adding to the present volumes an essay on the expression +of the various emotions by man and the lower animals. My attention was +called to this subject many years ago by Sir Charles Bell's admirable +work. This illustrious anatomist maintains that man is endowed with +certain muscles solely for the sake of expressing his emotions. As this +view is obviously opposed to the belief that man is descended from some +other and lower form, it was necessary for me to consider it. I likewise +wished to ascertain how far the emotions are expressed in the same +manner by the different races of man. But owing to the length of the +present work, I have thought it better to reserve my essay, which is +partially completed, for separate publication. + + + + +PART I. + +THE DESCENT OR ORIGIN OF MAN. + +PART I.--THE DESCENT OF MAN. + + + + +CHAPTER I. + +THE EVIDENCE OF THE DESCENT OF MAN FROM SOME LOWER FORM. + + + Nature of the evidence bearing on the origin of man--Homologous + structures in man and the lower animals--Miscellaneous points + of correspondence--Development--Rudimentary structures, + muscles, sense-organs, hair, bones, reproductive organs, + &c.--The bearing of these three great classes of facts on the + origin of man. + + +He who wishes to decide whether man is the modified descendant of some +pre-existing form, would probably first enquire whether man varies, +however slightly, in bodily structure and in mental faculties; and if +so, whether the variations are transmitted to his offspring in +accordance with the laws which prevail with the lower animals; such as +that of the transmission of characters to the same age or sex. Again, +are the variations the result, as far as our ignorance permits us to +judge, of the same general causes, and are they governed by the same +general laws, as in the case of other organisms; for instance by +correlation, the inherited effects of use and disuse, &c.? Is man +subject to similar malconformations, the result of arrested development, +of reduplication of parts, &c., and does he display in any of his +anomalies reversion to some former and ancient type of structure? It +might also naturally be enquired whether man, like so many other +animals, has given rise to varieties and sub-races, differing but +slightly from each other, or to races differing so much that they must +be classed as doubtful species? How are such races distributed over the +world; and how, when crossed, do they react on each other, both in the +first and succeeding generations? And so with many other points. + +The enquirer would next come to the important point, whether man tends +to increase at so rapid a rate, as to lead to occasional severe +struggles for existence, and consequently to beneficial variations, +whether in body or mind, being preserved, and injurious ones eliminated. +Do the races or species of men, whichever term may be applied, encroach +on and replace each other, so that some finally become extinct? We shall +see that all these questions, as indeed is obvious in respect to most of +them, must be answered in the affirmative, in the same manner as with +the lower animals. But the several considerations just referred to may +be conveniently deferred for a time; and we will first see how far the +bodily structure of man shows traces, more or less plain, of his descent +from some lower form. In the two succeeding chapters the mental powers +of man, in comparison with those of the lower animals, will be +considered. + + +_The Bodily Structure of Man._--It is notorious that man is constructed +on the same general type or model with other mammals. All the bones in +his skeleton can be compared with corresponding bones in a monkey, bat, +or seal. So it is with his muscles, nerves, blood-vessels and internal +viscera. The brain, the most important of all the organs, follows the +same law, as shewn by Huxley and other anatomists. Bischoff,[3] who is a +hostile witness, admits that every chief fissure and fold in the brain +of man has its analogy in that of the orang; but he adds that at no +period of development do their brains perfectly agree; nor could this be +expected, for otherwise their mental powers would have been the same. +Vulpian[4] remarks: "Les diffrences relles qui existent entre +l'encphale de l'homme et celui des singes suprieurs, sont bien +minimes. Il ne faut pas se faire d'illusions cet gard. L'homme est +bien plus prs des singes anthropomorphes par les caractres anatomiques +de son cerveau que ceux-ci ne le sont non-seulement des autres +mammifres, mais mmes de certains quadrumanes, des guenons et des +macaques." But it would be superfluous here to give further details on +the correspondence between man and the higher mammals in the structure +of the brain and all other parts of the body. + +It may, however, be worth while to specify a few points, not directly or +obviously connected with structure, by which this correspondence or +relationship is well shewn. + +Man is liable to receive from the lower animals, and to communicate to +them, certain diseases as hydrophobia, variola, the glanders, &c.; and +this fact proves the close similarity of their tissues and blood, both +in minute structure and composition, far more plainly than does their +comparison under the best microscope, or by the aid of the best chemical +analysis. Monkeys are liable to many of the same non-contagious diseases +as we are; thus Rengger,[5] who carefully observed for a long time the +_Cebus Azar_ in its native land, found it liable to catarrh, with the +usual symptoms, and which when often recurrent led to consumption. +These monkeys suffered also from apoplexy, inflammation of the bowels, +and cataract in the eye. The younger ones when shedding their milk-teeth +often died from fever. Medicines produced the same effect on them as on +us. Many kinds of monkeys have a strong taste for tea, coffee, and +spirituous liquors: they will also, as I have myself seen, smoke tobacco +with pleasure. Brehm asserts that the natives of north-eastern Africa +catch the wild baboons by exposing vessels with strong beer, by which +they are made drunk. He has seen some of these animals, which he kept in +confinement, in this state; and he gives a laughable account of their +behaviour and strange grimaces. On the following morning they were very +cross and dismal; they held their aching heads with both hands and wore +a most pitiable expression: when beer or wine was offered them, they +turned away with disgust, but relished the juice of lemons.[6] An +American monkey, an Ateles, after getting drunk on brandy, would never +touch it again, and thus was wiser than many men. These trifling facts +prove how similar the nerves of taste must be in monkeys and man, and +how similarly their whole nervous system is affected. + +Man is infested with internal parasites, sometimes causing fatal +effects, and is plagued by external parasites, all of which belong to +the same genera or families with those infesting other mammals. Man is +subject like other mammals, birds, and even insects, to that mysterious +law, which causes certain normal processes, such as gestation, as well +as the maturation and duration of various diseases, to follow lunar +periods.[7] His wounds are repaired by the same process of healing; and +the stumps left after the amputation of his limbs occasionally possess, +especially during an early embryonic period, some power of regeneration, +as in the lowest animals.[8] + +The whole process of that most important function, the reproduction of +the species, is strikingly the same in all mammals, from the first act +of courtship by the male[9] to the birth and nurturing of the young. +Monkeys are born in almost as helpless a condition as our own infants; +and in certain genera the young differ fully as much in appearance from +the adults, as do our children from their full-grown parents.[10] It has +been urged by some writers as an important distinction, that with man +the young arrive at maturity at a much later age than with any other +animal; but if we look to the races of mankind which inhabit tropical +countries the difference is not great, for the orang is believed not to +be adult till the age of from ten to fifteen years.[11] Man differs +from woman in size, bodily strength, hairyness, &c., as well as in mind, +in the same manner as do the two sexes of many mammals. It is, in short, +scarcely possible to exaggerate the close correspondence in general +structure, in the minute structure of the tissues, in chemical +composition and in constitution, between man and the higher animals, +especially the anthropomorphous apes. + + +_Embryonic Development._--Man is developed from an ovule, about the +125th of an inch in diameter, which differs in no respect from the +ovules of other animals. The embryo itself at a very early period can +hardly be distinguished from that of other members of the vertebrate +kingdom. At this period the arteries run in arch-like branches, as if to +carry the blood to branchi which are not present in the higher +vertebrata, though the slits on the sides of the neck still remain (_f_, +_g_, fig. 1), marking their former position. At a somewhat later period, +when the extremities are developed, "the feet of lizards and mammals," +as the illustrious Von Baer remarks, "the wings and feet of birds, no +less than the hands and feet of man, all arise from the same fundamental +form." It is, says Prof. Huxley,[12] "quite in the later stages of +development that the young human being presents marked differences from +the young ape, while the latter departs as much from the dog in its +developments, as the man does. Startling as this last assertion may +appear to be, it is demonstrably true." + +As some of my readers may never have seen a drawing of an embryo, I have +given one of man and another of a dog, at about the same early stage of +development, carefully copied from two works of undoubted accuracy.[13] + +[Illustration: Fig. 1. Upper figure human embryo, from Ecker. Lower +figure that of a dog, from Bischoff. + + _a._ Fore-brain, cerebral hemispheres, &c. + _b._ Mid-brain, corpora quadrigemina. + _c._ Hind-brain, cerebellum, medulla oblongata. + _d._ Eye. + _e._ Ear. + _f._ First visceral arch. + _g._ Second visceral arch. + H. Vertebral columns and muscles in process of development. + _i._ Anterior {~BOX DRAWINGS LIGHT DOWN AND LEFT~} + {~BOX DRAWINGS LIGHT VERTICAL~} extremities. + K. Posterior {~BOX DRAWINGS LIGHT UP AND LEFT~} + L. Tail or os coccyx.] + +After the foregoing statements made by such high authorities, it would +be superfluous on my part to give a number of borrowed details, shewing +that the embryo of man closely resembles that of other mammals. It may, +however, be added that the human embryo likewise resembles in various +points of structure certain low forms when adult. For instance, the +heart at first exists as a simple pulsating vessel; the excreta are +voided through a cloacal passage; and the os coccyx projects like a true +tail, "extending considerably beyond the rudimentary legs."[14] In the +embryos of all air-breathing vertebrates, certain glands called the +corpora Wolffiana, correspond with and act like the kidneys of mature +fishes.[15] Even at a later embryonic period, some striking resemblances +between man and the lower animals may be observed. Bischoff says that +the convolutions of the brain in a human foetus at the end of the +seventh month reach about the same stage of development as in a baboon +when adult.[16] The great toe, as Prof. Owen remarks,[17] "which forms +the fulcrum when standing or walking, is perhaps the most +characteristic peculiarity in the human structure;" but in an embryo, +about an inch in length, Prof. Wyman[18] found "that the great toe was +shorter than the others, and, instead of being parallel to them, +projected at an angle from the side of the foot, thus corresponding with +the permanent condition of this part in the quadrumana." I will conclude +with a quotation from Huxley,[19] who after asking, does man originate +in a different way from a dog, bird, frog or fish? says, "the reply is +not doubtful for a moment; without question, the mode of origin and the +early stages of the development of man are identical with those of the +animals immediately below him in the scale: without a doubt in these +respects, he is far nearer to apes, than the apes are to the dog." + + +_Rudiments._--This subject, though not intrinsically more important than +the two last, will for several reasons be here treated with more +fullness.[20] Not one of the higher animals can be named which does not +bear some part in a rudimentary condition; and man forms no exception to +the rule. Rudimentary organs must be distinguished from those that are +nascent; though in some cases the distinction is not easy. The former +are either absolutely useless, such as the mamm of male quadrupeds, or +the incisor teeth of ruminants which never cut through the gums; or they +are of such slight service to their present possessors, that we cannot +suppose that they were developed under the conditions which now exist. +Organs in this latter state are not strictly rudimentary, but they are +tending in this direction. Nascent organs, on the other hand, though not +fully developed, are of high service to their possessors, and are +capable of further development. Rudimentary organs are eminently +variable; and this is partly intelligible, as they are useless or nearly +useless, and consequently are no longer subjected to natural selection. +They often become wholly suppressed. When this occurs, they are +nevertheless liable to occasional reappearance through reversion; and +this is a circumstance well worthy of attention. + +Disuse at that period of life, when an organ is chiefly used, and this +is generally during maturity, together with inheritance at a +corresponding period of life, seem to have been the chief agents in +causing organs to become rudimentary. The term "disuse" does not relate +merely to the lessened action of muscles, but includes a diminished flow +of blood to a part or organ, from being subjected to fewer alternations +of pressure, or from becoming in any way less habitually active. +Rudiments, however, may occur in one sex of parts normally present in +the other sex; and such rudiments, as we shall hereafter see, have often +originated in a distinct manner. In some cases organs have been reduced +by means of natural selection, from having become injurious to the +species under changed habits of life. The process of reduction is +probably often aided through the two principles of compensation and +economy of growth; but the later stages of reduction, after disuse has +done all that can fairly be attributed to it, and when the saving to be +effected by the economy of growth would be very small,[21] are difficult +to understand. The final and complete suppression of a part, already +useless and much reduced in size, in which case neither compensation nor +economy can come into play, is perhaps intelligible by the aid of the +hypothesis of pangenesis, and apparently in no other way. But as the +whole subject of rudimentary organs has been fully discussed and +illustrated in my former works,[22] I need here say no more on this +head. + +Rudiments of various muscles have been observed in many parts of the +human body;[23] and not a few muscles, which are regularly present in +some of the lower animals can occasionally be detected in man in a +greatly reduced condition. Every one must have noticed the power which +many animals, especially horses, possess of moving or twitching their +skin; and this is effected by the panniculus carnosus. Remnants of this +muscle in an efficient state are found in various parts of our bodies; +for instance, on the forehead, by which the eyebrows are raised. The +_platysma myoides_, which is well developed on the neck, belongs to this +system, but cannot be voluntarily brought into action. Prof. Turner, of +Edinburgh, has occasionally detected, as he informs me, muscular +fasciculi in five different situations, namely in the axill, near the +scapul, &c., all of which must be referred to the system of the +panniculus. He has also shewn[24] that the _musculus sternalis_ or +_sternalis brutorum_, which is not an extension of the _rectus +abdominalis_, but is closely allied to the panniculus, occurred in the +proportion of about 3 per cent. in upwards of 600 bodies: he adds, that +this muscle affords "an excellent illustration of the statement that +occasional and rudimentary structures are especially liable to variation +in arrangement." + +Some few persons have the power of contracting the superficial muscles +on their scalps; and these muscles are in a variable and partially +rudimentary condition. M. A. de Candolle has communicated to me a +curious instance of the long-continued persistence or inheritance of +this power, as well as of its unusual development. He knows a family, in +which one member, the present head of a family, could, when a youth, +pitch several heavy books from his head by the movement of the scalp +alone; and he won wagers by performing this feat. His father, uncle, +grandfather, and all his three children possess the same power to the +same unusual degree. This family became divided eight generations ago +into two branches; so that the head of the above-mentioned branch is +cousin in the seventh degree to the head of the other branch. This +distant cousin resides in another part of France, and on being asked +whether he possessed the same faculty, immediately exhibited his power. +This case offers a good illustration how persistently an absolutely +useless faculty may be transmitted. + +The extrinsic muscles which serve to move the whole external ear, and +the intrinsic muscles which move the different parts, all of which +belong to the system of the panniculus, are in a rudimentary condition +in man; they are also variable in development, or at least in function. +I have seen one man who could draw his ears forwards, and another who +could draw them backwards;[25] and from what one of these persons told +me, it is probable that most of us by often touching our ears and thus +directing our attention towards them, could by repeated trials recover +some power of movement. The faculty of erecting the ears and of +directing them to different points of the compass, is no doubt of the +highest service to many animals, as they thus perceive the point of +danger; but I have never heard of a man who possessed the least power of +erecting his ears,--the one movement which might be of use to him. The +whole external shell of the ear may be considered a rudiment, together +with the various folds and prominences (helix and anti-helix, tragus and +anti-tragus, &c.) which in the lower animals strengthen and support the +ear when erect, without adding much to its weight. Some authors, +however, suppose that the cartilage of the shell serves to transmit +vibrations to the acoustic nerve; but Mr. Toynbee,[26] after collecting +all the known evidence on this head, concludes that the external shell +is of no distinct use. The ears of the chimpanzee and orang are +curiously like those of man, and I am assured by the keepers in the +Zoological Gardens that these animals never move or erect them; so that +they are in an equally rudimentary condition, as far as function is +concerned, as in man. Why these animals, as well as the progenitors of +man, should have lost the power of erecting their ears we cannot say. It +may be, though I am not quite satisfied with this view, that owing to +their arboreal habits and great strength they were but little exposed to +danger, and so during a lengthened period moved their ears but little, +and thus gradually lost the power of moving them. This would be a +parallel case with that of those large and heavy birds, which from +inhabiting oceanic islands have not been exposed to the attacks of +beasts of prey, and have consequently lost the power of using their +wings for flight. + +[Illustration: Fig. 2. Human Ear, modelled and drawn by Mr. Woolner. + +_a._ The projecting point.] + +The celebrated sculptor, Mr. Woolner, informs me of one little +peculiarity in the external ear, which he has often observed both in men +and women, and of which he perceived the full signification. His +attention was first called to the subject whilst at work on his figure +of Puck, to which he had given pointed ears. He was thus led to examine +the ears of various monkeys, and subsequently more carefully those of +man. The peculiarity consists in a little blunt point, projecting from +the inwardly folded margin, or helix. Mr. Woolner made an exact model of +one such case, and has sent me the accompanying drawing. (Fig. 2.) These +points not only project inwards, but often a little outwards, so that +they are visible when the head is viewed from directly in front or +behind. They are variable in size and somewhat in position, standing +either a little higher or lower; and they sometimes occur on one ear and +not on the other. Now the meaning of these projections is not, I think, +doubtful; but it may be thought that they offer too trifling a character +to be worth notice. This thought, however, is as false as it is natural. +Every character, however slight, must be the result of some definite +cause; and if it occurs in many individuals deserves consideration. The +helix obviously consists of the extreme margin of the ear folded +inwards; and this folding appears to be in some manner connected with +the whole external ear being permanently pressed backwards. In many +monkeys, which do not stand high in the order, as baboons and some +species of macacus,[27] the upper portion of the ear is slightly +pointed, and the margin is not at all folded inwards; but if the margin +were to be thus folded, a slight point would necessarily project inwards +and probably a little outwards. This could actually be observed in a +specimen of the _Ateles beelzebuth_ in the Zoological Gardens; and we +may safely conclude that it is a similar structure--a vestige of +formerly pointed ears--which occasionally reappears in man. + +The nictitating membrane, or third eyelid, with its accessory muscles +and other structures, is especially well developed in birds, and is of +much functional importance to them, as it can be rapidly drawn across +the whole eye-ball. It is found in some reptiles and amphibians, and in +certain fishes, as in sharks. It is fairly well developed in the two +lower divisions of the mammalian series, namely, in the monotremata and +marsupials, and in some few of the higher mammals, as in the walrus. But +in man, the quadrumana, and most other mammals, it exists, as is +admitted by all anatomists, as a mere rudiment, called the semilunar +fold.[28] + +The sense of smell is of the highest importance to the greater number of +mammals--to some, as the ruminants, in warning them of danger; to +others, as the carnivora, in finding their prey; to others, as the wild +boar, for both purposes combined. But the sense of smell is of extremely +slight service, if any, even to savages, in whom it is generally more +highly developed than in the civilised races. It does not warn them of +danger, nor guide them to their food; nor does it prevent the Esquimaux +from sleeping in the most fetid atmosphere, nor many savages from eating +half-putrid meat. Those who believe in the principle of gradual +evolution, will not readily admit that this sense in its present state +was originally acquired by man, as he now exists. No doubt he inherits +the power in _an_ enfeebled and so far rudimentary condition, from some +early progenitor, to whom it was highly serviceable and by whom it was +continually used. We can thus perhaps understand how it is, as Dr. +Maudsley has truly remarked,[29] that the sense of smell in man "is +singularly effective in recalling vividly the ideas and images of +forgotten scenes and places;" for we see in those animals, which have +this sense highly developed, such as dogs and horses, that old +recollections of persons and places are strongly associated with their +odour. + +Man differs conspicuously from all the other Primates in being almost +naked. But a few short straggling hairs are found over the greater part +of the body in the male sex, and fine down on that of the female sex, In +individuals belonging to the same race these hairs are highly variable, +not only in abundance, but likewise in position: thus the shoulders in +some Europeans are quite naked, whilst in others they bear thick tufts +of hair.[30] There can be little doubt that the hairs thus scattered +over the body are the rudiments of the uniform hairy coat of the lower +animals. This view is rendered all the more probable, as it is known +that fine, short, and pale-coloured hairs on the limbs and other parts +of the body occasionally become developed into "thickset, long, and +rather coarse dark hairs," when abnormally nourished near old-standing +inflamed surfaces.[31] + +I am informed by Mr. Paget that persons belonging to the same family +often have a few hairs in their eyebrows much longer than the others; so +that this slight peculiarity seems to be inherited. These hairs +apparently represent the vibriss, which are used as organs of touch by +many of the lower animals. In a young chimpanzee I observed that a few +upright, rather long, hairs, projected above the eyes, where the true +eyebrows, if present, would have stood. + +The fine wool-like hair, or so-called lanugo, with which the human +foetus during the sixth month is thickly covered, offers a more +curious case. It is first developed, during the fifth month, on the +eyebrows and face, and especially round the mouth, where it is much +longer than that on the head. A moustache of this kind was observed by +Eschricht[32] on a female foetus; but this is not so surprising a +circumstance as it may at first appear, for the two sexes generally +resemble each other in all external characters during an early period of +growth. The direction and arrangement of the hairs on all parts of the +foetal body are the same as in the adult, but are subject to much +variability. The whole surface, including even the forehead and ears, is +thus thickly clothed; but it is a significant fact that the palms of the +hands and the soles of the feet are quite naked, like the inferior +surfaces of all four extremities in most of the lower animals. As this +can hardly be an accidental coincidence, we must consider the woolly +covering of the foetus to be the rudimental representative of the +first permanent coat of hair in those mammals which are born hairy. This +representation is much more complete, in accordance with the usual law +of embryological development, than that afforded by the straggling hairs +on the body of the adult. + +It appears as if the posterior molar or wisdom-teeth were tending to +become rudimentary in the more civilised races of man. These teeth are +rather smaller than the other molars, as is likewise the case with the +corresponding teeth in the chimpanzee and orang; and they have only two +separate fangs. They do not cut through the gums till about the +seventeenth year, and I am assured by dentists that they are much more +liable to decay, and are earlier lost, than the other teeth. It is also +remarkable that they are much more liable to vary both in structure and +in the period of their development than the other teeth.[33] In the +Melanian races, on the other hand, the wisdom-teeth are usually +furnished with three separate fangs, and are generally sound: they also +differ from the other molars in size less than in the Caucasian +races.[34] Prof. Schaaffhausen accounts for this difference between the +races by "the posterior dental portion of the jaw being always +shortened" in those that are civilised,[35] and this shortening may, I +presume, be safely attributed to civilised men habitually feeding on +soft, cooked food, and thus using their jaws less. I am informed by Mr. +Brace that it is becoming quite a common practice in the United States +to remove some of the molar teeth of children, as the jaw does not grow +large enough for the perfect development of the normal number. + +With respect to the alimentary canal I have met with an account of only +a single rudiment, namely the vermiform appendage of the ccum. The +ccum is a branch or diverticulum of the intestine, ending in a +cul-de-sac, and it is extremely long in many of the lower +vegetable-feeding mammals. In the marsupial koala it is actually more +than thrice as long as the whole body.[36] It is sometimes produced into +a long gradually-tapering point, and is sometimes constricted in parts. +It appears as if, in consequence of changed diet or habits, the ccum +had become much shortened in various animals, the vermiform appendage +being left as a rudiment of the shortened part. That this appendage is a +rudiment, we may infer from its small size, and from the evidence which +Prof. Canestrini[37] has collected of its variability in man. It is +occasionally quite absent, or again is largely developed. The passage is +sometimes completely closed for half or two-thirds of its length, with +the terminal part consisting of a flattened solid expansion. In the +orang this appendage is long and convoluted: in man it arises from the +end of the short ccum, and is commonly from four to five inches in +length, being only about the third of an inch in diameter. Not only is +it useless, but it is sometimes the cause of death, of which fact I have +lately heard two instances: this is due to small hard bodies, such as +seeds, entering the passage and causing inflammation.[38] + +In the Quadrumana and some other orders of mammals, especially in the +Carnivora, there is a passage near the lower end of the humerus, called +the supra-condyloid foramen, through which the great nerve of the fore +limb passes, and often the great artery. Now in the humerus of man, as +Dr. Struthers[39] and others have shewn, there is generally a trace of +this passage, and it is sometimes fairly well developed, being formed by +a depending hook-like process of bone, completed by a band of ligament. +When present the great nerve invariably passes through it, and this +clearly indicates that it is the homologue and rudiment of the +supra-condyloid foramen of the lower animals. Prof. Turner estimates, as +he informs me, that it occurs in about one per cent. of recent +skeletons; but during ancient times it appears to have been much more +common. Mr. Busk[40] has collected the following evidence on this head: +Prof. Broca "noticed the perforation in four and a half per cent. of the +arm-bones collected in the 'Cimetire du Sud' at Paris; and in the +Grotto of Orrony, the contents of which are referred to the Bronze +period, as many as eight humeri out of thirty-two were perforated; but +this extraordinary proportion, he thinks, might be due to the cavern +having been a sort of 'family vault.' Again, M. Dupont found 30 per +cent. of perforated bones in the caves of the Valley of the Lesse, +belonging to the Reindeer period; whilst M. Leguay, in a sort of +_dolmen_ at Argenteuil, observed twenty-five per cent. to be perforated; +and M. Pruner-Bey found twenty-six per cent. in the same condition in +bones from Vaural. Nor should it be left unnoticed that M. Pruner-Bey +states that this condition is common in Guanche skeletons." The fact +that ancient races, in this and several other cases, more frequently +present structures which resemble those of the lower animals than do the +modern races, is interesting. One chief cause seems to be that ancient +races stand somewhat nearer than modern races in the long line of +descent to their remote animal-like progenitors. + +The os coccyx in man, though functionless as a tail, plainly represents +this part in other vertebrate animals. At an early embryonic period it +is free, and, as we have seen, projects beyond the lower extremities. In +certain rare and anomalous cases it has been known, according to Isidore +Geoffroy St.-Hilaire and others,[41] to form a small external rudiment +of a tail. The os coccyx is short, usually including only four vertebr: +and these are in a rudimental condition, for they consist, with the +exception of the basal one, of the centrum alone.[42] They are furnished +with some small muscles; one of which, as I am informed by Prof. Turner, +has been expressly described by Theile as a rudimentary repetition of +the extensor of the tail, which is so largely developed in many mammals. + +The spinal cord in man extends only as far downwards as the last dorsal +or first lumbar vertebra; but a thread-like structure (the _filum +terminale_) runs down the axis of the sacral part of the spinal canal, +and even along the back of the coccygeal bones. The upper part of this +filament, as Prof. Turner informs me, is undoubtedly homologous with the +spinal cord; but the lower part apparently consists merely of the _pia +mater_, or vascular investing membrane. Even in this case the os coccyx +may be said to possess a vestige of so important a structure as the +spinal cord, though no longer enclosed within a bony canal. The +following fact, for which I am also indebted to Prof. Turner, shews how +closely the os coccyx corresponds with the true tail in the lower +animals: Luschka has recently discovered at the extremity of the +coccygeal bones a very peculiar convoluted body, which is continuous +with the middle sacral artery; and this discovery led Krause and Meyer +to examine the tail of a monkey (Macacus) and of a cat, in both of which +they found, though not at the extremity, a similarly convoluted body. + +The reproductive system offers various rudimentary structures; but these +differ in one important respect from the foregoing cases. We are not +here concerned with a vestige of a part which does not belong to the +species in an efficient state; but with a part which is always present +and efficient in the one sex, being represented in the other by a mere +rudiment. Nevertheless, the occurrence of such rudiments is as difficult +to explain on the belief of the separate creation of each species, as in +the foregoing cases. Hereafter I shall have to recur to these rudiments, +and shall shew that their presence generally depends merely on +inheritance; namely, on parts acquired by one sex having been partially +transmitted to the other. Here I will only give some instances of such +rudiments. It is well known that in the males of all mammals, including +man, rudimentary mamm exist. These in several instances have become +well developed, and have yielded a copious supply of milk. Their +essential identity in the two sexes is likewise shewn by their +occasional sympathetic enlargement in both during an attack of the +measles. The _vesicula prostatica_, which has been observed in many +male mammals, is now universally acknowledged to be the homologue of the +female uterus, together with the connected passage. It is impossible to +read Leuckart's able description of this organ, and his reasoning, +without admitting the justness of his conclusion. This is especially +clear in the case of those mammals in which the true female uterus +bifurcates, for in the males of these the vesicula likewise +bifurcates.[43] Some additional rudimentary structures belonging to the +reproductive system might here have been adduced.[44] + + +The bearing of the three great classes of facts now given is +unmistakeable. But it would be superfluous here fully to recapitulate +the line of argument given in detail in my 'Origin of Species.' The +homological construction of the whole frame in the members of the same +class is intelligible, if we admit their descent from a common +progenitor, together with their subsequent adaptation to diversified +conditions. On any other view the similarity of pattern between the hand +of a man or monkey, the foot of a horse, the flipper of a seal, the wing +of a bat, &c., is utterly inexplicable. It is no scientific explanation +to assert that they have all been formed on the same ideal plan. With +respect to development, we can clearly understand, on the principle of +variations supervening at a rather late embryonic period, and being +inherited at a corresponding period, how it is that the embryos of +wonderfully different forms should still retain, more or less perfectly, +the structure of their common progenitor. No other explanation has ever +been given of the marvellous fact that the embryo of a man, dog, seal, +bat, reptile, &c., can at first hardly be distinguished from each other. +In order to understand the existence of rudimentary organs, we have only +to suppose that a former progenitor possessed the parts in question in a +perfect state, and that under changed habits of life they became greatly +reduced, either from simple disuse, or through the natural selection of +those individuals which were least encumbered with a superfluous part, +aided by the other means previously indicated. + +Thus we can understand how it has come to pass that man and all other +vertebrate animals have been constructed on the same general model, why +they pass through the same early stages of development, and why they +retain certain rudiments in common. Consequently we ought frankly to +admit their community of descent: to take any other view, is to admit +that our own structure and that of all the animals around us, is a mere +snare laid to entrap our judgment. This conclusion is greatly +strengthened, if we look to the members of the whole animal series, and +consider the evidence derived from their affinities or classification, +their geographical distribution and geological succession. It is only +our natural prejudice, and that arrogance which made our forefathers +declare that they were descended from demi-gods, which leads us to demur +to this conclusion. But the time will before long come when it will be +thought wonderful, that naturalists, who were well acquainted with the +comparative structure and development of man and other mammals, should +have believed that each was the work of a separate act of creation. + + + + +CHAPTER II. + +COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER ANIMALS. + + + The difference in mental power between the highest ape and the + lowest savage, immense--Certain instincts in common--The + emotions--Curiosity--Imitation--Attention--Memory--Imagination + --Reason--Progressive improvement--Tools and weapons used by + animals--Language--Self-consciousness--Sense of beauty--Belief + in God, spiritual agencies, superstitions. + + +We have seen in the last chapter that man bears in his bodily structure +clear traces of his descent from some lower form; but it may be urged +that, as man differs so greatly in his mental power from all other +animals, there must be some error in this conclusion. No doubt the +difference in this respect is enormous, even if we compare the mind of +one of the lowest savages, who has no words to express any number higher +than four, and who uses no abstract terms for the commonest objects or +affections,[45] with that of the most highly organised ape. The +difference would, no doubt, still remain immense, even if one of the +higher apes had been improved or civilised as much as a dog has been in +comparison with its parent-form, the wolf or jackal. The Fuegians rank +amongst the lowest barbarians; but I was continually struck with +surprise how closely the three natives on board H.M.S. "Beagle," who had +lived some years in England and could talk a little English, resembled +us in disposition and in most of our mental faculties. If no organic +being excepting man had possessed any mental power, or if his powers had +been of a wholly different nature from those of the lower animals, then +we should never have been able to convince ourselves that our high +faculties had been gradually developed. But it can be clearly shewn that +there is no fundamental difference of this kind. We must also admit that +there is a much wider interval in mental power between one of the lowest +fishes, as a lamprey or lancelet, and one of the higher apes, than +between an ape and man; yet this immense interval is filled up by +numberless gradations. + +Nor is the difference slight in moral disposition between a barbarian, +such as the man described by the old navigator Byron, who dashed his +child on the rocks for dropping a basket of sea-urchins, and a Howard or +Clarkson; and in intellect, between a savage who does not use any +abstract terms, and a Newton or Shakspeare. Differences of this kind +between the highest men of the highest races and the lowest savages, are +connected by the finest gradations. Therefore it is possible that they +might pass and be developed into each other. + +My object in this chapter is solely to shew that there is no fundamental +difference between man and the higher mammals in their mental faculties. +Each division of the subject might have been extended into a separate +essay, but must here be treated briefly. As no classification of the +mental powers has been universally accepted, I shall arrange my remarks +in the order most convenient for my purpose; and will select those facts +which have most struck me, with the hope that they may produce some +effect on the reader. + +With respect to animals very low in the scale, I shall have to give some +additional facts under Sexual Selection, shewing that their mental +powers are higher than might have been expected. The variability of the +faculties in the individuals of the same species is an important point +for us, and some few illustrations will here be given. But it would be +superfluous to enter into many details on this head, for I have found on +frequent enquiry, that it is the unanimous opinion of all those who have +long attended to animals of many kinds, including birds, that the +individuals differ greatly in every mental characteristic. In what +manner the mental powers were first developed in the lowest organisms, +is as hopeless an enquiry as how life first originated. These are +problems for the distant future, if they are ever to be solved by man. + +As man possesses the same senses with the lower animals, his fundamental +intuitions must be the same. Man has also some few instincts in common, +as that of self-preservation, sexual love, the love of the mother for +her new-born offspring, the power possessed by the latter of sucking, +and so forth. But man, perhaps, has somewhat fewer instincts than those +possessed by the animals which come next to him in the series. The orang +in the Eastern islands, and the chimpanzee in Africa, build platforms on +which they sleep; and, as both species follow the same habit, it might +be argued that this was due to instinct, but we cannot feel sure that it +is not the result of both animals having similar wants and possessing +similar powers of reasoning. These apes, as we may assume, avoid the +many poisonous fruits of the tropics, and man has no such knowledge; but +as our domestic animals, when taken to foreign lands and when first +turned out in the spring, often eat poisonous herbs, which they +afterwards avoid, we cannot feel sure that the apes do not learn from +their own experience or from that of their parents what fruits to +select. It is however certain, as we shall presently see, that apes +have an instinctive dread of serpents, and probably of other dangerous +animals. + +The fewness and the comparative simplicity of the instincts in the +higher animals are remarkable in contrast with those of the lower +animals. Cuvier maintained that instinct and intelligence stand in an +inverse ratio to each other; and some have thought that the intellectual +faculties of the higher animals have been gradually developed from their +instincts. But Pouchet, in an interesting essay,[46] has shewn that no +such inverse ratio really exists. Those insects which possess the most +wonderful instincts are certainly the most intelligent. In the +vertebrate series, the least intelligent members, namely fishes and +amphibians, do not possess complex instincts; and amongst mammals the +animal most remarkable for its instincts, namely the beaver, is highly +intelligent, as will be admitted by every one who has read Mr. Morgan's +excellent account of this animal.[47] + +Although the first dawnings of intelligence, according to Mr. Herbert +Spencer,[48] have been developed through the multiplication and +co-ordination of reflex actions, and although many of the simpler +instincts graduate into actions of this kind and can hardly be +distinguished from them, as in the case of young animals sucking, yet +the more complex instincts seem to have originated independently of +intelligence. I am, however, far from wishing to deny that instinctive +actions may lose their fixed and untaught character, and be replaced by +others performed by the aid of the free will. On the other hand, some +intelligent actions--as when, birds on oceanic islands first learn to +avoid man--after being performed during many generations, become +converted into instincts and are inherited. They may then be said to be +degraded in character, for they are no longer performed through reason +or from experience. But the greater number of the more complex instincts +appear to have been gained in a wholly different manner, through the +natural selection of variations of simpler instinctive actions. Such +variations appear to arise from the same unknown causes acting on the +cerebral organisation, which induce slight variations or individual +differences in other parts of the body; and these variations, owing to +our ignorance, are often said to arise spontaneously. We can, I think, +come to no other conclusion with respect to the origin of the more +complex instincts, when we reflect on the marvellous instincts of +sterile worker-ants and bees, which leave no offspring to inherit the +effects of experience and of modified habits. + +Although a high degree of intelligence is certainly compatible with the +existence of complex instincts, as we see in the insects just named and +in the beaver, it is not improbable that they may to a certain extent +interfere with each other's development. Little is known about the +functions of the brain, but we can perceive that as the intellectual +powers become highly developed, the various parts of the brain must be +connected by the most intricate channels of intercommunication; and as a +consequence each separate part would perhaps tend to become less well +fitted to answer in a definite and uniform, that is instinctive, manner +to particular sensations or associations. + +I have thought this digression worth giving, because we may easily +underrate the mental powers of the higher animals, and especially of +man, when we compare their actions founded on the memory of past events, +on foresight, reason, and imagination, with exactly similar actions +instinctively performed by the lower animals; in this latter case the +capacity of performing such actions having been gained, step by step, +through the variability of the mental organs and natural selection, +without any conscious intelligence on the part of the animal during each +successive generation. No doubt, as Mr. Wallace has argued,[49] much of +the intelligent work done by man is due to imitation and not to reason; +but there is this great difference between his actions and many of those +performed by the lower animals, namely, that man cannot, on his first +trial, make, for instance, a stone hatchet or a canoe, through his power +of imitation. He has to learn his work by practice; a beaver, on the +other hand, can make its dam or canal, and a bird its nest, as well, or +nearly as well, the first time it tries, as when old and experienced. + +To return to our immediate subject: the lower animals, like man, +manifestly feel pleasure and pain, happiness and misery. Happiness is +never better exhibited than by young animals, such as puppies, kittens, +lambs, &c., when playing together, like our own children. Even insects +play together, as has been described by that excellent observer, P. +Huber,[50] who saw ants chasing and pretending to bite each other, like +so many puppies. + +The fact that the lower animals are excited by the same emotions as +ourselves is so well established, that it will not be necessary to weary +the reader by many details. Terror acts in the same manner on them as on +us, causing the muscles to tremble, the heart to palpitate, the +sphincters to be relaxed, and the hair to stand on end. Suspicion, the +offspring of fear, is eminently characteristic of most wild animals. +Courage and timidity are extremely variable qualities in the +individuals of the same species, as is plainly seen in our dogs. Some +dogs and horses are ill-tempered and easily turn sulky; others are +good-tempered; and these qualities are certainly inherited. Every one +knows how liable animals are to furious rage, and how plainly they show +it. Many anecdotes, probably true, have been published on the +long-delayed and artful revenge of various animals. The accurate Rengger +and Brehm[51] state that the American and African monkeys which they +kept tame, certainly revenged themselves. The love of a dog for his +master is notorious; in the agony of death he has been known to caress +his master, and every one has heard of the dog suffering under +vivisection, who licked the hand of the operator; this man, unless he +had a heart of stone, must have felt remorse to the last hour of his +life. As Whewell[52] has remarked, "who that reads the touching +instances of maternal affection, related so often of the women of all +nations, and of the females of all animals, can doubt that the principle +of action is the same in the two cases?" + +We see maternal affection exhibited in the most trifling details; thus +Rengger observed an American monkey (a Cebus) carefully driving away the +flies which plagued her infant; and Duvaucel saw a Hylobates washing the +faces of her young ones in a stream. So intense is the grief of female +monkeys for the loss of their young, that it invariably caused the death +of certain kinds kept under confinement by Brehm in N. Africa. +Orphan-monkeys were always adopted and carefully guarded by the other +monkeys, both males and females. One female baboon had so capacious a +heart that she not only adopted young monkeys of other species, but +stole young dogs and cats, which she continually carried about. Her +kindness, however, did not go so far as to share her food with her +adopted offspring, at which Brehm was surprised, as his monkeys always +divided everything quite fairly with their own young ones. An adopted +kitten scratched the above-mentioned affectionate baboon, who certainly +had a fine intellect, for she was much astonished at being scratched, +and immediately examined the kitten's feet, and without more ado bit off +the claws. In the Zoological Gardens, I heard from the keeper that an +old baboon (_C. chacma_) had adopted a Rhesus monkey; but when a young +drill and mandrill were placed in the cage, she seemed to perceive that +these monkeys, though distinct species, were her nearer relatives, for +she at once rejected the Rhesus and adopted both of them. The young +Rhesus, as I saw, was greatly discontented at being thus rejected, and +it would, like a naughty child, annoy and attack the young drill and +mandrill whenever it could do so with safety; this conduct exciting +great indignation in the old baboon. Monkeys will also, according to +Brehm, defend their master when attacked by any one, as well as dogs to +whom they are attached, from the attacks of other dogs. But we here +trench on the subject of sympathy, to which I shall recur. Some of +Brehm's monkeys took much delight in teasing, in various ingenious ways, +a certain old dog whom they disliked, as well as other animals. + +Most of the more complex emotions are common to the higher animals and +ourselves. Every one has seen how jealous a dog is of his master's +affection, if lavished on any other creature; and I have observed the +same fact with monkeys. This shews that animals not only love, but have +the desire to be loved. Animals manifestly feel emulation. They love +approbation or praise; and a dog carrying a basket for his master +exhibits in a high degree self-complacency or pride. There can, I think, +be no doubt that a dog feels shame, as distinct from fear, and something +very like modesty when begging too often for food. A great dog scorns +the snarling of a little dog, and this may be called magnanimity. +Several observers have stated that monkeys certainly dislike being +laughed at; and they sometimes invent imaginary offences. In the +Zoological Gardens I saw a baboon who always got into a furious rage +when his keeper took out a letter or book and read it aloud to him; and +his rage was so violent that, as I witnessed on one occasion, he bit his +own leg till the blood flowed. + +We will now turn to the more intellectual emotions and faculties, which +are very important, as forming the basis for the development of the +higher mental powers. Animals manifestly enjoy excitement and suffer +from ennui, as may be seen with dogs, and, according to Rengger, with +monkeys. All animals feel Wonder, and many exhibit Curiosity. They +sometimes suffer from this latter quality, as when the hunter plays +antics and thus attracts them; I have witnessed this with deer, and so +it is with the wary chamois, and with some kinds of wild-ducks. Brehm +gives a curious account of the instinctive dread which his monkeys +exhibited towards snakes; but their curiosity was so great that they +could not desist from occasionally satiating their horror in a most +human fashion, by lifting up the lid of the box in which the snakes were +kept. I was so much surprised at his account, that I took a stuffed and +coiled-up snake into the monkey-house at the Zoological Gardens, and the +excitement thus caused was one of the most curious spectacles which I +ever beheld. Three species of Cercopithecus were the most alarmed; they +dashed about their cages and uttered sharp signal-cries of danger, which +were understood by the other monkeys. A few young monkeys and one old +Anubis baboon alone took no notice of the snake. I then placed the +stuffed specimen on the ground in one of the larger compartments. After +a time all the monkeys collected round it in a large circle, and staring +intently, presented a most ludicrous appearance. They became extremely +nervous; so that when a wooden ball, with which they were familiar as a +plaything, was accidently moved in the straw, under which it was partly +hidden, they all instantly started away. These monkeys behaved very +differently when a dead fish, a mouse, and some other new objects were +placed in their cages; for though at first frightened, they soon +approached, handled and examined them. I then placed a live snake in a +paper bag, with the mouth loosely closed, in one of the larger +compartments. One of the monkeys immediately approached, cautiously +opened the bag a little, peeped in, and instantly dashed away. Then I +witnessed what Brehm has described, for monkey after monkey, with head +raised high and turned on one side, could not resist taking momentary +peeps into the upright bag, at the dreadful object lying quiet at the +bottom. It would almost appear as if monkeys had some notion of +zoological affinities, for those kept by Brehm exhibited a strange, +though mistaken, instinctive dread of innocent lizards and frogs. An +orang, also, has been known to be much alarmed at the first sight of a +turtle.[53] + +The principle of _Imitation_ is strong in man, and especially in man in +a barbarous state. Desor[54] has remarked that no animal voluntarily +imitates an action performed by man, until in the ascending scale we +come to monkeys, which are well-known to be ridiculous mockers. Animals, +however, sometimes imitate each others' actions: thus two species of +wolves, which had been reared by dogs, learned to bark, as does +sometimes the jackal,[55] but whether this can be called voluntary +imitation is another question. From one account which I have read, there +is reason to believe that puppies nursed by cats sometimes learn to lick +their feet and thus to clean their faces: it is at least certain, as I +hear from a perfectly trustworthy friend, that some dogs behave in this +manner. Birds imitate the songs of their parents, and sometimes those of +other birds; and parrots are notorious imitators of any sound which they +often hear. + +Hardly any faculty is more important for the intellectual progress of +man than the power of _Attention_. Animals clearly manifest this power, +as when a cat watches by a hole and prepares to spring on its prey. Wild +animals sometimes become so absorbed when thus engaged, that they may be +easily approached. Mr. Bartlett has given me a curious proof how +variable this faculty is in monkeys. A man who trains monkeys to act +used to purchase common kinds from the Zoological Society at the price +of five pounds for each; but he offered to give double the price, if he +might keep three or four of them for a few days, in order to select one. +When asked how he could possibly so soon learn whether a particular +monkey would turn out a good actor, he answered that it all depended on +their power of attention. If when he was talking and explaining anything +to a monkey, its attention was easily distracted, as by a fly on the +wall or other trifling object, the case was hopeless. If he tried by +punishment to make an inattentive monkey act, it turned sulky. On the +other hand, a monkey which carefully attended to him could always be +trained. + +It is almost superfluous to state that animals have excellent _Memories_ +for persons and places. A baboon at the Cape of Good Hope, as I have +been informed by Sir Andrew Smith, recognised him with joy after an +absence of nine months. I had a dog who was savage and averse to all +strangers, and I purposely tried his memory after an absence of five +years and two days. I went near the stable where he lived, and shouted +to him in my old manner; he showed no joy, but instantly followed me out +walking and obeyed me, exactly as if I had parted with him only +half-an-hour before. A train of old associations, dormant during five +years, had thus been instantaneously awakened in his mind. Even ants, as +P. Huber[56] has clearly shewn, recognised their fellow-ants belonging +to the same community after a separation of four months. Animals can +certainly by some means judge of the intervals of time between recurrent +events. + +The _Imagination_ is one of the highest prerogatives of man. By this +faculty he unites, independently of the will, former images and ideas, +and thus creates brilliant and novel results. A poet, as Jean Paul +Richter remarks,[57] "who must reflect whether he shall make a +character say yes or no--to the devil with him; he is only a stupid +corpse." Dreaming gives us the best notion of this power; as Jean Paul +again says, "The dream is an involuntary art of poetry." The value of +the products of our imagination depends of course on the number, +accuracy, and clearness of our impressions; on our judgment and taste in +selecting or rejecting the involuntary combinations, and to a certain +extent on our power of voluntarily combining them. As dogs, cats, +horses, and probably all the higher animals, even birds, as is stated on +good authority,[58] have vivid dreams, and this is shewn by their +movements and voice, we must admit that they possess some power of +imagination. + +Of all the faculties of the human mind, it will, I presume, be admitted +that _Reason_ stands at the summit. Few persons any longer dispute that +animals possess some power of reasoning. Animals may constantly be seen +to pause, deliberate, and resolve. It is a significant fact, that the +more the habits of any particular animal are studied by a naturalist, +the more he attributes to reason and the less to unlearnt instincts.[59] +In future chapters we shall see that some animals extremely low in the +scale apparently display a certain amount of reason. No doubt it is +often difficult to distinguish between the power of reason and that of +instinct. Thus Dr. Hayes, in his work on 'The Open Polar Sea,' +repeatedly remarks that his dogs, instead of continuing to draw the +sledges in a compact body, diverged and separated when they came to thin +ice, so that their weight might be more evenly distributed. This was +often the first warning and notice which the travellers received that +the ice was becoming thin and dangerous. Now, did the dogs act thus from +the experience of each individual, or from the example of the older and +wiser dogs, or from an inherited habit, that is from an instinct? This +instinct might possibly have arisen since the time, long ago, when dogs +were first employed by the natives in drawing their sledges; or the +Arctic wolves, the parent-stock of the Esquimaux dog, may have acquired +this instinct, impelling them not to attack their prey in a close pack +when on thin ice. Questions of this kind are most difficult to answer. + +So many facts have been recorded in various works shewing that animals +possess some degree of reason, that I will here give only two or three +instances, authenticated by Rengger, and relating to American monkeys, +which stand low in their order. He states that when he first gave eggs +to his monkeys, they smashed them and thus lost much of their contents; +afterwards they gently hit one end against some hard body, and picked +off the bits of shell with their fingers. After cutting themselves only +once with any sharp tool, they would not touch it again, or would handle +it with the greatest care. Lumps of sugar were often given them wrapped +up in paper; and Rengger sometimes put a live wasp in the paper, so that +in hastily unfolding it they got stung; after this had once happened, +they always first held the packet to their ears to detect any movement +within. Any one who is not convinced by such facts as these, and by what +he may observe with his own dogs, that animals can reason, would not be +convinced by anything that I could add. Nevertheless I will give one +case with respect to dogs, as it rests on two distinct observers, and +can hardly depend on the modification of any instinct. + +Mr. Colquhoun[60] winged two wild-ducks, which fell on the opposite side +of a stream; his retriever tried to bring over both at once, but could +not succeed; she then, though never before known to ruffle a feather, +deliberately killed one, brought over the other, and returned for the +dead bird. Col. Hutchinson relates that two partridges were shot at +once, one being killed, the other wounded; the latter ran away, and was +caught by the retriever, who on her return came across the dead bird; +"she stopped, evidently greatly puzzled, and after one or two trials, +finding she could not take it up without permitting the escape of the +winged bird, she considered a moment, then deliberately murdered it by +giving it a severe crunch, and afterwards brought away both together. +This was the only known instance of her ever having wilfully injured any +game." Here we have reason, though not quite perfect, for the retriever +might have brought the wounded bird first and then returned for the dead +one, as in the case of the two wild-ducks. + +The muleteers in S. America say, "I will not give you the mule whose +step is easiest, but _la mas rational_,--the one that reasons best;" and +Humboldt[61] adds, "this popular expression, dictated by long +experience, combats the system of animated machines, better perhaps than +all the arguments of speculative philosophy." + + +It has, I think, now been shewn that man and the higher animals, +especially the Primates, have some few instincts in common. All have the +same senses, intuitions and sensations--similar passions, affections, +and emotions, even the more complex ones; they feel wonder and +curiosity; they possess the same faculties of imitation, attention, +memory, imagination, and reason, though in very different degrees. +Nevertheless many authors have insisted that man is separated through +his mental faculties by an impassable barrier from all the lower +animals. I formerly made a collection of above a score of such +aphorisms, but they are not worth giving, as their wide difference and +number prove the difficulty, if not the impossibility, of the attempt. +It has been asserted that man alone is capable of progressive +improvement; that he alone makes use of tools or fire, domesticates +other animals, possesses property, or employs language; that no other +animal is self-conscious, comprehends itself, has the power of +abstraction, or possesses general ideas; that man alone has a sense of +beauty, is liable to caprice, has the feeling of gratitude, mystery, +&c.; believes in God, or is endowed with a conscience. I will hazard a +few remarks on the more important and interesting of these points. + +Archbishop Sumner formerly maintained[62] that man alone is capable of +progressive improvement. With animals, looking first to the individual, +every one who has had any experience in setting traps knows that young +animals can be caught much more easily than old ones; and they can be +much more easily approached by an enemy. Even with respect to old +animals, it is impossible to catch many in the same place and in the +same kind of trap, or to destroy them by the same kind of poison; yet it +is improbable that all should have partaken of the poison, and +impossible that all should have been caught in the trap. They must learn +caution by seeing their brethren caught or poisoned. In North America, +where the fur-bearing animals have long been pursued, they exhibit, +according to the unanimous testimony of all observers, an almost +incredible amount of sagacity, caution, and cunning; but trapping has +been there so long carried on that inheritance may have come into play. + +If we look to successive generations, or to the race, there is no doubt +that birds and other animals gradually both acquire and lose caution in +relation to man or other enemies;[63] and this caution is certainly in +chief part an inherited habit or instinct, but in part the result of +individual experience. A good observer, Leroy,[64] states that in +districts where foxes are much hunted, the young when they first leave +their burrows are incontestably much more wary than the old ones in +districts where they are not much disturbed. + +Our domestic dogs are descended from wolves and jackals,[65] and though +they may not have gained in cunning, and may have lost in waryness and +suspicion, yet they have progressed in certain moral qualities, such as +in affection, trust-worthiness, temper, and probably in general +intelligence. The common rat has conquered and beaten several other +species throughout Europe, in parts of North America, New Zealand, and +recently in Formosa, as well as on the mainland of China. Mr. +Swinhoe,[66] who describes these latter cases, attributes the victory of +the common rat over the large _Mus coninga_ to its superior cunning; and +this latter quality may be attributed to the habitual exercise of all +its faculties in avoiding extirpation by man, as well as to nearly all +the less cunning or weak-minded rats having been successively destroyed +by him. To maintain, independently of any direct evidence, that no +animal during the course of ages has progressed in intellect or other +mental faculties, is to beg the question of the evolution of species. +Hereafter we shall see that, according to Lartet, existing mammals +belonging to several orders have larger brains than their ancient +tertiary prototypes. + +It has often been said that no animal uses any tool; but the chimpanzee +in a state of nature cracks a native fruit, somewhat like a walnut, with +a stone.[67] Rengger[68] easily taught an American monkey thus to break +open hard palm-nuts, and afterwards of its own accord it used stones to +open other kinds of nuts, as well as boxes. It thus also removed the +soft rind of fruit that had a disagreeable flavour. Another monkey was +taught to open the lid of a large box with a stick, and afterwards it +used the stick as a lever to move heavy bodies; and I have myself seen a +young orang put a stick into a crevice, slip his hand to the other end, +and use it in the proper manner as a lever. In the cases just mentioned +stones and sticks were employed as implements; but they are likewise +used as weapons. Brehm[69] states, on the authority of the well-known +traveller Schimper, that in Abyssinia when the baboons belonging to one +species (_C. gelada_) descend in troops from the mountains to plunder +the fields, they sometimes encounter troops of another species (_C. +hamadryas_), and then a fight ensues. The Geladas roll down great +stones, which the Hamadryas try to avoid, and then, both species, +making a great uproar, rush furiously against each other. Brehm, when +accompanying the Duke of Coburg-Gotha, aided in an attack with fire-arms +on a troop of baboons in the pass of Mensa in Abyssinia. The baboons in +return rolled so many stones down the mountain, some as large as a man's +head, that the attackers had to beat a hasty retreat; and the pass was +actually for a time closed against the caravan. It deserves notice that +these baboons thus acted in concert. Mr. Wallace[70] on three occasions +saw female orangs, accompanied by their young, "breaking off branches +and the great spiny fruit of the Durian tree, with every appearance of +rage; causing such a shower of missiles as effectually kept us from +approaching too near the tree." + +In the Zoological Gardens a monkey which had weak teeth used to break +open nuts with a stone; and I was assured by the keepers that this +animal, after using the stone, hid it in the straw, and would not let +any other monkey touch it. Here, then, we have the idea of property; but +this idea is common to every dog with a bone, and to most or all birds +with their nests. + +The Duke of Argyll[71] remarks, that the fashioning of an implement for +a special purpose is absolutely peculiar to man; and he considers that +this forms an immeasurable gulf between him and the brutes. It is no +doubt a very important distinction, but there appears to me much truth +in Sir J. Lubbock's suggestion,[72] that when primeval man first used +flint-stones for any purpose, he would have accidentally splintered +them, and would then have used the sharp fragments. From this step it +would be a small one to intentionally break the flints, and not a very +wide step to rudely fashion them. This latter advance, however, may have +taken long ages, if we may judge by the immense interval of time which +elapsed before the men of the neolithic period took to grinding and +polishing their stone tools. In breaking the flints, as Sir J. Lubbock +likewise remarks, sparks would have been emitted, and in grinding them +heat would have been evolved: "thus the two usual methods of obtaining +fire may have originated." The nature of fire would have been known in +the many volcanic regions where lava occasionally flows through forests. +The anthropomorphous apes, guided probably by instinct, build for +themselves temporary platforms; but as many instincts are largely +controlled by reason, the simpler ones, such as this of building a +platform, might readily pass into a voluntary and conscious act. The +orang is known to cover itself at night with the leaves of the Pandanus; +and Brehm states that one of his baboons used to protect itself from the +heat of the sun by throwing a straw-mat over its head. In these latter +habits, we probably see the first steps towards some of the simpler +arts; namely rude architecture and dress, as they arose amongst the +early progenitors of man. + + +_Language._--This faculty has justly been considered as one of the chief +distinctions between man and the lower animals. But man, as a highly +competent judge, Archbishop Whately remarks, "is not the only animal +that can make use of language to express what is passing in his mind, +and can understand, more or less, what is so expressed by another."[73] +In Paraguay the _Cebus Azar_ when excited utters at least six distinct +sounds, which excite in other monkeys similar emotions.[74] The +movements of the features and gestures of monkeys are understood by us, +and they partly understand ours, as Rengger and others declare. It is a +more remarkable fact that the dog, since being domesticated, has learnt +to bark[75] in at least four or five distinct tones. Although barking is +a new art, no doubt the wild species, the parents of the dog, expressed +their feelings by cries of various kinds. With the domesticated dog we +have the bark of eagerness, as in the chase; that of anger; the yelping +or howling bark of despair, as when shut up; that of joy, as when +starting on a walk with his master; and the very distinct one of demand +or supplication, as when wishing for a door or window to be opened. + +Articulate language is, however, peculiar to man; but he uses in common +with the lower animals inarticulate cries to express his meaning, aided +by gestures and the movements of the muscles of the face.[76] This +especially holds good with the more simple and vivid feelings, which are +but little connected with our higher intelligence. Our cries of pain, +fear, surprise, anger, together with their appropriate actions, and the +murmur of a mother to her beloved child, are more expressive than any +words. It is not the mere power of articulation that distinguishes man +from other animals, for as every one knows, parrots can talk; but it is +his large power of connecting definite sounds with definite ideas; and +this obviously depends on the development of the mental faculties. + +As Horne Tooke, one of the founders of the noble science of philology, +observes, language is an art, like brewing or baking; but writing would +have been a much more appropriate simile. It certainly is not a true +instinct, as every language has to be learnt. It differs, however, +widely from all ordinary arts, for man has an instinctive tendency to +speak, as we see in the babble of our young children; whilst no child +has an instinctive tendency to brew, bake, or write. Moreover, no +philologist now supposes that any language has been deliberately +invented; each has been slowly and unconsciously developed by many +steps. The sounds uttered by birds offer in several respects the nearest +analogy to language, for all the members of the same species utter the +same instinctive cries expressive of their emotions; and all the kinds +that have the power of singing exert this power instinctively; but the +actual song, and even the call-notes, are learnt from their parents or +foster-parents. These sounds, as Daines Barrington[77] has proved, "are +no more innate than language is in man." The first attempts to sing "may +be compared to the imperfect endeavour in a child to babble." The young +males continue practising, or, as the bird-catchers say, recording, for +ten or eleven months. Their first essays show hardly a rudiment of the +future song; but as they grow older we can perceive what they are aiming +at; and at last they are said "to sing their song round." Nestlings +which have learnt the song of a distinct species, as with the +canary-birds educated in the Tyrol, teach and transmit their new song to +their offspring. The slight natural differences of song in the same +species inhabiting different districts may be appositely compared, as +Barrington remarks, "to provincial dialects;" and the songs of allied, +though distinct species may be compared with the languages of distinct +races of man. I have given the foregoing details to shew that an +instinctive tendency to acquire an art is not a peculiarity confined to +man. + +With respect to the origin of articulate language, after having read on +the one side the highly interesting works of Mr. Hensleigh Wedgwood, the +Rev. F. Farrar, and Prof. Schleicher,[78] and the celebrated lectures of +Prof. Max Mller on the other side, I cannot doubt that language owes +its origin to the imitation and modification, aided by signs and +gestures, of various natural sounds, the voices of other animals, and +man's own instinctive cries. When we treat of sexual selection we shall +see that primeval man, or rather some early progenitor of man, probably +used his voice largely, as does one of the gibbon-apes at the present +day, in producing true musical cadences, that is in singing; we may +conclude from a widely-spread analogy that this power would have been +especially exerted during the courtship of the sexes, serving to express +various emotions, as love, jealousy, triumph, and serving as a challenge +to their rivals. The imitation by articulate sounds of musical cries +might have given rise to words expressive of various complex emotions. +As bearing on the subject of imitation, the strong tendency in our +nearest allies, the monkeys, in microcephalous idiots,[79] and in the +barbarous races of mankind, to imitate whatever they hear deserves +notice. As monkeys certainly understand much that is said to them by +man, and as in a state of nature they utter signal-cries of danger to +their fellows,[80] it does not appear altogether incredible, that some +unusually wise ape-like animal should have thought of imitating the +growl of a beast of prey, so as to indicate to his fellow monkeys the +nature of the expected danger. And this would have been a first step in +the formation of a language. + +As the voice was used more and more, the vocal organs would have been +strengthened and perfected through the principle of the inherited +effects of use; and this would have reacted on the power of speech. But +the relation between the continued use of language and the development +of the brain has no doubt been far more important. The mental powers in +some early progenitor of man must have been more highly developed than +in any existing ape, before even the most imperfect form of speech could +have come into use; but we may confidently believe that the continued +use and advancement of this power would have reacted on the mind by +enabling and encouraging it to carry on long trains of thought. A long +and complex train of thought can no more be carried on without the aid +of words, whether spoken or silent, than a long calculation without the +use of figures or algebra. It appears, also, that even ordinary trains +of thought almost require some form of language, for the dumb, deaf, and +blind girl, Laura Bridgman, was observed to use her fingers whilst +dreaming.[81] Nevertheless a long succession of vivid and connected +ideas, may pass through the mind without the aid of any form of +language, as we may infer from the prolonged dreams of dogs. We have, +also, seen that retriever-dogs are able to reason to a certain extent; +and this they manifestly do without the aid of language. The intimate +connection between the brain, as it is now developed in us, and the +faculty of speech, is well shewn by those curious cases of +brain-disease, in which speech is specially affected, as when the power +to remember substantives is lost, whilst other words can be correctly +used.[82] There is no more improbability in the effects of the continued +use of the vocal and mental organs being inherited, than in the case of +handwriting, which depends partly on the structure of the hand and +partly on the disposition of the mind; and handwriting is certainly +inherited.[83] + +Why the organs now used for speech should have been originally perfected +for this purpose, rather than any other organs, it is not difficult to +see. Ants have considerable powers of intercommunication by means of +their antenn, as shewn by Huber, who devotes a whole chapter to their +language. We might have used our fingers as efficient instruments, for a +person with practice can report to a deaf man every word of a speech +rapidly delivered at a public meeting; but the loss of our hands, whilst +thus employed, would have been a serious inconvenience. As all the +higher mammals possess vocal organs constructed on the same general +plan with ours, and which are used as a means of communication, it was +obviously probable, if the power of communication had to be improved, +that these same organs would have been still further developed; and this +has been effected by the aid of adjoining and well-adapted parts, namely +the tongue and lips.[84] The fact of the higher apes not using their +vocal organs for speech, no doubt depends on their intelligence not +having been sufficiently advanced. The possession by them of organs, +which with long-continued practice might have been used for speech, +although not thus used, is paralleled by the case of many birds which +possess organs fitted for singing, though they never sing. Thus, the +nightingale and crow have vocal organs similarly constructed, these +being used by the former for diversified song, and by the latter merely +for croaking.[85] + +The formation of different languages and of distinct species, and the +proofs that both have been developed through a gradual process, are +curiously the same.[86] But we can trace the origin of many words +further back than in the case of species, for we can perceive that they +have arisen from the imitation of various sounds, as in alliterative +poetry. We find in distinct languages striking homologies due to +community of descent, and analogies due to a similar process of +formation. The manner in which certain letters or sounds change when +others change is very like correlated growth. We have in both cases the +reduplication of parts, the effects of long-continued use, and so forth. +The frequent presence of rudiments, both in languages and in species, is +still more remarkable. The letter _m_ in the word _am_, means _I_; so +that in the expression _I am_, a superfluous and useless rudiment has +been retained. In the spelling also of words, letters often remain as +the rudiments of ancient forms of pronunciation. Languages, like organic +beings, can be classed in groups under groups; and they can be classed +either naturally according to descent, or artificially by other +characters. Dominant languages and dialects spread widely and lead to +the gradual extinction of other tongues. A language, like a species, +when once extinct, never, as Sir C. Lyell remarks, reappears. The same +language never has two birthplaces. Distinct languages may be crossed +or blended together.[87] We see variability in every tongue, and new +words are continually cropping up; but as there is a limit to the powers +of the memory, single words, like whole languages, gradually become +extinct. As Max Mller[88] has well remarked:--"A struggle for life is +constantly going on amongst the words and grammatical forms in each +language. The better, the shorter, the easier forms are constantly +gaining the upper hand, and they owe their success to their own inherent +virtue." To these more important causes of the survival of certain +words, mere novelty may, I think, be added; for there is in the mind of +man a strong love for slight changes in all things. The survival or +preservation of certain favoured words in the struggle for existence is +natural selection. + +The perfectly regular and wonderfully complex construction of the +languages of many barbarous nations has often been advanced as a proof, +either of the divine origin of these languages, or of the high art and +former civilisation of their founders. Thus F. von Schlegel writes: "In +those languages which appear to be at the lowest grade of intellectual +culture, we frequently observe a very high and elaborate degree of art +in their grammatical structure. This is especially the case with the +Basque and the Lapponian, and many of the American languages."[89] But +it is assuredly an error to speak of any language as an art in the sense +of its having been elaborately and methodically formed. Philologists now +admit that conjugations, declensions, &c., originally existed as +distinct words, since joined together; and as such words express the +most obvious relations between objects and persons, it is not surprising +that they should have been used by the men of most races during the +earliest ages. With respect to perfection, the following illustration +will best shew how easily we may err: a Crinoid sometimes consists of no +less than 150,000 pieces of shell,[90] all arranged with perfect +symmetry in radiating lines; but a naturalist does not consider an +animal of this kind as more perfect than a bilateral one with +comparatively few parts, and with none of these alike, excepting on the +opposite sides of the body. He justly considers the differentiation and +specialisation of organs as the test of perfection. So with languages, +the most symmetrical and complex ought not to be ranked above irregular, +abbreviated, and bastardised languages, which have borrowed expressive +words and useful forms of construction from various conquering, or +conquered, or immigrant races. + +From these few and imperfect remarks I conclude that the extremely +complex and regular construction of many barbarous languages, is no +proof that they owe their origin to a special act of creation.[91] Nor, +as we have seen, does the faculty of articulate speech in itself offer +any insuperable objection to the belief that man has been developed from +some lower form. + + +_Self-consciousness, Individuality, Abstraction, General Ideas, &c._--It +would be useless to attempt discussing these high faculties, which, +according to several recent writers, make the sole and complete +distinction between man and the brutes, for hardly two authors agree in +their definitions. Such faculties could not have been fully developed in +man until his mental powers had advanced to a high standard, and this +implies the use of a perfect language. No one supposes that one of the +lower animals reflects whence he comes or whither he goes,--what is +death or what is life, and so forth. But can we feel sure that an old +dog with an excellent memory and some power of imagination, as shewn by +his dreams, never reflects on his past pleasures in the chase? and this +would be a form of self-consciousness. On the other hand, as Bchner[92] +has remarked, how little can the hard-worked wife of a degraded +Australian savage, who uses hardly any abstract words and cannot count +above four, exert her self-consciousness, or reflect on the nature of +her own existence. + +That animals retain their mental individuality is unquestionable. When +my voice awakened a train of old associations in the mind of the +above-mentioned dog, he must have retained his mental individuality, +although every atom of his brain had probably undergone change more than +once during the interval of five years. This dog might have brought +forward the argument lately advanced to crush all evolutionists, and +said, "I abide amid all mental moods and all material changes.... The +teaching that atoms leave their impressions as legacies to other atoms +falling into the places they have vacated is contradictory of the +utterance of consciousness, and is therefore false; but it is the +teaching necessitated by evolutionism, consequently the hypothesis is a +false one."[93] + + +_Sense of Beauty._--This sense has been declared to be peculiar to man. +But when we behold male birds elaborately displaying their plumes and +splendid colours before the females, whilst other birds not thus +decorated make no such display, it is impossible to doubt that the +females admire the beauty of their male partners. As women everywhere +deck themselves with these plumes, the beauty of such ornaments cannot +be disputed. The Bower-birds by tastefully ornamenting their +playing-passages with gaily-coloured objects, as do certain +humming-birds their nests, offer additional evidence that they possess a +sense of beauty. So with the song of birds, the sweet strains poured +forth by the males during the season of love are certainly admired by +the females, of which fact evidence will hereafter be given. If female +birds had been incapable of appreciating the beautiful colours, the +ornaments, and voices of their male partners, all the labour and +anxiety exhibited by them in displaying their charms before the females +would have been thrown away; and this it is impossible to admit. Why +certain bright colours and certain sounds should excite pleasure, when +in harmony, cannot, I presume, be explained any more than why certain +flavours and scents are agreeable; but assuredly the same colours and +the same sounds are admired by us and by many of the lower animals. + +The taste for the beautiful, at least as far as female beauty is +concerned, is not of a special nature in the human mind; for it differs +widely in the different races of man, as will hereafter be shewn, and is +not quite the same even in the different nations of the same race. +Judging from the hideous ornaments and the equally hideous music admired +by most savages, it might be urged that their sthetic faculty was not +so highly developed as in certain animals, for instance, in birds. +Obviously no animal would be capable of admiring such scenes as the +heavens at night, a beautiful landscape, or refined music; but such high +tastes, depending as they do on culture and complex associations, are +not enjoyed by barbarians or by uneducated persons. + +Many of the faculties, which have been of inestimable service to man for +his progressive advancement, such as the powers of the imagination, +wonder, curiosity, an undefined sense of beauty, a tendency to +imitation, and the love of excitement or novelty, could not fail to have +led to the most capricious changes of customs and fashions. I have +alluded to this point, because a recent writer[94] has oddly fixed on +Caprice "as one of the most remarkable and typical differences between +savages and brutes." But not only can we perceive how it is that roan is +capricious, but the lower animals are, as we shall hereafter see, +capricious in their affections, aversions, and sense of beauty. There is +also good reason to suspect that they love novelty, for its own sake. + + +_Belief in God--Religion._--There is no evidence that man was +aboriginally endowed with the ennobling belief in the existence of an +Omnipotent God. On the contrary there is ample evidence, derived not +from hasty travellers, but from men who have long resided with savages, +that numerous races have existed and still exist, who have no idea of +one or more gods, and who have no words in their languages to express +such an idea.[95] The question is of course wholly distinct from that +higher one, whether there exists a Creator and Ruler of the universe; +and this has been answered in the affirmative by the highest intellects +that have ever lived. + +If, however, we include under the term "religion" the belief in unseen +or spiritual agencies, the case is wholly different; for this belief +seems to be almost universal with the less civilised races. Nor is it +difficult to comprehend how it arose. As soon as the important faculties +of the imagination, wonder, and curiosity, together with some power of +reasoning, had become partially developed, man would naturally have +craved to understand what was passing around him, and have vaguely +speculated on his own existence. As Mr. M'Lennan[96] has remarked, +"Some explanation of the phenomena of life, a man must feign for +himself; and to judge from the universality of it, the simplest +hypothesis, and the first to occur to men, seems to have been that +natural phenomena are ascribable to the presence in animals, plants, and +things, and in the forces of nature, of such spirits prompting to action +as men are conscious they themselves possess." It is probable, as Mr. +Tylor has clearly shewn, that dreams may have first given rise to the +notion of spirits; for savages do not readily distinguish between +subjective and objective impressions. When a savage dreams, the figures +which appear before him are believed to have come from a distance and to +stand over him; or "the soul of the dreamer goes out on its travels, and +comes home with a remembrance of what it has seen."[97] But until the +above-named faculties of imagination, curiosity, reason, &c., had been +fairly well developed in the mind of man, his dreams would not have led +him to believe in spirits, any more than in the case of a dog. + +The tendency in savages to imagine that natural objects and agencies +are animated by spiritual or living essences, is perhaps illustrated by +a little fact which I once noticed: my dog, a full-grown and very +sensible animal, was lying on the lawn during a hot and still day; but +at a little distance a slight breeze occasionally moved an open parasol, +which would have been wholly disregarded by the dog, had any one stood +near it. As it was, every time that the parasol slightly moved, the dog +growled fiercely and barked. He must, I think, have reasoned to himself +in a rapid and unconscious manner, that movement without any apparent +cause indicated the presence of some strange living agent, and no +stranger had a right to be on his territory. + +The belief in spiritual agencies would easily pass into the belief in +the existence of one or more gods. For savages would naturally attribute +to spirits the same passions, the same love of vengeance or simplest +form of justice, and the same affections which they themselves +experienced. The Fuegians appear to be in this respect in an +intermediate condition, for when the surgeon on board the "Beagle" shot +some young ducklings as specimens, York Minster declared in the most +solemn manner, "Oh! Mr. Bynoe, much rain, much snow, blow much;" and +this was evidently a retributive punishment for wasting human food. So +again he related how, when his brother killed a "wild man," storms long +raged, much rain and snow fell. Yet we could never discover that the +Fuegians believed in what we should call a God, or practised any +religious rites; and Jemmy Button, with justifiable pride, stoutly +maintained that there was no devil in his land. This latter assertion is +the more remarkable, as with savages the belief in bad spirits is far +more common than the belief in good spirits. + +The feeling of religious devotion is a highly complex one, consisting of +love, complete submission to an exalted and mysterious superior, a +strong sense of dependence,[98] fear, reverence, gratitude, hope for the +future, and perhaps other elements. No being could experience so complex +an emotion until advanced in his intellectual and moral faculties to at +least a moderately high level. Nevertheless we see some distant approach +to this state of mind, in the deep love of a dog for his master, +associated with complete submission, some fear, and perhaps other +feelings. The behaviour of a dog when returning to his master after an +absence, and, as I may add, of a monkey to his beloved keeper, is widely +different from that towards their fellows. In the latter case the +transports of joy appear to be somewhat less, and the sense of equality +is shewn in every action. Professor Braubach[99] goes so far as to +maintain that a dog looks on his master as on a god. + +The same high mental faculties which first led man to believe in unseen +spiritual agencies, then in fetishism, polytheism, and ultimately in +monotheism, would infallibly lead him, as long as his reasoning powers +remained poorly developed, to various strange superstitions and customs. +Many of these are terrible to think of--such as the sacrifice of human +beings to a blood-loving god; the trial of innocent persons by the +ordeal of poison or fire; witchcraft, &c.--yet it is well occasionally +to reflect on these superstitions, for they shew us what an infinite +debt of gratitude we owe to the improvement of our reason, to science, +and our accumulated knowledge.[100] As Sir J. Lubbock has well +observed, "it is not too much to say that the horrible dread of unknown +evil hangs like a thick cloud over savage life, and embitters every +pleasure." These miserable and indirect consequences of our highest +faculties may be compared with the incidental and occasional mistakes of +the instincts of the lower animals. + + + + +CHAPTER III. + +COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER +ANIMALS--_continued_. + + + The moral sense--Fundamental proposition--The qualities of + social animals--Origin of sociability--Struggle between opposed + instincts--Man a social animal--The more enduring social + instincts conquer other less persistent instincts--The social + virtues alone regarded by savages--The self-regarding virtues + acquired at a later stage of development--The importance of the + judgment of the members of the same community on + conduct--Transmission of moral tendencies--Summary. + + +I fully subscribe to the judgment of those writers[101] who maintain +that of all the differences between man and the lower animals, the moral +sense or conscience is by far the most important. This sense, as +Mackintosh[102] remarks, "has a rightful supremacy over every other +principle of human action;" it is summed up in that short but imperious +word _ought_, so full of high significance. It is the most noble of all +the attributes of man, leading him without a moment's hesitation to risk +his life for that of a fellow-creature; or after due deliberation, +impelled simply by the deep feeling of right or duty, to sacrifice it in +some great cause. Immanuel Kant exclaims, "Duty! Wondrous thought, that +workest neither by fond insinuation, flattery, nor by any threat, but +merely by holding up thy naked law in the soul, and so extorting for +thyself always reverence, if not always obedience; before whom all +appetites are dumb, however secretly they rebel; whence thy +original?"[103] + +This great question has been discussed by many writers[104] of +consummate ability; and my sole excuse for touching on it is the +impossibility of here passing it over, and because, as far as I know, no +one has approached it exclusively from the side of natural history. The +investigation possesses, also, some independent interest, as an attempt +to see how far the study of the lower animals can throw light on one of +the highest psychical faculties of man. + +The following proposition seems to me in a high degree probable--namely, +that any animal whatever, endowed with well-marked social +instincts,[105] would inevitably acquire a moral sense or conscience, as +soon as its intellectual powers had become as well developed, or nearly +as well developed, as in man. For, _firstly_, the social instincts lead +an animal to take pleasure in the society of its fellows, to feel a +certain amount of sympathy with them, and to perform various services +for them. The services may be of a definite and evidently instinctive +nature; or there may be only a wish and readiness, as with most of the +higher social animals, to aid their fellows in certain general ways. But +these feelings and services are by no means extended to all the +individuals of the same species, only to those of the same association. +_Secondly_, as soon as the mental faculties had become highly developed, +images of all past actions and motives would be incessantly passing +through the brain of each individual; and that feeling of +dissatisfaction which invariably results, as we shall hereafter see, +from any unsatisfied instinct, would arise, as often as it was perceived +that the enduring and always present social instinct had yielded to some +other instinct, at the time stronger, but neither enduring in its +nature, nor leaving behind it a very vivid impression. It is clear that +many instinctive desires, such as that of hunger, are in their nature of +short duration; and after being satisfied are not readily or vividly +recalled. _Thirdly_, after the power of language had been acquired and +the wishes of the members of the same community could be distinctly +expressed, the common opinion how each member ought to act for the +public good, would naturally become to a large extent the guide to +action. But the social instincts would still give the impulse to act for +the good of the community, this impulse being strengthened, directed, +and sometimes even deflected by public opinion, the power of which +rests, as we shall presently see, on instinctive sympathy. _Lastly_, +habit in the individual would ultimately play a very important part in +guiding the conduct of each member; for the social instincts and +impulses, like all other instincts, would be greatly strengthened by +habit, as would obedience to the wishes and judgment of the community. +These several subordinate propositions must now be discussed; and some +of them at considerable length. + +It may be well first to premise that I do not wish to maintain that any +strictly social animal, if its intellectual faculties were to become as +active and as highly developed as in man, would acquire exactly the same +moral sense as ours. In the same manner as various animals have some +sense of beauty, though they admire widely different objects, so they +might have a sense of right and wrong, though led by it to follow widely +different lines of conduct. If, for instance, to take an extreme case, +men were reared under precisely the same conditions as hive-bees, there +can hardly be a doubt that our unmarried females would, like the +worker-bees, think it a sacred duty to kill their brothers, and mothers +would strive to kill their fertile daughters; and no one would think of +interfering. Nevertheless the bee, or any other social animal, would in +our supposed case gain, as it appears to me, some feeling of right and +wrong, or a conscience. For each individual would have an inward sense +of possessing certain stronger or more enduring instincts, and others +less strong or enduring; so that there would often be a struggle which +impulse should be followed; and satisfaction or dissatisfaction would be +felt, as past impressions were compared during their incessant passage +through the mind. In this case an inward monitor would tell the animal +that it would have been better to have followed the one impulse rather +than the other. The one course ought to have been followed: the one +would have been right and the other wrong; but to these terms I shall +have to recur. + + +_Sociability._--Animals of many kinds are social; we find even distinct +species living together, as with some American monkeys, and with the +united flocks of rooks, jackdaws, and starlings. Man shows the same +feeling in his strong love for the dog, which the dog returns with +interest. Every one must have noticed how miserable horses, dogs, sheep, +&c. are when separated from their companions; and what affection at +least the two former kinds show on their reunion. It is curious to +speculate on the feelings of a dog, who will rest peacefully for hours +in a room with his master or any of the family, without the least notice +being taken of him; but if left for a short time by himself, barks or +howls dismally. We will confine our attention to the higher social +animals, excluding insects, although these aid each other in many +important ways. The most common service which the higher animals perform +for each other, is the warning each other of danger by means of the +united senses of all. Every sportsman knows, as Dr. Jaeger remarks,[106] +how difficult it is to approach animals in a herd or troop. Wild horses +and cattle do not, I believe, make any danger-signal; but the attitude +of any one who first discovers an enemy, warns the others. Rabbits stamp +loudly on the ground with their hind-feet as a signal: sheep and chamois +do the same, but with their fore-feet, uttering likewise a whistle. Many +birds and some mammals post sentinels, which in the case of seals are +said[107] generally to be the females. The leader of a troop of monkeys +acts as the sentinel, and utters cries expressive both of danger and of +safety.[108] Social animals perform many little services for each +other: horses nibble, and cows lick each other, on any spot which +itches: monkeys search for each other's external parasites; and Brehm +states that after a troop of the _Cercopithecus griseo-viridis_ has +rushed through a thorny brake, each monkey stretches itself on a branch, +and another monkey sitting by "conscientiously" examines its fur and +extracts every thorn or burr. + +Animals also render more important services to each other: thus wolves +and some other beasts of prey hunt in packs, and aid each other in +attacking their victims. Pelicans fish in concert. The Hamadryas baboons +turn over stones to find insects, &c.; and when they come to a large +one, as many as can stand round, turn it over together and share the +booty. Social animals mutually defend each other. The males of some +ruminants come to the front when there is danger and defend the herd +with their horns. I shall also in a future chapter give cases of two +young wild bulls attacking an old one in concert, and of two stallions +together trying to drive away a third stallion from a troop of mares. +Brehm encountered in Abyssinia a great troop of baboons which were +crossing a valley: some had already ascended the opposite mountain, and +some were still in the valley: the latter were attacked by the dogs, but +the old males immediately hurried down from the rocks, and with mouths +widely opened roared so fearfully, that the dogs precipitately +retreated. They were again encouraged to the attack; but by this time +all the baboons had reascended the heights, excepting a young one, about +six months old, who, loudly calling for aid, climbed on a block of rock +and was surrounded. Now one of the largest males, a true hero, came down +again from the mountain, slowly went to the young one, coaxed him, and +triumphantly led him away--the dogs being too much astonished to make an +attack. I cannot resist giving another scene which was witnessed by this +same naturalist; an eagle seized a young Cercopithecus, which, by +clinging to a branch, was not at once carried off; it cried loudly for +assistance, upon which the other members of the troop with much uproar +rushed to the rescue, surrounded the eagle, and pulled out so many +feathers, that he no longer thought of his prey, but only how to escape. +This eagle, as Brehm remarks, assuredly would never again attack a +monkey in a troop. + +It is certain that associated animals have a feeling of love for each +other which is not felt by adult and non-social animals. How far in most +cases they actually sympathise with each other's pains and pleasures is +more doubtful, especially with respect to the latter. Mr. Buxton, +however, who had excellent means of observation,[109] states that his +macaws, which lived free in Norfolk, took "an extravagant interest" in a +pair with a nest, and whenever the female left it, she was surrounded by +a troop "screaming horrible acclamations in her honour." It is often +difficult to judge whether animals have any feeling for each other's +sufferings. Who can say what cows feel, when they surround and stare +intently on a dying or dead companion? That animals sometimes are far +from feeling any sympathy is too certain; for they will expel a wounded +animal from the herd, or gore or worry it to death. This is almost the +blackest fact in natural history, unless indeed the explanation which +has been suggested is true, that their instinct or reason leads them to +expel an injured companion, lest beasts of prey, including man, should +be tempted to follow the troop. In this case their conduct is not much +worse than that of the North American Indians who leave their feeble +comrades to perish on the plains, or the Feegeans, who, when their +parents get old or fall ill, bury them alive.[110] + +Many animals, however, certainly sympathise with each other's distress +or danger. This is the case even with birds; Capt. Stansbury[111] found +on a salt lake in Utah an old and completely blind pelican, which was +very fat, and must have been long and well fed by his companions. Mr. +Blyth, as he informs me, saw Indian crows feeding two or three of their +companions which were blind; and I have heard of an analogous case with +the domestic cock. We may, if we choose, call these actions instinctive; +but such cases are much too rare for the development of any special +instinct.[112] I have myself seen a dog, who never passed a great friend +of his, a cat which lay sick in a basket, without giving her a few licks +with his tongue, the surest sign of kind feeling in a dog. + +It must be called sympathy that leads a courageous dog to fly at any one +who strikes his master, as he certainly will. I saw a person pretending +to beat a lady who had a very timid little dog on her lap, and the trial +had never before been made. The little creature instantly jumped away, +but after the pretended beating was over, it was really pathetic to see +how perseveringly he tried to lick his mistress's face and comfort her. +Brehm[113] states that when a baboon in confinement was pursued to be +punished, the others tried to protect him. It must have been sympathy in +the cases above given which led the baboons and Cercopitheci to defend +their young comrades from the dogs and the eagle. I will give only one +other instance of sympathetic and heroic conduct in a little American +monkey. Several years ago a keeper at the Zoological Gardens, showed me +some deep and scarcely healed wounds on the nape of his neck, inflicted +on him whilst kneeling on the floor by a fierce baboon. The little +American monkey, who was a warm friend of this keeper, lived in the same +large compartment, and was dreadfully afraid of the great baboon. +Nevertheless, as soon as he saw his friend the keeper in peril, he +rushed to the rescue, and by screams and bites so distracted the baboon +that the man was able to escape, after running great risk, as the +surgeon who attended him thought, of his life. + +Besides love and sympathy, animals exhibit other qualities which in us +would be called moral; and I agree with Agassiz[114] that dogs possess +something very like a conscience. They certainly possess some power of +self-command, and this does not appear to be wholly the result of fear. +As Braubach[115] remarks, a dog will refrain from stealing food in the +absence of his master. Dogs have long been accepted as the very type of +fidelity and obedience. All animals living in a body which defend each +other or attack their enemies in concert, must be in some degree +faithful to each other; and those that follow a leader must be in some +degree obedient. When the baboons in Abyssinia[116] plunder a garden, +they silently follow their leader; and if an imprudent young animal +makes a noise, he receives a slap from the others to teach him silence +and obedience; but as soon as they are sure that there is no danger, all +show their joy by much clamour. + +With respect to the impulse which leads certain animals to associate +together, and to aid each other in many ways, we may infer that in most +cases they are impelled by the same sense of satisfaction or pleasure +which they experience in performing other instinctive actions; or by the +same sense of dissatisfaction, as in other cases of prevented +instinctive actions. We see this in innumerable instances, and it is +illustrated in a striking manner by the acquired instincts of our +domesticated animals; thus a young shepherd-dog delights in driving and +running round a flock of sheep, but not in worrying them; a young +foxhound delights in hunting a fox, whilst some other kinds of dogs as I +have witnessed, utterly disregard foxes. What a strong feeling of inward +satisfaction must impel a bird, so full of activity, to brood day after +day over her eggs. Migratory birds are miserable if prevented from +migrating, and perhaps they enjoy starting on their long flight. Some +few instincts are determined solely by painful feelings, as by fear, +which leads to self-preservation, or is specially directed against +certain enemies. No one, I presume, can analyse the sensations of +pleasure or pain. In many cases, however, it is probable that instincts +are persistently followed from the mere force of inheritance, without +the stimulus of either pleasure or pain. A young pointer, when it first +scents game, apparently cannot help pointing. A squirrel in a cage who +pats the nuts which it cannot eat, as if to bury them in the ground, can +hardly be thought to act thus either from pleasure or pain. Hence the +common assumption that men must be impelled to every action by +experiencing some pleasure or pain may be erroneous. Although a habit +may be blindly and implicitly followed, independently of any pleasure or +pain felt at the moment, yet if it be forcibly and abruptly checked, a +vague sense of dissatisfaction is generally experienced; and this is +especially true in regard to persons of feeble intellect. + +It has often been assumed that animals were in the first place rendered +social, and that they feel as a consequence uncomfortable when separated +from each other, and comfortable whilst together; but it is a more +probable view that these sensations were first developed, in order that +those animals which would profit by living in society, should be induced +to live together. In the same manner as the sense of hunger and the +pleasure of eating were, no doubt, first acquired in order to induce +animals to eat. The feeling of pleasure from society is probably an +extension of the parental or filial affections; and this extension may +be in chief part attributed to natural selection, but perhaps in part to +mere habit. For with those animals which were benefited by living in +close association, the individuals which took the greatest pleasure in +society would best escape various dangers; whilst those that cared least +for their comrades and lived solitary would perish in greater numbers. +With respect to the origin of the parental and filial affections, which +apparently lie at the basis of the social affections, it is hopeless to +speculate; but we may infer that they have been to a large extent +gained through natural selection. So it has almost certainly been with +the unusual and opposite feeling of hatred between the nearest +relations, as with the worker-bees which kill their brother-drones, and +with the queen-bees which kill their daughter-queens; the desire to +destroy, instead of loving, their nearest relations having been here of +service to the community. + +The all-important emotion of sympathy is distinct from that of love. A +mother may passionately love her sleeping and passive infant, but she +can then hardly be said to feel sympathy for it. The love of a man for +his dog is distinct from sympathy, and so is that of a dog for his +master. Adam Smith formerly argued, as has Mr. Bain recently, that the +basis of sympathy lies in our strong retentiveness of former states of +pain or pleasure. Hence, "the sight of another person enduring hunger, +cold, fatigue, revives in us some recollection of these states, which +are painful even in idea." We are thus impelled to relieve the +sufferings of another, in order that our own painful feelings may be at +the same time relieved. In like manner we are led to participate in the +pleasures of others.[117] But I cannot see how this view explains the +fact that sympathy is excited in an immeasurably stronger degree by a +beloved than by an indifferent person. The mere sight of suffering, +independently of love, would suffice to call up in us vivid +recollections and associations. Sympathy may at first have originated in +the manner above suggested; but it seems now to have become an instinct, +which is especially directed towards beloved objects, in the same manner +as fear with animals is especially directed against certain enemies. As +sympathy is thus directed, the mutual love of the members of the same +community will extend its limits. No doubt a tiger or lion feels +sympathy for the sufferings of its own young, but not for any other +animal. With strictly social animals the feeling will be more or less +extended to all the associated members, as we know to be the case. With +mankind selfishness, experience, and imitation probably add, as Mr. Bain +has shewn, to the power of sympathy; for we are led by the hope of +receiving good in return to perform acts of sympathetic kindness to +others; and there can be no doubt that the feeling of sympathy is much +strengthened by habit. In however complex a manner this feeling may have +originated, as it is one of high importance to all those animals which +aid and defend each other, it will have been increased, through natural +selection; for those communities, which included the greatest number of +the most sympathetic members, would flourish best and rear the greatest +number of offspring. + +In many cases it is impossible to decide whether certain social +instincts have been acquired through natural selection, or are the +indirect result of other instincts and faculties, such as sympathy, +reason, experience, and a tendency to imitation; or again, whether they +are simply the result of long-continued habit. So remarkable an instinct +as the placing sentinels to warn the community of danger, can hardly +have been the indirect result of any other faculty; it must therefore +have been directly acquired. On the other hand, the habit followed by +the males of some social animals, of defending the community and of +attacking their enemies or their prey in concert, may perhaps have +originated from mutual sympathy; but courage, and in most cases +strength, must have been previously acquired, probably through natural +selection. + +Of the various instincts and habits, some are much stronger than others, +that is, some either give more pleasure in their performance and more +distress in their prevention than others; or, which is probably quite as +important, they are more persistently followed through inheritance +without exciting any special feeling of pleasure or pain. We are +ourselves conscious that some habits are much more difficult to cure or +change than others. Hence a struggle may often be observed in animals +between different instincts, or between an instinct and some habitual +disposition; as when a dog rushes after a hare, is rebuked, pauses, +hesitates, pursues again or returns ashamed to his master; or as between +the love of a female dog for her young puppies and for her master, for +she may be seen to slink away to them, as if half ashamed of not +accompanying her master. But the most curious instance known to me of +one instinct conquering another, is the migratory instinct conquering +the maternal instinct. The former is wonderfully strong; a confined bird +will at the proper season beat her breast against the wires of her cage, +until it is bare and bloody. It causes young salmon to leap out of the +fresh water, where they could still continue to live, and thus +unintentionally to commit suicide. Every one knows how strong the +maternal instinct is, leading even timid birds to face great danger, +though with hesitation and in opposition to the instinct of +self-preservation. Nevertheless the migratory instinct is so powerful +that late in the autumn swallows and house-martins frequently desert +their tender young, leaving them to perish miserably in their +nests.[118] + +We can perceive that an instinctive impulse, if it be in any way more +beneficial to a species than some other or opposed instinct, would be +rendered the more potent of the two through natural selection; for the +individuals which had it most strongly developed would survive in larger +numbers. Whether this is the case with the migratory in comparison with +the maternal instinct, may well be doubted. The great persistence or +steady action of the former at certain seasons of the year during the +whole day, may give it for a time paramount force. + + +_Man a social animal._--Most persons admit that man is a social being. +We see this in his dislike of solitude, and in his wish for society +beyond that of his own family. Solitary confinement is one of the +severest punishments which can be inflicted. Some authors suppose that +man primevally lived in single families; but at the present day, though +single families, or only two or three together, roam the solitudes of +some savage lands, they are always, as far as I can discover, friendly +with other families inhabiting the same district. Such families +occasionally meet in council, and they unite for their common defence. +It is no argument against savage man being a social animal, that the +tribes inhabiting adjacent districts are almost always at war with each +other; for the social instincts never extend to all the individuals of +the same species. Judging from the analogy of the greater number of the +Quadrumana, it is probable that the early ape-like progenitors of man +were likewise social; but this is not of much importance for us. +Although man, as he now exists, has few special instincts, having lost +any which his early progenitors may have possessed, this is no reason +why he should not have retained from an extremely remote period some +degree of instinctive love and sympathy for his fellows. We are indeed +all conscious that we do possess such sympathetic feelings;[119] but our +consciousness does not tell us whether they are instinctive, having +originated long ago in the same manner as with the lower animals, or +whether they have been acquired by each of us during our early years. As +man is a social animal, it is also probable that he would inherit a +tendency to be faithful to his comrades, for this quality is common to +most social animals. He would in like manner possess some capacity for +self-command, and perhaps of obedience to the leader of the community. +He would from an inherited tendency still be willing to defend, in +concert with others, his fellow-men, and would be ready to aid them in +any way which did not too greatly interfere with his own welfare or his +own strong desires. + +The social animals which stand at the bottom of the scale are guided +almost exclusively, and those which stand higher in the scale are +largely guided, in the aid which they give to the members of the same +community, by special instincts; but they are likewise in part impelled +by mutual love and sympathy, assisted apparently by some amount of +reason. Although man, as just remarked, has no special instincts to tell +him how to aid his fellow-men, he still has the impulse, and with his +improved intellectual faculties would naturally be much guided in this +respect by reason and experience. Instinctive sympathy would, also, +cause him to value highly the approbation of his fellow-men; for, as Mr. +Bain has clearly shewn,[120] the love of praise and the strong feeling +of glory, and the still stronger horror of scorn and infamy, "are due to +the workings of sympathy." Consequently man would be greatly influenced +by the wishes, approbation, and blame of his fellow-men, as expressed by +their gestures and language. Thus the social instincts, which must have +been acquired by man in a very rude state, and probably even by his +early ape-like progenitors, still give the impulse to many of his best +actions; but his actions are largely determined by the expressed wishes +and judgment of his fellow-men, and unfortunately still oftener by his +own strong, selfish desires. But as the feelings of love and sympathy +and the power of self-command become strengthened by habit, and as the +power of reasoning becomes clearer so that man can appreciate the +justice of the judgments of his fellow-men, he will feel himself +impelled, independently of any pleasure or pain felt at the moment, to +certain lines of conduct. He may then say, I am the supreme judge of my +own conduct, and in the words of Kant, I will not in my own person +violate the dignity of humanity. + + +_The more enduring Social Instincts conquer the less Persistent +Instincts._--We have, however, not as yet considered the main point, on +which the whole question of the moral sense hinges. Why should a man +feel that he ought to obey one instinctive desire rather than another? +Why does he bitterly regret if he has yielded to the strong sense of +self-preservation, and has not risked his life to save that of a +fellow-creature; or why does he regret having stolen food from severe +hunger? + +It is evident in the first place, that with mankind the instinctive +impulses have different degrees of strength; a young and timid mother +urged by the maternal instinct will, without a moment's hesitation, run +the greatest danger for her infant, but not for a mere fellow-creature. +Many a man, or even boy, who never before risked his life for another, +but in whom courage and sympathy were well developed, has, disregarding +the instinct of self-preservation, instantaneously plunged into a +torrent to save a drowning fellow-creature. In this case man is impelled +by the same instinctive motive, which caused the heroic little American +monkey, formerly described, to attack the great and dreaded baboon, to +save his keeper. Such actions as the above appear to be the simple +result of the greater strength of the social or maternal instincts than +of any other instinct or motive; for they are performed too +instantaneously for reflection, or for the sensation of pleasure or +pain; though if prevented distress would be caused. + +I am aware that some persons maintain that actions performed +impulsively, as in the above cases, do not come under the dominion of +the moral sense, and cannot be called moral. They confine this term to +actions done deliberately, after a victory over opposing desires, or to +actions prompted by some lofty motive. But it appears scarcely possible +to draw any clear line of distinction of this kind; though the +distinction may be real. As far as exalted motives are concerned, many +instances have been recorded of barbarians, destitute of any feeling of +general benevolence towards mankind, and not guided by any religious +motive, who have deliberately as prisoners sacrificed their lives,[121] +rather than betray their comrades; and surely their conduct ought to be +considered as moral. As far as deliberation and the victory over +opposing motives are concerned, animals may be seen doubting between +opposed instincts, as in rescuing their offspring or comrades from +danger; yet their actions, though done for the good of others, are not +called moral. Moreover, an action repeatedly performed by us, will at +last be done without deliberation or hesitation, and can then hardly be +distinguished from an instinct; yet surely no one will pretend that an +action thus done ceases to be moral. On the contrary, we all feel that +an act cannot be considered as perfect, or as performed in the most +noble manner, unless it be done impulsively, without deliberation or +effort, in the same manner as by a man in whom the requisite qualities +are innate. He who is forced to overcome his fear or want of sympathy +before he acts, deserves, however, in one way higher credit than the man +whose innate disposition leads him to a good act without effort. As we +cannot distinguish between motives, we rank all actions of a certain +class as moral, when they are performed by a moral being. A moral being +is one who is capable of comparing his past and future actions or +motives, and of approving or disapproving of them. We have no reason to +suppose that any of the lower animals have this capacity; therefore +when a monkey faces danger to rescue its comrade, or takes charge of an +orphan-monkey, we do not call its conduct moral. But in the case of man, +who alone can with certainty be ranked as a moral being, actions of a +certain class are called moral, whether performed deliberately after a +struggle with opposing motives, or from the effects of slowly-gained +habit, or impulsively through instinct. + +But to return to our more immediate subject; although some instincts are +more powerful than others, thus leading to corresponding actions, yet it +cannot be maintained that the social instincts are ordinarily stronger +in man, or have become stronger through long-continued habit, than the +instincts, for instance, of self-preservation, hunger, lust, vengeance, +&c. Why then does man regret, even though he may endeavour to banish any +such regret, that he has followed the one natural impulse, rather than +the other; and why does he further feel that he ought to regret his +conduct? Man in this respect differs profoundly from the lower animals. +Nevertheless we can, I think, see with some degree of clearness the +reason of this difference. + +Man, from the activity of his mental faculties, cannot avoid reflection: +past impressions and images are incessantly passing through his mind +with distinctness. Now with those animals which live permanently in a +body, the social instincts are ever present and persistent. Such animals +are always ready to utter the danger-signal, to defend the community, +and to give aid to their fellows in accordance with their habits; they +feel at all times, without the stimulus of any special passion or +desire, some degree of love and sympathy for them; they are unhappy if +long separated from them, and always happy to be in their company. So it +is with ourselves. A man who possessed no trace of such feelings would +be an unnatural monster. On the other hand, the desire to satisfy +hunger, or any passion, such as vengeance, is in its nature temporary, +and can for a time be fully satisfied. Nor is it easy, perhaps hardly +possible, to call up with complete vividness the feeling, for instance, +of hunger; nor indeed, as has often been remarked, of any suffering. The +instinct of self-preservation is not felt except in the presence of +danger; and many a coward has thought himself brave until he has met his +enemy face to face. The wish for another man's property is perhaps as +persistent a desire as any that can be named; but even in this case the +satisfaction of actual possession is generally a weaker feeling than the +desire: many a thief, if not an habitual one, after success has wondered +why he stole some article. + +Thus, as man cannot prevent old impressions continually repassing +through his mind, he will be compelled to compare the weaker impressions +of, for instance, past hunger, or of vengeance satisfied or danger +avoided at the cost of other men, with the instinct of sympathy and +good-will to his fellows, which is still present and ever in some degree +active in his mind. He will then feel in his imagination that a stronger +instinct has yielded to one which now seems comparatively weak; and then +that sense of dissatisfaction will inevitably be felt with which man is +endowed, like every other animal, in order that his instincts may be +obeyed. The case before given, of the swallow, affords an illustration, +though of a reversed nature, of a temporary though for the time strongly +persistent instinct conquering another instinct which is usually +dominant over all others. At the proper season these birds seem all day +long to be impressed with the desire to migrate; their habits change; +they become restless, are noisy, and congregate in flocks. Whilst the +mother-bird is feeding or brooding over her nestlings, the maternal +instinct is probably stronger than the migratory; but the instinct which +is more persistent gains the victory, and at last, at a moment when her +young ones are not in sight, she takes flight and deserts them. When +arrived at the end of her long journey, and the migratory instinct +ceases to act, what an agony of remorse each bird would feel, if, from +being endowed with great mental activity, she could not prevent the +image continually passing before her mind of her young ones perishing in +the bleak north from cold and hunger. + +At the moment of action, man will no doubt be apt to follow the stronger +impulse; and though this may occasionally prompt him to the noblest +deeds, it will far more commonly lead him to gratify his own desires at +the expense of other men. But after their gratification, when past and +weaker impressions are contrasted with the ever-enduring social +instincts, retribution will surely come. Man will then feel dissatisfied +with himself, and will resolve with more or less force to act +differently for the future. This is conscience; for conscience looks +backwards and judges past actions, inducing that kind of +dissatisfaction, which if weak we call regret, and if severe remorse. + +These sensations are, no doubt, different from those experienced when +other instincts or desires are left unsatisfied; but every unsatisfied +instinct has its own proper prompting sensation, as we recognise with +hunger, thirst, &c. Man thus prompted, will through long habit acquire +such perfect self-command, that his desires and passions will at last +instantly yield to his social sympathies, and there will no longer be a +struggle between them. The still hungry, or the still revengeful man +will not think of stealing food, or of wreaking his vengeance. It is +possible, or, as we shall hereafter see, even probable, that the habit +of self-command may, like other habits, be inherited. Thus at last man +comes to feel, through acquired and perhaps inherited habit, that it is +best for him to obey his more persistent instincts. The imperious word +_ought_ seems merely to imply the consciousness of the existence of a +persistent instinct, either innate or partly acquired, serving him as a +guide, though liable to be disobeyed. We hardly use the word _ought_ in +a metaphorical sense, when we say hounds ought to hunt, pointers to +point, and retrievers to retrieve their game. If they fail thus to act, +they fail in their duty and act wrongly. + +If any desire or instinct, leading to an action opposed to the good of +others, still appears to a man, when recalled to mind, as strong as, or +stronger than, his social instinct, he will feel no keen regret at +having followed it; but he will be conscious that if his conduct were +known to his fellows, it would meet with their disapprobation; and few +are so destitute of sympathy as not to feel discomfort when this is +realised. If he has no such sympathy, and if his desires leading to bad +actions are at the time strong, and when recalled are not overmastered +by the persistent social instincts, then he is essentially a bad +man;[122] and the sole restraining motive left is the fear of +punishment, and the conviction that in the long run it would be best for +his own selfish interests to regard the good of others rather than his +own. + +It is obvious that every one may with an easy conscience gratify his own +desires, if they do not interfere with his social instincts, that is +with the good of others; but in order to be quite free from +self-reproach, or at least of anxiety, it is almost necessary for him to +avoid the disapprobation, whether reasonable or not, of his fellow men. +Nor must he break through the fixed habits of his life, especially if +these are supported by reason; for if he does, he will assuredly feel +dissatisfaction. He must likewise avoid the reprobation of the one God +or gods, in whom according to his knowledge or superstition he may +believe; but in this case the additional fear of divine punishment often +supervenes. + + +_The strictly Social Virtues at first alone regarded._--The above view +of the first origin and nature of the moral sense, which tells us what +we ought to do, and of the conscience which reproves us if we disobey +it, accords well with what we see of the early and undeveloped condition +of this faculty in mankind. The virtues which must be practised, at +least generally, by rude men, so that they may associate in a body, are +those which are still recognised as the most important. But they are +practised almost exclusively in relation to the men of the same tribe; +and their opposites are not regarded as crimes in relation to the men of +other tribes. No tribe could hold together if murder, robbery, +treachery, &c., were common; consequently such crimes within the limits +of the same tribe "are branded with everlasting infamy;"[123] but excite +no such sentiment beyond these limits. A North-American Indian is well +pleased with himself, and is honoured by others, when he scalps a man of +another tribe; and a Dyak cuts off the head of an unoffending person +and dries it as a trophy. The murder of infants has prevailed on the +largest scale throughout the world,[124] and has met with no reproach; +but infanticide, especially of females, has been thought to be good for +the tribe, or at least not injurious. Suicide during former times was +not generally considered as a crime,[125] but rather from the courage +displayed as an honourable act; and it is still largely practised by +some semi-civilised nations without reproach, for the loss to a nation +of a single individual is not felt: whatever the explanation may be, +suicide, as I hear from Sir J. Lubbock, is rarely practised by the +lowest barbarians. It has been recorded that an Indian Thug +conscientiously regretted that he had not strangled and robbed as many +travellers as did his father before him. In a rude state of civilisation +the robbery of strangers is, indeed, generally considered as honourable. + +The great sin of Slavery has been almost universal, and slaves have +often been treated in an infamous manner. As barbarians do not regard +the opinion of their women, wives are commonly treated like slaves. Most +savages are utterly indifferent to the sufferings of strangers, or even +delight in witnessing them. It is well known that the women and children +of the North-American Indians aided in torturing their enemies. Some +savages take a horrid pleasure in cruelty to animals,[126] and humanity +with them is an unknown virtue. Nevertheless, feelings of sympathy and +kindness are common, especially during sickness, between the members of +the same tribe, and are sometimes extended beyond the limits of the +tribe. Mungo Park's touching account of the kindness of the negro women +of the interior to him is well known. Many instances could be given of +the noble fidelity of savages towards each other, but not to strangers; +common experience justifies the maxim of the Spaniard, "Never, never +trust an Indian." There cannot be fidelity without truth; and this +fundamental virtue is not rare between the members of the same tribe: +thus Mungo Park heard the negro women teaching their young children to +love the truth. This, again, is one of the virtues which becomes so +deeply rooted in the mind that it is sometimes practised by savages even +at a high cost, towards strangers; but to lie to your enemy has rarely +been thought a sin, as the history of modern diplomacy too plainly +shews. As soon as a tribe has a recognised leader, disobedience becomes +a crime, and even abject submission is looked at as a sacred virtue. + +As during rude times no man can be useful or faithful to his tribe +without courage, this quality has universally been placed in the highest +rank; and although, in civilised countries, a good, yet timid, man may +be far more useful to the community than a brave one, we cannot help +instinctively honouring the latter above a coward, however benevolent. +Prudence, on the other hand, which does not concern the welfare of +others, though a very useful virtue, has never been highly esteemed. As +no man can practise the virtues necessary for the welfare of his tribe +without self-sacrifice, self-command, and the power of endurance, these +qualities have been at all times highly and most justly valued. The +American savage voluntarily submits without a groan to the most horrid +tortures to prove and strengthen his fortitude and courage; and we +cannot help admiring him, or even an Indian Fakir, who, from a foolish +religious motive, swings suspended by a hook buried in his flesh. + +The other self-regarding virtues, which do not obviously, though they +may really, affect the welfare of the tribe, have never been esteemed by +savages, though now highly appreciated by civilised nations. The +greatest intemperance with savages is no reproach. Their utter +licentiousness, not to mention unnatural crimes, is something +astounding.[127] As soon, however, as marriage, whether polygamous or +monogamous, becomes common, jealousy will lead to the inculcation of +female virtue; and this being honoured will tend to spread to the +unmarried females. How slowly it spreads to the male sex we see at the +present day. Chastity eminently requires self-command; therefore it has +been honoured from a very early period in the moral history of civilised +man. As a consequence of this, the senseless practice of celibacy has +been ranked from a remote period as a virtue.[128] The hatred of +indecency, which appears to us so natural as to be thought innate, and +which is so valuable an aid to chastity, is a modern virtue, +appertaining exclusively, as Sir G. Staunton remarks,[129] to civilised +life. This is shewn by the ancient religious rites of various nations, +by the drawings on the walls of Pompeii, and by the practices of many +savages. + +We have now seen that actions are regarded by savages, and were probably +so regarded by primeval man, as good or bad, solely as they affect in an +obvious manner the welfare of the tribe,--not that of the species, nor +that of man as an individual member of the tribe. This conclusion +agrees well with the belief that the so-called moral sense is +aboriginally derived from the social instincts, for both relate at first +exclusively to the community. The chief causes of the low morality of +savages, as judged by our standard, are, firstly, the confinement of +sympathy to the same tribe. Secondly, insufficient powers of reasoning, +so that the bearing of many virtues, especially of the self-regarding +virtues, on the general welfare of the tribe is not recognised. Savages, +for instance, fail to trace the multiplied evils consequent on a want of +temperance, chastity, &c. And, thirdly, weak power of self-command; for +this power has not been strengthened through long-continued, perhaps +inherited, habit, instruction and religion. + +I have entered into the above details on the immorality of savages,[130] +because some authors have recently taken a high view of their moral +nature, or have attributed most of their crimes to mistaken +benevolence.[131] These authors appear to rest their conclusion on +savages possessing, as they undoubtedly do possess, and often in a high +degree, those virtues which are serviceable, or even necessary, for the +existence of a tribal community. + + +_Concluding Remarks._--Philosophers of the derivative[132] school of +morals formerly assumed that the foundation of morality lay in a form of +Selfishness; but more recently in the "Greatest Happiness principle." +According to the view given above, the moral sense is fundamentally +identical with the social instincts; and in the case of the lower +animals it would be absurd to speak of these instincts as having been +developed from selfishness, or for the happiness of the community. They +have, however, certainly been developed for the general good of the +community. The term, general good, may be defined as the means by which +the greatest possible number of individuals can be reared in full vigour +and health, with all their faculties perfect, under the conditions to +which they are exposed. As the social instincts both of man and the +lower animals have no doubt been developed by the same steps, it would +be advisable, if found practicable, to use the same definition in both +cases, and to take as the test of morality, the general good or welfare +of the community, rather than the general happiness; but this definition +would perhaps require some limitation on account of political ethics. + +When a man risks his life to save that of a fellow-creature, it seems +more appropriate to say that he acts for the general good or welfare, +rather than for the general happiness of mankind. No doubt the welfare +and the happiness of the individual usually coincide; and a contented, +happy tribe will flourish better than one that is discontented and +unhappy. We have seen that at an early period in the history of man, the +expressed wishes of the community will have naturally influenced to a +large extent the conduct of each member; and as all wish for happiness, +the "greatest happiness principle" will have become a most important +secondary guide and object; the social instincts, including sympathy, +always serving as the primary impulse and guide. Thus the reproach of +laying the foundation of the most noble part of our nature in the base +principle of selfishness is removed; unless indeed the satisfaction +which every animal feels when it follows its proper instincts, and the +dissatisfaction felt when prevented, be called selfish. + +The expression of the wishes and judgment of the members of the same +community, at first by oral and afterwards by written language, serves, +as just remarked, as a most important secondary guide of conduct, in aid +of the social instincts, but sometimes in opposition to them. This +latter fact is well exemplified by the _Law of Honour_, that is the law +of the opinion of our equals, and not of all our countrymen. The breach +of this law, even when the breach is known to be strictly accordant with +true morality, has caused many a man more agony than a real crime. We +recognise the same influence in the burning sense of shame which most of +us have felt even after the interval of years, when calling to mind some +accidental breach of a trifling though fixed rule of etiquette. The +judgment of the community will generally be guided by some rude +experience of what is best in the long run for all the members; but this +judgment will not rarely err from ignorance and from weak powers of +reasoning. Hence the strangest customs and superstitions, in complete +opposition to the true welfare and happiness of mankind, have become +all-powerful throughout the world. We see this in the horror felt by a +Hindoo who breaks his caste, in the shame of a Mahometan woman who +exposes her face, and in innumerable other instances. It would be +difficult to distinguish between the remorse felt by a Hindoo who has +eaten unclean food, from that felt after committing a theft; but the +former would probably be the more severe. + +How so many absurd rules of conduct, as well as so many absurd religious +beliefs, have originated we do not know; nor how it is that they have +become, in all quarters of the world, so deeply impressed on the mind +of men; but it is worthy of remark that a belief constantly inculcated +during the early years of life, whilst the brain is impressible, appears +to acquire almost the nature of an instinct; and the very essence of an +instinct is that it is followed independently of reason. Neither can we +say why certain admirable virtues, such as the love of truth, are much +more highly appreciated by some savage tribes than by others;[133] nor, +again, why similar differences prevail even amongst civilised nations. +Knowing how firmly fixed many strange customs and superstitions have +become, we need feel no surprise that the self-regarding virtues should +now appear to us so natural, supported as they are by reason, as to be +thought innate, although they were not valued by man in his early +condition. + +Notwithstanding many sources of doubt, man can generally and readily +distinguish between the higher and lower moral rules. The higher are +founded on the social instincts, and relate to the welfare of others. +They are supported by the approbation of our fellow-men and by reason. +The lower rules, though some of them when implying self-sacrifice hardly +deserve to be called lower, relate chiefly to self, and owe their origin +to public opinion, when matured by experience and cultivated; for they +are not practised by rude tribes. + +As man advances in civilisation, and small tribes are united into larger +communities, the simplest reason would tell each individual that he +ought to extend his social instincts and sympathies to all the members +of the same nation, though personally unknown to him. This point being +once reached, there is only an artificial barrier to prevent his +sympathies extending to the men of all nations and races. If, indeed, +such men are separated from him by great differences in appearance or +habits, experience unfortunately shews us how long it is before we look +at them as our fellow-creatures. Sympathy beyond the confines of man, +that is humanity to the lower animals, seems to be one of the latest +moral acquisitions. It is apparently unfelt by savages, except towards +their pets. How little the old Romans knew of it is shewn by their +abhorrent gladiatorial exhibitions. The very idea of humanity, as far as +I could observe, was new to most of the Gauchos of the Pampas. This +virtue, one of the noblest with which man is endowed, seems to arise +incidentally from our sympathies becoming more tender and more widely +diffused, until they are extended to all sentient beings. As soon as +this virtue is honoured and practised by some few men, it spreads +through instruction and example to the young, and eventually through +public opinion. + +The highest stage in moral culture at which we can arrive, is when we +recognise that we ought to control our thoughts, and "not even in inmost +thought to think again the sins that made the past so pleasant to +us."[134] Whatever makes any bad action familiar to the mind, renders +its performance by so much the easier. As Marcus Aurelius long ago said, +"Such as are thy habitual thoughts, such also will be the character of +thy mind; for the soul is dyed by the thoughts."[135] + +Our great philosopher, Herbert Spencer, has recently explained his views +on the moral sense. He says,[136] "I believe that the experiences of +utility organised and consolidated through all past generations of the +human race, have been producing corresponding modifications, which, by +continued transmission and accumulation, have become in us certain +faculties of moral intuition--certain emotions responding to right and +wrong conduct, which have no apparent basis in the individual +experiences of utility." There is not the least inherent improbability, +as it seems to me, in virtuous tendencies being more or less strongly +inherited; for, not to mention the various dispositions and habits +transmitted by many of our domestic animals, I have heard of cases in +which a desire to steal and a tendency to lie appeared to run in +families of the upper ranks; and as stealing is so rare a crime in the +wealthy classes, we can hardly account by accidental coincidence for the +tendency occurring in two or three members of the same family. If bad +tendencies are transmitted, it is probable that good ones are likewise +transmitted. Excepting through the principle of the transmission of +moral tendencies, we cannot understand the differences believed to exist +in this respect between the various races of mankind. We have, however, +as yet, hardly sufficient evidence on this head. + +Even the partial transmission of virtuous tendencies would be an immense +assistance to the primary impulse derived directly from the social +instincts, and indirectly from the approbation of our fellow-men. +Admitting for the moment that virtuous tendencies are inherited, it +appears probable, at least in such cases as chastity, temperance, +humanity to animals, &c., that they become first impressed on the mental +organisation through habit, instruction, and example, continued during +several generations in the same family, and in a quite subordinate +degree, or not at all, by the individuals possessing such virtues, +having succeeded best in the struggle for life. My chief source of doubt +with respect to any such inheritance, is that senseless customs, +superstitions, and tastes, such as the horror of a Hindoo for unclean +food, ought on the same principle to be transmitted. Although this in +itself is perhaps not less probable than that animals should acquire +inherited tastes for certain kinds of food or fear of certain foes, I +have not met with any evidence in support of the transmission of +superstitious customs or senseless habits. + + +Finally, the social instincts which no doubt were acquired by man, as by +the lower animals, for the good of the community, will from the first +have given to him some wish to aid his fellows, and some feeling of +sympathy. Such impulses will have served him at a very early period as a +rude rule of right and wrong. But as man gradually advanced in +intellectual power and was enabled to trace the more remote consequences +of his actions; as he acquired sufficient knowledge to reject baneful +customs and superstitions; as he regarded more and more not only the +welfare but the happiness of his fellow-men; as from habit, following on +beneficial experience, instruction, and example, his sympathies became +more tender and widely diffused, so as to extend to the men of all +races, to the imbecile, the maimed, and other useless members of +society, and finally to the lower animals,--so would the standard of his +morality rise higher and higher. And it is admitted by moralists of the +derivative school and by some intuitionists, that the standard of +morality has risen since an early period in the history of man.[137] + +As a struggle may sometimes be seen going on between the various +instincts of the lower animals, it is not surprising that there should +be a struggle in man between his social instincts, with their derived +virtues, and his lower, though at the moment, stronger impulses or +desires. This, as Mr. Galton[138] has remarked, is all the less +surprising, as man has emerged from a state of barbarism within a +comparatively recent period. After having yielded to some temptation we +feel a sense of dissatisfaction, analogous to that felt from other +unsatisfied instincts, called in this case conscience; for we cannot +prevent past images and impressions continually passing through our +minds, and these in their weakened state we compare with the +ever-present social instincts, or with habits gained in early youth and +strengthened during our whole lives, perhaps inherited, so that they are +at last rendered almost as strong as instincts. Looking to future +generations, there is no cause to fear that the social instincts will +grow weaker, and we may expect that virtuous habits will grow stronger, +becoming perhaps fixed by inheritance. In this case the struggle between +our higher and lower impulses will be less severe, and virtue will be +triumphant. + + +_Summary of the two last Chapters._--There can be no doubt that the +difference between the mind of the lowest man and that of the highest +animal is immense. An anthropomorphous ape, if he could take a +dispassionate view of his own case, would admit that though he could +form an artful plan to plunder a garden--though he could use stones for +fighting or for breaking open nuts, yet that the thought of fashioning +a stone into a tool was quite beyond his scope. Still less, as he would +admit, could he follow out a train of metaphysical reasoning, or solve a +mathematical problem, or reflect on God, or admire a grand natural +scene. Some apes, however, would probably declare that they could and +did admire the beauty of the coloured skin and fur of their partners in +marriage. They would admit, that though they could make other apes +understand by cries some of their perceptions and simpler wants, the +notion of expressing definite ideas by definite sounds had never crossed +their minds. They might insist that they were ready to aid their +fellow-apes of the same troop in many ways, to risk their lives for +them, and to take charge of their orphans; but they would be forced to +acknowledge that disinterested love for all living creatures, the most +noble attribute of man, was quite beyond their comprehension. + +Nevertheless the difference in mind between man and the higher animals, +great as it is, is certainly one of degree and not of kind. We have seen +that the senses and intuitions, the various emotions and faculties, such +as love, memory, attention, curiosity, imitation, reason, &c., of which +man boasts, may be found in an incipient, or even sometimes in a +well-developed condition, in the lower animals. They are also capable of +some inherited improvement, as we see in the domestic dog compared with +the wolf or jackal. If it be maintained that certain powers, such as +self-consciousness, abstraction, &c., are peculiar to man, it may well +be that these are the incidental results of other highly-advanced +intellectual faculties; and these again are mainly the result of the +continued use of a highly developed language. At what age does the +new-born infant possess the power of abstraction, or become +self-conscious and reflect on its own existence? We cannot answer; nor +can we answer in regard to the ascending organic scale. The half-art and +half-instinct of language still bears the stamp of its gradual +evolution. The ennobling belief in God is not universal with man; and +the belief in active spiritual agencies naturally follows from his other +mental powers. The moral sense perhaps affords the best and highest +distinction between man and the lower animals; but I need not say +anything on this head, as I have so lately endeavoured to shew +that the social instincts,--the prime principle of man's moral +constitution[139]--with the aid of active intellectual powers and the +effects of habit, naturally lead to the golden rule, "As ye would that +men should do to you, do ye to them likewise;" and this lies at the +foundation of morality. + +In a future chapter I shall make some few remarks on the probable steps +and means by which the several mental and moral faculties of man have +been gradually evolved. That this at least is possible ought not to be +denied, when we daily see their development in every infant; and when we +may trace a perfect gradation from the mind of an utter idiot, lower +than that of the lowest animal, to the mind of a Newton. + + + + +CHAPTER IV. + +ON THE MANNER OF DEVELOPMENT OF MAN FROM SOME LOWER FORM. + + + Variability of body and mind in man--Inheritance--Causes of + variability--Laws of variation the same in man as in the lower + animals--Direct action of the conditions of life--Effects of + the increased use and disuse of parts--Arrested + development--Reversion--Correlated variation--Rate of + increase--Checks to increase--Natural selection--Man the most + dominant animal in the world--Importance of his corporeal + structure--The causes which have led to his becoming + erect--Consequent changes of structure--Decrease in size of the + canine teeth--Increased size and altered shape of the + skull--Nakedness--Absence of a tail--Defenceless condition of + man. + + +We have seen in the first chapter that the homological structure of man, +his embryological development and the rudiments which he still retains, +all declare in the plainest manner that he is descended from some lower +form. The possession of exalted mental powers is no insuperable +objection to this conclusion. In order that an ape-like creature should +have been transformed into man, it is necessary that this early form, as +well as many successive links, should all have varied in mind and body. +It is impossible to obtain direct evidence on this head; but if it can +be shewn that man now varies--that his variations are induced by the +same general causes, and obey the same general laws, as in the case of +the lower animals--there can be little doubt that the preceding +intermediate links varied in a like manner. The variations at each +successive stage of descent must, also, have been in some manner +accumulated and fixed. + +The facts and conclusions to be given in this chapter relate almost +exclusively to the probable means by which the transformation of man has +been effected, as far as his bodily structure is concerned. The +following chapter will be devoted to the development of his intellectual +and moral faculties. But the present discussion likewise bears on the +origin of the different races or species of mankind, whichever term may +be preferred. + +It is manifest that man is now subject to much variability. No two +individuals of the same race are quite alike. We may compare millions of +faces, and each will be distinct. There is an equally great amount of +diversity in the proportions and dimensions of the various parts of the +body; the length of the legs being one of the most variable points.[140] +Although in some quarters of the world an elongated skull, and in other +quarters a short skull prevails, yet there is great diversity of shape +even within the limits of the same race, as with the aborigines of +America and South Australia,--the latter a race "probably as pure and +homogeneous in blood, customs, and language as any in existence"--and +even with the inhabitants of so confined an area as the Sandwich +Islands.[141] An eminent dentist assures me that there is nearly as much +diversity in the teeth, as in the features. The chief arteries so +frequently run in abnormal courses, that it has been found useful for +surgical purposes to calculate from 12,000 corpses how often each +course prevails.[142] The muscles are eminently variable: thus those of +the foot were found by Prof. Turner[143] not to be strictly alike in any +two out of fifty bodies; and in some the deviations were considerable. +Prof. Turner adds that the power of performing the appropriate movements +must have been modified in accordance with the several deviations. Mr. +J. Wood has recorded[144] the occurrence of 295 muscular variations in +thirty-six subjects, and in another set of the same number no less than +558 variations, reckoning both sides of the body as one. In the last +set, not one body out of the thirty-six was "found totally wanting in +departures from the standard descriptions of the muscular system given +in anatomical text-books." A single body presented the extraordinary +number of twenty-five distinct abnormalities. The same muscle sometimes +varies in many ways: thus Prof. Macalister describes[145] no less than +twenty distinct variations in the _palmaris accessorius_. + +The famous old anatomist, Wolff,[146] insists that the internal viscera +are more variable than the external parts: _Nulla particula est qu non +aliter et aliter in aliis se habeat hominibus._ He has even written a +treatise on the choice of typical examples of the viscera for +representation. A discussion on the beau-ideal of the liver, lungs, +kidneys, &c., as of the human face divine, sounds strange in our ears. + +The variability or diversity of the mental faculties in men of the same +race, not to mention the greater differences between the men of +distinct races, is so notorious that not a word need here be said. So it +is with the lower animals, as has been illustrated by a few examples in +the last chapter. All who have had charge of menageries admit this fact, +and we see it plainly in our dogs and other domestic animals. Brehm +especially insists that each individual monkey of those which he kept +under confinement in Africa had its own peculiar disposition and temper: +he mentions one baboon remarkable for its high intelligence; and the +keepers in the Zoological Gardens pointed out to me a monkey, belonging +to the New World division, equally remarkable for intelligence. Rengger, +also, insists on the diversity in the various mental characters of the +monkeys of the same species which he kept in Paraguay; and this +diversity, as he adds, is partly innate, and partly the result of the +manner in which they have been treated or educated.[147] + +I have elsewhere[148] so fully discussed the subject of Inheritance that +I need here add hardly anything. A greater number of facts have been +collected with respect to the transmission of the most trifling, as well +as of the most important characters in man than in any of the lower +animals; though the facts are copious enough with respect to the latter. +So in regard to mental qualities, their transmission is manifest in our +dogs, horses, and other domestic animals. Besides special tastes and +habits, general intelligence, courage, bad and good temper, &c., are +certainly transmitted. With man we see similar facts in almost every +family; and we now know through the admirable labours of Mr. Galton[149] +that genius, which implies a wonderfully complex combination of high +faculties, tends to be inherited; and, on the other hand, it is too +certain that insanity and deteriorated mental powers likewise run in the +same families. + +With respect to the causes of variability we are in all cases very +ignorant; but we can see that in man as in the lower animals, they stand +in some relation with the conditions to which each species has been +exposed during several generations. Domesticated animals vary more than +those in a state of nature; and this is apparently due to the +diversified and changing nature of their conditions. The different races +of man resemble in this respect domesticated animals, and so do the +individuals of the same race when inhabiting a very wide area, like that +of America. We see the influence of diversified conditions in the more +civilised nations, the members of which belong to different grades of +rank and follow different occupations, presenting a greater range of +character than the members of barbarous nations. But the uniformity of +savages has often been exaggerated, and in some cases can hardly be said +to exist.[150] It is nevertheless an error to speak of man, even if we +look only to the conditions to which he has been subjected, as "far more +domesticated"[151] than any other animal. Some savage races, such as +the Australians, are not exposed to more diversified conditions than are +many species which have very wide ranges. In another and much more +important respect, man differs widely from any strictly domesticated +animal; for his breeding has not been controlled, either through +methodical or unconscious selection. No race or body of men has been so +completely subjugated by other men, that certain individuals have been +preserved and thus unconsciously selected, from being in some way more +useful to their masters. Nor have certain male and female individuals +been intentionally picked out and matched, except in the well-known case +of the Prussian grenadiers; and in this case man obeyed, as might have +been expected, the law of methodical selection; for it is asserted that +many tall men were reared in the villages inhabited by the grenadiers +with their tall wives. + +If we consider all the races of man, as forming a single species, his +range is enormous; but some separate races, as the Americans and +Polynesians, have very wide ranges. It is a well-known law that +widely-ranging species are much more variable than species with +restricted ranges; and the variability of man may with more truth be +compared with that of widely-ranging species, than with that of +domesticated animals. + +Not only does variability appear to be induced in man and the lower +animals by the same general causes, but in both the same characters are +affected in a closely analogous manner. This has been proved in such +full detail by Godron and Quatrefages, that I need here only refer to +their works.[152] Monstrosities, which graduate into slight variations, +are likewise so similar in man and the lower animals, that the same +classification and the same terms can be used for both, as may be seen +in Isidore Geoffroy St.-Hilaire's great work.[153] This is a necessary +consequence of the same laws of change prevailing throughout the animal +kingdom. In my work on the variation of domestic animals, I have +attempted to arrange in a rude fashion the laws of variation under the +following heads:--The direct and definite action of changed conditions, +as shewn by all or nearly all the individuals of the same species +varying in the same manner under the same circumstances. The effects of +the long-continued use or disuse of parts. The cohesion of homologous +parts. The variability of multiple parts. Compensation of growth; but of +this law I have found no good instances in the case of man. The effects +of the mechanical pressure of one part on another; as of the pelvis on +the cranium of the infant in the womb. Arrests of development, leading +to the diminution or suppression of parts. The reappearance of long-lost +characters through reversion. And lastly, correlated variation. All +these so-called laws apply equally to man and the lower animals; and +most of them even to plants. It would be superfluous here to discuss all +of them;[154] but several are so important for us, that they must be +treated at considerable length. + + +_The direct and definite action of changed conditions._--This is a most +perplexing subject. It cannot be denied that changed conditions produce +some effect, and occasionally a considerable effect, on organisms of all +kinds; and it seems at first probable that if sufficient time were +allowed this would be the invariable result. But I have failed to obtain +clear evidence in favour of this conclusion; and valid reasons may be +urged on the other side, at least as far as the innumerable structures +are concerned, which are adapted for special ends. There can, however, +be no doubt that changed conditions induce an almost indefinite amount +of fluctuating variability, by which the whole organisation is rendered +in some degree plastic. + +In the United States, above 1,000,000 soldiers, who served in the late +war, were measured, and the States in which they were born and reared +recorded.[155] From this astonishing number of observations it is proved +that local influences of some kind act directly on stature; and we +further learn that "the State where the physical growth has in great +measure taken place, and the State of birth, which indicates the +ancestry, seem to exert a marked influence on the stature." For instance +it is established, "that residence in the Western States, during the +years of growth, tends to produce increase of stature." On the other +hand, it is certain that with sailors, their manner of life delays +growth, as shewn "by the great difference between the statures of +soldiers and sailors at the ages of 17 and 18 years." Mr. B. A. Gould +endeavoured to ascertain the nature of the influences which thus act on +stature; but he arrived only at negative results, namely, that they did +not relate to climate, the elevation of the land, soil, nor even "in any +controlling degree" to the abundance or need of the comforts of life. +This latter conclusion is directly opposed to that arrived at by +Villerm from the statistics of the height of the conscripts in +different parts of France. When we compare the differences in stature +between the Polynesian chiefs and the lower orders within the same +islands, or between the inhabitants of the fertile volcanic and low +barren coral islands of the same ocean,[156] or again between the +Fuegians on the eastern and western shores of their country, where the +means of subsistence are very different, it is scarcely possible to +avoid the conclusion that better food and greater comfort do influence +stature. But the preceding statements shew how difficult it is to arrive +at any precise result. Dr. Beddoe has lately proved that, with the +inhabitants of Britain, residence in towns and certain occupations have +a deteriorating influence on height; and he infers that the result is to +a certain extent inherited, as is likewise the case in the United +States. Dr. Beddoe further believes that wherever a "race attains its +maximum of physical development, it rises highest in energy and moral +vigour."[157] + +Whether external conditions produce any other direct effect on man is +not known. It might have been expected that differences of climate would +have had a marked influence, as the lungs and kidneys are brought into +fuller activity under a low temperature, and the liver and skin under a +high one.[158] It was formerly thought that the colour of the skin and +the character of the hair were determined by light or heat; and +although it can hardly be denied that some effect is thus produced, +almost all observers now agree that the effect has been very small, even +after exposure during many ages. But this subject will be more properly +discussed when we treat of the different races of mankind. With our +domestic animals there are grounds for believing that cold and damp +directly affect the growth of the hair; but I have not met with any +evidence on this head in the case of man. + + +_Effects of the increased Use and Disuse of Parts._--It is well known +that use strengthens the muscles in the individual, and complete disuse, +or the destruction of the proper nerve, weakens them. When the eye is +destroyed the optic nerve often becomes atrophied. When an artery is +tied, the lateral channels increase not only in diameter, but in the +thickness and strength of their coats. When one kidney ceases acting +from disease, the other increases in size and does double work. Bones +increase not only in thickness, but in length, from carrying a greater +weight.[159] Different occupations habitually followed lead to changed +proportions in various parts of the body. Thus it was clearly +ascertained by the United States Commission[160] that the legs of the +sailors employed in the late war were longer by 0.217 of an inch than +those of the soldiers, though the sailors were on an average shorter +men; whilst their arms were shorter by 1.09 of an inch, and therefore +out of proportion shorter in relation to their lesser height. This +shortness of the arms is apparently due to their greater use, and is an +unexpected result; but sailors chiefly use their arms in pulling and not +in supporting weights. The girth of the neck and the depth of the instep +are greater, whilst the circumference of the chest, waist, and hips is +less in sailors than in soldiers. + +Whether the several foregoing modifications would become hereditary, if +the same habits of life were followed during many generations, is not +known, but is probable. Rengger[161] attributes the thin legs and thick +arms of the Payaguas Indians to successive generations having passed +nearly their whole lives in canoes, with their lower extremities +motionless. Other writers have come to a similar conclusion in other +analogous cases. According to Cranz,[162] who lived for a long time with +the Esquimaux, "the natives believe that ingenuity and dexterity in +seal-catching (their highest art and virtue) is hereditary; there is +really something in it, for the son of a celebrated seal-catcher will +distinguish himself though he lost his father in childhood." But in this +case it is mental aptitude, quite as much as bodily structure, which +appears to be inherited. It is asserted that the hands of English +labourers are at birth larger than those of the gentry.[163] From the +correlation which exists, at least in some cases,[164] between the +development of the extremities and of the jaws, it is possible that in +those classes which do not labour much with their hands and feet, the +jaws would be reduced in size from this cause. That they are generally +smaller in refined and civilised men than in hard-working men or +savages, is certain. But with savages, as Mr. Herbert Spencer[165] has +remarked, the greater use of the jaws in chewing coarse, uncooked food, +would act in a direct manner on the masticatory muscles and on the bones +to which they are attached. In infants long before birth, the skin on +the soles of the feet is thicker than on any other part of the +body;[166] and it can hardly be doubted that this is due to the +inherited effects of pressure during a long series of generations. + +It is familiar to every one that watchmakers and engravers are liable to +become short-sighted, whilst sailors and especially savages are +generally long-sighted. Short-sight and long-sight certainly tend to be +inherited.[167] The inferiority of Europeans, in comparison with +savages, in eyesight and in the other senses, is no doubt the +accumulated and transmitted effect of lessened use during many +generations; for Rengger[168] states that he has repeatedly observed +Europeans, who had been brought up and spent their whole lives with the +wild Indians, who nevertheless did not equal them in the sharpness of +their senses. The same naturalist observes that the cavities in the +skull for the reception of the several sense-organs are larger in the +American aborigines than in Europeans; and this no doubt indicates a +corresponding difference in the dimensions of the organs themselves. +Blumenbach has also remarked on the large size of the nasal cavities in +the skulls of the American aborigines, and connects this fact with their +remarkably acute power of smell. The Mongolians of the plains of +Northern Asia, according to Pallas, have wonderfully perfect senses; and +Prichard believes that the great breadth of their skulls across the +zygomas follows from their highly-developed sense-organs.[169] + +The Quechua Indians inhabit the lofty plateaux of Peru, and Alcide +d'Orbigny states[170] that from continually breathing a highly rarefied +atmosphere they have acquired chests and lungs of extraordinary +dimensions. The cells, also, of the lungs are larger and more numerous +than in Europeans. These observations have been doubted; but Mr. D. +Forbes carefully measured many Aymaras, an allied race, living at the +height of between ten and fifteen thousand feet; and he informs me[171] +that they differ conspicuously from the men of all other races seen by +him, in the circumference and length of their bodies. In his table of +measurements, the stature of each man is taken at 1000, and the other +measurements are reduced to this standard. It is here seen that the +extended arms of the Aymaras are shorter than those of Europeans, and +much shorter than those of Negroes. The legs are likewise shorter, and +they present this remarkable peculiarity, that in every Aymara measured +the femur is actually shorter than the tibia. On an average the length +of the femur to that of the tibia is as 211 to 252; whilst in two +Europeans measured at the same time, the femora to the tibi were as +244 to 230; and in three Negroes as 258 to 241. The humerus is likewise +shorter relatively to the fore-arm. This shortening of that part of the +limb which is nearest to the body, appears to be, as suggested to me by +Mr. Forbes, a case of compensation in relation with the greatly +increased length of the trunk. The Aymaras present some other singular +points of structure, for instance, the very small projection of the +heel. + +These men are so thoroughly acclimatised to their cold and lofty abode, +that when formerly carried down by the Spaniards to the low Eastern +plains, and when now tempted down by high wages to the gold-washings, +they suffer a frightful rate of mortality. Nevertheless Mr. Forbes found +a few pure families which had survived during two generations; and he +observed that they still inherited their characteristic peculiarities. +But it was manifest, even without measurement, that these peculiarities +had all decreased; and on measurement their bodies were found not to be +so much elongated as those of the men on the high plateau; whilst their +femora had become somewhat lengthened, as had their tibi but in a less +degree. The actual measurements may be seen by consulting Mr. Forbes' +memoir. From these valuable observations, there can, I think, be no +doubt that residence during many generations at a great elevation tends, +both directly and indirectly, to induce inherited modifications in the +proportions of the body.[172] + +Although man may not have been much modified during the latter stages of +his existence through the increased or decreased use of parts, the +facts now given shew that his liability in this respect has not been +lost; and we positively know that the same law holds good with the lower +animals. Consequently we may infer, that when at a remote epoch the +progenitors of man were in a transitional state, and were changing from +quadrupeds into bipeds, natural selection would probably have been +greatly aided by the inherited effects of the increased or diminished +use of the different parts of the body. + + +_Arrests of Development._--Arrested development differs from arrested +growth, as parts in the former state continue to grow whilst still +retaining their early condition. Various monstrosities come under this +head, and some are known to be occasionally inherited, as a +cleft-palate. It will suffice for our purpose to refer to the arrested +brain-development of microcephalous idiots, as described in Vogt's great +memoir.[173] Their skulls are smaller, and the convolutions of the brain +are less complex than in normal men. The frontal sinus, or the +projection over the eyebrows, is largely developed, and the jaws are +prognathous to an "_effrayant_" degree; so that these idiots somewhat +resemble the lower types of mankind. Their intelligence and most of +their mental faculties are extremely feeble. They cannot acquire the +power of speech, and are wholly incapable of prolonged attention, but +are much given to imitation. They are strong and remarkably active, +continually gamboling and jumping about, and making grimaces. They often +ascend stairs on all-fours; and are curiously fond of climbing up +furniture or trees. We are thus reminded of the delight shewn by almost +all boys in climbing trees; and this again reminds us how lambs and +kids, originally alpine animals, delight to frisk on any hillock, +however small. + + +_Reversion._--Many of the cases to be here given might have been +introduced under the last heading. Whenever a structure is arrested in +its development, but still continues growing until it closely resembles +a corresponding structure in some lower and adult member of the same +group, we may in one sense consider it as a case of reversion. The lower +members in a group give us some idea how the common progenitor of the +group was probably constructed; and it is hardly credible that a part +arrested at an early phase of embryonic development should be enabled to +continue growing so as ultimately to perform its proper function, unless +it had acquired this power of continued growth during some earlier state +of existence, when the present exceptional or arrested structure was +normal. The simple brain of a microcephalous idiot, in as far as it +resembles that of an ape, may in this sense be said to offer a case of +reversion. There are other cases which come more strictly under our +present heading of reversion. Certain structures, regularly occurring in +the lower members of the group to which man belongs, occasionally make +their appearance in him, though not found in the normal human embryo; +or, if present in the normal human embryo, they become developed in an +abnormal manner, though this manner of development is proper to the +lower members of the same group. These remarks will be rendered clearer +by the following illustrations. + +In various mammals the uterus graduates from a double organ with two +distinct orifices and two passages, as in the marsupials, into a single +organ, showing no signs of doubleness except a slight internal fold, as +in the higher apes and man. The rodents exhibit a perfect series of +gradations between these two extreme states. In all mammals the uterus +is developed from two simple primitive tubes, the inferior portions of +which form the cornua; and it is in the words of Dr. Farre "by the +coalescence of the two cornua at their lower extremities that the body +of the uterus is formed in man; while in those animals in which no +middle portion or body exists, the cornua remain ununited. As the +development of the uterus proceeds, the two cornua become gradually +shorter, until at length they are lost, or, as it were, absorbed into +the body of the uterus." The angles of the uterus are still produced +into cornua, even so high in the scale as in the lower apes, and their +allies the lemurs. + +Now in women anomalous cases are not very infrequent, in which the +mature uterus is furnished with cornua, or is partially divided into two +organs; and such cases, according to Owen, repeat "the grade of +concentrative development," attained by certain rodents. Here perhaps we +have an instance of a simple arrest of embryonic development, with +subsequent growth and perfect functional development, for either side of +the partially double uterus is capable of performing the proper office +of gestation. In other and rarer cases, two distinct uterine cavities +are formed, each having its proper orifice and passage.[174] No such +stage is passed through during the ordinary development of the embryo, +and it is difficult to believe, though perhaps not impossible, that the +two simple, minute, primitive tubes could know how (if such an +expression may be used) to grow into two distinct uteri, each with a +well-constructed orifice and passage, and each furnished with numerous +muscles, nerves, glands and vessels, if they had not formerly passed +through a similar course of development, as in the case of existing +marsupials. No one will pretend that so perfect a structure as the +abnormal double uterus in woman could be the result of mere chance. But +the principle of reversion, by which long-lost dormant structures are +called back into existence, might serve as the guide for the full +development of the organ, even after the lapse of an enormous interval +of time. + +Professor Canestrini,[175] after discussing the foregoing and various +analogous cases, arrives at the same conclusion as that just given. He +adduces, as another instance, the malar bone, which, in some of the +Quadrumana and other mammals, normally consists of two portions. This is +its condition in the two-months-old human foetus; and thus it +sometimes remains, through arrested development, in man when adult, more +especially in the lower prognathous races. Hence Canestrini concludes +that some ancient progenitor of man must have possessed this bone +normally divided into two portions, which subsequently became fused +together. In man the frontal bone consists of a single piece, but in the +embryo and in children, and in almost all the lower mammals, it consists +of two pieces separated by a distinct suture. This suture occasionally +persists, more or less distinctly, in man after maturity, and more +frequently in ancient than in recent crania, especially as Canestrini +has observed in those exhumed from the Drift and belonging to the +brachycephalic type. Here again he comes to the same conclusion as in +the analogous case of the malar bones. In this and other instances +presently to be given, the cause of ancient races approaching the lower +animals in certain characters more frequently than do the modern races, +appears to be that the latter stand at a somewhat greater distance in +the long line of descent from their early semi-human progenitors. + +Various other anomalies in man, more or less analogous with the +foregoing, have been advanced by different authors[176] as cases of +reversion; but these seem not a little doubtful, for we have to descend +extremely low in the mammalian series before we find such structures +normally present.[177] + +In man the canine teeth are perfectly efficient instruments for +mastication. But their true canine character, as Owen[178] remarks, "is +indicated by the conical form of the crown, which terminates in an +obtuse point, is convex outward and flat or sub-concave within, at the +base of which surface there is a feeble prominence. The conical form is +best expressed in the Melanian races, especially the Australian. The +canine is more deeply implanted, and by a stronger fang than the +incisors." Nevertheless this tooth no longer serves man as a special +weapon for tearing his enemies or prey; it may, therefore, as far as its +proper function is concerned, be considered as rudimentary. In every +large collection of human skulls some may be found, as Hckel[179] +observes, with the canine teeth projecting considerably beyond the +others in the same manner, but in a less degree, as in the +anthropomorphous apes. In these cases, open spaces between the teeth in +the one jaw are left for the reception of the canines belonging to the +opposite jaw. An interspace of this kind in a Kaffir skull, figured by +Wagner, is surprisingly wide.[180] Considering how few ancient skulls +have been examined in comparison with recent skulls, it is an +interesting fact that in at least three cases the canines project +largely; and in the Naulette jaw they are spoken of as enormous.[181] + +The males alone of the anthropomorphous apes have their canines fully +developed; but in the female gorilla, and in a less degree in the female +orang, these teeth project considerably beyond the others; therefore the +fact that women sometimes have, as I have been assured, considerably +projecting canines, is no serious objection to the belief that their +occasional great development in man is a case of reversion to an +ape-like progenitor. He who rejects with scorn the belief that the shape +of his own canines, and their occasional great development in other men, +are due to our early progenitors having been provided with these +formidable weapons, will probably reveal by sneering the line of his +descent. For though he no longer intends, nor has the power, to use +these teeth as weapons, he will unconsciously retract his "snarling +muscles" (thus named by Sir C. Bell)[182] so as to expose them ready for +action, like a dog prepared to fight. + +Many muscles are occasionally developed in man, which are proper to the +Quadrumana or other mammals. Professor Vlacovich[183] examined forty +male subjects, and found a muscle, called by him the ischiopubic, in +nineteen of them; in three others there was a ligament which represented +this muscle; and in the remaining eighteen no trace of it. Out of thirty +female subjects this muscle was developed on both sides in only two, but +in three others the rudimentary ligament was present. This muscle, +therefore, appears to be much more common in the male than in the female +sex; and on the principle of the descent of man from some lower form, +its presence can be understood; for it has been detected in several of +the lower animals, and in all of these it serves exclusively to aid the +male in the act of reproduction. + +Mr. J. Wood, in his valuable series of papers,[184] has minutely +described a vast number of muscular variations in man, which resemble +normal structures in the lower animals. Looking only to the muscles +which closely resemble those regularly present in our nearest allies, +the Quadrumana, they are too numerous to be here even specified. In a +single male subject, having a strong bodily frame and well-formed skull, +no less than seven muscular variations were observed, all of which +plainly represented muscles proper to various kinds of apes. This man, +for instance, had on both sides of his neck a true and powerful +"_levator clavicul_," such as is found in all kinds of apes, and which +is said to occur in about one out of sixty human subjects.[185] Again, +this man had "a special abductor of the metatarsal bone of the fifth +digit, such as Professor Huxley and Mr. Flower have shewn to exist +uniformly in the higher and lower apes." The hands and arms of man are +eminently characteristic structures, but their muscles are extremely +liable to vary, so as to resemble the corresponding muscles in the lower +animals.[186] Such resemblances are either complete and perfect or +imperfect, yet in this latter case manifestly of a transitional nature. +Certain variations are more common in man, and others in woman, without +our being able to assign any reason. Mr. Wood, after describing numerous +cases, makes the following pregnant remark: "Notable departures from the +ordinary type of the muscular structures run in grooves or directions, +which must be taken to indicate some unknown factor, of much importance +to a comprehensive knowledge of general and scientific anatomy."[187] + +That this unknown factor is reversion to a former state of existence may +be admitted as in the highest degree probable. It is quite incredible +that a man should through mere accident abnormally resemble, in no less +than seven of his muscles, certain apes, if there had been no genetic +connection between them. On the other hand, if man is descended from +some ape-like creature, no valid reason can be assigned why certain +muscles should not suddenly reappear after an interval of many thousand +generations, in the same manner as with horses, asses, and mules, +dark-coloured stripes suddenly reappear on the legs and shoulders, after +an interval of hundreds, or more probably thousands, of generations. + +These various cases of reversion are so closely related to those of +rudimentary organs given in the first chapter, that many of them might +have been indifferently introduced in either chapter. Thus a human +uterus furnished with cornua may be said to represent in a rudimentary +condition the same organ in its normal state in certain mammals. Some +parts which are rudimental in man, as the os coccyx in both sexes and +the mamm in the male sex, are always present; whilst others, such as +the supra-condyloid foramen, only occasionally appear, and therefore +might have been introduced under the head of reversion. These several +reversionary, as well as the strictly rudimentary, structures reveal the +descent of man from some lower form in an unmistakeable manner. + + +_Correlated Variation._--In man, as in the lower animals, many +structures are so intimately related, that when one part varies so does +another, without our being able, in most cases, to assign any reason. We +cannot say whether the one part governs the other, or whether both are +governed by some earlier developed part. Various monstrosities, as I. +Geoffroy repeatedly insists, are thus intimately connected. Homologous +structures are particularly liable to change together, as we see on the +opposite sides of the body, and in the upper and lower extremities. +Meckel long ago remarked that when the muscles of the arm depart from +their proper type, they almost always imitate those of the leg; and so +conversely with the muscles of the legs. The organs of sight and +hearing, the teeth and hair, the colour of the skin and hair, colour and +constitution, are more or less correlated.[188] Professor Schaaffhausen +first drew attention to the relation apparently existing between a +muscular frame and strongly-pronounced supra-orbital ridges, which are +so characteristic of the lower races of man. + +Besides the variations which can be grouped with more or less +probability under the foregoing heads, there is a large class of +variations which may be provisionally called spontaneous, for they +appear, owing to our ignorance, to arise without any exciting cause. It +can, however, be shewn that such variations, whether consisting of +slight individual differences, or of strongly-marked and abrupt +deviations of structure, depend much more on the constitution of the +organism than on the nature of the conditions to which it has been +subjected.[189] + + +_Rate of Increase._--Civilised populations have been known under +favourable conditions, as in the United States, to double their number +in twenty-five years; and according to a calculation by Euler, this +might occur in a little over twelve years.[190] At the former rate the +present population of the United States, namely, thirty millions, would +in 657 years cover the whole terraqueous globe so thickly, that four men +would have to stand on each square yard of surface. The primary or +fundamental check to the continued increase of man is the difficulty of +gaining subsistence and of living in comfort. We may infer that this is +the case from what we see, for instance, in the United States, where +subsistence is easy and there is plenty of room. If such means were +suddenly doubled in Great Britain, our number would be quickly doubled. +With civilised nations the above primary check acts chiefly by +restraining marriages. The greater death-rate of infants in the poorest +classes is also very important; as well as the greater mortality at all +ages, and from various diseases, of the inhabitants of crowded and +miserable houses. The effects of severe epidemics and wars are soon +counterbalanced, and more than counterbalanced, in nations placed under +favourable conditions. Emigration also comes in aid as a temporary +check, but not to any great extent with the extremely poor classes. + +There is reason to suspect, as Malthus has remarked, that the +reproductive power is actually less in barbarous than in civilised +races. We know nothing positively on this head, for with savages no +census has been taken; but from the concurrent testimony of +missionaries, and of others who have long resided with such people, it +appears that their families are usually small, and large ones rare. This +may be partly accounted for, as it is believed, by the women suckling +their infants for a prolonged period; but it is highly probable that +savages, who often suffer much hardship, and who do not obtain so much +nutritious food as civilised men, would be actually less prolific. I +have shewn in a former work,[191] that all our domesticated quadrupeds +and birds, and all our cultivated plants, are more fertile than the +corresponding species in a state of nature. It is no valid objection to +this conclusion that animals suddenly supplied with an excess of food, +or when rendered very fat, and that most plants when suddenly removed +from very poor to very rich soil, are rendered more or less sterile. We +might, therefore, expect that civilised men, who in one sense are highly +domesticated, would be more prolific than wild men. It is also probable +that the increased fertility of civilised nations would become, as with +our domestic animals, an inherited character: it is at least known that +with mankind a tendency to produce twins runs in families.[192] + +Notwithstanding that savages appear to be less prolific than civilised +people, they would no doubt rapidly increase if their numbers were not +by some means rigidly kept down. The Santali, or hill-tribes of India, +have recently afforded a good illustration of this fact; for they have +increased, as shewn by Mr. Hunter,[193] at an extraordinary rate since +vaccination has been introduced, other pestilences mitigated, and war +sternly repressed. This increase, however, would not have been possible +had not these rude people spread into the adjoining districts and worked +for hire. Savages almost always marry; yet there is some prudential +restraint, for they do not commonly marry at the earliest possible age. +The young men are often required to show that they can support a wife, +and they generally have first to earn the price with which to purchase +her from her parents. With savages the difficulty of obtaining +subsistence occasionally limits their number in a much more direct +manner than with civilised people, for all tribes periodically suffer +from severe famines. At such times savages are forced to devour much bad +food, and their health can hardly fail to be injured. Many accounts have +been published of their protruding stomachs and emaciated limbs after +and during famines. They are then, also, compelled to wander much about, +and their infants, as I was assured in Australia, perish in large +numbers. As famines are periodical, depending chiefly on extreme +seasons, all tribes must fluctuate in number. They cannot steadily and +regularly increase, as there is no artificial increase in the supply of +food. Savages when hardly pressed encroach on each other's territories, +and war is the result; but they are indeed almost always at war with +their neighbours. They are liable to many accidents on land and water in +their search for food; and in some countries they must suffer much from +the larger beasts of prey. Even in India, districts have been +depopulated by the ravages of tigers. + +Malthus has discussed these several checks, but he does not lay stress +enough on what is probably the most important of all, namely +infanticide, especially of female infants, and the habit of procuring +abortion. These practices now prevail in many quarters of the world, and +infanticide seems formerly to have prevailed, as Mr. M'Lennan[194] has +shewn, on a still more extensive scale. These practices appear to have +originated in savages recognising the difficulty, or rather the +impossibility of supporting all the infants that are born. +Licentiousness may also be added to the foregoing checks; but this does +not follow from failing means of subsistence; though there is reason to +believe that in some cases (as in Japan) it has been intentionally +encouraged as a means of keeping down the population. + +If we look back to an extremely remote epoch, before man had arrived at +the dignity of manhood, he would have been guided more by instinct and +less by reason than are savages at the present time. Our early +semi-human progenitors would not have practised infanticide, for the +instincts of the lower animals are never so perverted as to lead them +regularly to destroy their own offspring. There would have been no +prudential restraint from marriage, and the sexes would have freely +united at an early age. Hence the progenitors of man would have tended +to increase rapidly, but checks of some kind, either periodical or +constant, must have kept down their numbers, even more severely than +with existing savages. What the precise nature of these checks may have +been, we cannot say, any more than with most other animals. We know that +horses and cattle, which are not highly prolific animals, when first +turned loose in South America, increased at an enormous rate. The +slowest breeder of all known animals, namely the elephant, would in a +few thousand years stock the whole world. The increase of every species +of monkey must be checked by some means; but not, as Brehm remarks, by +the attacks of beasts of prey. No one will assume that the actual power +of reproduction in the wild horses and cattle of America, was at first +in any sensible degree increased; or that, as each district became fully +stocked, this same power was diminished. No doubt in this case and in +all others, many checks concur, and different checks under different +circumstances; periodical dearths, depending on unfavourable seasons, +being probably the most important of all. So it will have been with the +early progenitors of man. + + +_Natural Selection._--We have now seen that man is variable in body and +mind; and that the variations are induced, either directly or +indirectly, by the same general causes, and obey the same general laws, +as with the lower animals. Man has spread widely over the face of the +earth, and must have been exposed, during his incessant migrations,[195] +to the most diversified conditions. The inhabitants of Tierra del +Fuego, the Cape of Good Hope, and Tasmania in the one hemisphere, and of +the Arctic regions in the other, must have passed through many climates +and changed their habits many times, before they reached their present +homes.[196] The early progenitors of man must also have tended, like all +other animals, to have increased beyond their means of subsistence; they +must therefore occasionally have been exposed to a struggle for +existence, and consequently to the rigid law of natural selection. +Beneficial variations of all kinds will thus, either occasionally or +habitually, have been preserved, and injurious ones eliminated. I do not +refer to strongly-marked deviations of structure, which occur only at +long intervals of time, but to mere individual differences. We know, for +instance, that the muscles of our hands and feet, which determine our +powers of movement, are liable, like those of the lower animals,[197] to +incessant variability. If then the ape-like progenitors of man which +inhabited any district, especially one undergoing some change in its +conditions, were divided into two equal bodies, the one half which +included all the individuals best adapted by their powers of movement +for gaining subsistence or for defending themselves, would on an average +survive in greater number and procreate more offspring than the other +and less well endowed half. + +Man in the rudest state in which he now exists is the most dominant +animal that has ever appeared on the earth. He has spread more widely +than any other highly organised form; and all others have yielded +before him. He manifestly owes this immense superiority to his +intellectual faculties, his social habits, which lead him to aid and +defend his fellows, and to his corporeal structure. The supreme +importance of these characters has been proved by the final arbitrament +of the battle for life. Through his powers of intellect, articulate +language has been evolved; and on this his wonderful advancement has +mainly depended. He has invented and is able to use various weapons, +tools, traps, &c., with which he defends himself, kills or catches prey, +and otherwise obtains food. He has made rafts or canoes on which to fish +or cross over to neighbouring fertile islands. He has discovered the art +of making fire, by which hard and stringy roots can be rendered +digestible, and poisonous roots or herbs innocuous. This last discovery, +probably the greatest, excepting language, ever made by man, dates from +before the dawn of history. These several inventions, by which man in +the rudest state has become so preeminent, are the direct result of the +development of his powers of observation, memory, curiosity, +imagination, and reason. I cannot, therefore, understand how it is that +Mr. Wallace[198] maintains, that "natural selection could only have +endowed the savage with a brain a little superior to that of an ape." + +Although the intellectual powers and social habits of man are of +paramount importance to him, we must not underrate the importance of his +bodily structure, to which subject the remainder of this chapter will be +devoted. The development of the intellectual and social or moral +faculties will be discussed in the following chapter. + +Even to hammer with precision is no easy matter, as every one who has +tried to learn carpentry will admit. To throw a stone with as true an +aim as can a Fuegian in defending himself, or in killing birds, requires +the most consummate perfection in the correlated action of the muscles +of the hand, arm, and shoulder, not to mention a fine sense of touch. In +throwing a stone or spear, and in many other actions, a man must stand +firmly on his feet; and this again demands the perfect coadaptation of +numerous muscles. To chip a flint into the rudest tool, or to form a +barbed spear or hook from a bone, demands the use of a perfect hand; +for, as a most capable judge, Mr. Schoolcraft,[199] remarks, the shaping +fragments of stone into knives, lances, or arrow-heads, shews +"extraordinary ability and long practice." We have evidence of this in +primeval men having practised a division of labour; each man did not +manufacture his own flint tools or rude pottery; but certain individuals +appear to have devoted themselves to such work, no doubt receiving in +exchange the produce of the chase. Archologists are convinced that an +enormous interval of time elapsed before our ancestors thought of +grinding chipped flints into smooth tools. A man-like animal who +possessed a hand and arm sufficiently perfect to throw a stone with +precision or to form a flint into a rude tool, could, it can hardly be +doubted, with sufficient practice make almost anything, as far as +mechanical skill alone is concerned, which a civilised man can make. The +structure of the hand in this respect may be compared with that of the +vocal organs, which in the apes are used for uttering various +signal-cries, or, as in one species, musical cadences; but in man +closely similar vocal organs have become adapted through the inherited +effects of use for the utterance of articulate language. + +Turning now to the nearest allies of man, and therefore to the best +representatives of our early progenitors, we find that the hands in the +Quadrumana are constructed on the same general pattern as in us, but are +far less perfectly adapted for diversified uses. Their hands do not +serve so well as the feet of a dog for locomotion; as may be seen in +those monkeys which walk on the outer margins of the palms, or on the +backs of their bent fingers, as in the chimpanzee and orang.[200] Their +hands, however, are admirably adapted for climbing trees. Monkeys seize +thin branches or ropes, with the thumb on one side and the fingers and +palm on the other side, in the same manner as we do. They can thus also +carry rather large objects, such as the neck of a bottle, to their +mouths. Baboons turn over stones and scratch up roots with their hands. +They seize nuts, insects, or other small objects with the thumb in +opposition to the fingers, and no doubt they thus extract eggs and the +young from the nests of birds. American monkeys beat the wild oranges on +the branches until the rind is cracked, and then tear it off with the +fingers of the two hands. Other monkeys open mussel-shells with the two +thumbs. With their fingers they pull out thorns and burrs, and hunt for +each other's parasites. In a state of nature they break open hard fruits +with the aid of stones. They roll down stones or throw them at their +enemies; nevertheless, they perform these various actions clumsily, and +they are quite unable, as I have myself seen, to throw a stone with +precision. + +It seems to me far from true that because "objects are grasped clumsily" +by monkeys, "a much less specialised organ of prehension" would have +served them[201] as well as their present hands. On the contrary, I see +no reason to doubt that a more perfectly constructed hand would have +been an advantage to them, provided, and it is important to note this, +that their hands had not thus been rendered less well adapted for +climbing trees. We may suspect that a perfect hand would have been +disadvantageous for climbing; as the most arboreal monkeys in the world, +namely Ateles in America and Hylobates in Asia, either have their thumbs +much reduced in size and even rudimentary, or their fingers +partially coherent, so that their hands are converted into mere +grasping-hooks.[202] + +As soon as some ancient member in the great series of the Primates came, +owing to a change in its manner of procuring subsistence, or to a change +in the conditions of its native country, to live somewhat less on trees +and more on the ground, its manner of progression would have been +modified; and in this case it would have had to become either more +strictly quadrupedal or bipedal. Baboons frequent hilly and rocky +districts, and only from necessity climb up high trees;[203] and they +have acquired almost the gait of a dog. Man alone has become a biped; +and we can, I think, partly see how he has come to assume his erect +attitude, which forms one of the most conspicuous differences between +him and his nearest allies. Man could not have attained his present +dominant position in the world without the use of his hands which are so +admirably adapted to act in obedience to his will. As Sir C. Bell[204] +insists "the hand supplies all instruments, and by its correspondence +with the intellect gives him universal dominion." But the hands and arms +could hardly have become perfect enough to have manufactured weapons, or +to have hurled stones and spears with a true aim, as long as they were +habitually used for locomotion and for supporting the whole weight of +the body, or as long as they were especially well adapted, as previously +remarked, for climbing trees. Such rough treatment would also have +blunted the sense of touch, on which their delicate use largely depends. +From these causes alone it would have been an advantage to man to have +become a biped; but for many actions it is almost necessary that both +arms and the whole upper part of the body should be free; and he must +for this end stand firmly on his feet. To gain this great advantage, the +feet have been rendered flat, and the great toe peculiarly modified, +though this has entailed the loss of the power of prehension. It accords +with the principle of the division of physiological labour, which +prevails throughout the animal kingdom, that as the hands became +perfected for prehension, the feet should have become perfected for +support and locomotion. With some savages, however, the foot has not +altogether lost its prehensile power, as shewn by their manner of +climbing trees and of using them in other ways.[205] + +If it be an advantage to man to have his hands and arms free and to +stand firmly on his feet, of which there can be no doubt from his +preeminent success in the battle of life, then I can see no reason why +it should not have been advantageous to the progenitors of man to have +become more and more erect or bipedal. They would thus have been better +able to have defended themselves with stones or clubs, or to have +attacked their prey, or otherwise obtained food. The best constructed +individuals would in the long run have succeeded best, and have survived +in larger numbers. If the gorilla and a few allied forms had become +extinct, it might have been argued with great force and apparent truth, +that an animal could not have been gradually converted from a quadruped +into a biped; as all the individuals in an intermediate condition would +have been miserably ill-fitted for progression. But we know (and this is +well worthy of reflection) that several kinds of apes are now actually +in this intermediate condition; and no one doubts that they are on the +whole well adapted for their conditions of life. Thus the gorilla runs +with a sidelong shambling gait, but more commonly progresses by resting +on its bent hands. The long-armed apes occasionally use their arms like +crutches, swinging their bodies forward between them, and some kinds of +Hylobates, without having been taught, can walk or run upright with +tolerable quickness; yet they move awkwardly, and much less securely +than man. We see, in short, with existing monkeys various gradations +between a form of progression strictly like that of a quadruped and that +of a biped or man. + +As the progenitors of man became more and more erect, with their hands +and arms more and more modified for prehension and other purposes, with +their feet and legs at the same time modified for firm support and +progression, endless other changes of structure would have been +necessary. The pelvis would have had to be made broader, the spine +peculiarly curved and the head fixed in an altered position, and all +these changes have been attained by man. Prof. Schaaffhausen[206] +maintains that "the powerful mastoid processes of the human skull are +the result of his erect position;" and these processes are absent in the +orang, chimpanzee, &c., and are smaller in the gorilla than in man. +Various other structures might here have been specified, which appear +connected with man's erect position. It is very difficult to decide how +far all these correlated modifications are the result of natural +selection, and how far of the inherited effects of the increased use of +certain parts, or of the action of one part on another. No doubt these +means of change act and react on each other: thus when certain muscles, +and the crests of bone to which they are attached, become enlarged by +habitual use, this shews that certain fictions are habitually performed +and must be serviceable. Hence the individuals which performed them +best, would tend to survive in greater numbers. + +The free use of the arms and hands, partly the cause and partly the +result of man's erect position, appears to have led in an indirect +manner to other modifications of structure. The early male progenitors +of man were, as previously stated, probably furnished with great canine +teeth; but as they gradually acquired the habit of using stones, clubs, +or other weapons, for fighting with their enemies, they would have used +their jaws and teeth less and less. In this case, the jaws, together +with the teeth, would have become reduced in size, as we may feel sure +from innumerable analogous cases. In a future chapter we shall meet with +a closely-parallel case, in the reduction or complete disappearance of +the canine teeth in male ruminants, apparently in relation with the +development of their horns; and in horses, in relation with their habit +of fighting with their incisor teeth and hoofs. + +In the adult male anthropomorphous apes, as Rtimeyer,[207] and others +have insisted, it is precisely the effect which the jaw-muscles by their +great development have produced on the skull, that causes it to differ +so greatly in many respects from that of man, and has given to it "a +truly frightful physiognomy." Therefore as the jaws and teeth in the +progenitors of man gradually become reduced in size, the adult skull +would have presented nearly the same characters which it offers in the +young of the anthropomorphous apes, and would thus have come to resemble +more nearly that of existing man. A great reduction of the canine teeth +in the males would almost certainly, as we shall hereafter see, have +affected through inheritance the teeth of the females. + +As the various mental faculties were gradually developed, the brain +would almost certainly have become larger. No one, I presume, doubts +that the large size of the brain in man, relatively to his body, in +comparison with that of the gorilla or orang, is closely connected with +his higher mental powers. We meet with closely analogous facts with +insects, in which the cerebral ganglia are of extraordinary dimensions +in ants; these ganglia in all the Hymenoptera being many times larger +than in the less intelligent orders, such as beetles.[208] On the other +hand, no one supposes that the intellect of any two animals or of any +two men can be accurately gauged by the cubic contents of their skulls. +It is certain that there may be extraordinary mental activity with an +extremely small absolute mass of nervous matter: thus the wonderfully +diversified instincts, mental powers, and affections of ants are +generally known, yet their cerebral ganglia are not so large as the +quarter of a small pin's head. Under this latter point of view, the +brain of an ant is one of the most marvellous atoms of matter in the +world, perhaps more marvellous than the brain of man. + +The belief that there exists in man some close relation between the size +of the brain and the development of the intellectual faculties is +supported by the comparison of the skulls of savage and civilised races, +of ancient and modern people, and by the analogy of the whole +vertebrate series. Dr. J. Barnard Davis has proved[209] by many careful +measurements, that the mean internal capacity of the skull in Europeans +is 923 cubic inches; in Americans 875; in Asiatics 871; and in +Australians only 819 inches. Professor Broca[210] found that skulls +from graves in Paris of the nineteenth century, were larger than those +from vaults of the twelfth century, in the proportion of 1484 to 1426; +and Prichard is persuaded that the present inhabitants of Britain have +"much more capacious brain-cases" than the ancient inhabitants. +Nevertheless it must be admitted that some skulls of very high +antiquity, such as the famous one of Neanderthal, are well developed and +capacious. With respect to the lower animals, M. E. Lartet,[211] by +comparing the crania of tertiary and recent mammals, belonging to the +same groups, has come to the remarkable conclusion that the brain is +generally larger and the convolutions more complex in the more recent +form. On the other hand I have shewn[212] that the brains of domestic +rabbits are considerably reduced in bulk, in comparison with those of +the wild rabbit or hare; and this may be attributed to their having been +closely confined during many generations, so that they have exerted but +little their intellect, instincts, senses, and voluntary movements. + +The gradually increasing weight of the brain and skull in man must have +influenced the development of the supporting spinal column, more +especially whilst he was becoming erect. As this change of position was +being brought about, the internal pressure of the brain, will, also, +have influenced the form of the skull; for many facts shew how easily +the skull is thus affected. Ethnologists believe that it is modified by +the kind of cradle in which infants sleep. Habitual spasms of the +muscles and a cicatrix from a severe burn have permanently modified the +facial bones. In young persons whose heads from disease have become +fixed either sideways or backwards, one of the eyes has changed its +position, and the bones of the skull have been modified; and this +apparently results from the brain pressing in a new direction.[213] I +have shewn that with long-eared rabbits, even so trifling a cause as the +lopping forward of one ear drags forward on that side almost every bone +of the skull; so that the bones on the opposite sides no longer strictly +correspond. Lastly, if any animal were to increase or diminish much in +general size, without any change in its mental powers; or if the mental +powers were to be much increased or diminished without any great change +in the size of the body; the shape of the skull would almost certainly +be altered. I infer this from my observations on domestic rabbits, some +kinds of which have become very much larger than the wild animal, whilst +others have retained nearly the same size, but in both cases the brain +has been much reduced relatively to the size of the body. Now I was at +first much surprised by finding that in all these rabbits the skull had +become elongated or dolichocephalic; for instance, of two skulls of +nearly equal breadth, the one from a wild rabbit and the other from a +large domestic kind, the former was only 315 and the latter 43 inches +in length.[214] One of the most marked distinctions in different races +of man is that the skull in some is elongated, and in others rounded; +and here the explanation suggested by the case of the rabbits may +partially hold good; for Welcker finds that short "men incline more to +brachycephaly, and tall men to dolichocephaly;"[215] and tall men may be +compared with the larger and longer-bodied rabbits, all of which have +elongated skulls, or are dolichocephalic. + +From these several facts we can to a certain extent understand the means +through which the great size and more or less rounded form of the skull +has been acquired by man; and these are characters eminently distinctive +of him in comparison with the lower animals. + +Another most conspicuous difference between man and the lower animals is +the nakedness of his skin. Whales and dolphins (Cetacea), dugongs +(Sirenia) and the hippopotamus are naked; and this may be advantageous +to them for gliding through the water; nor would it be injurious to them +from the loss of warmth, as the species which inhabit the colder regions +are protected by a thick layer of blubber, serving the same purpose as +the fur of seals and otters. Elephants and rhinoceroses are almost +hairless; and as certain extinct species which formerly lived under an +arctic climate were covered with long wool or hair, it would almost +appear as if the existing species of both genera had lost their hairy +covering from exposure to heat. This appears the more probable, as the +elephants in India which live on elevated and cool districts are more +hairy[216] than those on the lowlands. May we then infer that man became +divested of hair from having aboriginally inhabited some tropical land? +The fact of the hair being chiefly retained in the male sex on the chest +and face, and in both sexes at the junction of all four limbs with the +trunk, favours this inference, assuming that the hair was lost before +man became erect; for the parts which now retain most hair would then +have been most protected from the heat of the sun. The crown of the +head, however, offers a curious exception, for at all times it must have +been one of the most exposed parts, yet it is thickly clothed with hair. +In this respect man agrees with the great majority of quadrupeds, which +generally have their upper and exposed surfaces more thickly clothed +than the lower surface. Nevertheless, the fact that the other members of +the order of Primates, to which man belongs, although inhabiting various +hot regions, are well clothed with hair, generally thickest on the upper +surface,[217] is strongly opposed to the supposition that man became +naked through the action of the sun. I am inclined to believe, as we +shall see under sexual selection, that man, or rather primarily woman, +became divested of hair for ornamental purposes; and according to this +belief it is not surprising that man should differ so greatly in +hairiness from all his lower brethren, for characters gained through +sexual selection often differ in closely-related forms to an +extraordinary degree. + +According to a popular impression, the absence of a tail is eminently +distinctive of man; but as those apes which come nearest to man are +destitute of this organ, its disappearance does not especially concern +us. Nevertheless it may be well to own that no explanation, as far as I +am aware, has ever been given of the loss of the tail by certain apes +and man. Its loss, however, is not surprising, for it sometimes differs +remarkably in length in species of the same genera: thus in some species +of Macacus the tail is longer than the whole body, consisting of +twenty-four vertebr; in others it consists of a scarcely visible stump, +containing only three or four vertebr. In some kinds of baboons there +are twenty-five, whilst in the mandrill there are ten very small stunted +caudal vertebr, or, according to Cuvier,[218] sometimes only five. This +great diversity in the structure and length of the tail in animals +belonging to the same genera, and following nearly the same habits of +life, renders it probable that the tail is not of much importance to +them; and if so, we might have expected that it would sometimes have +become more or less rudimentary, in accordance with what we incessantly +see with other structures. The tail almost always tapers towards the end +whether it be long or short; and this, I presume, results from the +atrophy, through disuse, of the terminal muscles together with their +arteries and nerves, leading to the atrophy of the terminal bones. With +respect to the os coccyx, which in man and the higher apes manifestly +consists of the few basal and tapering segments of an ordinary tail, I +have heard it asked how could these have become completely embedded +within the body; but there is no difficulty in this respect, for in many +monkeys the basal segments of the true tail are thus embedded. For +instance, Mr. Murie informs me that in the skeleton of a not full-grown +_Macacus inornatus_, he counted nine or ten caudal vertebr, which +altogether were only 18 inch in length. Of these the three basal ones +appeared to have been embedded; the remainder forming the free part of +the tail, which was only one inch in length, and half an inch in +diameter. Here, then, the three embedded caudal vertebr plainly +correspond with the four coalesced vertebr of the human os coccyx. + + +I have now endeavoured to shew that some of the most distinctive +characters of man have in all probability been acquired, either +directly, or more commonly indirectly, through natural selection. We +should bear in mind that modifications in structure or constitution, +which are of no service to an organism in adapting it to its habits of +life, to the food which it consumes, or passively to the surrounding +conditions, cannot have been thus acquired. We must not, however, be too +confident in deciding what modifications are of service to each being: +we should remember how little we know about the use of many parts, or +what changes in the blood or tissues may serve to fit an organism for a +new climate or some new kind of food. Nor must we forget the principle +of correlation, by which, as Isidore Geoffroy has shewn in the case of +man, many fit-range deviations of structure are tied together. +Independently of correlation, a change in one part often leads through +the increased or decreased use of other parts, to other changes of a +quite unexpected nature. It is also well to reflect on such facts, as +the wonderful growth of galls on plants caused by the poison of an +insect, and on the remarkable changes of colour in the plumage of +parrots when fed on certain fishes, or inoculated with the poison of +toads;[219] for we can thus see that the fluids of the system, if +altered for some special purpose, might induce other strange changes. We +should especially bear in mind that modifications acquired and +continually used during past ages for some useful purpose would probably +become firmly fixed and might be long inherited. + +Thus a very large yet undefined extension may safely be given to the +direct and indirect results of natural selection; but I now admit, after +reading the essay by Ngeli on plants, and the remarks by various +authors with respect to animals, more especially those recently made by +Professor Broca, that in the earlier editions of my 'Origin of Species' +I probably attributed too much to the action of natural selection or the +survival of the fittest. I have altered the fifth edition of the Origin +so as to confine my remarks to adaptive changes of structure. I had not +formerly sufficiently considered the existence of many structures which +appear to be, as far as we can judge, neither beneficial nor injurious; +and this I believe to be one of the greatest oversights as yet detected +in my work. I may be permitted to say as some excuse, that I had two +distinct objects in view, firstly, to shew that species had not been +separately created, and secondly, that natural selection had been the +chief agent of change, though largely aided by the inherited effects of +habit, and slightly by the direct action of the surrounding conditions. +Nevertheless I was not able to annul the influence of my former belief, +then widely prevalent, that each species had been purposely created; and +this led to my tacitly assuming that every detail of structure, +excepting rudiments, was of some special, though unrecognised, service. +Any one with this assumption in his mind would naturally extend the +action of natural selection, either during past or present times, too +far. Some of those who admit the principle of evolution, but reject +natural selection, seem to forget, when criticising my book, that I had +the above two objects in view; hence if I have erred in giving to +natural selection great power, which I am far from admitting, or in +having exaggerated its power, which is in itself probable, I have at +least, as I hope, done good service in aiding to overthrow the dogma of +separate creations. + +That all organic beings, including man, present many modifications of +structure which are of no service to them at present, nor have been +formerly, is, as I can now see, probable. We know not what produces the +numberless slight differences between the individuals of each species, +for reversion only carries the problem a few steps backwards; but each +peculiarity must have had its own efficient cause. If these causes, +whatever they may be, were to act more uniformly and energetically +during a lengthened period (and no reason can be assigned why this +should not sometimes occur), the result would probably be not mere +slight individual differences, but well-marked, constant modifications. +Modifications which are in no way beneficial cannot have been kept +uniform through natural selection, though any which were injurious would +have been thus eliminated. Uniformity of character would, however, +naturally follow from, the assumed uniformity of the exciting causes, +and likewise from the free intercrossing of many individuals. The same +organism might acquire in this manner during successive periods +successive modifications, and these would be transmitted in a nearly +uniform state as long as the exciting causes remained the same and there +was free intercrossing. With respect to the exciting causes we can only +say, as when speaking of so-called spontaneous variations, that they +relate much more closely to the constitution of the varying organism, +than to the nature of the conditions to which it has been subjected. + + +_Conclusion._--In this chapter we have seen that as man at the present +day is liable, like every other animal, to multiform individual +differences or slight variations, so no doubt were the early progenitors +of man; the variations being then as now induced by the same general +causes, and governed by the same general and complex laws. As all +animals tend to multiply beyond their means of subsistence, so it must +have been with the progenitors of man; and this will inevitably have led +to a struggle for existence and to natural selection. This latter +process will have been greatly aided by the inherited effects of the +increased use of parts; these two processes incessantly reacting on each +other. It appears, also, as we shall hereafter see, that various +unimportant characters have been acquired by man through sexual +selection. An unexplained residuum of change, perhaps a large one, must +be left to the assumed uniform action of those unknown agencies, which +occasionally induce strongly-marked and abrupt deviations of structure +in our domestic productions. + +Judging from the habits of savages and of the greater number of the +Quadrumana, primeval men, and even the ape-like progenitors of man, +probably lived in society. With strictly social animals, natural +selection sometimes acts indirectly on the individual, through the +preservation of variations which are beneficial only to the community. A +community including a large number of well-endowed individuals increases +in number and is victorious over other and less well-endowed +communities; although each separate member may gain no advantage over +the other members of the same community. With associated insects many +remarkable structures, which are of little or no service to the +individual or its own offspring, such as the pollen-collecting +apparatus, or the sting of the worker-bee, or the great jaws of +soldier-ants, have been thus acquired. With the higher social animals, I +am not aware that any structure has been modified solely for the good of +the community, though some are of secondary service to it. For instance, +the horns of ruminants and the great canine teeth of baboons appear to +have been acquired by the males as weapons for sexual strife, but they +are used in defence of the herd or troop. In regard to certain mental +faculties the case, as we shall see in the following chapter, is wholly +different; for these faculties have been chiefly, or even exclusively, +gained for the benefit of the community; the individuals composing the +community being at the same time indirectly benefited. + +It has often been objected to such views as the foregoing, that man is +one of the most helpless and defenceless creatures in the world; and +that during his early and less well-developed condition he would have +been still more helpless. The Duke of Argyll, for instance, insists[220] +that "the human frame has diverged from the structure of brutes, in the +direction of greater physical helplessness and weakness. That is to say, +it is a divergence which of all others it is most impossible to ascribe +to mere natural selection." He adduces the naked and unprotected state +of the body, the absence of great teeth or claws for defence, the little +strength of man, his small speed in running, and his slight power of +smell, by which to discover food or to avoid danger. To these +deficiencies there might have been added the still more serious loss of +the power of quickly climbing trees, so as to escape from enemies. +Seeing that the unclothed Fuegians can exist under their wretched +climate, the loss of hair would not have been a great injury to primeval +man, if he inhabited a warm country. When we compare defenceless man +with the apes, many of which are provided with formidable canine teeth, +we must remember that these in their fully-developed condition are +possessed by the males alone, being chiefly used by them for fighting +with their rivals; yet the females which are not thus provided, are able +to survive. + +In regard to bodily size or strength, we do not know whether man is +descended from some comparatively small species, like the chimpanzee, or +from one as powerful as the gorilla; and, therefore, we cannot say +whether man has become larger and stronger, or smaller and weaker, in +comparison with his progenitors. We should, however, bear in mind that +an animal possessing great size, strength, and ferocity, and which, like +the gorilla, could defend itself from all enemies, would probably, +though not necessarily, have failed to become social; and this would +most effectually have checked the acquirement by man of his higher +mental qualities, such as sympathy and the love of his fellow-creatures. +Hence it might have been an immense advantage to man to have sprung +from some comparatively weak creature. + +The slight corporeal strength of man, his little speed, his want of +natural weapons, &c., are more than counterbalanced, firstly by his +intellectual powers, through which he has, whilst still remaining in a +barbarous state, formed for himself weapons, tools, &c., and secondly by +his social qualities which lead him to give aid to his fellow-men and to +receive it in return. No country in the world abounds in a greater +degree with dangerous beasts than Southern Africa; no country presents +more fearful physical hardships than the Arctic regions; yet one of the +puniest races, namely, the Bushmen, maintain themselves in Southern +Africa, as do the dwarfed Esquimaux in the Arctic regions. The early +progenitors of man were, no doubt, inferior in intellect, and probably +in social disposition, to the lowest existing savages; but it is quite +conceivable that they might have existed, or even flourished, if, whilst +they gradually lost their brute-like powers, such as climbing trees, +&c., they at the same time advanced in intellect. But granting that the +progenitors of man were far more helpless and defenceless than any +existing savages, if they had inhabited some warm continent or large +island, such as Australia or New Guinea, or Borneo (the latter island +being now tenanted by the orang), they would not have been exposed to +any special danger. In an area as large as one of these islands, the +competition between tribe and tribe would have been sufficient, under +favourable conditions, to have raised man, through the survival of the +fittest, combined with the inherited effects of habit, to his present +high position in the organic scale. + + + + +CHAPTER V. + +ON THE DEVELOPMENT OF THE INTELLECTUAL AND MORAL FACULTIES DURING +PRIMEVAL AND CIVILISED TIMES. + + + The advancement of the intellectual powers through natural + selection--Importance of imitation--Social and moral + faculties--Their development within the limits of the same + tribe--Natural selection as affecting civilised + nations--Evidence that civilised nations were once barbarous. + + +The subjects to be discussed in this chapter are of the highest +interest, but are treated by me in a most imperfect and fragmentary +manner. Mr. Wallace, in an admirable paper before referred to,[221] +argues that man after he had partially acquired those intellectual and +moral faculties which distinguish him from the lower animals, would have +been but little liable to have had his bodily structure modified through +natural selection or any other means. For man is enabled through his +mental faculties "to keep with an unchanged body in harmony with the +changing universe." He has great power of adapting his habits to new +conditions of life. He invents weapons, tools and various stratagems, by +which he procures food and defends himself. When he migrates into a +colder climate he uses clothes, builds sheds, and makes fires; and, by +the aid of fire, cooks food otherwise indigestible. He aids his +fellow-men in many ways, and anticipates future events. Even at a remote +period he practised some subdivision of labour. + +The lower animals, on the other hand, must have their bodily structure +modified in order to survive under greatly changed conditions. They must +be rendered stronger, or acquire more effective teeth or claws, in order +to defend themselves from new enemies; or they must be reduced in size +so as to escape detection and danger. When they migrate into a colder +climate they must become clothed with thicker fur, or have their +constitutions altered. If they fail to be thus modified, they will cease +to exist. + +The case, however, is widely different, as Mr. Wallace has with justice +insisted, in relation to the intellectual and moral faculties of man. +These faculties are variable; and we have every reason to believe that +the variations tend to be inherited. Therefore, if they were formerly of +high importance to primeval man and to his ape-like progenitors, they +would have been perfected or advanced through natural selection. Of the +high importance of the intellectual faculties there can be no doubt, for +man mainly owes to them his preeminent position in the world. We can +see that, in the rudest state of society, the individuals who were the +most sagacious, who invented and used the best weapons or traps, and who +were best able to defend themselves, would rear the greatest number of +offspring. The tribes which included the largest number of men thus +endowed would increase in number and supplant other tribes. Numbers +depend primarily on the means of subsistence, and this, partly on the +physical nature of the country, but in a much higher degree on the arts +which are there practised. As a tribe increases and is victorious, it is +often still further increased by the absorption of other tribes.[222] +The stature and strength of the men of a tribe are likewise of some +importance for its success, and these depend in part on the nature and +amount of the food which can be obtained. In Europe the men of the +Bronze period were supplanted by a more powerful and, judging from their +sword-handles, larger-handed race;[223] but their success was probably +due in a much higher degree to their superiority in the arts. + +All that we know about savages, or may infer from their traditions and +from old monuments, the history of which is quite forgotten by the +present inhabitants, shew that from the remotest times successful tribes +have supplanted other tribes. Relics of extinct or forgotten tribes have +been discovered throughout the civilised regions of the earth, on the +wild plains of America, and on the isolated islands in the Pacific +Ocean. At the present day civilised nations are everywhere supplanting +barbarous nations, excepting where the climate opposes a deadly barrier; +and they succeed mainly, though not exclusively, through their arts, +which are the products of the intellect. It is, therefore, highly +probable that with mankind the intellectual faculties have been +gradually perfected through natural selection; and this conclusion is +sufficient for our purpose. Undoubtedly it would have been very +interesting to have traced the development of each separate faculty from +the state in which it exists in the lower animals to that in which it +exists in man; but neither my ability nor knowledge permit the attempt. + +It deserves notice that as soon as the progenitors of man became social +(and this probably occurred at a very early period), the advancement of +the intellectual faculties will have been aided and modified in an +important manner, of which we see only traces in the lower animals, +namely, through the principle of imitation, together with reason and +experience. Apes are much given to imitation, as are the lowest savages; +and the simple fact previously referred to, that after a time no animal +can be caught in the same place by the same sort of trap, shews that +animals learn by experience, and imitate each others' caution. Now, if +some one man in a tribe, more sagacious than the others, invented a new +snare or weapon, or other means of attack or defence, the plainest +self-interest, without the assistance of much reasoning power, would +prompt the other members to imitate him; and all would thus profit. The +habitual practice of each new art must likewise in some slight degree +strengthen the intellect. If the new invention were an important one, +the tribe would increase in number, spread, and supplant other tribes. +In a tribe thus rendered more numerous there would always be a rather +better chance of the birth of other superior and inventive members. If +such men left children to inherit their mental superiority, the chance +of the birth of still more ingenious members would be somewhat better, +and in a very small tribe decidedly better. Even if they left no +children, the tribe would still include their blood-relations; and it +has been ascertained by agriculturists[224] that by preserving and +breeding from the family of an animal, which when slaughtered was found +to be valuable, the desired character has been obtained. + + +Turning now to the social and moral faculties. In order that primeval +men, or the ape-like progenitors of man, should have become social, they +must have acquired the same instinctive feelings which impel other +animals to live in a body; and they no doubt exhibited the same general +disposition. They would have felt uneasy when separated from their +comrades, for whom they would have felt some degree of love; they would +have warned each other of danger, and have given mutual aid in attack or +defence. All this implies some degree of sympathy, fidelity, and +courage. Such social qualities, the paramount importance of which to the +lower animals is disputed by no one, were no doubt acquired by the +progenitors of man in a similar manner, namely, through natural +selection, aided by inherited habit. When two tribes of primeval man, +living in the same country, came into competition, if the one tribe +included (other circumstances being equal) a greater number of +courageous, sympathetic, and faithful members, who were always ready to +warn each other of danger, to aid and defend each other, this tribe +would without doubt succeed best and conquer the other. Let it be borne +in mind how all-important, in the never-ceasing wars of savages, +fidelity and courage must be. The advantage which disciplined soldiers +have over undisciplined hordes follows chiefly from the confidence which +each man feels in his comrades. Obedience, as Mr. Bagehot has well +shewn,[225] is of the highest value, for any form of government is +better than none. Selfish and contentious people will not cohere, and +without coherence nothing can be effected. A tribe possessing the above +qualities in a high degree would spread and be victorious over other +tribes; but in the course of time it would, judging from all past +history, be in its turn overcome by some other and still more highly +endowed tribe. Thus the social and moral qualities would tend slowly to +advance and be diffused throughout the world. + +But it may be asked, how within the limits of the same tribe did a large +number of members first become endowed with these social and moral +qualities, and how was the standard of excellence raised? It is +extremely doubtful whether the offspring of the more sympathetic and +benevolent parents, or of those which were the most faithful to their +comrades, would be reared in greater number than the children of selfish +and treacherous parents of the same tribe. He who was ready to sacrifice +his life, as many a savage has been, rather than betray his comrades, +would often leave no offspring to inherit his noble nature. The bravest +men, who were always willing to come to the front in war, and who freely +risked their lives for others, would on an average perish in larger +number than other men. Therefore it seems scarcely possible (bearing in +mind that we are not here speaking of one tribe being victorious over +another) that the number of men gifted with such virtues, or that the +standard of their excellence, could be increased through natural +selection, that is, by the survival of the fittest. + +Although the circumstances which lead to an increase in the number of +men thus endowed within the same tribe are too complex to be clearly +followed out, we can trace some of the probable steps. In the first +place, as the reasoning powers and foresight of the members became +improved, each man would soon learn from experience that if he aided his +fellow-men, he would commonly receive aid in return. From this low +motive he might acquire the habit of aiding his fellows; and the habit +of performing benevolent actions certainly strengthens the feeling of +sympathy, which gives the first impulse to benevolent actions. Habits, +moreover, followed during many generations probably tend to be +inherited. + +But there is another and much more powerful stimulus to the development +of the social virtues, namely, the praise and the blame of our +fellow-men. The love of approbation and the dread of infamy, as well as +the bestowal of praise or blame, are primarily due, as we have seen in +the third chapter, to the instinct of sympathy; and this instinct no +doubt was originally acquired, like all the other social instincts, +through natural selection. At how early a period the progenitors of man, +in the course of their development, became capable of feeling and being +impelled by the praise or blame of their fellow-creatures, we cannot, of +course, say. But it appears that even dogs appreciate encouragement, +praise, and blame. The rudest savages feel the sentiment of glory, as +they clearly show by preserving the trophies of their prowess, by their +habit of excessive boasting, and even by the extreme care which they +take of their personal appearance and decorations; for unless they +regarded the opinion of their comrades, such habits would be senseless. + +They certainly feel shame at the breach of some of their lesser rules; +but how far they experience remorse is doubtful. I was at first +surprised that I could not recollect any recorded instances of this +feeling in savages; and Sir J. Lubbock[226] states that he knows of +none. But if we banish from our minds all cases given in novels and +plays and in death-bed confessions made to priests, I doubt whether many +of us have actually witnessed remorse; though we may have often seen +shame and contrition for smaller offences. Remorse is a deeply hidden +feeling. It is incredible that a savage, who will sacrifice his life +rather than betray his tribe, or one who will deliver himself up as a +prisoner rather than break his parole,[227] would not feel remorse in +his inmost soul, though he might conceal it, if he had failed in a duty +which he held sacred. + +We may therefore conclude that primeval man, at a very remote period, +would have been influenced by the praise and blame of his fellows. It is +obvious, that the members of the same tribe would approve of conduct +which appeared to them to be for the general good, and would reprobate +that which appeared evil. To do good unto others--to do unto others as +ye would they should do unto you,--is the foundation-stone of morality. +It is, therefore, hardly possible to exaggerate the importance during +rude times of the love of praise and the dread of blame. A man who was +not impelled by any deep, instinctive feeling, to sacrifice his life for +the good of others, yet was roused to such actions by a sense of glory, +would by his example excite the same wish for glory in other men, and +would strengthen by exercise the noble feeling of admiration. He might +thus do far more good to his tribe than by begetting offspring with a +tendency to inherit his own high character. + +With increased experience and reason, man perceives the more remote +consequences of his actions, and the self-regarding virtues, such as +temperance, chastity, &c., which during early times are, as we have +before seen, utterly disregarded, come to be highly esteemed or even +held sacred. I need not, however, repeat what I have said on this head +in the third chapter. Ultimately a highly complex sentiment, having its +first origin in the social instincts, largely guided by the approbation +of our fellow-men, ruled by reason, self-interest, and in later times by +deep religious feelings, confirmed by instruction and habit, all +combined, constitute our moral sense or conscience. + +It must not be forgotten that although a high standard of morality gives +but a slight or no advantage to each individual man and his children +over the other men of the same tribe, yet that an advancement in the +standard of morality and an increase in the number of well-endowed men +will certainly give an immense advantage to one tribe over another. +There can be no doubt that a tribe including many members who, from +possessing in a high degree the spirit of patriotism, fidelity, +obedience, courage, and sympathy, were always ready to give aid to each +other and to sacrifice themselves for the common good, would be +victorious over most other tribes; and this would be natural selection. +At all times throughout the world tribes have supplanted other tribes; +and as morality is one element in their success, the standard of +morality and the number of well-endowed men will thus everywhere tend to +rise and increase. + +It is, however, very difficult to form any judgment why one particular +tribe and not another has been successful and has risen in the scale of +civilisation. Many savages are in the same condition as when first +discovered several centuries ago. As Mr. Bagehot has remarked, we are +apt to look at progress as the normal rule in human society; but history +refutes this. The ancients did not even entertain the idea; nor do the +oriental nations at the present day. According to another high +authority, Mr. Maine,[228] "the greatest part of mankind has never +shewn a particle of desire that its civil institutions should be +improved." Progress seems to depend on many concurrent favourable +conditions, far too complex to be followed out. But it has often been +remarked, that a cool climate from leading to industry and the various +arts has been highly favourable, or even indispensable for this end. The +Esquimaux, pressed by hard necessity, have succeeded in many ingenious +inventions, but their climate has been too severe for continued +progress. Nomadic habits, whether over wide plains, or through the dense +forests of the tropics, or along the shores of the sea, have in every +case been highly detrimental. Whilst observing the barbarous inhabitants +of Tierra del Fuego, it struck me that the possession of some property, +a fixed abode, and the union of many families under a chief, were the +indispensable requisites for civilisation. Such habits almost +necessitate the cultivation of the ground; and the first steps in +cultivation would probably result, as I have elsewhere shewn,[229] from +some such accident as the seeds of a fruit-tree falling on a heap of +refuse and producing an unusually fine variety. The problem, however, of +the first advance of savages towards civilisation is at present much too +difficult to be solved. + + +_Natural Selection as affecting Civilised Nations._--In the last and +present chapters I have considered the advancement of man from a former +semi-human condition to his present state as a barbarian. But some +remarks on the agency of natural selection on civilised nations may be +here worth adding. This subject has been ably discussed by Mr. W. R. +Greg,[230] and previously by Mr. Wallace and Mr. Galton.[231] Most of +my remarks are taken from these three authors. With savages, the weak in +body or mind are soon eliminated; and those that survive commonly +exhibit a vigorous state of health. We civilised men, on the other hand, +do our utmost to check the process of elimination; we build asylums for +the imbecile, the maimed, and the sick; we institute poor-laws; and our +medical men exert their utmost skill to save the life of every one to +the last moment. There is reason to believe that vaccination has +preserved thousands, who from a weak constitution would formerly have +succumbed to small-pox. Thus the weak members of civilised societies +propagate their kind. No one who has attended to the breeding of +domestic animals will doubt that this must be highly injurious to the +race of man. It is surprising how soon a want of care, or care wrongly +directed, leads to the degeneration of a domestic race; but excepting in +the case of man himself, hardly any one is so ignorant as to allow his +worst animals to breed. + +The aid which we feel impelled to give to the helpless is mainly an +incidental result of the instinct of sympathy, which was originally +acquired as part of the social instincts, but subsequently rendered, in +the manner previously indicated, more tender and more widely diffused. +Nor could we check our sympathy, if so urged by hard reason, without +deterioration in the noblest part of our nature. The surgeon may harden +himself whilst performing an operation, for he knows that he is acting +for the good of his patient; but if we were intentionally to neglect the +weak and helpless, it could only be for a contingent benefit, with a +certain and great present evil. Hence we must bear without complaining +the undoubtedly bad effects of the weak surviving and propagating their +kind; but there appears to be at least one check in steady action, +namely the weaker and inferior members of society not marrying so freely +as the sound; and this check might be indefinitely increased, though +this is more to be hoped for than expected, by the weak in body or mind +refraining from marriage. + +In all civilised countries man accumulates property and bequeaths it to +his children. So that the children in the same country do not by any +means start fair in the race for success. But this is far from an +unmixed evil; for without the accumulation of capital the arts could not +progress; and it is chiefly through their power that the civilised races +have extended, and are now everywhere extending, their range, so as to +take the place of the lower races. Nor does the moderate accumulation of +wealth interfere with the process of selection. When a poor man becomes +rich, his children enter trades or professions in which there is +struggle enough, so that the able in body and mind succeed best. The +presence of a body of well-instructed men, who have not to labour for +their daily bread, is important to a degree which cannot be +over-estimated; as all high intellectual work is carried on by them, and +on such work material progress of all kinds mainly depends, not to +mention other and higher advantages. No doubt wealth when very great +tends to convert men into useless drones, but their number is never +large; and some degree of elimination here occurs, as we daily see rich +men, who happen to be fools or profligate, squandering away all their +wealth. + +Primogeniture with entailed estates is a more direct evil, though it may +formerly have been a great advantage by the creation of a dominant +class, and any government is better than anarchy. The eldest sons, +though they may be weak in body or mind, generally marry, whilst the +younger sons, however superior in these respects, do not so generally +marry. Nor can worthless eldest sons with entailed estates squander +their wealth. But here, as elsewhere, the relations of civilised life +are so complex that some compensatory checks intervene. The men who are +rich through primogeniture are able to select generation after +generation the more beautiful and charming women; and these must +generally be healthy in body and active in mind. The evil consequences, +such as they may be, of the continued preservation of the same line of +descent, without any selection, are checked by men of rank always +wishing to increase their wealth and power; and this they effect by +marrying heiresses. But the daughters of parents who have produced +single children, are themselves, as Mr. Galton has shewn,[232] apt to be +sterile; and thus noble families are continually cut off in the direct +line, and their wealth flows into some side channel; but unfortunately +this channel is not determined by superiority of any kind. + +Although civilisation thus checks in many ways the action of natural +selection, it apparently favours, by means of improved food and the +freedom from occasional hardships, the better development of the body. +This may be inferred from civilised men having been found, wherever +compared, to be physically stronger than savages. They appear also to +have equal powers of endurance, as has been proved in many adventurous +expeditions. Even the great luxury of the rich can be but little +detrimental; for the expectation of life of our aristocracy, at all ages +and of both sexes, is very little inferior to that of healthy English +lives in the lower classes.[233] + +We will now look to the intellectual faculties alone. If in each grade +of society the members were divided into two equal bodies, the one +including the intellectually superior and the other the inferior, there +can be little doubt that the former would succeed best in all +occupations and rear a greater number of children. Even in the lowest +walks of life, skill and ability must be of some advantage, though in +many occupations, owing to the great division of labour, a very small +one. Hence in civilised nations there will be some tendency to an +increase both in the number and in the standard of the intellectually +able. But I do not wish to assert that this tendency may not be more +than counterbalanced in other ways, as by the multiplication of the +reckless and improvident; but even to such as these, ability must be +some advantage. + +It has often been objected to views like the foregoing, that the most +eminent men who have ever lived have left no offspring to inherit their +great intellect. Mr. Galton says,[234] "I regret I am unable to solve +the simple question whether, and how far, men and women who are +prodigies of genius are infertile. I have, however, shewn that men of +eminence are by no means so." + +Great lawgivers, the founders of beneficent religions, great +philosophers and discoverers in science, aid the progress of mankind in +a far higher degree by their works than by leaving a numerous progeny. +In the case of corporeal structures, it is the selection of the slightly +better-endowed and the elimination of the slightly less well-endowed +individuals, and not the preservation of strongly-marked and rare +anomalies, that leads to the advancement of a species.[235] So it will +be with the intellectual faculties, namely from the somewhat more able +men in each grade of society succeeding rather better than the less +able, and consequently increasing in number, if not otherwise prevented. +When in any nation the standard of intellect and the number of +intellectual men have increased, we may expect from the law of the +deviation from an average, as shewn by Mr. Galton, that prodigies of +genius will appear somewhat more frequently than before. + +In regard to the moral qualities, some elimination of the worst +dispositions is always in progress even in the most civilised nations. +Malefactors are executed, or imprisoned for long periods, so that they +cannot freely transmit their bad qualities. Melancholic and insane +persons are confined, or commit suicide. Violent and quarrelsome men +often come to a bloody end. Restless men who will not follow any steady +occupation--and this relic of barbarism is a great check to +civilisation[236]--emigrate to newly-settled countries, where they prove +useful pioneers. Intemperance is so highly destructive, that the +expectation of life of the intemperate, at the age, for instance, of +thirty, is only 13.8 years; whilst for the rural labourers of England at +the same age it is 4059 years.[237] Profligate women bear few children, +and profligate men rarely marry; both suffer from disease. In the +breeding of domestic animals, the elimination of those individuals, +though few in number, which are in any marked manner inferior, is by no +means an unimportant element towards success. This especially holds good +with injurious characters which tend to reappear through reversion, such +as blackness in sheep; and with mankind some of the worst dispositions, +which occasionally without any assignable cause make their appearance in +families, may perhaps be reversions to a savage state, from which we are +not removed by very many generations. This view seems indeed recognised +in the common expression that such men are the black sheep of the +family. + +With civilised nations, as far as an advanced standard of morality, and +an increased number of fairly well-endowed men are concerned, natural +selection apparently effects but little; though the fundamental social +instincts were originally thus gained. But I have already said enough, +whilst treating of the lower races, on the causes which lead to the +advance of morality, namely, the approbation of our fellow-men--the +strengthening of our sympathies by habit--example and imitation-- +reason--experience and even self-interest--instruction during +youth, and religious feelings. + +A most important obstacle in civilised countries to an increase in the +number of men of a superior class has been strongly urged by Mr. Greg +and Mr. Galton,[238] namely, the fact that the very poor and reckless, +who are often degraded by vice, almost invariably marry early, whilst +the careful and frugal, who are generally otherwise virtuous, marry late +in life, so that they may be able to support themselves and their +children in comfort. Those who marry early produce within a given period +not only a greater number of generations, but, as shewn by Dr. +Duncan,[239] they produce many more children. The children, moreover, +that are born by mothers during the prime of life are heavier and +larger, and therefore probably more vigorous, than those born at other +periods. Thus the reckless, degraded, and often vicious members of +society, tend to increase at a quicker rate than the provident and +generally virtuous members. Or as Mr. Greg puts the case: "The careless, +squalid, unaspiring Irishman multiplies like rabbits: the frugal, +foreseeing, self-respecting, ambitious Scot, stern in his morality, +spiritual in his faith, sagacious and disciplined in his intelligence, +passes his best years in struggle and in celibacy, marries late, and +leaves few behind him. Given a land originally peopled by a thousand +Saxons and a thousand Celts--and in a dozen generations five-sixths of +the population would be Celts, but five-sixths of the property, of the +power, of the intellect, would belong to the one-sixth of Saxons that +remained. In the eternal 'struggle for existence,' it would be the +inferior and _less_ favoured race that had prevailed--and prevailed by +virtue not of its good qualities but of its faults." + +There are, however, some checks to this downward tendency. We have seen +that the intemperate suffer from a high rate of mortality, and the +extremely profligate leave few offspring. The poorest classes crowd into +towns, and it has been proved by Dr. Stark from the statistics of ten +years in Scotland,[240] that at all ages the death-rate is higher in +towns than in rural districts, "and during the first five years of life +the town death-rate is almost exactly double that of the rural +districts." As these returns include both the rich and the poor, no +doubt more than double the number of births would be requisite to keep +up the number of the very poor inhabitants in the towns, relatively to +those in the country. With women, marriage at too early an age is highly +injurious; for it has been found in France that, "twice as many wives +under twenty die in the year, as died out of the same number of the +unmarried." The mortality, also, of husbands under twenty is +"excessively high,"[241] but what the cause of this may be seems +doubtful. Lastly, if the men who prudently delay marrying until they can +bring up their families in comfort, were to select, as they often do, +women in the prime of life, the rate of increase in the better class +would be only slightly lessened. + +It was established from an enormous body of statistics, taken during +1853, that the unmarried men throughout France, between the ages of +twenty and eighty, die in a much larger proportion than the married: for +instance, out of every 1000 unmarried men, between the ages of twenty +and thirty, 113 annually died, whilst of the married only 65 +died.[242] A similar law was proved to hold good, during the years 1863 +and 1864, with the entire population above the age of twenty in +Scotland: for instance, out of every 1000 unmarried men, between the +ages of twenty and thirty, 1497 annually died, whilst of the married +only 724 died, that is less than half.[243] Dr. Stark remarks on this, +"Bachelorhood is more destructive to life than the most unwholesome +trades, or than residence in an unwholesome house or district where +there has never been the most distant attempt at sanitary improvement." +He considers that the lessened mortality is the direct result of +"marriage, and the more regular domestic habits which attend that +state." He admits, however, that the intemperate, profligate, and +criminal classes, whose duration of life is low, do not commonly marry; +and it must likewise be admitted that men with a weak constitution, +ill health, or any great infirmity in body or mind, will often not wish +to marry, or will be rejected. Dr. Stark seems to have come to the +conclusion that marriage in itself is a main cause of prolonged life, +from finding that aged married men still have a considerable advantage +in this respect over the unmarried of the same advanced age; but every +one must have known instances of men, who with weak health during youth +did not marry, and yet have survived to old age, though remaining weak +and therefore always with a lessened chance of life. There is another +remarkable circumstance which seems to support Dr. Stark's conclusion, +namely, that widows and widowers in France suffer in comparison with the +married a very heavy rate of mortality; but Dr. Farr attributes this to +the poverty and evil habits consequent on the disruption of the family, +and to grief. On the whole we may conclude with Dr. Farr that the lesser +mortality of married than of unmarried men, which seems to be a general +law, "is mainly due to the constant elimination of imperfect types, and +to the skilful selection of the finest individuals out of each +successive generation;" the selection relating only to the marriage +state, and acting on all corporeal, intellectual, and moral qualities. +We may, therefore, infer that sound and good men who out of prudence +remain for a time unmarried do not suffer a high rate of mortality. + +If the various checks specified in the two last paragraphs, and perhaps +others as yet unknown, do not prevent the reckless, the vicious and +otherwise inferior members of society from increasing at a quicker rate +than the better class of men, the nation will retrograde, as has +occurred too often in the history of the world. We must remember that +progress is no invariable rule. It is most difficult to say why one +civilised nation rises, becomes more powerful, and spreads more widely, +than another; or why the same nation progresses more at one time than at +another. We can only say that it depends on an increase in the actual +number of the population, on the number of the men endowed with high +intellectual and moral faculties, as well as on their standard of +excellence. Corporeal structure, except so far as vigour of body leads +to vigour of mind, appears to have little influence. + +It has been urged by several writers that as high intellectual powers +are advantageous to a nation, the old Greeks, who stood some grades +higher in intellect than any race that has ever existed,[244] ought to +have risen, if the power of natural selection were real, still higher +in the scale, increased in number, and stocked the whole of Europe. Here +we have the tacit assumption, so often made with respect to corporeal +structures, that there is some innate tendency towards continued +development in mind and body. But development of all kinds depends on +many concurrent favourable circumstances. Natural selection acts only in +a tentative manner. Individuals and races may have acquired certain +indisputable advantages, and yet have perished from failing in other +characters. The Greeks may have retrograded from a want of coherence +between the many small states, from the small size of their whole +country, from the practice of slavery, or from extreme sensuality; for +they did not succumb until "they were enervated and corrupt to the very +core."[245] The western nations of Europe, who now so immeasurably +surpass their former savage progenitors and stand at the summit of +civilisation, owe little or none of their superiority to direct +inheritance from the old Greeks; though they owe much to the written +works of this wonderful people. + +Who can positively say why the Spanish nation, so dominant at one time, +has been distanced in the race. The awakening of the nations of Europe +from the dark ages is a still more perplexing problem. At this early +period, as Mr. Galton[246] has remarked, almost all the men of a gentle +nature, those given to meditation or culture of the mind, had no refuge +except in the bosom of the Church which demanded celibacy; and this +could hardly fail to have had a deteriorating influence on each +successive generation. During this same period the Holy Inquisition +selected with extreme care the freest and boldest men in order to burn +or imprison them. In Spain alone some of the best men--those who doubted +and questioned, and without doubting there can be no progress--were +eliminated during three centuries at the rate of a thousand a year. The +evil which the Catholic Church has thus effected, though no doubt +counterbalanced to a certain, perhaps large extent in other ways, is +incalculable; nevertheless, Europe has progressed at an unparalleled +rate. + +The remarkable success of the English as colonists over other European +nations, which is well illustrated by comparing the progress of the +Canadians of English and French extraction, has been ascribed to their +"daring and persistent energy;" but who can say how the English gained +their energy. There is apparently much truth in the belief that the +wonderful progress of the United States, as well as the character of the +people, are the results of natural selection; the more energetic, +restless, and courageous men from all parts of Europe having emigrated +during the last ten or twelve generations to that great country, and +having there succeeded best.[247] Looking to the distant future, I do +not think that the Rev. Mr. Zincke takes an exaggerated view when he +says:[248] "All other series of events--as that which resulted in the +culture of mind in Greece, and that which resulted in the empire of +Rome--only appear to have purpose and value when viewed in connection +with, or rather as subsidiary to ... the great stream of Anglo-Saxon +emigration to the west." + +Obscure as is the problem of the advance of civilisation, we can at +least see that a nation which produced during a lengthened period the +greatest number of highly intellectual, energetic, brave, patriotic, and +benevolent men, would generally prevail over less favoured nations. + +Natural selection follows from the struggle for existence; and this from +a rapid rate of increase. It is impossible not bitterly to regret, but +whether wisely is another question, the rate at which man tends to +increase; for this leads in barbarous tribes to infanticide and many +other evils, and in civilised nations to abject poverty, celibacy, and +to the late marriages of the prudent. But as man suffers from the same +physical evils with the lower animals, he has no right to expect an +immunity from the evils consequent on the struggle for existence. Had he +not been subjected to natural selection, assuredly he would never have +attained to the rank of manhood. When we see in many parts of the world +enormous areas of the most fertile land peopled by a few wandering +savages, but which are capable of supporting numerous happy homes, it +might be argued that the struggle for existence had not been +sufficiently severe to force man upwards to his highest standard. +Judging from all that we know of man and the lower animals, there has +always been sufficient variability in the intellectual and moral +faculties, for their steady advancement through natural selection. No +doubt such advancement demands many favourable concurrent circumstances; +but it may well be doubted whether the most favourable would have +sufficed, had not the rate of increase been rapid, and the consequent +struggle for existence severe to an extreme degree. + + +_On the evidence that all civilised nations were once barbarous._--As we +have had to consider the steps by which some semi-human creature has +been gradually raised to the rank of man in his most perfect state, the +present subject cannot be quite passed over. But it has been treated in +so full and admirable a manner by Sir J. Lubbock,[249] Mr. Tylor, Mr. +M'Lennan, and others, that I need here give only the briefest summary of +their results. The arguments recently advanced by the Duke of +Argyll[250] and formerly by Archbishop Whately, in favour of the belief +that man came into the world as a civilised being and that all savages +have since undergone degradation, seem to me weak in comparison with +those advanced on the other side. Many nations, no doubt, have fallen +away in civilisation, and some may have lapsed into utter barbarism, +though on this latter head I have not met with any evidence. The +Fuegians were probably compelled by other conquering hordes to settle in +their inhospitable country, and they may have become in consequence +somewhat more degraded; but it would be difficult to prove that they +have fallen much below the Botocudos who inhabit the finest parts of +Brazil. + +The evidence that all civilised nations are the descendants of +barbarians, consists, on the one side, of clear traces of their former +low condition in still-existing customs, beliefs, language, &c.; and on +the other side, of proofs that savages are independently able to raise +themselves a few steps in the scale of civilisation, and have actually +thus risen. The evidence on the first head is extremely curious, but +cannot be here given: I refer to such cases as that, for instance, of +the art of enumeration, which, as Mr. Tylor clearly shews by the words +still used in some places, originated in counting the fingers, first of +one hand and then of the other, and lastly of the toes. We have traces +of this in our own decimal system, and in the Roman numerals, which +after reaching to the number V., change into VI., &c., when the other +hand no doubt was used. So again, "when we speak of three-score and ten, +we are counting by the vigesimal system, each score thus ideally made, +standing for 20--for 'one man' as a Mexican or Carib would put it."[251] +According to a large and increasing school of philologists, every +language bears the marks of its slow and gradual evolution. So it is +with the art of writing, as letters are rudiments of pictorial +representations. It is hardly possible to read Mr. M'Lennan's work[252] +and not admit that almost all civilised nations still retain some traces +of such rude habits as the forcible capture of wives. What ancient +nation, as the same author asks, can be named that was originally +monogamous? The primitive idea of justice, as shewn by the law of battle +and other customs of which traces still remain, was likewise most rude. +Many existing superstitions are the remnants of former false religious +beliefs. The highest form of religion--the grand idea of God hating sin +and loving righteousness--was unknown during primeval times. + +Turning to the other kind of evidence: Sir J. Lubbock has shewn that +some savages have recently improved a little in some of their simpler +arts. From the extremely curious account which he gives of the weapons, +tools, and arts, used or practised by savages in various parts of the +world, it cannot be doubted that these have nearly all been independent +discoveries, excepting perhaps the art of making fire.[253] The +Australian boomerang is a good instance of one such independent +discovery. The Tahitians when first visited had advanced in many +respects beyond the inhabitants of most of the other Polynesian islands. +There are no just grounds for the belief that the high culture of the +native Peruvians and Mexicans was derived from any foreign source;[254] +many native plants were there cultivated, and a few native animals +domesticated. We should bear in mind that a wandering crew from some +semi-civilised land, if washed to the shores of America, would not, +judging from the small influence of most missionaries, have produced any +marked effect on the natives, unless they had already become somewhat +advanced. Looking to a very remote period in the history of the world, +we find, to use Sir J. Lubbock's well-known terms, a paleolithic and +neolithic period; and no one will pretend that the art of grinding rough +flint tools was a borrowed one. In all parts of Europe, as far east as +Greece, in Palestine, India, Japan, New Zealand, and Africa, including +Egypt, flint tools have been discovered in abundance; and of their use +the existing inhabitants retain no tradition. There is also indirect +evidence of their former use by the Chinese and ancient Jews. Hence +there can hardly be a doubt that the inhabitants of these many +countries, which include nearly the whole civilised world, were once in +a barbarous condition. To believe that man was aboriginally civilised +and then suffered utter degradation in so many regions, is to take a +pitiably low view of human nature. It is apparently a truer and more +cheerful view that progress has been much more general than +retrogression; that man has risen, though by slow and interrupted steps, +from a lowly condition to the highest standard as yet attained by him in +knowledge, morals, and religion. + + + + +CHAPTER VI. + +ON THE AFFINITIES AND GENEALOGY OF MAN. + + + Position of man in the animal series--The natural system + genealogical--Adaptive characters of slight value--Various + small points of resemblance between man and the + Quadrumana--Rank of man in the natural system--Birthplace and + antiquity of man--Absence of fossil connecting-links--Lower + stages in the genealogy of man, as inferred, firstly from his + affinities and secondly from his structure--Early androgynous + condition of the Vertebrata--Conclusion. + + +Even if it be granted that the difference between man and his nearest +allies is as great in corporeal structure as some naturalists maintain, +and although we must grant that the difference between them is immense +in mental power, yet the facts given in the previous chapters declare, +as it appears to me, in the plainest manner, that man is descended from +some lower form, notwithstanding that connecting-links have not hitherto +been discovered. + +Man is liable to numerous, slight, and diversified variations, which are +induced by the same general causes, are governed and transmitted in +accordance with the same general laws, as in the lower animals. Man +tends to multiply at so rapid a rate that his offspring are necessarily +exposed to a struggle for existence, and consequently to natural +selection. He has given rise to many races, some of which are so +different that they have often been ranked by naturalists as distinct +species. His body is constructed on the same homological plan as that of +other mammals, independently of the uses to which the several parts may +be put. He passes through the same phases of embryological development. +He retains many rudimentary and useless structures, which no doubt were +once serviceable. Characters occasionally make their reappearance in +him, which we have every reason to believe were possessed by his early +progenitors. If the origin of man had been wholly different from that of +all other animals, these various appearances would be mere empty +deceptions; but such an admission is incredible. These appearances, on +the other hand, are intelligible, at least to a large extent, if man is +the co-descendant with other mammals of some unknown and lower form. + +Some naturalists, from being deeply impressed with the mental and +spiritual powers of man, have divided the whole organic world into three +kingdoms, the Human, the Animal, and the Vegetable, thus giving to man a +separate kingdom.[255] Spiritual powers cannot be compared or classed by +the naturalist; but he may endeavour to shew, as I have done, that the +mental faculties of man and the lower animals do not differ in kind, +although immensely in degree. A difference in degree, however great, +does not justify us in placing man in a distinct kingdom, as will +perhaps be best illustrated by comparing the mental powers of two +insects, namely, a coccus or scale-insect and an ant, which undoubtedly +belong to the same class. The difference is here greater, though of a +somewhat different kind, than that between man and the highest mammal. +The female coccus, whilst young, attaches itself by its proboscis to a +plant; sucks the sap but never moves again; is fertilised and lays eggs; +and this is its whole history. On the other hand, to describe the habits +and mental powers of a female ant, would require, as Pierre Huber has +shewn, a large volume; I may, however, briefly specify a few points. +Ants communicate information to each other, and several unite for the +same work, or games of play. They recognise their fellow-ants after +months of absence. They build great edifices, keep them clean, close the +doors in the evening, and post sentries. They make roads, and even +tunnels under rivers. They collect food for the community, and when an +object, too large for entrance, is brought to the nest, they enlarge the +door, and afterwards build it up again.[256] They go out to battle in +regular bands, and freely sacrifice their lives for the common weal. +They emigrate in accordance with a preconcerted plan. They capture +slaves. They keep Aphides as milch-cows. They move the eggs of their +aphides, as well as their own eggs and cocoons, into warm parts of the +nest, in order that they may be quickly hatched; and endless similar +facts could be given. On the whole, the difference in mental power +between an ant and a coccus is immense; yet no one has ever dreamed of +placing them in distinct classes, much less in distinct kingdoms. No +doubt this interval is bridged over by the intermediate mental powers of +many other insects; and this is not the case with man and the higher +apes. But we have every reason to believe that breaks in the series are +simply the result of many forms having become extinct. + +Professor Owen, relying chiefly on the structure of the brain, has +divided the mammalian series into four sub-classes. One of these he +devotes to man; in another he places both the marsupials and the +monotremata; so that he makes man as distinct from all other mammals as +are these two latter groups conjoined. This view has not been accepted, +as far as I am aware, by any naturalist capable of forming an +independent judgment, and therefore need not here be further considered. + +We can understand why a classification founded on any single character +or organ--even an organ so wonderfully complex and important as the +brain--or on the high development of the mental faculties, is almost +sure to prove unsatisfactory. This principle has indeed been tried with +hymenopterous insects; but when thus classed by their habits or +instincts, the arrangement proved thoroughly artificial.[257] +Classifications may, of course, be based on any character whatever, as +on size, colour, or the element inhabited; but naturalists have long +felt a profound conviction that there is a natural system. This system, +it is now generally admitted, must be, as far as possible, genealogical +in arrangement,--that is, the co-descendants of the same form must be +kept together in one group, separate from the co-descendants of any +other form; but if the parent-forms are related, so will be their +descendants, and the two groups together will form a larger group. The +amount of difference between the several groups--that is the amount of +modification which each has undergone--will be expressed by such terms +as genera, families, orders, and classes. As we have no record of the +lines of descent, these lines can be discovered only by observing the +degrees of resemblance between the beings which are to be classed. For +this object numerous points of resemblance are of much more importance +than the amount of similarity or dissimilarity in a few points. If two +languages were found to resemble each other in a multitude of words and +points of construction, they would be universally recognised as having +sprung from a common source, notwithstanding that they differed greatly +in some few words or points of construction. But with organic beings the +points of resemblance must not consist of adaptations to similar habits +of life: two animals may, for instance, have had their whole frames +modified for living in the water, and yet they will not be brought any +nearer to each other in the natural system. Hence we can see how it is +that resemblances in unimportant structures, in useless and rudimentary +organs, and in parts not as yet fully developed or functionally active, +are by far the most serviceable for classification; for they can hardly +be due to adaptations within a late period; and thus they reveal the old +lines of descent or of true affinity. + +We can further see why a great amount of modification in some one +character ought not to lead us to separate widely any two organisms. A +part which already differs much from the same part in other allied forms +has already, according to the theory of evolution, varied much; +consequently it would (as long as the organism remained exposed to the +same exciting conditions) be liable to further variations of the same +kind; and these, if beneficial, would be preserved, and thus continually +augmented. In many cases the continued development of a part, for +instance, of the beak of a bird, or of the teeth of a mammal, would not +be advantageous to the species for gaining its food, or for any other +object; but with man we can see no definite limit, as far as advantage +is concerned, to the continued development of the brain and mental +faculties. Therefore in determining the position of man in the natural +or genealogical system, the extreme development of his brain ought not +to outweigh a multitude of resemblances in other less important or +quite unimportant points. + +The greater number of naturalists who have taken into consideration the +whole structure of man, including his mental faculties, have followed +Blumenbach and Cuvier, and have placed man in a separate Order, under +the title of the Bimana, and therefore on an equality with the Orders of +the Quadrumana, Carnivora, &c. Recently many of our best naturalists +have recurred to the view first propounded by Linnus, so remarkable for +his sagacity, and have placed man in the same Order with the Quadrumana, +under the title of the Primates. The justice of this conclusion will be +admitted if, in the first place, we bear in mind the remarks just made +on the comparatively small importance for classification of the great +development of the brain in man; bearing, also, in mind that the +strongly-marked differences between the skulls of man and the Quadrumana +(lately insisted upon by Bischoff, Aeby, and others) apparently follow +from their differently developed brains. In the second place, we must +remember that nearly all the other and more important differences +between man and the Quadrumana are manifestly adaptive in their nature, +and relate chiefly to the erect position of man; such as the structure +of his hand, foot, and pelvis, the curvature of his spine, and the +position of his head. The family of seals offers a good illustration of +the small importance of adaptive characters for classification. These +animals differ from all other Carnivora in the form of their bodies and +in the structure of their limbs, far more than does man from the higher +apes; yet in every system, from that of Cuvier to the most recent one by +Mr. Flower,[258] seals are ranked as a mere family in the Order of the +Carnivora. If man had not been his own classifier, he would never have +thought of founding a separate order for his own reception. + +It would be beyond my limits, and quite beyond my knowledge, even to +name the innumerable points of structure in which man agrees with the +other Primates. Our great anatomist and philosopher, Prof. Huxley, has +fully discussed this subject,[259] and has come to the conclusion that +man in all parts of his organisation differs less from the higher apes, +than these do from the lower members of the same group. Consequently +there "is no justification for placing man in a distinct order." + +In an early part of this volume I brought forward various facts, shewing +how closely man agrees in constitution with the higher mammals; and this +agreement, no doubt, depends on our close similarity in minute structure +and chemical composition. I gave, as instances, our liability to the +same diseases, and to the attacks of allied parasites; our tastes in +common for the same stimulants, and the similar effects thus produced, +as well as by various drugs; and other such facts. + +As small unimportant points of resemblance between man and the higher +apes are not commonly noticed in systematic works, and as, when +numerous, they clearly reveal our relationship, I will specify a few +such points. The relative position of the features are manifestly the +same in man and the Quadrumana; and the various emotions are displayed +by nearly similar movements of the muscles and skin, chiefly above the +eyebrows and round the mouth. Some few expressions are, indeed, almost +the same, as in the weeping of certain kinds of monkeys, and in the +laughing noise made by others, during which the corners of the mouth are +drawn backwards, and the lower eyelids wrinkled. The external ears are +curiously alike. In man the nose is much more prominent than in most +monkeys; but we may trace the commencement of an aquiline curvature in +the nose of the Hoolock Gibbon; and this in the _Semnopithecus nasica_ +is carried to a ridiculous extreme. + +The faces of many monkeys are ornamented with beards, whiskers, or +moustaches. The hair on the head grows to a great length in some species +of Semnopithecus;[260] and in the Bonnet monkey (_Macacus radiatus_) it +radiates from a point on the crown, with a parting down the middle, as +in man. It is commonly said that the forehead gives to man his noble and +intellectual appearance; but the thick hair on the head of the Bonnet +monkey terminates abruptly downwards, and is succeeded by such short and +fine hair, or down, that at a little distance the forehead, with the +exception of the eyebrows, appears quite naked. It has been erroneously +asserted that eyebrows are not present in any monkey. In the species +just named the degree of nakedness of the forehead differs in different +individuals; and Eschricht states[261] that in our children the limit +between the hairy scalp and the naked forehead is sometimes not well +defined; so that here we seem to have a trifling case of reversion to a +progenitor, in whom the forehead had not as yet become quite naked. + +It is well known that the hair on our arms tends to converge from above +and below to a point at the elbow. This curious arrangement, so unlike +that in most of the lower mammals, is common to the gorilla, chimpanzee, +orang, some species of Hylobates, and even to some few American monkeys. +But in _Hylobates agilis_ the hair on the fore-arm is directed +downwards or towards the wrist in the ordinary manner; and in _H. lar_ +it is nearly erect, with only a very slight forward inclination; so that +in this latter species it is in a transitional state. It can hardly be +doubted that with most mammals the thickness of the hair and its +direction on the back is adapted to throw off the rain; even the +transverse hairs on the fore-legs of a dog may serve for this end when +he is coiled up asleep. Mr. Wallace remarks that the convergence of the +hair towards the elbow on the arms of the orang (whose habits he has so +carefully studied) serves to throw off the rain, when, as is the custom +of this animal, the arms are bent, with the hands clasped round a branch +or over its own head. We should, however, bear in mind that the attitude +of an animal may perhaps be in part determined by the direction of the +hair; and not the direction of the hair by the attitude. If the above +explanation is correct in the case of the orang, the hair on our +fore-arms offers a curious record of our former state; for no one +supposes that it is now of any use in throwing off the rain, nor in our +present erect condition is it properly directed for this purpose. + +It would, however, be rash to trust too much to the principle of +adaptation in regard to the direction of the hair in man or his early +progenitors; for it is impossible to study the figures given by +Eschricht of the arrangement of the hair on the human foetus (this +being the same as in the adult) and not agree with this excellent +observer that other and more complex causes have intervened. The points +of convergence seem to stand in some relation to those points in the +embryo which are last closed in during development. There appears, also, +to exist some relation between the arrangement of the hair on the +limbs, and the course of the medullary arteries.[262] + +It must not be supposed that the resemblances between man and certain +apes in the above and many other points--such as in having a naked +forehead, long tresses on the head, &c.--are all necessarily the result +of unbroken inheritance from a common progenitor thus characterised, or +of subsequent reversion. Many of these resemblances are more probably +due to analogous variation, which follows, as I have elsewhere attempted +to shew,[263] from co-descended organisms having a similar constitution +and having been acted on by similar causes inducing variability. With +respect to the similar direction of the hair on the fore-arms of man and +certain monkeys, as this character is common to almost all the +anthropomorphous apes, it may probably be attributed to inheritance; but +not certainly so, as some very distinct American monkeys are thus +characterised. The same remark is applicable to the tailless condition +of man; for the tail is absent in all the anthropomorphous apes. +Nevertheless this character cannot with certainty be attributed to +inheritance, as the tail, though not absent, is rudimentary in several +other Old World and in some New World species, and is quite absent in +several species belonging to the allied group of Lemurs. + +Although, as we have now seen, man has no just right to form a separate +Order for his own reception, he may perhaps claim a distinct Sub-order +or Family. Prof. Huxley, in his last work,[264] divides the Primates +into three Sub-orders; namely, the Anthropid with man alone, the +Simiad including monkeys of all kinds, and the Lemurid with the +diversified genera of lemurs. As far as differences in certain important +points of structure are concerned, man may no doubt rightly claim the +rank of a Sub-order; and this rank is too low, if we look chiefly to his +mental faculties. Nevertheless, under a genealogical point of view it +appears that this rank is too high, and that man ought to form merely a +Family, or possibly even only a Sub-family. If we imagine three lines of +descent proceeding from a common source, it is quite conceivable that +two of them might after the lapse of ages be so slightly changed as +still to remain as species of the same genus; whilst the third line +might become so greatly modified as to deserve to rank as a distinct +Sub-family, Family, or even Order. But in this case it is almost certain +that the third line would still retain through inheritance numerous +small points of resemblance with the other two lines. Here then would +occur the difficulty, at present insoluble, how much weight we ought to +assign in our classifications to strongly-marked differences in some few +points,--that is to the amount of modification undergone; and how much +to close resemblance in numerous unimportant points, as indicating the +lines of descent or genealogy. The former alternative is the most +obvious, and perhaps the safest, though the latter appears the most +correct as giving a truly natural classification. + +To form a judgment on this head, with reference to man we must glance at +the classification of the Simiad. This family is divided by almost all +naturalists into the Catarhine group, or Old World monkeys, all of which +are characterised (as their name expresses) by the peculiar structure of +their nostrils and by having four premolars in each jaw; and into the +Platyrhine group or New World monkeys (including two very distinct +sub-groups), all of which are characterised by differently-constructed +nostrils and by having six premolars in each jaw. Some other small +differences might be mentioned. Now man unquestionably belongs in his +dentition, in the structure of his nostrils, and some other respects, to +the Catarhine or Old World division; nor does he resemble the +Platyrhines more closely than the Catarhines in any characters, +excepting in a few of not much importance and apparently of an adaptive +nature. Therefore it would be against all probability to suppose that +some ancient New World species had varied, and had thus produced a +man-like creature with all the distinctive characters proper to the Old +World division; losing at the same time all its own distinctive +characters. There can consequently hardly be a doubt that man is an +offshoot from the Old World Simian stem; and that under a genealogical +point of view, he must be classed with the Catarhine division.[265] + +The anthropomorphous apes, namely the gorilla, chimpanzee, orang, and +hylobates, are separated as a distinct sub-group from the other Old +World monkeys by most naturalists. I am aware that Gratiolet, relying on +the structure of the brain, does not admit the existence of this +sub-group, and no doubt it is a broken one; thus the orang, as Mr. St. +G. Mivart remarks,[266] "is one of the most peculiar and aberrant forms +to be found in the Order." The remaining, non-anthropomorphous, Old +World monkeys, are again divided by some naturalists into two or three +smaller sub-groups; the genus Semnopithecus, with its peculiar +sacculated stomach, being the type of one such sub-group. But it appears +from M. Gaudry's wonderful discoveries in Attica, that during the +Miocene period a form existed there, which connected Semnopithecus and +Macacus; and this probably illustrates the manner in which the other and +higher groups were once blended together. + +If the anthropomorphous apes be admitted to form a natural sub-group, +then as man agrees with them, not only in all those characters which he +possesses in common with the whole Catarhine group, but in other +peculiar characters, such as the absence of a tail and of callosities +and in general appearance, we may infer that some ancient member of the +anthropomorphous sub-group gave birth to man. It is not probable that a +member of one of the other lower sub-groups should, through the law of +analogous variation, have given rise to a man-like creature, resembling +the higher anthropomorphous apes in so many respects. No doubt man, in +comparison with most of his allies, has undergone an extraordinary +amount of modification, chiefly in consequence of his greatly developed +brain and erect position; nevertheless we should bear in mind that he +"is but one of several exceptional forms of Primates."[267] + +Every naturalist, who believes in the principle of evolution, will +grant that the two main divisions of the Simiad, namely the Catarhine +and Platyrhine monkeys, with their sub-groups, have all proceeded from +some one extremely ancient progenitor. The early descendants of this +progenitor, before they had diverged to any considerable extent from +each other, would still have formed a single natural group; but some of +the species or incipient genera would have already begun to indicate by +their diverging characters the future distinctive marks of the Catarhine +and Platyrhine divisions. Hence the members of this supposed ancient +group would not have been so uniform in their dentition or in the +structure of their nostrils, as are the existing Catarhine monkeys in +one way and the Platyrhines in another way, but would have resembled in +this respect the allied Lemurid which differ greatly from each other in +the form of their muzzles,[268] and to an extraordinary degree in their +dentition. + +The Catarhine and Platyrhine monkeys agree in a multitude of characters, +as is shewn by their unquestionably belonging to one and the same Order. +The many characters which they possess in common can hardly have been +independently acquired by so many distinct species; so that these +characters must have been inherited. But an ancient form which possessed +many characters common to the Catarhine and Platyrhine monkeys, and +others in an intermediate condition, and some few perhaps distinct from +those now present in either group, would undoubtedly have been ranked, +if seen by a naturalist, as an ape or monkey. And as man under a +genealogical point of view belongs to the Catarhine or Old World stock, +we must conclude, however much the conclusion may revolt our pride, +that our early progenitors would have been properly thus +designated.[269] But we must not fall into the error of supposing that +the early progenitor of the whole Simian stock, including man, was +identical with, or even closely resembled, any existing ape or monkey. + + +_On the Birthplace and Antiquity of Man._--We are naturally led to +enquire where was the birthplace of man at that stage of descent when +our progenitors diverged from the Catarhine stock. The fact that they +belonged to this stock clearly shews that they inhabited the Old World; +but not Australia nor any oceanic island, as we may infer from the laws +of geographical distribution. In each great region of the world the +living mammals are closely related to the extinct species of the same +region. It is therefore probable that Africa was formerly inhabited by +extinct apes closely allied to the gorilla and chimpanzee; and as these +two species are now man's nearest allies, it is somewhat more probable +that our early progenitors lived on the African continent than +elsewhere. But it is useless to speculate on this subject, for an ape +nearly as large as a man, namely the Dryopithecus of Lartet, which was +closely allied to the anthropomorphous Hylobates, existed in Europe +during the Upper Miocene period; and since so remote a period the earth +has certainly undergone many great revolutions, and there has been ample +time for migration on the largest scale. + +At the period and place, whenever and wherever it may have been, when +man first lost his hairy covering, he probably inhabited a hot country; +and this would have been favourable for a frugiferous diet, on which, +judging from analogy, he subsisted. We are far from knowing how long ago +it was when man first diverged from the Catarhine stock; but this may +have occurred at an epoch as remote as the Eocene period; for the higher +apes had diverged from the lower apes as early as the Upper Miocene +period, as shewn by the existence of the Dryopithecus. We are also quite +ignorant at how rapid a rate organisms, whether high or low in the +scale, may under favourable circumstances be modified: we know, however, +that some have retained the same form during an enormous lapse of time. +From what we see going on under domestication, we learn that within the +same period some of the co-descendants of the same species may be not at +all changed, some a little, and some greatly changed. Thus it may have +been with man, who has undergone a great amount of modification in +certain characters in comparison with the higher apes. + +The great break in the organic chain between man and his nearest allies, +which cannot be bridged over by any extinct or living species, has often +been advanced as a grave objection to the belief that man is descended +from some lower form; but this objection will not appear of much weight +to those who, convinced by general reasons, believe in the general +principle of evolution. Breaks incessantly occur in all parts of the +series, some being wide, sharp and defined, others less so in various +degrees; as between the orang and its nearest allies--between the +Tarsius and the other Lemurid--between the elephant and in a more +striking manner between the Ornithorhynchus or Echidna, and other +mammals. But all these breaks depend merely on the number of related +forms which have become extinct. At some future period, not very distant +as measured by centuries, the civilised races of man will almost +certainly exterminate and replace throughout the world the savage races. +At the same time the anthropomorphous apes, as Professor Schaaffhausen +has remarked,[270] will no doubt be exterminated. The break will then be +rendered wider, for it will intervene between man in a more civilised +state, as we may hope, than the Caucasian, and some ape as low as a +baboon, instead of as at present between the negro or Australian and the +gorilla. + +With respect to the absence of fossil remains, serving to connect man +with his ape-like progenitors, no one will lay much stress on this fact, +who will read Sir C. Lyell's discussion,[271] in which he shews that in +all the vertebrate classes the discovery of fossil remains has been an +extremely slow and fortuitous process. Nor should it be forgotten that +those regions which are the most likely to afford remains connecting man +with some extinct ape-like creature, have not as yet been searched by +geologists. + + +_Lower Stages in the Genealogy of Man._--We have seen that man appears +to have diverged from the Catarhine or Old World division of the +Simiad, after these had diverged from the New World division. We will +now endeavour to follow the more remote traces of his genealogy, +trusting in the first place to the mutual affinities between the various +classes and orders, with some slight aid from the periods, as far as +ascertained, of their successive appearance on the earth. The Lemurid +stand below and close to the Simiad, constituting a very distinct +family of the Primates, or, according to Hckel, a distinct Order. This +group is diversified and broken to an extraordinary degree, and includes +many aberrant forms. It has, therefore, probably suffered much +extinction. Most of the remnants survive on islands, namely in +Madagascar and in the islands of the Malayan archipelago, where they +have not been exposed to such severe competition as they would have been +on well-stocked continents. This group likewise presents many +gradations, leading, as Huxley remarks,[272] "insensibly from the crown +and summit of the animal creation down to creatures from which there is +but a step, as it seems, to the lowest, smallest, and least intelligent +of the placental mammalia." From these various considerations it is +probable that the Simiad were originally developed from the progenitors +of the existing Lemurid; and these in their turn from forms standing +very low in the mammalian series. + +The Marsupials stand in many important characters below the placental +mammals. They appeared at an earlier geological period, and their range +was formerly much more extensive than what it now is. Hence the +Placentata are generally supposed to have been derived from the +Implacentata or Marsupials; not, however, from forms closely like the +existing Marsupials, but from their early progenitors. The Monotremata +are plainly allied to the Marsupials; forming a third and still lower +division in the great mammalian series. They are represented at the +present day solely by the Ornithorhynchus and Echidna; and these two +forms may be safely considered as relics of a much larger group which +have been preserved in Australia through some favourable concurrence of +circumstances. The Monotremata are eminently interesting, as in several +important points of structure they lead towards the class of reptiles. + +In attempting to trace the genealogy of the Mammalia, and therefore of +man, lower down in the series, we become involved in greater and greater +obscurity. He who wishes to see what ingenuity and knowledge can effect, +may consult Prof. Hckel's works.[273] I will content myself with a few +general remarks. Every evolutionist will admit that the five great +vertebrate classes, namely, mammals, birds, reptiles, amphibians, and +fishes, are all descended from some one prototype; for they have much in +common, especially during their embryonic state. As the class of fishes +is the most lowly organised and appeared before the others, we may +conclude that all the members of the vertebrate kingdom are derived from +some fish-like animal, less highly organised than any as yet found in +the lowest known formations. The belief that animals so distinct as a +monkey or elephant and a humming-bird, a snake, frog, and fish, &c., +could all have sprung from the same parents, will appear monstrous to +those who have not attended to the recent progress of natural history. +For this belief implies the former existence of links closely binding +together all these forms, now so utterly unlike. + +Nevertheless it is certain that groups of animals have existed, or do +now exist, which serve to connect more or less closely the several great +vertebrate classes. We have seen that the Ornithorhynchus graduates +towards reptiles; and Prof. Huxley has made the remarkable discovery, +confirmed by Mr. Cope and others, that the old Dinosaurians are +intermediate in many important respects between certain reptiles and +certain birds--the latter consisting of the ostrich-tribe (itself +evidently a widely-diffused remnant of a larger group) and of the +Archeopteryx, that strange Secondary bird having a long tail like that +of the lizard. Again, according to Prof. Owen,[274] the +Ichthyosaurians--great sea-lizards furnished with paddles--present many +affinities with fishes, or rather, according to Huxley, with amphibians. +This latter class (including in its highest division frogs and toads) is +plainly allied to the Ganoid fishes. These latter fishes swarmed during +the earlier geological periods, and were constructed on what is called a +highly generalised type, that is they presented diversified affinities +with other groups of organisms. The amphibians and fishes are also so +closely united by the Lepidosiren, that naturalists long disputed in +which of these two classes it ought to be placed. The Lepidosiren and +some few Ganoid fishes have been preserved from utter extinction by +inhabiting our rivers, which are harbours of refuge, bearing the same +relation to the great waters of the ocean that islands bear to +continents. + +Lastly, one single member of the immense and diversified class of +fishes, namely the lancelet or amphioxus, is so different from all other +fishes, that Hckel maintains that it ought to form a distinct class in +the vertebrate kingdom. This fish is remarkable for its negative +characters; it can hardly be said to possess a brain, vertebral column, +or heart, &c.; so that it was classed by the older naturalists amongst +the worms. Many years ago Prof. Goodsir perceived that the lancelet +presented some affinities with the Ascidians, which are invertebrate, +hermaphrodite, marine creatures permanently attached to a support. They +hardly appear like animals, and consist of a simple, tough, leathery +sack, with two small projecting orifices. They belong to the Molluscoida +of Huxley--a lower division of the great kingdom of the Mollusca; but +they have recently been placed by some naturalists amongst the Vermes or +worms. Their larv somewhat resemble tadpoles in shape,[275] and have +the power of swimming freely about. Some observations lately made by M. +Kowalevsky,[276] since confirmed by Prof. Kuppfer, will form a discovery +of extraordinary interest, if still further extended, as I hear from M. +Kowalevsky in Naples he has now effected. The discovery is that the +larv of Ascidians are related to the Vertebrata, in their manner of +development, in the relative position of the nervous system, and in +possessing a structure closely like the _chorda dorsalis_ of vertebrate +animals. It thus appears, if we may rely on embryology, which has always +proved the safest guide in classification, that we have at last gained a +clue to the source whence the Vertebrata have been derived. We should +thus be justified in believing that at an extremely remote period a +group of animals existed, resembling in many respects the larv of our +present Ascidians, which diverged into two great branches--the one +retrograding in development and producing the present class of +Ascidians, the other rising to the crown and summit of the animal +kingdom by giving birth to the Vertebrata. + + +We have thus far endeavoured rudely to trace the genealogy of the +Vertebrata by the aid of their mutual affinities. We will now look to +man as he exists; and we shall, I think, be able partially to restore +during successive periods, but not in due order of time, the structure +of our early progenitors. This can be effected by means of the rudiments +which man still retains, by the characters which occasionally make their +appearance in him through reversion, and by the aid of the principles of +morphology and embryology. The various facts, to which I shall here +allude, have been given in the previous chapters. The early progenitors +of man were no doubt once covered with hair, both sexes having beards; +their ears were pointed and capable of movement; and their bodies were +provided with a tail, having the proper muscles. Their limbs and bodies +were also acted on by many muscles which now only occasionally reappear, +but are normally present in the Quadrumana. The great artery and nerve +of the humerus ran through a supra-condyloid foramen. At this or some +earlier period, the intestine gave forth a much larger diverticulum or +ccum than that now existing. The foot, judging from the condition of +the great toe in the foetus, was then prehensile; and our progenitors, +no doubt, were arboreal in their habits, frequenting some warm, +forest-clad land. The males were provided with great canine teeth, +which served them as formidable weapons. + +At a much earlier period the uterus was double; the excreta were voided +through a cloaca; and the eye was protected by a third eyelid or +nictitating membrane. At a still earlier period the progenitors of man +must have been aquatic in their habits; for morphology plainly tells us +that our lungs consist of a modified swim-bladder, which once served as +a float. The clefts on the neck in the embryo of man show where the +branchi once existed. At about this period the true kidneys were +replaced by the corpora Wolffiana. The heart existed as a simple +pulsating vessel; and the chorda dorsalis took the place of a vertebral +column. These early predecessors of man, thus seen in the dim recesses +of time, must have been as lowly organised as the lancelet or amphioxus, +or even still more lowly organised. + +There is one other point deserving a fuller notice. It has long been +known that in the vertebrate kingdom one sex bears rudiments of various +accessory parts, appertaining to the reproductive system, which properly +belong to the opposite sex; and it has now been ascertained that at a +very early embryonic period both sexes possess true male and female +glands. Hence some extremely remote progenitor of the whole vertebrate +kingdom appears to have been hermaphrodite or androgynous.[277] But here +we encounter a singular difficulty. In the mammalian class the males +possess in their vesicul prostatic rudiments of a uterus with the +adjacent passage; they bear also rudiments of mamm, and some male +marsupials have rudiments of a marsupial sack.[278] Other analogous +facts could be added. Are we, then, to suppose that some extremely +ancient mammal possessed organs proper to both sexes, that is, continued +androgynous after it had acquired the chief distinctions of its proper +class, and therefore after it had diverged from the lower classes of the +vertebrate kingdom? This seems improbable in the highest degree; for had +this been the case, we might have expected that some few members of the +two lower classes, namely fishes[279] and amphibians, would still have +remained androgynous. We must, on the contrary, believe that when the +five vertebrate classes diverged from their common progenitor the sexes +had already become separated. To account, however, for male mammals +possessing rudiments of the accessory female organs, and for female +mammals possessing rudiments of the masculine organs, we need not +suppose that their early progenitors were still androgynous after they +had assumed their chief mammalian characters. It is quite possible that +as the one sex gradually acquired the accessory organs proper to it, +some of the successive steps or modifications were transmitted to the +opposite sex. When we treat of sexual selection, we shall meet with +innumerable instances of this form of transmission,--as in the case of +the spurs, plumes, and brilliant colours, acquired by male birds for +battle or ornament, and transferred to the females in an imperfect or +rudimentary condition. + +The possession by male mammals of functionally imperfect mammary organs +is, in some respects, especially curious. The Monotremata have the +proper milk-secreting glands with orifices, but no nipples; and as these +animals stand at the very base of the mammalian series, it is probable +that the progenitors of the class possessed, in like manner, the +milk-secreting glands, but no nipples. This conclusion is supported by +what is known of their manner of development; for Professor Turner +informs me, on the authority of Klliker and Lauger, that in the embryo +the mammary glands can be distinctly traced before the nipples are in +the least visible; and it should be borne in mind that the development +of successive parts in the individual generally seems to represent and +accord with the development of successive beings in the same line of +descent. The Marsupials differ from the Monotremata by possessing +nipples; so that these organs were probably first acquired by the +Marsupials after they had diverged from, and risen above, the +Monotremata, and were then transmitted to the placental mammals. No one +will suppose that after the Marsupials had approximately acquired their +present structure, and therefore at a rather late period in the +development of the mammalian series, any of its members still remained +androgynous. We seem, therefore, compelled to recur to the foregoing +view, and to conclude that the nipples were first developed in the +females of some very early marsupial form, and were then, in accordance +with a common law of inheritance, transferred in a functionally +imperfect condition to the males. + +Nevertheless a suspicion has sometimes crossed my mind that long after +the progenitors of the whole mammalian class had ceased to be +androgynous, both sexes might have yielded milk and thus nourished their +young; and in the case of the Marsupials, that both sexes might have +carried their young in marsupial sacks. This will not appear utterly +incredible, if we reflect that the males of syngnathous fishes receive +the eggs of the females in their abdominal pouches, hatch them, and +afterwards, as some believe, nourish the young;[280]--that certain other +male fishes hatch the eggs within their mouths or branchial +cavities;--that certain male toads take the chaplets of eggs from the +females and wind them round their own thighs, keeping them there until +the tadpoles are born;--that certain male birds undertake the whole duty +of incubation, and that male pigeons, as well as the females, feed their +nestlings with a secretion from their crops. But the above suspicion +first occurred to me from the mammary glands in male mammals being +developed so much more perfectly than the rudiments of those other +accessory reproductive parts, which are found in the one sex though +proper to the other. The mammary glands and nipples, as they exist in +male mammals, can indeed hardly be called rudimentary; they are simply +not fully developed and not functionally active. They are +sympathetically affected under the influence of certain diseases, like +the same organs in the female. At birth they often secrete a few drops +of milk; and they have been known occasionally in man and other mammals +to become well developed, and to yield a fair supply of milk. Now if we +suppose that during a former prolonged period male mammals aided the +females in nursing their offspring, and that afterwards from some cause, +as from a smaller number of young being produced, the males ceased +giving this aid, disuse of the organs during maturity would lead to +their becoming inactive; and from two well-known principles of +inheritance this state of inactivity would probably be transmitted to +the males at the corresponding age of maturity. But at all earlier ages +these organs would be left unaffected, so that they would be equally +well developed in the young of both sexes. + + +_Conclusion._--The best definition of advancement or progress in the +organic scale ever given, is that by Von Baer; and this rests on the +amount of differentiation and specialisation of the several parts of the +same being, when arrived, as I should be inclined to add, at maturity. +Now as organisms have become slowly adapted by means of natural +selection for diversified lines of life, their parts will have become, +from the advantage gained by the division of physiological labour, more +and more differentiated and specialised for various functions. The same +part appears often to have been modified first for one purpose, and then +long afterwards for some other and quite distinct purpose; and thus all +the parts are rendered more and more complex. But each organism will +still retain the general type of structure of the progenitor from which +it was aboriginally derived. In accordance with this view it seems, if +we turn to geological evidence, that organisation on the whole has +advanced throughout the world by slow and interrupted steps. In the +great kingdom of the Vertebrata it has culminated in man. It must not, +however, be supposed that groups of organic beings are always supplanted +and disappear as soon as they have given birth to other and more perfect +groups. The latter, though victorious over their predecessors, may not +have become better adapted for all places in the economy of nature. Some +old forms appear to have survived from inhabiting protected sites, where +they have not been exposed to very severe competition; and these often +aid us in constructing our genealogies, by giving us a fair idea of +former and lost populations. But we must not fall into the error of +looking at the existing members of any lowly-organised group as perfect +representatives of their ancient predecessors. + +The most ancient progenitors in the kingdom of the Vertebrata, at which +we are able to obtain an obscure glance, apparently consisted of a group +of marine animals,[281] resembling the larv of existing Ascidians. +These animals probably gave rise to a group of fishes, as lowly +organised as the lancelet; and from these the Ganoids, and other fishes +like the Lepidosiren, must have been developed. From such fish a very +small advance would carry us on to the amphibians. We have seen that +birds and reptiles were once intimately connected together; and the +Monotremata now, in a slight degree, connect mammals with reptiles. But +no one can at present say by what line of descent the three higher and +related classes, namely, mammals, birds, and reptiles, were derived from +either of the two lower vertebrate classes, namely amphibians and +fishes. In the class of mammals the steps are not difficult to conceive +which led from the ancient Monotremata to the ancient Marsupials; and +from these to the early progenitors of the placental mammals. We may +thus ascend to the Lemurid; and the interval is not wide from these to +the Simiad. The Simiad then branched off into two great stems, the New +World and Old World monkeys; and from the latter, at a remote period, +Man, the wonder and glory of the Universe, proceeded. + +Thus we have given to man a pedigree of prodigious length, but not, it +may be said, of noble quality. The world, it has often been remarked, +appears as if it had long been preparing for the advent of man; and +this, in one sense is strictly true, for he owes his birth to a long +line of progenitors. If any single link in this chain had never existed, +man would not have been exactly what he now is. Unless we wilfully close +our eyes, we may, with our present knowledge, approximately recognise +our parentage; nor need we feel ashamed of it. The most humble organism +is something much higher than the inorganic dust under our feet; and no +one with an unbiassed mind can study any living creature, however +humble, without being struck with enthusiasm at its marvellous structure +and properties. + + + + +CHAPTER VII. + +ON THE RACES OF MAN. + + + The nature and value of specific characters--Application to the + races of man--Arguments in favour of, and opposed to, ranking + the so-called races of man as distinct + species--Sub-species--Monogenists and polygenists--Convergence + of character--Numerous points of resemblance in body and mind + between the most distinct races of man--The state of man when + he first spread over the earth--Each race not descended from a + single pair--The extinction of races--The formation of + races--The effects of crossing--Slight influence of the direct + action of the conditions of life--Slight or no influence of + natural selection--Sexual selection. + + +It is not my intention here to describe the several so-called races of +men; but to inquire what is the value of the differences between them +under a classificatory point of view, and how they have originated. In +determining whether two or more allied forms ought to be ranked as +species or varieties, naturalists are practically guided by the +following considerations; namely, the amount of difference between them, +and whether such differences relate to few or many points of structure, +and whether they are of physiological importance; but more especially +whether they are constant. Constancy of character is what is chiefly +valued and sought for by naturalists. Whenever it can be shewn, or +rendered probable, that the forms in question have remained distinct for +a long period, this becomes an argument of much weight in favour of +treating them as species. Even a slight degree of sterility between any +two forms when first crossed, or in their offspring, is generally +considered as a decisive test of their specific distinctness; and their +continued persistence without blending within the same area, is usually +accepted as sufficient evidence, either of some degree of mutual +sterility, or in the case of animals of some repugnance to mutual +pairing. + +Independently of blending from intercrossing, the complete absence, in a +well-investigated region, of varieties linking together any two +closely-allied forms, is probably the most important of all the +criterions of their specific distinctness; and this is a somewhat +different consideration from mere constancy of character, for two forms +may be highly variable and yet not yield intermediate varieties. +Geographical distribution is often unconsciously and sometimes +consciously brought into play; so that forms living in two widely +separated areas, in which most of the other inhabitants are specifically +distinct, are themselves usually looked at as distinct; but in truth +this affords no aid in distinguishing geographical races from so-called +good or true species. + +Now let us apply these generally-admitted principles to the races of +man, viewing him in the same spirit as a naturalist would any other +animal. In regard to the amount of difference between the races, we must +make some allowance for our nice powers of discrimination gained by the +long habit of observing ourselves. In India, as Elphinstone +remarks,[282] although a newly-arrived European cannot at first +distinguish the various native races, yet they soon appear to him +extremely dissimilar; and the Hindoo cannot at first perceive any +difference between the several European nations. Even the most distinct +races of man, with the exception of certain negro tribes, are much more +like each other in form than would at first be supposed. This is well +shewn by the French photographs in the Collection Anthropologique du +Musum of the men belonging to various races, the greater number of +which, as many persons to whom I have shown them have remarked, might +pass for Europeans. Nevertheless, these men if seen alive would +undoubtedly appear very distinct, so that we are clearly much influenced +in our judgment by the mere colour of the skin and hair, by slight +differences in the features, and by expression. + +There is, however, no doubt that the various races, when carefully +compared and measured, differ much from each other,--as in the texture +of the hair, the relative proportions of all parts of the body,[283] the +capacity of the lungs, the form and capacity of the skull, and even in +the convolutions of the brain.[284] But it would be an endless task to +specify the numerous points of structural difference. The races differ +also in constitution, in acclimatisation, and in liability to certain +diseases. Their mental characteristics are likewise very distinct; +chiefly as it would appear in their emotional, but partly in their +intellectual, faculties. Every one who has had the opportunity of +comparison, must have been struck with the contrast between the +taciturn, even morose, aborigines of S. America and the light-hearted, +talkative negroes. There is a nearly similar contrast between the Malays +and the Papuans,[285] who live under the same physical conditions, and +are separated from each other only by a narrow space of sea. + +We will first consider the arguments which may be advanced in favour of +classing the races of man as distinct species, and then those on the +other side. If a naturalist, who had never before seen such beings, were +to compare a Negro, Hottentot, Australian, or Mongolian, he would at +once perceive that they differed in a multitude of characters, some of +slight and some of considerable importance. On inquiry he would find +that they were adapted to live under widely different climates, and that +they differed somewhat in bodily constitution and mental disposition. If +he were then told that hundreds of similar specimens could be brought +from the same countries, he would assuredly declare that they were as +good species as many to which he had been in the habit of affixing +specific names. This conclusion would be greatly strengthened as soon as +he had ascertained that these forms had all retained the same character +for many centuries; and that negroes, apparently identical with existing +negroes, had lived at least 4000 years ago.[286] He would also hear from +an excellent observer, Dr. Lund,[287] that the human skulls found in +the caves of Brazil, entombed with many extinct mammals, belonged to the +same type as that now prevailing throughout the American Continent. + +Our naturalist would then perhaps turn to geographical distribution, and +he would probably declare that forms differing not only in appearance, +but fitted for the hottest and dampest or driest countries, as well as +for the arctic regions, must be distinct species. He might appeal to the +fact that no one species in the group next to man, namely the +Quadrumana, can resist a low temperature or any considerable change of +climate; and that those species which come nearest to man have never +been reared to maturity, even under the temperate climate of Europe. He +would be deeply impressed with the fact, first noticed by Agassiz,[288] +that the different races of man are distributed over the world in the +same zoological provinces, as those inhabited by undoubtedly distinct +species and genera of mammals. This is manifestly the case with the +Australian, Mongolian, and Negro races of man; in a less well-marked +manner with the Hottentots; but plainly with the Papuans and Malays, who +are separated, as Mr. Wallace has shewn, by nearly the same line which +divides the great Malayan and Australian zoological provinces. The +aborigines of America range throughout the Continent; and this at first +appears opposed to the above rule, for most of the productions of the +Southern and Northern halves differ widely; yet some few living forms, +as the opossum, range from the one into the other, as did formerly some +of the gigantic Edentata. The Esquimaux, like other Arctic animals, +extend round the whole polar regions. It should be observed that the +mammalian forms which inhabit the several zoological provinces, do not +differ from each other in the same degree; so that it can hardly be +considered as an anomaly that the Negro differs more, and the American +much less, from the other races of man than do the mammals of the same +continents from those of the other provinces. Man, it may be added, does +not appear to have aboriginally inhabited any oceanic island; and in +this respect he resembles the other members of his class. + +In determining whether the varieties of the same kind of domestic animal +should be ranked as specifically distinct, that is, whether any of them +are descended from distinct wild species, every naturalist would lay +much stress on the fact, if established, of their external parasites +being specifically distinct. All the more stress would be laid on this +fact, as it would be an exceptional one, for I am informed by Mr. Denny +that the most different kinds of dogs, fowls, and pigeons, in England, +are infested by the same species of Pediculi or lice. Now Mr. A. Murray +has carefully examined the Pediculi collected in different countries +from the different races of man;[289] and he finds that they differ, not +only in colour, but in the structure of their claws and limbs. In every +case in which numerous specimens were obtained the differences were +constant. The surgeon of a whaling ship in the Pacific assured me that +when the Pediculi, with which some Sandwich Islanders on board swarmed, +strayed on to the bodies of the English sailors, they died in the course +of three or four days. These Pediculi were darker coloured and appeared +different from those proper to the natives of Chiloe in South America, +of which he gave me specimens. These, again, appeared larger and much +softer than European lice. Mr. Murray procured four kinds from Africa, +namely from the Negroes of the Eastern and Western coasts, from the +Hottentots and Caffres; two kinds from the natives of Australia; two +from North, and two from South America. In these latter cases it may be +presumed that the Pediculi came from natives inhabiting different +districts. With insects slight structural differences, if constant, are +generally esteemed of specific value: and the fact of the races of man +being infested by parasites, which appear to be specifically distinct, +might fairly be urged as an argument that the races themselves ought to +be classed as distinct species. + +Our supposed naturalist having proceeded thus far in his investigation, +would next inquire whether the races of men, when crossed, were in any +degree sterile. He might consult the work[290] of a cautious and +philosophical observer, Professor Broca; and in this he would find good, +evidence that some races were quite fertile together; but evidence of an +opposite nature in regard to other races. Thus it has been asserted that +the native women of Australia and Tasmania rarely produce children to +European men; the evidence, however, on this head has now been shewn to +be almost valueless. The half-castes are killed by the pure blacks; and +an account has lately been published of eleven half-caste youths +murdered and burnt at the same time, whose remains were found by the +police.[291] Again, it has often been said that when mulattoes +intermarry they produce few children; on the other hand, Dr. Bachman of +Charlestown[292] positively asserts that he has known mulatto families +which have intermarried for several generations, and have continued on +an average as fertile as either pure whites or pure blacks. Inquiries +formerly made by Sir C. Lyell on this subject led him, as he informs me, +to the same conclusion. In the United States the census for the year +1854 included, according to Dr. Bachman, 405,751 mulattoes; and this +number, considering all the circumstances of the case, seems small; but +it may partly be accounted for by the degraded and anomalous position of +the class, and by the profligacy of the women. A certain amount of +absorption of mulattoes into negroes must always be in progress; and +this would lead to an apparent diminution of the former. The inferior +vitality of mulattoes is spoken of in a trustworthy work[293] as a +well-known phenomenon; but this is a different consideration from their +lessened fertility; and can hardly be advanced as a proof of the +specific distinctness of the parent races. No doubt both animal and +vegetable hybrids, when produced from extremely distinct species, are +liable to premature death; but the parents of mulattoes cannot be put +under the category of extremely distinct species. The common Mule, so +notorious for long life and vigour, and yet so sterile, shews how little +necessary connection there is in hybrids between lessened fertility and +vitality: other analogous cases could be added. + +Even if it should hereafter be proved that all the races of men were +perfectly fertile together, he who was inclined from other reasons to +rank them as distinct species, might with justice argue that fertility +and sterility are not safe criterions of specific distinctness. We know +that these qualities are easily affected by changed conditions of life +or by close inter-breeding, and that they are governed by highly complex +laws, for instance that of the unequal fertility of reciprocal crosses +between the same two species. With forms which must be ranked as +undoubted species, a perfect series exists from those which are +absolutely sterile when crossed, to those which are almost or quite +fertile. The degrees of sterility do not coincide strictly with the +degrees of difference in external structure or habits of life. Man in +many respects may be compared with those animals which have long been +domesticated, and a large body of evidence can be advanced in favour of +the Pallasian doctrine[294] that domestication tends to eliminate the +sterility which is so general a result of the crossing of species in a +state of nature. From these several considerations, it may be justly +urged that the perfect fertility of the intercrossed races of man, if +established, would not absolutely preclude us from ranking them as +distinct species. + +Independently of fertility, the character of the offspring from a cross +has sometimes been thought to afford evidence whether the parent-forms +ought to be ranked as species or varieties; but after carefully studying +the evidence, I have come to the conclusion that no general rules of +this kind can be trusted. Thus with mankind the offspring of distinct +races resemble in all respects the offspring of true species and of +varieties. This is shewn, for instance, by the manner in which the +characters of both parents are blended, and by one form absorbing +another through repeated crosses. In this latter case the progeny both +of crossed species and varieties retain for a long period a tendency to +revert to their ancestors, especially to that one which is prepotent in +transmission. When any character has suddenly appeared in a race or +species as the result of a single act of variation, as is general with +monstrosities,[295] and this race is crossed with another not thus +characterised, the characters in question do not commonly appear in a +blended condition in the young, but are transmitted to them either +perfectly developed or not at all. As with the crossed races of man +cases of this kind rarely or never occur, this may be used as an +argument against the view suggested by some ethnologists, namely that +certain characters, for instance the blackness of the negro, first +appeared as a sudden variation or sport. Had this occurred, it is +probable that mulattoes would often have been born, either completely +black or completely white. + +We have now seen that a naturalist might feel himself fully justified in +ranking the races of man as distinct species; for he has found that they +are distinguished by many differences in structure and constitution, +some being of importance. These differences have, also, remained nearly +constant for very long periods of time. He will have been in some degree +influenced by the enormous range of man, which is a great anomaly in the +class of mammals, if mankind be viewed as a single species. He will have +been struck with the distribution of the several so-called races, in +accordance with that of other undoubtedly distinct species of mammals. +Finally he might urge that the mutual fertility of all the races has not +as yet been fully proved; and even if proved would not be an absolute +proof of their specific identity. + + +On the other side of the question, if our supposed naturalist were to +enquire whether the forms of man kept distinct like ordinary species, +when mingled together in large numbers in the same country, he would +immediately discover that this was by no means the case. In Brazil he +would behold an immense mongrel population of Negroes and Portuguese; in +Chiloe and other parts of South America, he would behold the whole +population consisting of Indians and Spaniards blended in various +degrees.[296] In many parts of the same continent he would meet with the +most complex crosses between Negroes, Indians, and Europeans; and such +triple crosses afford the severest test, judging from the vegetable +kingdom, of the mutual fertility of the parent-forms. In one island of +the Pacific he would find a small population of mingled Polynesian and +English blood; and in the Viti Archipelago a population of Polynesians +and Negritos crossed in all degrees. Many analogous cases could be +added, for instance, in South Africa. Hence the races of man are not +sufficiently distinct to co-exist without fusion; and this it is, which +in all ordinary cases affords the usual test of specific distinctness. + +Our naturalist would likewise be much disturbed as soon as he perceived +that the distinctive characters of every race of man were highly +variable. This strikes every one when he first beholds the negro-slaves +in Brazil, who have been imported from all parts of Africa. The same +remark holds good with the Polynesians, and with many other races. It +may be doubted whether any character can be named which is distinctive +of a race and is constant. Savages, even within the limits of the same +tribe, are not nearly so uniform in character, as has often been said. +Hottentot women offer certain peculiarities, more strongly marked than +those occurring in any other race, but these are known not to be of +constant occurrence. In the several American tribes, colour and +hairyness differ considerably; as does colour to a certain degree, and +the shape of the features greatly, in the Negroes of Africa. The shape +of the skull varies much in some races;[297] and so it is with every +other character. Now all naturalists have learnt by dearly-bought +experience, how rash it is to attempt to define species by the aid of +inconstant characters. + +But the most weighty of all the arguments against treating the races of +man as distinct species, is that they graduate into each other, +independently in many cases, as far as we can judge, of their having +intercrossed. Man has been studied more carefully than any other organic +being, and yet there is the greatest possible diversity amongst capable +judges whether he should be classed as a single species or race, or as +two (Virey), as three (Jacquinot), as four (Kant), five (Blumenbach), +six (Buffon), seven (Hunter), eight (Agassiz), eleven (Pickering), +fifteen (Bory St. Vincent), sixteen (Desmoulins), twenty-two (Morton), +sixty (Crawfurd), or as sixty-three, according to Burke.[298] This +diversity of judgment does not prove that the races ought not to be +ranked as species, but it shews that they graduate into each other, and +that it is hardly possible to discover clear distinctive characters +between them. + +Every naturalist who has had the misfortune to undertake the +description of a group of highly varying organisms, has encountered +cases (I speak after experience) precisely like that of man; and if of a +cautious disposition, he will end by uniting all the forms which +graduate into each other as a single species; for he will say to himself +that he has no right to give names to objects which he cannot define. +Cases of this kind occur in the Order which includes man, namely in +certain genera of monkeys; whilst in other genera, as in Cercopithecus, +most of the species can be determined with certainty. In the American +genus Cebus, the various forms are ranked by some naturalists as +species, by others as mere geographical races. Now if numerous specimens +of Cebus were collected from all parts of South America, and those forms +which at present appear to be specifically distinct, were found to +graduate into each other by close steps, they would be ranked by most +naturalists as mere varieties or races; and thus the greater number of +naturalists have acted with respect to the races of man. Nevertheless it +must be confessed that there are forms, at least in the vegetable +kingdom,[299] which we cannot avoid naming as species, but which are +connected together, independently of intercrossing, by numberless +gradations. + +Some naturalists have lately employed the term "sub-species" to +designate forms which possess many of the characteristics of true +species, but which hardly deserve so high a rank. Now if we reflect on +the weighty arguments, above given, for raising the races of man to the +dignity of species, and the insuperable difficulties on the other side +in defining them, the term "sub-species" might here be used with much +propriety. But from long habit the term "race" will perhaps always be +employed. The choice of terms is only so far important as it is highly +desirable to use, as far as that may be possible, the same terms for the +same degrees of difference. Unfortunately this is rarely possible; for +within the same family the larger genera generally include +closely-allied forms, which can be distinguished only with much +difficulty, whilst the smaller genera include forms that are perfectly +distinct; yet all must equally be ranked as species. So again the +species within the same large genus by no means resemble each other to +the same degree: on the contrary, in most cases some of them can be +arranged in little groups round other species, like satellites round +planets.[300] + + +The question whether mankind consists of one or several species has of +late years been much agitated by anthropologists, who are divided into +two schools of monogenists and polygenists. Those who do not admit the +principle of evolution, must look at species either as separate +creations or as in some manner distinct entities; and they must decide +what forms to rank as species by the analogy of other organic beings +which are commonly thus received. But it is a hopeless endeavour to +decide this point on sound grounds, until some definition of the term +"species" is generally accepted; and the definition must not include an +element which cannot possibly be ascertained, such as an act of +creation. We might as well attempt without any definition to decide +whether a certain number of houses should be called a village, or town, +or city. We have a practical illustration of the difficulty in the +never-ending doubts whether many closely-allied mammals, birds, +insects, and plants, which represent each other in North America and +Europe, should be ranked species or geographical races; and so it is +with the productions of many islands situated at some little distance +from the nearest continent. + +Those naturalists, on the other hand, who admit the principle of +evolution, and this is now admitted by the greater number of rising men, +will feel no doubt that all the races of man are descended from a single +primitive stock; whether or not they think fit to designate them as +distinct species, for the sake of expressing their amount of +difference.[301] With our domestic animals the question whether the +various races have arisen from one or more species is different. +Although all such races, as well as all the natural species within the +same genus, have undoubtedly sprung from the same primitive stock, yet +it is a fit subject for discussion, whether, for instance, all the +domestic races of the dog have acquired their present differences since +some one species was first domesticated and bred by man; or whether they +owe some of their characters to inheritance from distinct species, which +had already been modified in a state of nature. With mankind no such +question can arise, for he cannot be said to have been domesticated at +any particular period. + +When the races of man diverged at an extremely remote epoch from their +common progenitor, they will have differed but little from each other, +and been few in number; consequently they will then, as far as their +distinguishing characters are concerned, have had less claim to rank as +distinct species, than the existing so-called races. Nevertheless such +early races would perhaps have been ranked by some naturalists as +distinct species, so arbitrary is the term, if their differences, +although extremely slight, had been more constant than at present, and +had not graduated into each other. + +It is, however, possible, though far from probable, that the early +progenitors of man might at first have diverged much in character, until +they became more unlike each other than are any existing races; but that +subsequently, as suggested by Vogt,[302] they converged in character. +When man selects for the same object the offspring of two distinct +species, he sometimes induces, as far as general appearance is +concerned, a considerable amount of convergence. This is the case, as +shewn by Von Nathusius,[303] with the improved breeds of pigs, which are +descended from two distinct species; and in a less well-marked manner +with the improved breeds of cattle. A great anatomist, Gratiolet, +maintains that the anthropomorphous apes do not form a natural +sub-group; but that the orang is a highly developed gibbon or +semnopithecus; the chimpanzee a highly developed macacus; and the +gorilla a highly developed mandrill. If this conclusion, which rests +almost exclusively on brain-characters, be admitted, we should have a +case of convergence at least in external characters, for the +anthropomorphous apes are certainly more like each other in many points +than they are to other apes. All analogical resemblances, as of a whale +to a fish, may indeed be said to be cases of convergence; but this term +has never been applied to superficial and adaptive resemblances. It +would be extremely rash in most cases to attribute to convergence close +similarity in many points of structure in beings which had once been +widely different. The form of a crystal is determined solely by the +molecular forces, and it is not surprising that dissimilar substances +should sometimes assume the same form; but with organic beings we should +bear in mind that the form of each depends on an infinitude of complex +relations, namely on the variations which have arisen, these being due +to causes far too intricate to be followed out,--on the nature of the +variations which have been preserved, and this depends on the +surrounding physical conditions, and in a still higher degree on the +surrounding organisms with which each has come into competition,--and +lastly, on inheritance (in itself a fluctuating element) from +innumerable progenitors, all of which have had their forms determined +through equally complex relations. It appears utterly incredible that +two organisms, if differing in a marked manner, should ever afterwards +converge so closely as to lead to a near approach to identity throughout +their whole organisation. In the case of the convergent pigs above +referred to, evidence of their descent from two primitive stocks is +still plainly retained, according to Von Nathusius, in certain bones of +their skulls. If the races of man were descended, as supposed by some +naturalists, from two or more distinct species, which had differed as +much, or nearly as much, from each other, as the orang differs from the +gorilla, it can hardly be doubted that marked differences in the +structure of certain bones would still have been discoverable in man as +he now exists. + + +Although the existing races of man differ in many respects, as in +colour, hair, shape of skull, proportions of the body, &c., yet if their +whole organisation be taken into consideration they are found to +resemble each other closely in a multitude of points. Many of these +points are of so unimportant or of so singular a nature, that it is +extremely improbable that they should have been independently acquired +by aboriginally distinct species or races. The same remark holds good +with equal or greater force with respect to the numerous points of +mental similarity between the most distinct races of man. The American +aborigines, Negroes and Europeans differ as much from each other in mind +as any three races that can be named; yet I was incessantly struck, +whilst living with the Fuegians on board the "Beagle," with the many +little traits of character, shewing how similar their minds were to +ours; and so it was with a full-blooded negro with whom I happened once +to be intimate. + +He who will carefully read Mr. Tylor's and Sir J. Lubbock's interesting +works[304] can hardly fail to be deeply impressed with the close +similarity between the men of all races in tastes, dispositions and +habits. This is shewn by the pleasure which they all take in dancing, +rude music, acting, painting, tattooing, and otherwise decorating +themselves,--in their mutual comprehension of gesture-language--and, as +I shall be able to shew in a future essay, by the same expression in +their features, and by the same inarticulate cries, when they are +excited by various emotions. This similarity, or rather identity, is +striking, when contrasted with the different expressions which may be +observed in distinct species of monkeys. There is good evidence that the +art of shooting with bows and arrows has not been handed down from any +common progenitor of mankind, yet the stone arrow-heads, brought from +the most distant parts of the world and manufactured at the most remote +periods, are, as Nilsson has shewn,[305] almost identical; and this fact +can only be accounted for by the various races having similar inventive +or mental powers. The same observation has been made by +archologists[306] with respect to certain widely-prevalent ornaments, +such as zigzags, &c.; and with respect to various simple beliefs and +customs, such as the burying of the dead under megalithic structures. I +remember observing in South America,[307] that there, as in so many +other parts of the world, man has generally chosen the summits of lofty +hills, on which to throw up piles of stones, either for the sake of +recording some remarkable event, or for burying his dead. + +Now when naturalists observe a close agreement in numerous small details +of habits, tastes and dispositions between two or more domestic races, +or between nearly-allied natural forms, they use this fact as an +argument that all are descended from a common progenitor who was thus +endowed; and consequently that all should be classed under the same +species. The same argument may be applied with much force to the races +of man. + +As it is improbable that the numerous and unimportant points of +resemblance between the several races of man in bodily structure and +mental faculties (I do not here refer to similar customs) should all +have been independently acquired, they must have been inherited from +progenitors who were thus characterised. We thus gain some insight into +the early state of man, before he had spread step by step over the face +of the earth. The spreading of man to regions widely separated by the +sea, no doubt, preceded any considerable amount of divergence of +character in the several races; for otherwise we should sometimes meet +with the same race in distinct continents; and this is never the case. +Sir J. Lubbock, after comparing the arts now practised by savages in all +parts of the world, specifies those which man could not have known, when +he first wandered from his original birthplace; for if once learnt they +would never have been forgotten.[308] He thus shews that "the spear, +which is but a development of the knife-point, and the club, which is +but a long hammer, are the only things left." He admits, however, that +the art of making fire probably had already been discovered, for it is +common to all the races now existing, and was known to the ancient +cave-inhabitants of Europe. Perhaps the art of making rude canoes or +rafts was likewise known; but as man existed at a remote epoch, when the +land in many places stood at a very different level, he would have been +able, without the aid of canoes, to have spread widely. Sir J. Lubbock +further remarks how improbable it is that our earliest ancestors could +have "counted as high as ten, considering that so many races now in +existence cannot get beyond four." Nevertheless, at this early period, +the intellectual and social faculties of man could hardly have been +inferior in any extreme degree to those now possessed by the lowest +savages; otherwise primeval man could not have been so eminently +successful in the struggle for life, as proved by his early and wide +diffusion. + +From the fundamental differences between certain languages, some +philologists have inferred that when man first became widely diffused he +was not a speaking animal; but it may be suspected that languages, far +less perfect than any now spoken, aided by gestures, might have been +used, and yet have left no traces on subsequent and more +highly-developed tongues. Without the use of some language, however +imperfect, it appears doubtful whether man's intellect could have risen +to the standard implied by his dominant position at an early period. + +Whether primeval man, when he possessed very few arts of the rudest +kind, and when his power of language was extremely imperfect, would have +deserved to be called man, must depend on the definition which we +employ. In a series of forms graduating insensibly from some ape-like +creature to man as he now exists, it would be impossible to fix on any +definite point when the term "man" ought to be used. But this is a +matter of very little importance. So again it is almost a matter of +indifference whether the so-called races of man are thus designated, or +are ranked as species or sub-species; but the latter term appears the +most appropriate. Finally, we may conclude that when the principles of +evolution are generally accepted, as they surely will be before long, +the dispute between the monogenists and the polygenists will die a +silent and unobserved death. + + +One other question ought not to be passed over without notice, namely, +whether, as is sometimes assumed, each sub-species or race of man has +sprung from a single pair of progenitors. With our domestic animals a +new race can readily be formed from a single pair possessing some new +character, or even from a single individual thus characterised, by +carefully matching the varying offspring; but most of our races have +been formed, not intentionally from a selected pair, but unconsciously +by the preservation of many individuals which have varied, however +slightly, in some useful or desired manner. If in one country stronger +and heavier horses, and in another country lighter and fleeter horses, +were habitually preferred, we may feel sure that two distinct sub-breeds +would, in the course of time, be produced, without any particular pairs +or individuals having been separated and bred from in either country. +Many races have been thus formed, and their manner of formation is +closely analogous with that of natural species. We know, also, that the +horses which have been brought to the Falkland Islands have become, +during successive generations, smaller and weaker, whilst those which +have run wild on the Pampas have acquired larger and coarser heads; and +such changes are manifestly due, not to any one pair, but to all the +individuals having been subjected to the same conditions, aided, +perhaps, by the principle of reversion. The new sub-breeds in none of +these cases are descended from any single pair, but from many +individuals which have varied in different degrees, but in the same +general manner; and we may conclude that the races of man have been +similarly produced, the modifications being either the direct result of +exposure to different conditions, or the indirect result of some form of +selection. But to this latter subject we shall presently return. + + +_On the Extinction of the Races of Man._--The partial and complete +extinction of many races and sub-races of man are historically known +events. Humboldt saw in South America a parrot which was the sole living +creature that could speak the language of a lost tribe. Ancient +monuments and stone implements found in all parts of the world, of which +no tradition is preserved by the present inhabitants, indicate much +extinction. Some small and broken tribes, remnants of former races, +still survive in isolated and generally mountainous districts. In Europe +the ancient races were all, according to Schaaffhausen,[309] "lower in +the scale than the rudest living savages;" they must therefore have +differed, to a certain extent, from any existing race. The remains +described by Professor Broca[310] from Les Eyzies, though they +unfortunately appear to have belonged to a single family, indicate a +race with a most singular combination of low or simious and high +characteristics, and is "entirely different from any other race, ancient +or modern, that we have ever heard of." It differed, therefore, from the +quaternary race of the caverns of Belgium. + +Unfavourable physical conditions appear to have had but little effect in +the extinction of races.[311] Man has long lived in the extreme regions +of the North, with no wood wherewith to make his canoes or other +implements, and with blubber alone for burning and giving him warmth, +but more especially for melting the snow. In the Southern extremity of +America the Fuegians survive without the protection of clothes, or of +any building worthy to be called a hovel. In South Africa the aborigines +wander over the most arid plains, where dangerous beasts abound. Man can +withstand the deadly influence of the Terai at the foot of the Himalaya, +and the pestilential shores of tropical Africa. + +Extinction follows chiefly from the competition of tribe with tribe, +and race with race. Various checks are always in action, as specified in +a former chapter, which serve to keep down the numbers of each savage +tribe,--such as periodical famines, the wandering of the parents and the +consequent deaths of infants, prolonged suckling, the stealing of women, +wars, accidents, sickness, licentiousness, especially infanticide, and, +perhaps, lessened fertility from less nutritious food, and many +hardships. If from any cause any one of these checks is lessened, even +in a slight degree, the tribe thus favoured will tend to increase; and +when one of two adjoining tribes becomes more numerous and powerful than +the other, the contest is soon settled by war, slaughter, cannibalism, +slavery, and absorption. Even when a weaker tribe is not thus abruptly +swept away, if it once begins to decrease, it generally goes on +decreasing until it is extinct.[312] + +When civilised nations come into contact with barbarians the struggle is +short, except where a deadly climate gives its aid to the native race. +Of the causes which lead to the victory of civilised nations, some are +plain and some very obscure. We can see that the cultivation of the land +will be fatal in many ways to savages, for they cannot, or will not, +change their habits. New diseases and vices are highly destructive; and +it appears that in every nation a new disease causes much death, until +those who are most susceptible to its destructive influence are +gradually weeded out;[313] and so it may be with the evil effects from +spirituous liquors, as well as with the unconquerably strong taste for +them shewn by so many savages. It further appears, mysterious as is the +fact, that the first meeting of distinct and separated people generates +disease.[314] Mr. Sproat, who in Vancouver Island closely attended to +the subject of extinction, believes that changed habits of life, which +always follow from the advent of Europeans, induces much ill-health. He +lays, also, great stress on so trifling a cause as that the natives +become "bewildered and dull by the new life around them; they lose the +motives for exertion, and get no new ones in their place."[315] + +The grade of civilisation seems a most important element in the success +of nations which come in competition. A few centuries ago Europe feared +the inroads of Eastern barbarians; now, any such fear would be +ridiculous. It is a more curious fact, that savages did not formerly +waste away, as Mr. Bagehot has remarked, before the classical nations, +as they now do before modern civilised nations; had they done so, the +old moralists would have mused over the event; but there is no lament in +any writer of that period over the perishing barbarians.[316] + +Although the gradual decrease and final extinction of the races of man +is an obscure problem, we can see that it depends on many causes, +differing in different places and at different times. It is the same +difficult problem as that presented by the extinction of one of the +higher animals--of the fossil horse, for instance, which disappeared +from South America, soon afterwards to be replaced, within the same +districts, by countless troops of the Spanish horse. The New Zealander +seems conscious of this parallelism, for he compares his future fate +with that of the native rat almost exterminated by the European rat. The +difficulty, though great to our imagination, and really great if we wish +to ascertain the precise causes, ought not to be so to our reason, as +long as we keep steadily in mind that the increase of each species and +each race is constantly hindered by various checks; so that if any new +check, or cause of destruction, even a slight one, be superadded, the +race will surely decrease in number; and as it has everywhere been +observed that savages are much opposed to any change of habits, by which +means injurious checks could be counterbalanced, decreasing numbers will +sooner or later lead to extinction; the end, in most cases, being +promptly determined by the inroads of increasing and conquering tribes. + + +_On the Formation of the Races of Man._--It may be premised that when we +find the same race, though broken up into distinct tribes, ranging over +a great area, as over America, we may attribute their general +resemblance to descent from a common stock. In some cases the crossing +of races already, distinct has led to the formation of new races. The +singular fact that Europeans and Hindoos, who belong to the same Aryan +stock and speak a language fundamentally the same, differ widely in +appearance, whilst Europeans differ but little from Jews, who belong to +the Semitic stock and speak quite another language, has been accounted +for by Broca[317] through the Aryan branches having been largely crossed +during their wide diffusion by various indigenous tribes. When two races +in close contact cross, the first result is a heterogeneous mixture: +thus Mr. Hunter, in describing the Santali or hill-tribes of India, says +that hundreds of imperceptible gradations may be traced "from the black, +squat tribes of the mountains to the tall olive-coloured Brahman, with +his intellectual brow, calm eyes, and high but narrow head;" so that it +is necessary in courts of justice to ask the witnesses whether they are +Santalis or Hindoos.[318] Whether a heterogeneous people, such as the +inhabitants of some of the Polynesian islands, formed by the crossing of +two distinct races, with few or no pure members left, would ever become +homogeneous, is not known from direct evidence. But as with our +domesticated animals, a crossed breed can certainly, in the course of a +few generations, be fixed and made uniform by careful selection,[319] we +may infer that the free and prolonged intercrossing during many +generations of a heterogeneous mixture would supply the place of +selection, and overcome any tendency to reversion, so that a crossed +race would ultimately become homogeneous, though it might not partake in +an equal degree of the characters of the two parent-races. + +Of all the differences between the races of man, the colour of the skin +is the most conspicuous and one of the best marked. Differences of this +kind, it was formerly thought, could be accounted for by long exposure +under different climates; but Pallas first shewed that this view is not +tenable, and he has been followed by almost all anthropologists.[320] +The view has been rejected chiefly because the distribution of the +variously coloured races, most of whom must have long inhabited their +present homes, does not coincide with corresponding differences of +climate. Weight must also be given to such cases as that of the Dutch +families, who, as we hear on excellent authority,[321] have not +undergone the least change of colour, after residing for three centuries +in South Africa. The uniform appearance in various parts of the world of +gypsies and Jews, though the uniformity of the latter has been somewhat +exaggerated,[322] is likewise an argument on the same side. A very damp +or a very dry atmosphere has been supposed to be more influential in +modifying the colour of the skin than mere heat; but as D'Orbigny in +South America, and Livingstone in Africa, arrived at diametrically +opposite conclusions with respect to dampness and dryness, any +conclusion on this head must be considered as very doubtful.[323] + +Various facts, which I have elsewhere given, prove that the colour of +the skin and hair is sometimes correlated in a surprising manner with a +complete immunity from the action of certain vegetable poisons and from +the attacks of certain parasites. Hence it occurred to me, that negroes +and other dark races might have acquired their dark tints by the darker +individuals escaping during a long series of generations from the deadly +influence of the miasmas of their native countries. + +I afterwards found that the same idea had long ago occurred to Dr. +Wells.[324] That negroes, and even mulattoes, are almost completely +exempt from the yellow-fever, which is so destructive in tropical +America, has long been known.[325] They likewise escape to a large +extent the fatal intermittent fevers that prevail along, at least, 2600 +miles of the shores of Africa, and which annually cause one-fifth of the +white settlers to die, and another fifth to return home invalided.[326] +This immunity in the negro seems to be partly inherent, depending on +some unknown peculiarity of constitution, and partly the result of +acclimatisation. Pouchet[327] states that the negro regiments, borrowed +from the Viceroy of Egypt for the Mexican war, which had been recruited +near the Soudan, escaped the yellow-fever almost equally well with the +negroes originally brought from various parts of Africa, and accustomed +to the climate of the West Indies. That acclimatisation plays a part is +shewn by the many cases in which negroes, after having resided for some +time in a colder climate, have become to a certain extent liable to +tropical fevers.[328] The nature of the climate under which the white +races have long resided, likewise has some influence on them; for during +the fearful epidemic of yellow-fever in Demerara during 1837, Dr. Blair +found that the death-rate of the immigrants was proportional to the +latitude of the country whence they had come. With the negro the +immunity, as far as it is the result of acclimatisation, implies +exposure during a prodigious length of time; for the aborigines of +tropical America, who have resided there from time immemorial, are not +exempt from yellow-fever; and the Rev. B. Tristram states, that there +are districts in Northern Africa which the native inhabitants are +compelled annually to leave, though the negroes can remain with safety. + +That the immunity of the negro is in any degree correlated with the +colour of his skin is a mere conjecture: it may be correlated with some +difference in his blood, nervous system, or other tissues. Nevertheless, +from the facts above alluded to, and from some connection apparently +existing between complexion and a tendency to consumption, the +conjecture seemed to me not improbable. Consequently I endeavoured, with +but little success,[329] to ascertain how far it held good. The late +Dr. Daniell, who had long lived on the West Coast of Africa, told me +that he did not believe in any such relation. He was himself unusually +fair, and had withstood the climate in a wonderful manner. When he first +arrived as a boy on the coast, an old and experienced negro chief +predicted from his appearance that this would prove the case. Dr. +Nicholson, of Antigua, after having attended to this subject, wrote to +me that he did not think that dark-coloured Europeans escaped the +yellow-fever better than those that were light-coloured. Mr. J. M. +Harris altogether denies[330] that Europeans with dark hair withstand a +hot climate better than other men; on the contrary, experience has +taught him in making a selection of men for service on the coast of +Africa, to choose those with red hair. As far, therefore, as these +slight indications serve, there seems no foundation for the hypothesis, +which has been accepted by several writers, that the colour of the black +races may have resulted from darker and darker individuals having +survived in greater numbers, during their exposure to the +fever-generating miasmas of their native countries. + +Although with our present knowledge we cannot account for the +strongly-marked differences in colour between the races of man, either +through correlation with constitutional peculiarities, or through the +direct action of climate; yet we must not quite ignore the latter +agency, for there is good reason to believe that some inherited effect +is thus produced.[331] + +We have seen in our third chapter that the conditions of life, such as +abundant food and general comfort, affect in a direct manner the +development of the bodily frame, the effects being transmitted. Through +the combined influences of climate and changed habits of life, European +settlers, in the United States undergo, as is generally admitted, a +slight but extraordinarily rapid change of appearance. There is, also, a +considerable body of evidence shewing that in the Southern States the +house-slaves of the third generation present a markedly different +appearance from the field-slaves.[332] + +If, however, we look to the races of man, as distributed over the world, +we must infer that their characteristic differences cannot be accounted +for by the direct action of different conditions of life, even after +exposure to them for an enormous period of time. The Esquimaux live +exclusively on animal food; they are clothed in thick fur, and are +exposed to intense cold and to prolonged darkness; yet they do not +differ in any extreme degree from the inhabitants of Southern China, who +live entirely on vegetable food and are exposed almost naked to a hot, +glaring climate. The unclothed Fuegians live on the marine productions +of their inhospitable shores; the Botocudos of Brazil wander about the +hot forests of the interior and live chiefly on vegetable productions; +yet these tribes resemble each other so closely that the Fuegians on +board the "Beagle" were mistaken by some Brazilians for Botocudos. The +Botocudos again, as well as the other inhabitants of tropical America, +are wholly different from the Negroes who inhabit the opposite shores of +the Atlantic, are exposed to a nearly similar climate, and follow nearly +the same habits of life. + +Nor can the differences between the races of man be accounted for, +except to a quite insignificant degree, by the inherited effects of the +increased or decreased use of parts. Men who habitually live in canoes, +may have their legs somewhat stunted; those who inhabit lofty regions +have their chests enlarged; and those who constantly use certain +sense-organs have the cavities in which they are lodged somewhat +increased in size, and their features consequently a little modified. +With civilised nations, the reduced size of the jaws from lessened use, +the habitual play of different muscles serving to express different +emotions, and the increased size of the brain from greater intellectual +activity, have together produced a considerable effect on their general +appearance in comparison with savages.[333] It is also possible that +increased bodily stature, with no corresponding increase in the size of +the brain, may have given to some races (judging from the previously +adduced cases of the rabbits) an elongated skull of the dolichocephalic +type. + +Lastly, the little-understood principle of correlation will almost +certainly have come into action, as in the case of great muscular +development and strongly projecting supra-orbital ridges. It is not +improbable that the texture of the hair, which differs much in the +different races, may stand in some kind of correlation with the +structure of the skin; for the colour of the hair and skin are certainly +correlated, as is its colour and texture with the Mandans.[334] The +colour of the skin and the odour emitted by it are likewise in some +manner connected. With the breeds of sheep the number of hairs within a +given space and the number of the excretory pores stand in some relation +to each other.[335] If we may judge from the analogy of our domesticated +animals, many modifications of structure in man probably come under this +principle of correlated growth. + + +We have now seen that the characteristic differences between the races +of man cannot be accounted for in a satisfactory manner by the direct +action of the conditions of life, nor by the effects of the continued +use of parts, nor through the principle of correlation. We are therefore +led to inquire whether slight individual differences, to which man is +eminently liable, may not have been preserved and augmented during a +long series of generations through natural selection. But here we are at +once met by the objection that beneficial variations alone can be thus +preserved; and as far as we are enabled to judge (although always liable +to error on this head) not one of the external differences between the +races of man are of any direct or special service to him. The +intellectual and moral or social faculties must of course be excepted +from this remark; but differences in these faculties can have had little +or no influence on external characters. The variability of all the +characteristic differences between the races, before referred to, +likewise indicates that these differences cannot be of much importance; +for, had they been important, they would long ago have been either fixed +and preserved, or eliminated. In this respect man resembles those forms, +called by naturalists protean or polymorphic, which have remained +extremely variable, owing, as it seems, to their variations being of an +indifferent nature, and consequently to their having escaped the action +of natural selection. + +We have thus far been baffled in all our attempts to account for the +differences between the races of man; but there remains one important +agency, namely Sexual Selection, which appears to have acted as +powerfully on man, as on many other animals. I do not intend to assert +that sexual selection will account for all the differences between the +races. An unexplained residuum is left, about which we can in our +ignorance only say, that as individuals are continually born with, for +instance, heads a little rounder or narrower, and with noses a little +longer or shorter, such slight differences might become fixed and +uniform, if the unknown agencies which induced them were to act in a +more constant manner, aided by long-continued intercrossing. Such +modifications come under the provisional class, alluded to in our fourth +chapter, which for the want of a better term have been called +spontaneous variations. Nor do I pretend that the effects of sexual +selection can be indicated with scientific precision; but it can be +shewn that it would be an inexplicable fact if man had not been modified +by this agency, which has acted so powerfully on innumerable animals, +both high and low in the scale. It can further be shewn that the +differences between the races of man, as in colour, hairyness, form of +features, &c., are of the nature which it might have been expected would +have been acted on by sexual selection. But in order to treat this +subject in a fitting manner, I have found it necessary to pass the whole +animal kingdom in review; I have therefore devoted to it the Second Part +of this work. At the close I shall return to man, and, after attempting +to shew how far he has been modified through sexual selection, will give +a brief summary of the chapters in this First Part. + + + + +PART II.--SEXUAL SELECTION. + + +CHAPTER VIII. + +PRINCIPLES OF SEXUAL SELECTION. + + + Secondary sexual characters--Sexual selection--Manner of + action--Excess of males--Polygamy--The male alone generally + modified through sexual selection--Eagerness of the + male--Variability of the male--Choice exerted by the + female--Sexual compared with natural selection--Inheritance, at + corresponding periods of life, at corresponding seasons of the + year, and as limited by sex--Relations between the several + forms of inheritance--Causes why one sex and the young are not + modified through sexual selection--Supplement on the + proportional numbers of the two sexes throughout the animal + kingdom--On the limitation of the numbers of the two sexes + through natural selection. + + +With animals which have their sexes separated, the males necessarily +differ from the females in their organs of reproduction; and these +afford the primary sexual characters. But the sexes often differ in what +Hunter has called secondary sexual characters, which are not directly +connected with the act of reproduction; for instance, in the male +possessing certain organs of sense or locomotion, of which the female is +quite destitute, or in having them more highly-developed, in order that +he may readily find or reach her; or again, in the male having special +organs of prehension so as to hold her securely. These latter organs of +infinitely diversified kinds graduate into, and in some cases can hardly +be distinguished from, those which are commonly ranked as primary, such +as the complex appendages at the apex of the abdomen in male insects. +Unless indeed we confine the term "primary" to the reproductive glands, +it is scarcely possible to decide, as far as the organs of prehension +are concerned, which ought to be called primary and which secondary. + +The female often differs from the male in having organs for the +nourishment or protection of her young, as the mammary glands of +mammals, and the abdominal sacks of the marsupials. The male, also, in +some few cases differs from the female in possessing analogous organs, +as the receptacles for the ova possessed by the males of certain fishes, +and those temporarily developed in certain male frogs. Female bees have +a special apparatus for collecting and carrying pollen, and their +ovipositor is modified into a sting for the defence of their larv and +the community. In the females of many insects the ovipositor is modified +in the most complex manner for the safe placing of the eggs. Numerous +similar cases could be given, but they do not here concern us. There +are, however, other sexual differences quite disconnected with the +primary organs with which we are more especially concerned--such as the +greater size, strength, and pugnacity of the male, his weapons of +offence or means of defence against rivals, his gaudy colouring and +various ornaments, his power of song, and other such characters. + +Besides the foregoing primary and secondary sexual differences, the male +and female sometimes differ in structures connected with different +habits of life, and not at all, or only indirectly, related to the +reproductive functions. Thus the females of certain flies (Culicid and +Tabanid) are blood-suckers, whilst the males live on flowers and have +their mouths destitute of mandibles.[336] The males alone of certain +moths and of some crustaceans (_e.g._ Tanais) have imperfect, closed +mouths, and cannot feed. The Complemental males of certain cirripedes +live like epiphytic plants either on the female or hermaphrodite form, +and are destitute of a mouth and prehensile limbs. In these cases it is +the male which has been modified and has lost certain important organs, +which the other members of the same group possess. In other cases it is +the female which has lost such parts; for instance, the female glow-worm +is destitute of wings, as are many female moths, some of which never +leave their cocoons. Many female parasitic crustaceans have lost their +natatory legs. In some weevil-beetles (Curculionid) there is a great +difference between the male and female in the length of the rostrum or +snout;[337] but the meaning of this and of many analogous differences, +is not at all understood. Differences of structure between the two sexes +in relation to different habits of life are generally confined to the +lower animals; but with some few birds the beak of the male differs from +that of the female. No doubt in most, but apparently not in all these +cases, the differences are indirectly connected with the propagation of +the species: thus a female which has to nourish a multitude of ova will +require more food than the male, and consequently will require special +means for procuring it. A male animal which lived for a very short time +might without detriment lose through disuse its organs for procuring +food; but he would retain his locomotive organs in a perfect state, so +that he might reach the female. The female, on the other hand, might +safely lose her organs for flying, swimming, or walking, if she +gradually acquired habits which rendered such powers useless. + +We are, however, here concerned only with that kind of selection, which +I have called sexual selection. This depends on the advantage which +certain individuals have over other individuals of the same sex and +species, in exclusive relation to reproduction. When the two sexes +differ in structure in relation to different habits of life, as in the +cases above mentioned, they have no doubt been modified through natural +selection, accompanied by inheritance limited to one and the same sex. +So again the primary sexual organs, and those for nourishing or +protecting the young, come under this same head; for those individuals +which generated or nourished their offspring best, would leave, _cteris +paribus_, the greatest number to inherit their superiority; whilst those +which generated or nourished their offspring badly, would leave but few +to inherit their weaker powers. As the male has to search for the +female, he requires for this purpose organs of sense and locomotion, but +if these organs are necessary for the other purposes of life, as is +generally the case, they will have been developed through natural +selection. When the male has found the female he sometimes absolutely +requires prehensile organs to hold her; thus Dr. Wallace informs me that +the males of certain moths cannot unite with the females if their tarsi +or feet are broken. The males of many oceanic crustaceans have their +legs and antenn modified in an extraordinary manner for the prehension +of the female; hence we may suspect that owing to these animals being +washed about by the waves of the open sea, they absolutely require these +organs in order to propagate their kind, and if so their development +will have been the result of ordinary or natural selection. + +When the two sexes follow exactly the same habits of life, and the male +has more highly developed sense or locomotive organs than the female, it +may be that these in their perfected state are indispensable to the male +for finding the female; but in the vast majority of cases, they serve +only to give one male an advantage over another, for the less +well-endowed males, if time were allowed them, would succeed in pairing +with the females; and they would in all other respects, judging from the +structure of the female, be equally well adapted for their ordinary +habits of life. In such cases sexual selection must have come into +action, for the males have acquired their present structure, not from +being better fitted to survive in the struggle for existence, but from +having gained an advantage over other males, and from having transmitted +this advantage to their male offspring alone. It was the importance of +this distinction which led me to designate this form of selection as +sexual selection. So again, if the chief service rendered to the male by +his prehensile organs is to prevent the escape of the female before the +arrival of other males, or when assaulted by them, these organs will +have been perfected through sexual selection, that is by the advantage +acquired by certain males over their rivals. But in most cases it is +scarcely possible to distinguish between the effects of natural and +sexual selection. Whole chapters could easily be filled with details on +the differences between the sexes in their sensory, locomotive, and +prehensile organs. As, however, these structures are not more +interesting than others adapted for the ordinary purposes of life, I +shall almost pass them over, giving only a few instances under each +class. + +There are many other structures and instincts which must have been +developed through sexual selection--such as the weapons of offence and +the means of defence possessed by the males for fighting with and +driving away their rivals--their courage and pugnacity--their ornaments +of many kinds--their organs for producing vocal or instrumental +music--and their glands for emitting odours; most of these latter +structures serving only to allure or excite the female. That these +characters are the result of sexual and not of ordinary selection is +clear, as unarmed, unornamented, or unattractive males would succeed +equally well in the battle for life and in leaving a numerous progeny, +if better endowed males were not present. We may infer that this would +be the case, for the females, which are unarmed and unornamented, are +able to survive and procreate their kind. Secondary sexual characters of +the kind just referred to, will be fully discussed in the following +chapters, as they are in many respects interesting, but more especially +as they depend on the will, choice, and rivalry of the individuals of +either sex. When we behold two males fighting for the possession of the +female, or several male birds displaying their gorgeous plumage, and +performing the strangest antics before an assembled body of females, we +cannot doubt that, though led by instinct, they know what they are +about, and consciously exert their mental and bodily powers. + +In the same manner as man can improve the breed of his game-cocks by the +selection of those birds which are victorious in the cockpit, so it +appears that the strongest and most vigorous males, or those provided +with the best weapons, have prevailed under nature, and have led to the +improvement of the natural breed or species. Through repeated deadly +contests, a slight degree of variability, if it led to some advantage, +however slight, would suffice for the work of sexual selection; and it +is certain that secondary sexual characters are eminently variable. In +the same manner as man can give beauty, according to his standard of +taste, to his male poultry--can give to the Sebright bantam a new and +elegant plumage, an erect and peculiar carriage--so it appears that in a +state of nature female birds, by having long selected the more +attractive males, have added to their beauty. No doubt this implies +powers of discrimination and taste on the part of the female which will +at first appear extremely improbable; but I hope hereafter to shew that +this is not the case. + +From our ignorance on several points, the precise manner in which sexual +selection acts is to a certain extent uncertain. Nevertheless if those +naturalists who already believe in the mutability of species, will read +the following chapters, they will, I think, agree with me that sexual +selection has played an important part in the history of the organic +world. It is certain that with almost all animals there is a struggle +between the males for the possession of the female. This fact is so +notorious that it would be superfluous to give instances. Hence the +females, supposing that their mental capacity sufficed for the exertion +of a choice, could select one out of several males. But in numerous +cases it appears as if it had been specially arranged that there should +be a struggle between many males. Thus with migratory birds, the males +generally arrive before the females at their place of breeding, so that +many males are ready to contend for each female. The bird-catchers +assert that this is invariably the case with the nightingale and +blackcap, as I am informed by Mr. Jenner Weir, who confirms the +statement with respect to the latter species. + +Mr. Swaysland of Brighton, who has been in the habit, during the last +forty years, of catching our migratory birds on their first arrival, +writes to me that he has never known the females of any species to +arrive before their males. During one spring he shot thirty-nine males +of Ray's wagtail (_Budytes Raii_) before he saw a single female. Mr. +Gould has ascertained by dissection, as he informs me, that male snipes +arrive in this country before the females. In the case of fish, at the +period when the salmon ascend our rivers, the males in large numbers are +ready to breed before the females. So it apparently is with frogs and +toads. Throughout the great class of insects the males almost always +emerge from the pupal state before the other sex, so that they generally +swarm for a time before any females can be seen.[338] The cause of this +difference between the males and females in their periods of arrival and +maturity is sufficiently obvious. Those males which annually first +migrated into any country, or which in the spring were first ready to +breed, or were the most eager, would leave the largest number of +offspring; and these would tend to inherit similar instincts and +constitutions. On the whole there can be no doubt that with almost all +animals, in which the sexes are separate, there is a constantly +recurrent struggle between the males for the possession of the females. + +Our difficulty in regard to sexual selection lies in understanding how +it is that the males which conquer other males, or those which prove the +most attractive to the females, leave a greater number of offspring to +inherit their superiority than the beaten and less attractive males. +Unless this result followed, the characters which gave to certain males +an advantage over others, could not be perfected and augmented through +sexual selection. When the sexes exist in exactly equal numbers, the +worst-endowed males will ultimately find females (excepting where +polygamy prevails), and leave as many offspring, equally well fitted for +their general habits of life, as the best-endowed males. From various +facts and considerations, I formerly inferred that with most animals, in +which secondary sexual characters were well developed, the males +considerably exceeded the females in number; and this does hold good in +some few cases. If the males were to the females as two to one, or as +three to two, or even in a somewhat lower ratio, the whole affair would +be simple; for the better-armed or more attractive males would leave the +largest number of offspring. But after investigating, as far as +possible, the numerical proportions of the sexes, I do not believe that +any great inequality in number commonly exists. In most cases sexual +selection appears to have been effective in the following manner. + +Let us take any species, a bird for instance, and divide the females +inhabiting a district into two equal bodies: the one consisting of the +more vigorous and better-nourished individuals, and the other of the +less vigorous and healthy. The former, there can be little doubt, would +be ready to breed in the spring before the others; and this is the +opinion of Mr. Jenner Weir, who has during many years carefully attended +to the habits of birds. There can also be no doubt that the most +vigorous, healthy, and best-nourished females would on an average +succeed in rearing the largest number of offspring. The males, as we +have seen, are generally ready to breed before the females; of the males +the strongest, and with some species the best armed, drive away the +weaker males; and the former would then unite with the more vigorous and +best-nourished females, as these are the first to breed. Such vigorous +pairs would surely rear a larger number of offspring than the retarded +females, which would be compelled, supposing the sexes to be numerically +equal, to unite with the conquered and less powerful males; and this is +all that is wanted to add, in the course of successive generations, to +the size, strength and courage of the males, or to improve their +weapons. + +But in a multitude of cases the males which conquer other males, do not +obtain possession of the females, independently of choice on the part of +the latter. The courtship of animals is by no means so simple and short +an affair as might be thought. The females are most excited by, or +prefer pairing with, the more ornamented males, or those which are the +best songsters, or play the best antics; but it is obviously probable, +as has been actually observed in some cases, that they would at the same +time prefer the more vigorous and lively males.[339] Thus the more +vigorous females, which are the first to breed, will have the choice of +many males; and though they may not always select the strongest or best +armed, they will select those which are vigorous and well armed, and in +other respects the most attractive. Such early pairs would have the same +advantage in rearing offspring on the female side as above explained, +and nearly the same advantage on the male side. And this apparently has +sufficed during a long course of generations to add not only to the +strength and fighting-powers of the males, but likewise to their +various ornaments or other attractions. + +In the converse and much rarer case of the males selecting particular +females, it is plain that those which were the most vigorous and had +conquered others, would have the freest choice; and it is almost certain +that they would select vigorous as well as attractive females. Such +pairs would have an advantage in rearing offspring, more especially if +the male had the power to defend the female during the pairing-season, +as occurs with some of the higher animals, or aided in providing for the +young. The same principles would apply if both sexes mutually preferred +and selected certain individuals of the opposite sex; supposing that +they selected not only the more attractive, but likewise the more +vigorous individuals. + + +_Numerical Proportion of the Two Sexes._--I have remarked that sexual +selection would be a simple affair if the males considerably exceeded in +number the females. Hence I was led to investigate, as far as I could, +the proportions between the two sexes of as many animals as possible; +but the materials are scanty. I will here give only a brief abstract of +the results, retaining the details for a supplementary discussion, so as +not to interfere with the course of my argument. Domesticated animals +alone afford the opportunity of ascertaining the proportional numbers at +birth; but no records have been specially kept for this purpose. By +indirect means, however, I have collected a considerable body of +statistical data, from which it appears that with most of our domestic +animals the sexes are nearly equal at birth. Thus with race-horses, +25,560 births have been recorded during twenty-one years, and the male +births have been to the female births as 997 to 100. With greyhounds +the inequality is greater than with any other animal, for during twelve +years, out of 6878 births, the male births have been as 1101 to 100 +female births. It is, however, in some degree doubtful whether it is +safe to infer that the same proportional numbers would hold good under +natural conditions as under domestication; for slight and unknown +differences in the conditions affect to a certain extent the proportion +of the sexes. Thus with mankind, the male births in England are as +1045, in Russia as 1089, and with the Jews of Livornia as 120 _to_ 100 +females. The proportion is also mysteriously affected by the +circumstance of the births being legitimate or illegitimate. + +For our present purpose we are concerned with the proportion of the +sexes, not at birth, but at maturity, and this adds another element of +doubt; for it is a well ascertained fact that with man a considerably +larger proportion of males than of females die before or during birth, +and during the first few years of infancy. So it almost certainly is +with male lambs, and so it may be with the males of other animals. The +males of some animals kill each other by fighting; or they drive each +other about until they become greatly emaciated. They must, also, whilst +wandering about in eager search for the females, be often exposed to +various dangers. With many kinds of fish the males are much smaller than +the females, and they are believed often to be devoured by the latter or +by other fishes. With some birds the females appear to die in larger +proportion than the males: they are also liable to be destroyed on their +nests, or whilst in charge of their young. With insects the female larv +are often larger than those of the males, and would consequently be more +likely to be devoured: in some cases the mature females are less active +and less rapid in their movements than the males, and would not be so +well able to escape from danger. Hence, with animals in a state of +nature, in order to judge of the proportions of the sexes at maturity, +we must rely on mere estimation; and this, except perhaps when the +inequality is strongly marked, is but little trustworthy. Nevertheless, +as far as a judgment can be formed, we may conclude from the facts given +in the supplement, that the males of some few mammals, of many birds, of +some fish and insects, considerably exceed in number the females. + +The proportion between the sexes fluctuates slightly during successive +years: thus with race-horses, for every 100 females born, the males +varied from 107.1 in one year to 92.6 in another year, and with +greyhounds from 116.3 to 95.3. But had larger numbers been tabulated +throughout a more extensive area than England, these fluctuations would +probably have disappeared; and such as they are, they would hardly +suffice to lead under a state of nature to the effective action of +sexual selection. Nevertheless with some few wild animals, the +proportions seem, as shewn in the supplement, to fluctuate either during +different seasons or in different localities in a sufficient degree to +lead to such action. For it should be observed that any advantage gained +during certain years or in certain localities by those males which were +able to conquer other males, or were the most attractive to the females, +would probably be transmitted to the offspring and would not +subsequently be eliminated. During the succeeding seasons, when from the +equality of the sexes every male was everywhere able to procure a +female, the stronger or more attractive males previously produced would +still have at least as good a chance of leaving offspring as the less +strong or less attractive. + + +_Polygamy._--The practice of polygamy leads to the same results as +would follow from an actual inequality in the number of the sexes; for +if each male secures two or more females, many males will not be able to +pair; and the latter assuredly will be the weaker or less attractive +individuals. Many mammals and some few birds are polygamous, but with +animals belonging to the lower classes I have found no evidence of this +habit. The intellectual powers of such animals are, perhaps, not +sufficient to lead them to collect and guard a harem of females. That +some relation exists between polygamy and the development of secondary +sexual characters, appears nearly certain; and this supports the view +that a numerical preponderance of males would be eminently favourable to +the action of sexual selection. Nevertheless many animals, especially +birds, which are strictly monogamous, display strongly-marked secondary +sexual characters; whilst some few animals, which are polygamous, are +not thus characterised. + +We will first briefly run through the class of mammals, and then turn to +birds. The gorilla seems to be a polygamist, and the male differs +considerably from the female; so it is with some baboons which live in +herds containing twice as many adult females as males. In South America +the _Mycetes caraya_ presents well-marked sexual differences in colour, +beard, and vocal organs, and the male generally lives with two or three +wives: the male of the _Cebus capucinus_ differs somewhat from the +female, and appears to be polygamous.[340] Little is known on this head +with respect to most other monkeys, but some species are strictly +monogamous. The ruminants are eminently polygamous, and they more +frequently present sexual differences than almost any other group of +mammals, especially in their weapons, but likewise in other characters. +Most deer, cattle, and sheep are polygamous; as are most antelopes, +though some of the latter are monogamous. Sir Andrew Smith, in speaking +of the antelopes of South Africa, says that in herds of about a dozen +there was rarely more than one mature male. The Asiatic _Antilope saiga_ +appears to be the most inordinate polygamist in the world; for +Pallas[341] states that the male drives away all rivals, and collects a +herd of about a hundred, consisting of females and kids: the female is +hornless and has softer hair, but does not otherwise differ much from +the male. The horse is polygamous, but, except in his greater size and +in the proportions of his body, differs but little from the mare. The +wild boar, in his great tusks and some other characters, presents +well-marked sexual characters; in Europe and in India he leads a +solitary life, except during the breeding-season; but at this season he +consorts in India with several females, as Sir W. Elliot, who has had +large experience in observing this animal, believes: whether this holds +good in Europe is doubtful, but is supported by some statements. The +adult male Indian elephant, like the boar, passes much of his time in +solitude; but when associating with others, "it is rare to find," as Dr. +Campbell states, "more than one male with a whole herd of females." The +larger males expel or kill the smaller and weaker ones. The male differs +from the female by his immense tusks and greater size, strength, and +endurance; so great is the difference in these latter respects, that +the males when caught are valued at twenty per cent. above the +females.[342] With other pachydermatous animals the sexes differ very +little or not at all, and they are not, as far as known, polygamists. +Hardly a single species amongst the Cheiroptera and Edentata, or in the +great Orders of the Rodents and Insectivora, presents well-developed +secondary sexual differences; and I can find no account of any species +being polygamous, excepting, perhaps, the common rat, the males of +which, as some rat-catchers affirm, live with several females. + +The lion in South Africa, as I hear from Sir Andrew Smith, sometimes +lives with a single female, but generally with more than one, and, in +one case, was found with as many as five females, so that he is +polygamous. He is, as far as I can discover, the sole polygamist in the +whole group of the terrestrial Carnivora, and he alone presents +well-marked sexual characters. If, however, we turn to the marine +Carnivora, the case is widely different; for many species of seals +offer, as we shall hereafter see, extraordinary sexual differences, and +they are eminently polygamous. Thus the male sea-elephant of the +Southern Ocean, always possesses, according to Pron, several females, +and the sea-lion of Forster is said to be surrounded by from twenty to +thirty females. In the North, the male sea-bear of Steller is +accompanied by even a greater number of females. + +With respect to birds, many species, the sexes of which differ greatly +from each other, are certainly monogamous. In Great Britain we see +well-marked sexual differences in, for instance, the wild-duck which +pairs with a single female, with the common blackbird, and with the +bullfinch which is said to pair for life. So it is, as I am informed by +Mr. Wallace, with the Chatterers or Cotingid of South America, and +numerous other birds. In several groups I have not been able to discover +whether the species are polygamous or monogamous. Lesson says that birds +of paradise, so remarkable for their sexual differences, are polygamous, +but Mr. Wallace doubts whether he had sufficient evidence. Mr. Salvin +informs me that he has been led to believe that humming-birds are +polygamous. The male widow-bird; remarkable for his caudal plumes, +certainly seems to be a polygamist.[343] I have been assured by Mr. +Jenner Weir and by others, that three starlings not rarely frequent the +same nest; but whether this is a case of polygamy or polyandry has not +been ascertained. + +The Gallinace present almost as strongly marked sexual differences as +birds of paradise or humming-birds, and many of the species are, as is +well known, polygamous; others being strictly monogamous. What a +contrast is presented between the sexes by the polygamous peacock or +pheasant, and the monogamous guinea-fowl or partridge! Many similar +cases could be given, as in the grouse tribe, in which the males of the +polygamous capercailzie and black-cock differ greatly from the females; +whilst the sexes of the monogamous red grouse and ptarmigan differ very +little. Amongst the Cursores, no great number of species offer +strongly-marked sexual differences, except the bustards, and the great +bustard (_Otis tarda_), is said to be polygamous. With the Grallatores, +extremely few species differ sexually, but the ruff (_Machetes pugnax_) +affords a strong exception, and this species is believed by Montagu to +be a polygamist. Hence it appears that with birds there often exists a +close relation between polygamy and the development of strongly-marked +sexual differences. On asking Mr. Bartlett, at the Zoological Gardens, +who has had such large experience with birds, whether the male tragopan +(one of the Gallinace) was polygamous, I was struck by his answering, +"I do not know, but should think so from his splendid colours." + +It deserves notice that the instinct of pairing with a single female is +easily lost under domestication. The wild-duck is strictly monogamous, +the domestic-duck highly polygamous. The Rev. W. D. Fox informs me that +with some half-tamed wild-ducks, kept on a large pond in his +neighbourhood, so many mallards were shot by the gamekeeper that only +one was left for every seven or eight females; yet unusually large +broods were reared. The guinea-fowl is strictly monogamous; but Mr. Fox +finds that his birds succeed best when he keeps one cock to two or three +hens.[344] Canary-birds pair in a state of nature, but the breeders in +England successfully put one male to four or five females; nevertheless +the first female, as Mr. Fox has been assured, is alone treated as the +wife, she and her young ones being fed by him; the others are treated as +concubines. I have noticed these cases, as it renders it in some degree +probable that monogamous species, in a state of nature, might readily +become either temporarily or permanently polygamous. + +With respect to reptiles and fishes, too little is known of their habits +to enable us to speak of their marriage arrangements. The stickle-back +Gasterosteus, however, is said to be a polygamist;[345] and the male +during the breeding-season differs conspicuously from the female. + +To sum up on the means through which, as far as we can judge, sexual +selection has led to the development of secondary sexual characters. It +has been shewn that the largest number of vigorous offspring will be +reared from the pairing of the strongest and best-armed males, which +have conquered other males, with the most vigorous and best-nourished +females, which are the first to breed in the spring. Such females, if +they select the more attractive, and at the same time vigorous, males, +will rear a larger number of offspring than the retarded females, which +must pair with the less vigorous and less attractive males. So it will +be if the more vigorous males select the more attractive and at the same +time healthy and vigorous females; and this will especially hold good if +the male defends the female, and aids in providing food for the young. +The advantage thus gained by the more vigorous pairs in rearing a larger +number of offspring has apparently sufficed to render sexual selection +efficient. But a large preponderance in number of the males over the +females would be still more efficient; whether the preponderance was +only occasional and local, or permanent; whether it occurred at birth, +or subsequently from the greater destruction of the females; or whether +it indirectly followed from the practice of polygamy. + + +_The Male generally more modified than the Female._--Throughout the +animal kingdom, when the sexes differ from each other in external +appearance, it is the male which, with rare exceptions, has been chiefly +modified; for the female still remains more like the young of her own +species, and more like the other members of the same group. The cause of +this seems to lie in the males of almost all animals having stronger +passions than the females. Hence it is the males that fight together and +sedulously display their charms before the females; and those which are +victorious transmit their superiority to their male offspring. Why the +males do not transmit their characters to both sexes will hereafter be +considered. That the males of all mammals eagerly pursue the females is +notorious to every one. So it is with birds; but many male birds do not +so much pursue the female, as display their plumage, perform strange +antics, and pour forth their song, in her presence. With the few fish +which have been observed, the male seems much more eager than the +female; and so it is with alligators, and apparently with Batrachians. +Throughout the enormous class of insects, as Kirby remarks,[346] "the +law is, that the male shall seek the female." With spiders and +crustaceans, as I hear from two great authorities, Mr. Blackwall and Mr. +C. Spence Bate, the males are more active and more erratic in their +habits than the females. With insects and crustaceans, when the organs +of sense or locomotion are present in the one sex and absent in the +other, or when, as is frequently the case, they are more highly +developed in the one than the other, it is almost invariably the male, +as far as I can discover, which retains such organs, or has them most +developed; and this shews that the male is the more active member in the +courtship of the sexes.[347] + +The female, on the other hand, with the rarest exception, is less eager +than the male. As the illustrious Hunter[348] long ago observed, she +generally "requires to be courted;" she is coy, and may often be seen +endeavouring for a long time to escape from the male. Every one who has +attended to the habits of animals will be able to call to mind instances +of this kind. Judging from various facts, hereafter to be given, and +from the results which may fairly be attributed to sexual selection, the +female, though comparatively passive, generally exerts some choice and +accepts one male in preference to others. Or she may accept, as +appearances would sometimes lead us to believe, not the male which is +the most attractive to her, but the one which is the least distasteful. +The exertion of some choice on the part of the female seems almost as +general a law as the eagerness of the male. + +We are naturally led to enquire why the male in so many and such widely +distinct classes has been rendered more eager than the female, so that +he searches for her and plays the more active part in courtship. It +would be no advantage and some loss of power if both sexes were mutually +to search for each other; but why should the male almost always be the +seeker? With plants, the ovules after fertilisation have to be nourished +for a time; hence the pollen is necessarily brought to the female +organs--being placed on the stigma, through the agency of insects or of +the wind, or by the spontaneous movements of the stamens; and with the +Alg, &c., by the locomotive power of the antherozooids. With +lowly-organised animals permanently affixed to the same spot and having +their sexes separate, the male element is invariably brought to the +female; and we can see the reason; for the ova, even if detached before +being fertilised and not requiring subsequent nourishment or protection, +would be, from their larger relative size, less easily transported than +the male element. Hence plants[349] and many of the lower animals are, +in this respect, analogous. In the case of animals not affixed to the +same spot, but enclosed within a shell with no power of protruding any +part of their bodies, and in the case of animals having little power of +locomotion, the males must trust the fertilising element to the risk of +at least a short transit through the waters of the sea. It would, +therefore, be a great advantage to such animals, as their organisation +became perfected, if the males when ready to emit the fertilising +element, were to acquire the habit of approaching the female as closely +as possible. The males of various lowly-organised animals having thus +aboriginally acquired the habit of approaching and seeking the females, +the same habit would naturally be transmitted to their more highly +developed male descendants; and in order that they should become +efficient seekers, they would have to be endowed with strong passions. +The acquirement of such passions would naturally follow from the more +eager males leaving a larger number of offspring than the less eager. + +The great eagerness of the male has thus indirectly led to the much +more frequent development of secondary sexual characters in the male +than in the female. But the development of such characters will have +been much aided, if the conclusion at which I arrived after studying +domesticated animals, can be trusted, namely, that the male is more +liable to vary than the female. I am aware how difficult it is to verify +a conclusion of this kind. Some slight evidence, however, can be gained +by comparing the two sexes in mankind, as man has been more carefully +observed than any other animal. During the Novara Expedition[350] a vast +number of measurements of various parts of the body in different races +were made, and the men were found in almost every case to present a +greater range of variation than the women; but I shall have to recur to +this subject in a future chapter. Mr. J. Wood,[351] who has carefully +attended to the variation of the muscles in man, puts in italics the +conclusion that "the greatest number of abnormalities in each subject is +found in the males." He had previously remarked that "altogether in 102 +subjects the varieties of redundancy were found to be half as many again +as in females, contrasting widely with the greater frequency of +deficiency in females before described." Professor Macalister like wise +remarks[352] that variations in the muscles "are probably more common in +males than females." Certain muscles which are not normally present in +mankind are also more frequently developed in the male than in the +female sex, although exceptions to this rule are said to occur. Dr. +Burt Wilder[353] has tabulated the cases of 152 individuals with +supernumerary digits, of which 86 were males, and 39, or less than half, +females; the remaining 27 being of unknown sex. It should not, however, +be overlooked that women would more frequently endeavour to conceal a +deformity of this kind than men. Whether the large proportional number +of deaths of the male offspring of man and apparently of sheep, compared +with the female offspring, before, during, and shortly after birth (see +supplement), has any relation to a stronger tendency in the organs of +the male to vary and thus to become abnormal in structure or function, I +will not pretend to conjecture. + +In various classes of animals a few exceptional cases occur, in which +the female instead of the male has acquired well pronounced secondary +sexual characters, such as brighter colours, greater size, strength, or +pugnacity. With birds, as we shall hereafter see, there has sometimes +been a complete transposition of the ordinary characters proper to each +sex; the females having become the more eager in courtship, the males +remaining comparatively passive, but apparently selecting, as we may +infer from the results, the more attractive females. Certain female +birds have thus been rendered more highly coloured or otherwise +ornamented, as well as more powerful and pugnacious than the males, +these characters being transmitted to the female offspring alone. + +It may be suggested that in some cases a double process of selection has +been carried on; the males having selected the more attractive females, +and the latter the more attractive males. This process however, though +it might lead to the modification of both sexes, would not make the one +sex different from the other, unless indeed their taste for the +beautiful differed; but this is a supposition too improbable in the case +of any animal, excepting man, to be worth considering. There are, +however, many animals, in which the sexes resemble each other, both +being furnished with the same ornaments, which analogy would lead us to +attribute to the agency of sexual selection. In such cases it may be +suggested with more plausibility, that there has been a double or mutual +process of sexual selection; the more vigorous and precocious females +having selected the more attractive and vigorous males, the latter +having rejected all except the more attractive females. But from what we +know of the habits of animals, this view is hardly probable, the male +being generally eager to pair with any female. It is more probable that +the ornaments common to both sexes were acquired by one sex, generally +the male, and then transmitted to the offspring of both sexes. If, +indeed, during a lengthened period the males of any species were greatly +to exceed the females in number, and then during another lengthened +period under different conditions the reverse were to occur, a double, +but not simultaneous, process of sexual selection might easily be +carried on, by which the two sexes might be rendered widely different. + +We shall hereafter see that many animals exist, of which neither sex is +brilliantly coloured or provided with special ornaments, and yet the +members of both sexes or of one alone have probably been modified +through sexual selection. The absence of bright tints or other ornaments +may be the result of variations of the right kind never having occurred, +or of the animals themselves preferring simple colours, such as plain +black or white. Obscure colours have often been acquired through natural +selection for the sake of protection, and the acquirement through +sexual selection of conspicuous colours, may have been checked from the +danger thus incurred. But in other cases the males have probably +struggled together during long ages, through brute force, or by the +display of their charms, or by both means combined, and yet no effect +will have been produced unless a larger number of offspring were left by +the more successful males to inherit their superiority, than by the less +successful males; and this, as previously shewn, depends on various +complex contingencies. + +Sexual selection acts in a less rigorous manner than natural selection. +The latter produces its effects by the life or death at all ages of the +more or less successful individuals. Death, indeed, not rarely ensues +from the conflicts of rival males. But generally the less successful +male merely fails to obtain a female, or obtains later in the season a +retarded and less vigorous female, or, if polygamous, obtains fewer +females; so that they leave fewer, or less vigorous, or no offspring. In +regard to structures acquired through ordinary or natural selection, +there is in most cases, as long as the conditions of life remain the +same, a limit to the amount of advantageous modification in relation to +certain special ends; but in regard to structures adapted to make one +male victorious over another, either in fighting or in charming the +female, there is no definite limit to the amount of advantageous +modification; so that as long as the proper variations arise the work of +sexual selection will go on. This circumstance may partly account for +the frequent and extraordinary amount of variability presented by +secondary sexual characters. Nevertheless, natural selection will +determine that characters of this kind shall not be acquired by the +victorious males, which would be injurious to them in any high degree, +either by expending too much of their vital powers, or by exposing them +to any great danger. The development, however, of certain structures--of +the horns, for instance, in certain stags--has been carried to a +wonderful extreme; and in some instances to an extreme which, as far as +the general conditions of life are concerned, must be slightly injurious +to the male. From this fact we learn that the advantages which favoured +males have derived from conquering other males in battle or courtship, +and thus leaving a numerous progeny, have been in the long run greater +than those derived from rather more perfect adaptation to the external +conditions of life. We shall further see, and this could never have been +anticipated, that the power to charm the female has been in some few +instances more important than the power to conquer other males in +battle. + + +LAWS OF INHERITANCE. + +In order to understand how sexual selection has acted, and in the course +of ages has produced conspicuous results with many animals of many +classes, it is necessary to bear in mind the laws of inheritance, as far +as they are known. Two distinct elements are included under the term +"inheritance," namely the transmission and the development of +characters; but as these generally go together, the distinction is often +overlooked. We see this distinction in those characters which are +transmitted through the early years of life, but are developed only at +maturity or during old age. We see the same distinction more clearly +with secondary sexual characters, for these are transmitted through both +sexes, though developed in one alone. That they are present in both +sexes, is manifest when two species, having strongly-marked sexual +characters, are crossed, for each transmits the characters proper to +its own male and female sex to the hybrid offspring of both sexes. The +same fact is likewise manifest, when characters proper to the male are +occasionally developed in the female when she grows old or becomes +diseased; and so conversely with the male. Again, characters +occasionally appear, as if transferred from the male to the female, as +when, in certain breeds of the fowl, spurs regularly appear in the young +and healthy females; but in truth they are simply developed in the +female; for in every breed each detail in the structure of the spur is +transmitted through the female to her male offspring. In all cases of +reversion, characters are transmitted through two, three, or many +generations, and are then under certain unknown favourable conditions +developed. This important distinction between transmission and +development will be easiest kept in mind by the aid of the hypothesis of +pangenesis, whether or not it be accepted as true. According to this +hypothesis, every unit or cell of the body throws off gemmules or +undeveloped atoms, which are transmitted to the offspring of both sexes, +and are multiplied by self-division. They may remain undeveloped during +the early years of life or during successive generations; their +development into units or cells, like those from which they were +derived, depending on their affinity for, and union with, other units or +cells previously developed in the due order of growth. + + +_Inheritance at Corresponding Periods of Life._--This tendency is well +established. If a new character appears in an animal whilst young, +whether it endures throughout life or lasts only for a time, it will +reappear, as a general rule, at the same age and in the same manner in +the offspring. If, on the other hand, a new character appears at +maturity, or even during old age, it tends to reappear in the offspring +at the same advanced age. When deviations from this rule occur, the +transmitted characters much oftener appear before than after the +corresponding age. As I have discussed this subject at sufficient length +in another work,[354] I will here merely give two or three instances, +for the sake of recalling the subject to the reader's mind. In several +breeds of the Fowl, the chickens whilst covered with down, in their +first true plumage, and in their adult plumage, differ greatly from each +other, as well as from their common parent-form, the _Gallus bankiva_; +and these characters are faithfully transmitted by each breed to their +offspring at the corresponding period of life. For instance, the +chickens of spangled Hamburghs, whilst covered with down, have a few +dark spots on the head and rump, but are not longitudinally striped, as +in many other breeds; in their first true plumage, "they are beautifully +pencilled," that is each feather is transversely marked by numerous dark +bars; but in their second plumage the feathers all become spangled or +tipped with a dark round spot.[355] Hence in this breed variations have +occurred and have been transmitted at three distinct periods of life. +The Pigeon offers a more remarkable case, because the aboriginal +parent-species does not undergo with advancing age any change of +plumage, excepting that at maturity the breast becomes more iridescent; +yet there are breeds which do not acquire their characteristic colours +until they have moulted two, three, or four times; and these +modifications of plumage are regularly transmitted. + + +_Inheritance at Corresponding Seasons of the Year._--With animals in a +state of nature innumerable instances occur of characters periodically +appearing at different seasons. We see this with the horns of the stag, +and with the fur of arctic animals which becomes thick and white during +the winter. Numerous birds acquire bright colours and other decorations +during the breeding-season alone. I can throw but little light on this +form of inheritance from facts observed under domestication. Pallas +states,[356] that in Siberia domestic cattle and horses periodically +become lighter-coloured during the winter; and I have observed a similar +marked change of colour in certain ponies in England. Although I do not +know that this tendency to assume a differently coloured coat during +different seasons of the year is transmitted, yet it probably is so, as +all shades of colour are strongly inherited by the horse. Nor is this +form of inheritance, as limited by season, more remarkable than +inheritance as limited by age or sex. + + +_Inheritance as Limited by Sex._--The equal transmission of characters +to both, sexes is the commonest form of inheritance, at least with those +animals which do not present strongly-marked sexual differences, and +indeed with many of these. But characters are not rarely transferred +exclusively to that sex, in which they first appeared. Ample evidence on +this head has been advanced in my work on Variation under +Domestication; but a few instances may here be given. There are breeds +of the sheep and goat, in which the horns of the male differ greatly in +shape from those of the female; and these differences, acquired under +domestication, are regularly transmitted to the same sex. With +tortoise-shell cats the females alone, as a general rule, are thus +coloured, the males being rusty-red. With most breeds of the fowl, the +characters proper to each sex are transmitted to the same sex alone. So +general is this form of transmission that it is an anomaly when we see +in certain breeds variations transmitted equally to both sexes. There +are also certain sub-breeds of the fowl in which the males can hardly be +distinguished from each other, whilst the females differ considerably in +colour. With the pigeon the sexes of the parent-species do not differ in +any external character; nevertheless in certain domesticated breeds the +male is differently coloured from the female.[357] The wattle in the +English Carrier pigeon and the crop in the Pouter are more highly +developed in the male than in the female; and although these characters +have been gained through long-continued selection by man, the difference +between the two sexes is wholly due to the form of inheritance which has +prevailed; for it has arisen, not from, but rather in opposition to, the +wishes of the breeder. + +Most of our domestic races have been formed by the accumulation of many +slight variations; and as some of the successive steps have been +transmitted to one sex alone, and some to both sexes, we find in the +different breeds of the same species all gradations between great sexual +dissimilarity and complete similarity. Instances have already been +given with the breeds of the fowl and pigeon; and under nature analogous +cases are of frequent occurrence. With animals under domestication, but +whether under nature I will not venture to say, one sex may lose +characters proper to it, and may thus come to resemble to a certain +extent the opposite sex; for instance, the males of some breeds of the +fowl have lost their masculine plumes and hackles. On the other hand the +differences between the sexes may be increased under domestication, as +with merino sheep, in which the ewes have lost their horns. Again, +characters proper to one sex may suddenly appear in the other sex; as +with those sub-breeds of the fowl in which the hens whilst young acquire +spurs; or, as in certain Polish sub-breeds, in which the females, as +there is reason to believe, originally acquired a crest, and +subsequently transferred it to the males. All these cases are +intelligible on the hypothesis of pangenesis; for they depend on the +gemmules of certain units of the body, although present in both sexes, +becoming through the influence of domestication dormant in the one sex; +or if naturally dormant, becoming developed. + +There is one difficult question which it will be convenient to defer to +a future chapter; namely, whether a character at first developed in both +sexes, can be rendered through selection limited in its development to +one sex alone. If, for instance, a breeder observed that some of his +pigeons (in which species characters are usually transferred in an equal +degree to both sexes) varied into pale blue; could he by long-continued +selection make a breed, in which the males alone should be of this tint, +whilst the females remained unchanged? I will here only say, that this, +though perhaps not impossible, would be extremely difficult; for the +natural result of breeding from the pale-blue males would be to change +his whole stock, including both sexes, into this tint. If, however, +variations of the desired tint appeared, which were from the first +limited in their development to the male sex, there would not be the +least difficulty in making a breed characterised by the two sexes being +of a different colour, as indeed has been effected with a Belgian breed, +in which the males alone are streaked with black. In a similar manner, +if any variation appeared in a female pigeon, which was from the first +sexually limited in its development, it would be easy to make a breed +with the females alone thus characterised; but if the variation was not +thus originally limited, the process would be extremely difficult, +perhaps impossible. + + +_On the Relation between the period of Development of a Character and +its transmission to one sex or to both sexes._--Why certain characters +should be inherited by both sexes, and other characters by one sex +alone, namely by that sex in which the character first appeared, is in +most cases quite unknown. We cannot even conjecture why with certain +sub-breeds of the pigeon, black stri, though transmitted through the +female, should be developed in the male alone, whilst every other +character is equally transferred to both sexes. Why, again, with cats, +the tortoise-shell colour should, with rare exceptions, be developed in +the female alone. The very same characters, such as deficient or +supernumerary digits, colour-blindness, &c., may with mankind be +inherited by the males alone of one family, and in another family by the +females alone, though in both cases transmitted through the opposite as +well as the same sex.[358] Although we are thus ignorant, two rules +often hold good, namely that variations which, first appear in either +sex at a late period of life, tend to be developed in the same sex +alone; whilst variations which first appear early in life in either sex +tend to be developed in both sexes. I am, however, far from supposing +that this is the sole determining cause. As I have not elsewhere +discussed this subject, and as it has an important bearing on sexual +selection, I must here enter into lengthy and somewhat intricate +details. + +It is in itself probable that any character appearing at an early age +would tend to be inherited equally by both sexes, for the sexes do not +differ much in constitution, before the power of reproduction is gained. +On the other hand, after this power has been gained and the sexes have +come to differ in constitution, the gemmules (if I may again use the +language of pangenesis) which are cast off from each varying part in the +one sex would be much more likely to possess the proper affinities for +uniting with the tissues of the same sex, and thus becoming developed, +than with those of the opposite sex. + +I was first led to infer that a relation of this kind exists, from the +fact that whenever and in whatever manner the adult male has come to +differ from the adult female, he differs in the same manner from the +young of both sexes. The generality of this fact is quite remarkable: it +holds good with almost all mammals, birds, amphibians, and fishes; also +with many crustaceans, spiders and some few insects, namely certain +orthoptera and libellul. In all these cases the variations, through the +accumulation of which the male acquired his proper masculine characters, +must have occurred at a somewhat late period of life; otherwise the +young males would have been similarly characterised; and conformably +with our rule, they are transmitted to and developed in the adult males +alone. When, on the other hand, the adult male closely resembles the +young of both sexes (these, with rare exceptions, being alike), he +generally resembles the adult female; and in most of these cases the +variations through which the young and old acquired their present +characters, probably occurred in conformity with our rule during youth. +But there is here room for doubt, as characters are sometimes +transferred to the offspring at an earlier age than that at which they +first appeared in the parents, so that the parents may have varied when +adult, and have transferred their characters to their offspring whilst +young. There are, moreover, many animals, in which the two sexes closely +resemble each other, and yet both differ from their young; and here the +characters of the adults must have been acquired late in life; +nevertheless, these characters in apparent contradiction to our rule, +are transferred to both sexes. We must not, however, overlook the +possibility or even probability of successive variations of the same +nature sometimes occurring, under exposure to similar conditions, +simultaneously in both sexes at a rather late period of life; and in +this case the variations would be transferred to the offspring of both +sexes at a corresponding late age; and there would be no real +contradiction to our rule of the variations which occur late in life +being transferred exclusively to the sex in which they first appeared. +This latter rule seems to hold true more generally than the second rule, +namely, that variations which occur in either sex early in life tend to +be transferred to both sexes. As it was obviously impossible even to +estimate in how large a number of cases throughout the animal kingdom +these two propositions hold good, it occurred to me to investigate some +striking or crucial instances, and to rely on the result. + +An excellent case for investigation is afforded by the Deer Family. In +all the species, excepting one, the horns are developed in the male +alone, though certainly transmitted through the female, and capable of +occasional abnormal development in her. In the reindeer, on the other +hand, the female is provided with horns; so that in this species, the +horns ought, according to our rule, to appear early in life, long before +the two sexes had arrived at maturity and had come to differ much in +constitution. In all the other species of deer the horns ought to appear +later in life, leading to their development in that sex alone, in which +they first appeared in the progenitor of the whole Family. Now in seven +species, belonging to distinct sections of the family and inhabiting +different regions, in which the stags alone bear horns, I find that the +horns first appear at periods varying from nine months after birth in +the roebuck to ten or twelve or even more months in the stags of the six +other larger species.[359] But with the reindeer the case is widely +different, for as I hear from Prof. Nilsson, who kindly made special +enquiries for me in Lapland, the horns appear in the young animals +within four or five weeks after birth, and at the same time in both +sexes. So that here we have a structure, developed at a most unusually +early age in one species of the family, and common to both sexes in this +one species. + +In several kinds of antelopes the males alone are provided with horns, +whilst in the greater number both sexes have horns. With respect to the +period of development, Mr. Blyth informs me that there lived at one time +in the Zoological Gardens a young koodoo (_Ant. strepsiceros_), in which +species the males alone are horned, and the young of a closely-allied +species, viz. the eland (_Ant. oreas_), in which both sexes are horned. +Now in strict conformity with our rule, in the young male koodoo, +although arrived at the age of ten months, the horns were remarkably +small considering the size ultimately attained by them: whilst in the +young male eland, although only three months old, the horns were already +very much larger than in the koodoo. It is also worth notice that in the +prong-horned antelope,[360] in which species the horns, though present +in both sexes, are almost rudimentary in the female, they do not appear +until about five or six months after birth. With sheep, goats, and +cattle, in which the horns are well developed in both sexes, though not +quite equal in size, they can be felt, or even seen, at birth or soon +afterwards.[361] Our rule, however, fails in regard to some breeds of +sheep, for instance merinos, in which the rams alone are horned; for I +cannot find on enquiry,[362] that the horns are developed later in life +in this breed than in ordinary sheep in which both sexes are horned. But +with domesticated sheep the presence or absence of horns is not a firmly +fixed character; a certain proportion of the merino ewes bearing small +horns, and some of the rams being hornless; whilst with ordinary sheep +hornless ewes are occasionally produced. + +In most of the species of the splendid family of the Pheasants, the +males differ conspicuously from the females, and they acquire their +ornaments at a rather late period of life. The eared pheasant +(_Crossoptilon auritum_), however, offers a remarkable exception, for +both sexes possess the fine caudal plumes, the large ear-tufts and the +crimson velvet about the head; and I find on enquiry in the Zoological +Gardens that all these characters, in accordance with our rule, appear +very early in life. The adult male can, however, be distinguished from +the adult female by one character, namely by the presence of spurs; and +conformably with our rule, these do not begin to be developed, as I am +assured by Mr. Bartlett, before the age of six months, and even at this +age, can hardly be distinguished in the two sexes.[363] The male and +female Peacock differ conspicuously from each other in almost every +part of their plumage, except in the elegant head-crest, which is common +to both sexes; and this is developed very early in life, long before the +other ornaments which are confined to the male. The wild-duck offers an +analogous case, for the beautiful green speculum on the wings is common +to both sexes, though duller and somewhat smaller in the female, and it +is developed early in life, whilst the curled tail-feathers and other +ornaments peculiar to the male are developed later.[364] Between such +extreme cases of close sexual resemblance and wide dissimilarity, as +those of the Crossoptilon and peacock, many intermediate ones could be +given, in which the characters follow in their order of development our +two rules. + +As most insects emerge from their pupal state in a mature condition, it +is doubtful whether the period of development determines the +transference of their characters to one or both sexes. But we do not +know that the coloured scales, for instance, in two species of +butterflies, in one of which the sexes differ in colour, whilst in the +other they are alike, are developed at the same relative age in the +cocoon. Nor do we know whether all the scales are simultaneously +developed on the wings of the same species of butterfly, in which +certain coloured marks are confined to one sex, whilst other marks are +common to both sexes. A difference of this kind in the period of +development is not so improbable as it may at first appear; for with the +Orthoptera, which assume their adult state, not by a single +metamorphosis, but by a succession of moults, the young males of some +species at first resemble the females, and acquire their distinctive +masculine characters only during a later moult. Strictly analogous cases +occur during the successive moults of certain male crustaceans. + +We have as yet only considered the transference of characters, +relatively to their period of development, with species in a natural +state; we will now turn to domesticated animals; first touching on +monstrosities and diseases. The presence of supernumerary digits, and +the absence of certain phalanges, must be determined at an early +embryonic period--the tendency to profuse bleeding is at least +congenital, as is probably colour-blindness--yet these peculiarities, +and other similar ones, are often limited in their transmission to one +sex; so that the rule that characters which are developed at an early +period tend to be transmitted to both sexes, here wholly fails. But this +rule, as before remarked, does not appear to be nearly so generally true +as the converse proposition, namely, that characters which appear late +in life in one sex are transmitted exclusively to the same sex. From the +fact of the above abnormal peculiarities becoming attached to one sex, +long before the sexual functions are active, we may infer that there +must be a difference of some kind between the sexes at an extremely +early age. With respect to sexually-limited diseases, we know too little +of the period at which they originate, to draw any fair conclusion. +Gout, however, seems to fall under our rule; for it is generally caused +by intemperance after early youth, and is transmitted from the father to +his sons in a much more marked manner than to his daughters. + +In the various domestic breeds of sheep, goats, and cattle, the males +differ from their respective females in the shape or development of +their horns, forehead, mane, dewlap, tail, and hump on the shoulders; +and these peculiarities, in accordance with our rule, are not fully +developed until rather late in life. With dogs, the sexes do not differ, +except that in certain breeds, especially in the Scotch deer-hound, the +male is much larger and heavier than the female; and as we shall see in +a future chapter, the male goes on increasing in size to an unusually +late period of life, which will account, according to our rule, for his +increased size being transmitted to his male offspring alone. On the +other hand, the tortoise-shell colour of the hair, which is confined to +female cats, is quite distinct at birth, and this case violates our +rule. There is a breed of pigeons in which the males alone are streaked +with black, and the streaks can be detected even in the nestlings; but +they become more conspicuous at each successive moult, so that this case +partly opposes and partly supports the rule. With the English Carrier +and Pouter pigeon the full development of the wattle and the crop occurs +rather late in life, and these characters, conformably with our rule, +are transmitted in full perfection to the males alone. The following +cases perhaps come within the class previously alluded to, in which the +two sexes have varied in the same manner at a rather late period of +life, and have consequently transferred their new characters to both +sexes at a corresponding late period; and if so, such cases are not +opposed to our rule. Thus there are sub-breeds of the pigeon, described +by Neumeister,[365] both sexes of which change colour after moulting +twice or thrice, as does likewise the Almond Tumbler; nevertheless these +changes, though occurring rather late in life, are common to both sexes. +One variety of the Canary-bird, namely the London Prize, offers a nearly +analogous case. + +With the breeds of the Fowl the inheritance of various characters by one +sex or by both sexes, seems generally determined by the period at which +such characters are developed. Thus in all the many breeds in which the +adult male differs greatly in colour from the female and from the adult +male parent-species, he differs from the young male, so that the newly +acquired characters must have appeared at a rather late period of life. +On the other hand with most of the breeds in which the two sexes +resemble each other, the young are coloured in nearly the same manner as +their parents, and this renders it probable that their colours first +appeared early in life. We have instances of this fact in all black and +white breeds, in which the young and old of both sexes are alike; nor +can it be maintained that there is something peculiar in a black or +white plumage, leading to its transference to both sexes; for the males +alone of many natural species are either black or white, the females +being very differently coloured. With the so-called Cuckoo sub-breeds of +the fowl, in which the feathers are transversely pencilled with dark +stripes, both sexes and the chickens are coloured in nearly the same +manner. The laced plumage of the Sebright bantam is the same in both +sexes, and in the chickens the feathers are tipped with black, which +makes a near approach to lacing. Spangled Hamburghs, however, offer a +partial exception, for the two sexes, though not quite alike, resemble +each other more closely than do the sexes of the aboriginal +parent-species, yet they acquire their characteristic plumage late in +life, for the chickens are distinctly pencilled. Turning to other +characters besides colour: the males alone of the wild parent-species +and of most domestic breeds possess a fairly well developed comb, but in +the young of the Spanish fowl it is largely developed at a very early +age, and apparently in consequence of this it is of unusual size in the +adult females. In the Game breeds pugnacity is developed at a +wonderfully early age, of which curious proofs could be given; and this +character is transmitted to both sexes, so that the hens, from their +extreme pugnacity, are now generally exhibited in separate pens. With +the Polish breeds the bony protuberance of the skull which supports the +crest is partially developed even before the chickens are hatched, and +the crest itself soon begins to grow, though at first feebly;[366] and +in this breed a great bony protuberance and an immense crest +characterise the adults of both sexes. + +Finally, from what we have now seen of the relation which exists in many +natural species and domesticated races, between the period of the +development of their characters and the manner of their +transmission--for example the striking fact of the early growth of the +horns in the reindeer, in which both sexes have horns, in comparison +with their much later growth in the other species in which the male +alone bears horns--we may conclude that one cause, though not the sole +cause, of characters being exclusively inherited by one sex, is their +development at a late age. And secondly, that one, though apparently a +less efficient, cause of characters being inherited by both sexes is +their development at an early age, whilst the sexes differ but little in +constitution. It appears, however, that some difference must exist +between the sexes even during an early embryonic period, for characters +developed at this age not rarely become attached to one sex. + + +_Summary and concluding remarks._--From the foregoing discussion on the +various laws of inheritance, we learn that characters often or even +generally tend to become developed in the same sex, at the same age, and +periodically at the same season of the year, in which they first +appeared in the parents. But these laws, from unknown causes, are very +liable to change. Hence the successive steps in the modification of a +species might readily be transmitted in different ways; some of the +steps being transmitted to one sex, and some to both; some to the +offspring at one age, and some at all ages. Not only are the laws of +inheritance extremely complex, but so are the causes which induce and +govern variability. The variations thus caused are preserved and +accumulated by sexual selection, which is in itself an extremely complex +affair, depending, as it does, on ardour in love, courage, and the +rivalry of the males, and on the powers of perception, taste, and will +of the female. Sexual selection will also be dominated by natural +selection for the general welfare of the species. Hence the manner in +which the individuals of either sex or of both sexes are affected +through sexual selection cannot fail to be complex in the highest +degree. + +When variations occur late in life in one sex, and are transmitted to +the same sex at the same age, the other sex and the young are +necessarily left unmodified. When they occur late in life, but are +transmitted to both sexes at the same age, the young alone are left +unmodified. Variations, however, may occur at any period of life in one +sex or in both, and be transmitted to both sexes at all ages, and then +all the individuals of the species will be similarly modified. In the +following chapters it will be seen that all these cases frequently occur +under nature. + +Sexual selection can never act on any animal whilst young, before the +age for reproduction has arrived. From the great eagerness of the male +it has generally acted on this sex and not on the females. The males +have thus become provided with weapons for fighting with their rivals, +or with organs for discovering and securely holding the female, or for +exciting and charming her. When the sexes differ in these respects, it +is also, as we have seen, an extremely general law that the adult male +differs more or less from the young male; and we may conclude from this +fact that the successive variations, by which the adult male became +modified, cannot have occurred much before the age for reproduction. How +then are we to account for this general and remarkable coincidence +between the period of variability and that of sexual selection,-- +principles which are quite independent of each other? I +think we can see the cause: it is not that the males have never varied +at an early age, but that such variations have commonly been lost, +whilst those occurring at a later age have been preserved. + +All animals produce more offspring than can survive to maturity; and we +have every reason to believe that death falls heavily on the weak and +inexperienced young. If then a certain proportion of the offspring were +to vary at birth or soon afterwards, in some manner which at this age +was of no service to them, the chance of the preservation of such +variations would be small. We have good evidence under domestication how +soon variations of all kinds are lost, if not selected. But variations +which occurred at or near maturity, and which were of immediate service +to either sex, would probably be preserved; as would similar variations +occurring at an earlier period in any individuals which happened to +survive. As this principle has an important bearing on sexual selection, +it may be advisable to give an imaginary illustration. We will take a +pair of animals, neither very fertile nor the reverse, and assume that +after arriving at maturity they live on an average for five years, +producing each year five young. They would thus produce 25 offspring; +and it would not, I think, be an unfair estimate to assume that 18 or 20 +out of the 25 would perish before maturity, whilst still young and +inexperienced; the remaining seven or five sufficing to keep up the +stock of mature individuals. If so, we can see that variations which +occurred during youth, for instance in brightness, and which were not of +the least service to the young, would run a good chance of being utterly +lost. Whilst similar variations, which occurring at or near maturity in +the comparatively few individuals surviving to this age, and which +immediately gave an advantage to certain males, by rendering them more +attractive to the females, would be likely to be preserved. No doubt +some of the variations in brightness which occurred at an earlier age +would by chance be preserved, and eventually give to the male the same +advantage as those which appeared later; and this will account for the +young males commonly partaking to a certain extent (as may be observed +with many birds) of the bright colours of their adult male parents. If +only a few of the successive variations in brightness were to occur at a +late age, the adult male would be only a little brighter than the young +male; and such cases are common. + +In this illustration I have assumed that the young varied in a manner +which was of no service to them; but many characters proper to the adult +male would be actually injurious to the young,--as bright colours from +making them conspicuous, or horns of large size from expending much +vital force. Such variations in the young would promptly be eliminated +through natural selection. With the adult and experienced males, on the +other hand, the advantage thus derived in their rivalry with other males +would often more than counterbalance exposure to some degree of danger. +Thus we can understand how it is that variations which must originally +have appeared rather late in life have alone or in chief part been +preserved for the development of secondary sexual characters; and the +remarkable coincidence between the periods of variability and of sexual +selection is intelligible. + +As variations which give to the male an advantage in lighting with other +males, or in finding, securing, or charming the female, would be of no +use to the female, they will not have been preserved in this sex either +during youth or maturity. Consequently such variations would be +extremely liable to be lost; and the female, as far as these characters +are concerned, would be left unmodified, excepting in so far as she may +have received them by transference from the male. No doubt if the female +varied and transferred serviceable characters to her male offspring, +these would be favoured through sexual selection; and then both sexes +would thus far be modified in the same manner. But I shall hereafter +have to recur to these more intricate contingencies. + +In the following chapters, I shall treat of the secondary sexual +characters in animals of all classes, and shall endeavour in each case +to apply the principles explained in the present chapter. The lowest +classes will detain us for a very short time, but the higher animals, +especially birds, must be treated at considerable length. It should be +borne in mind that for reasons already assigned, I intend to give only a +few illustrative instances of the innumerable structures by the aid of +which the male finds the female, or, when found, holds her. On the other +hand, all structures and instincts by which the male conquers other +males, and by which he allures or excites the female, will be fully +discussed, as these are in many ways the most interesting. + + +_Supplement on the proportional numbers of the two sexes in animals +belonging to various classes._ + + +As no one, as far as I can discover, has paid attention to the relative +numbers of the two sexes throughout the animal kingdom, I will here give +such materials as I have been able to collect, although they are +extremely imperfect. They consist in only a few instances of actual +enumeration, and the numbers are not very large. As the proportions are +known with certainty on a large scale in the case of man alone, I will +first give them, as a standard of comparison. + + +_Man._--In England during ten years (from 1857 to 1866) 707,120 children +on an annual average have been born alive, in the proportion of 104.5 +males to 100 females. But in 1857 the male births throughout England +were as 105.2, and in 1865 as 104.0 to 100. Looking to separate +districts, in Buckinghamshire (where on an average 5000 children are +annually born) the _mean_ proportion of male to female births, during +the whole period of the above ten years, was as 102.8 to 100; whilst in +N. Wales (where the average annual births are 12,873) it was as high as +106.2 to 100. Taking a still smaller district, viz., Rutlandshire (where +the annual births average only 739), in 1864 the male births were as +114.6, and in 1862 as 97.0 to 100; but even in this small district the +average of the 7385 births during the whole ten years was as 104.5 to +100; that is in the same ratio as throughout England.[367] The +proportions are sometimes slightly disturbed by unknown causes; thus +Prof. Faye states "that in some districts of Norway there has been +during a decennial period a steady deficiency of boys, whilst in others +the opposite condition has existed." In France during forty-four years +the male to the female births have been as 106.2 to 100; but during this +period it has occurred five times in one department, and six times in +another, that the female births have exceeded the males. In Russia the +average proportion is as high as 108.9 to 100.[368] It is a singular +fact that with Jews the proportion of male births is decidedly larger +than with Christians: thus in Prussia the proportion is as 113, in +Breslau as 114, and in Livonia as 120 to 100; the Christian births in +these countries being the same as usual, for instance, in Livonia as 104 +to 100.[369] It is a still more singular fact that in different nations, +under different conditions and climates, in Naples, Prussia, Westphalia, +France and England, the excess of male over female births is less when +they are illegitimate than when legitimate.[370] + +In various parts of Europe, according to Prof. Faye and other authors, +"a still greater preponderance of males would be met with, if death +struck both sexes in equal proportion in the womb and during birth. But +the fact is, that for every 100 still-born females, we have in several +countries from 134.6 to 144.9 still-born males." Moreover during the +first four or five years of life more male children die than females; +"for example in England, during the first year, 126 boys die +for every 100 girls,--a proportion which in France is still more +unfavourable."[371] As a consequence of this excess in the death-rate of +male children, and of the exposure of men when adult to various dangers, +and of their tendency to emigrate, the females in all old-settled +countries, where statistical records have been kept,[372] are found to +preponderate considerably over the males. + +It has often been supposed that the relative ages of the parents +determine the sex of the offspring; and Prof. Leuckart[373] has advanced +what he considers sufficient evidence, with respect to man and certain +domesticated animals, to shew that this is one important factor in the +result. So again the period of impregnation has been thought to be the +efficient cause; but recent observations discountenance this belief. +Again, with mankind polygamy has been supposed to lead to the birth of a +greater proportion of female infants; but Dr. J. Campbell[374] carefully +attended to this subject in the harems of Siam, and he concludes that +the proportion of male to female births is the same as from monogamous +unions. Hardly any animal has been rendered so highly polygamous as our +English race-horses, and we shall immediately see that their male and +female offspring are almost exactly equal in number. + + + _Horses._--Mr. Tegetmeier has been so kind as to tabulate for + me from the 'Racing Calendar' the births of race-horses during + a period of twenty-one years, viz. from 1846 to 1867; 1849 + being omitted, as no returns were that year published. The + total births have been 25,560,[375] consisting of 12,763 males + and 12,797 females, or in the proportion of 99.7 males to 100 + females. As these numbers are tolerably large, and as they are + drawn from all parts of England, during several years, we may + with much confidence conclude that with the domestic horse, or + at least with the race-horse, the two sexes are produced in + almost equal numbers. The fluctuations in the proportions + during successive years are closely like those which occur with + mankind, when a small and thinly-populated area is considered: + thus in 1856 the male horses were as 107.1, and in 1867 as only + 92.6 to 100 females. In the tabulated returns the proportions + vary in cycles, for the males exceeded the females during six + successive years; and the females exceeded the males during two + periods each of four years: this, however, may be accidental; + at least I can detect nothing of the kind with man in the + decennial table in the Registrar's Report for 1866. I may add + that certain, mares, and this holds good with certain cows and + with women, tend to produce more of one sex than of the other; + Mr. Wright of Yeldersley House, informs me that one of his Arab + mares, though put seven times to different horses, produced + seven fillies. + + + _Dogs._--During a period of twelve years, from 1857 to 1868, + the births of a large number of greyhounds, throughout England, + have been sent to the 'Field' newspaper; and I am again + indebted to Mr. Tegetmeier for carefully tabulating the + results. The recorded, births have been 6878, consisting of + 3605 males and 3273 females, that is, in the proportion of + 110.1 males to 100 females. The greatest fluctuations occurred + in 1864, when the proportion was as 95.3 males, and in 1867, as + 116.3 males to 100 females. The above average proportion of + 110.1 to 100 is probably nearly correct in the case of the + greyhound, but whether it would hold with other domesticated + breeds is in some degree doubtful. Mr. Cupples has enquired + from several great breeders of dogs, and finds that all without + exception believe that females are produced in excess; he + suggests that this belief may have arisen from females being + less valued and the consequent disappointment producing a + stronger impression on the mind. + + + _Sheep._--The sexes of sheep are not ascertained by + agriculturists until several months after birth, at the period + when the males are castrated; so that the following returns do + not give the proportions at birth. Moreover, I find that + several great breeders in Scotland, who annually raise some + thousand sheep, are firmly convinced that a larger proportion + of males than of females die during the first one or two years; + therefore the proportion of males would be somewhat greater at + birth than at the age of castration. This is a remarkable + coincidence with what occurs, as we have seen, with mankind, + and both cases probably depend on some common cause. I have + received returns from four gentlemen in England who have bred + lowland sheep, chiefly Leicesters, during the last ten or + sixteen years; they amount altogether to 8965 births, + consisting of 4407 males and 4558 females; that is in the + proportion of 96.7 males to 100 females. With respect to + Cheviot and black-faced sheep bred in Scotland, I have received + returns from six breeders, two of them on a large scale, + chiefly for the years 1867-1869, but some of the returns + extending back to 1862. The total number recorded amounts to + 50,685, consisting of 25,071 males and 25,614 females, or in + the proportion of 97.9 males to 100 females. If we take the + English and Scotch returns together, the total number amounts + to 59,650, consisting of 29,478 males and 30,172 females, or + as 977 to 100. So that with sheep at the age of castration the + females are certainly in excess of the males; but whether this + would hold good at birth is doubtful, owing to the greater + liability in the males to early death.[376] + + Of _Cattle_ I have received returns from nine gentlemen of 982 + births, too few to be trusted; these consisted of 477 + bull-calves and 505 cow-calves; _i.e._ in the proportion of + 944 males to 100 females. The Rev. W. D. Fox informs me that + in 1867 out of 34 calves born on a farm in Derbyshire only one + was a bull. Mr. Harrison Weir writes to me that he has enquired + from several breeders of _Pigs_, and most of them estimate the + male to the female births as about 7 to 6. This same gentleman + has bred _Rabbits_ for many years, and has noticed that a far + greater number of bucks are produced than does. + + Of mammalia in a state of nature I have been able to learn very + little. In regard to the common rat, I have received + conflicting statements. Mr. R. Elliot of Laighwood, informs me + that a rat-catcher assured him that he had always found the + males in great excess, even with the young in the nest. In + consequence of this, Mr. Elliot himself subsequently examined + some hundred old ones, and found the statement true. Mr. F. + Buckland has bred a large number of white rats, and he also + believes that the males greatly exceed the females. In regard + to Moles, it is said that "the males are much more numerous + than the females;"[377] and as the catching of these animals is + a special occupation, the statement may perhaps be trusted. Sir + A. Smith, in describing an antelope of S. Africa[378] (_Kobus + ellipsiprymnus_), remarks, that in the herds of this and other + species, the males are few in number compared with the females: + the natives believe that they are born in this proportion; + others believe that the younger males are expelled from the + herds, and Sir A. Smith says, that though he has himself never + seen herds consisting of young males alone, others affirm that + this does occur. It appears probable that the young males when + expelled from the herd, would be likely to fell a prey to the + many beasts of prey of the country. + + + BIRDS. + + With respect to the _Fowl_, I have received only one account, + namely, that out of 1001 chickens of a highly-bred stock of + Cochins, reared during eight years by Mr. Stretch, 487 proved + males and 514 females: _i.e._ as 94.7 to 100. In regard to + domestic pigeons there is good evidence that the males are + produced in excess, or that their lives are longer; for these + birds invariably pair, and single males, as Mr. Tegetmeier + informs me, can always be purchased cheaper than females. + Usually the two birds reared from the two eggs laid in the same + nest consist of a male and female; but Mr. Harrison Weir, who + has been so large a breeder, says that he has often bred two + cocks from the same nest, and seldom two hens; moreover the hen + is generally the weaker of the two, and more liable to perish. + + With respect to birds in a state of nature, Mr. Gould and + others[379] are convinced that the males are generally the more + numerous; and as the young males of many species resemble the + females, the latter would naturally appear to be the most + numerous. Large numbers of pheasants are reared by Mr. Baker of + Leadenhall from eggs laid by wild birds, and he informs Mr. + Jenner Weir that four or five males to one female are generally + produced. An experienced observer remarks[380] that in + Scandinavia the broods of the capercailzie and black-cock + contain more males than females; and that with the Dal-ripa (a + kind of ptarmigan) more males than females attend the _leks_ or + places of courtship; but this latter circumstance is accounted + for by some observers by a greater number of hen birds being + killed by vermin. From various facts given by White of + Selbourne,[381] it seems clear that the males of the partridge + must be in considerable excess in the south of England; and I + have been assured that this is the case in Scotland. Mr. Weir + on enquiring from the dealers who receive at certain seasons + large numbers of ruffs (_Machetes pugnax_) was told that the + males are much the most numerous. This same naturalist has also + enquired for me from the bird-catchers, who annually catch an + astonishing number of various small species alive for the + London market, and he was unhesitatingly answered by an old and + trustworthy man, that with the chaffinch the males are in large + excess; he thought as high as 2 males to 1 female, or at least + as high as 5 to 3.[382] The males of the blackbird, he likewise + maintained, were by far the most numerous, whether caught by + traps or by netting at night. These statements may apparently + be trusted, because the same man said that the sexes are about + equal with the lark, the twite (_Linaria montana_), and + goldfinch. On the other hand he is certain that with the common + linnet, the females preponderate greatly, but unequally during + different years; during some years he has found the females to + the males as four to one. It should, however, be borne in mind, + that the chief season for catching birds does not begin till + September, so that with some species partial migrations may + have begun, and the flocks at this period often consist of hens + alone. Mr. Salvin paid particular attention to the sexes of the + humming-birds in Central America, and he is convinced that with + most of the species the males are in excess; thus one year he + procured 204 specimens belonging to ten species, and these + consisted of 166 males and of 38 females. With two other + species the females were in excess: but the proportions + apparently vary either during different seasons or in different + localities; for on one occasion the males of _Campylopterus + hemileucurus_ were to the females as five to two, and on + another occasion[383] in exactly the reversed ratio. As bearing + on this latter point, I may add, that Mr. Powys found in Corfu + and Epirus the sexes of the chaffinch keeping apart, and "the + females by far the most numerous;" whilst in Palestine Mr. + Tristram found "the male flocks appearing greatly to exceed the + female in number."[384] So again with the _Quiscalus major_, + Mr. G. Taylor[385] says, that in Florida there were "very few + females in proportion to the males," whilst in Honduras the + proportion was the other way, the species there having the + character of a polygamist. + + + FISH. + + With Fish the proportional numbers of the sexes can be + ascertained only by catching them in the adult or nearly adult + state; and there are many difficulties in arriving at any just + conclusion.[386] Infertile females might readily be mistaken + for males, as Dr. Gnther has remarked to me in regard to + trout. With some species the males are believed to die soon + after fertilising the ova. With many species the males are of + much smaller size than the females, so that a large number of + males would escape from the same net by which the females were + caught. M. Carbonnier,[387] who has especially attended to the + natural history of the pike (_Esox lucius_) states that many + males, owing to their small size, are devoured by the larger + females; and he believes that the males of almost all fish are + exposed from the same cause to greater danger than the females. + Nevertheless in the few cases in which the proportional numbers + have been actually observed, the males appear to be largely in + excess. Thus Mr. R. Buist, the superintendent of the + Stormontfield experiments, says that in 1865, out of 70 salmon + first landed for the purpose of obtaining the ova, upwards of + 60 were males. In 1867 he again "calls attention to the vast + disproportion of the males to the females. We had at the outset + at least ten males to one female." Afterwards sufficient + females for obtaining ova were procured. He adds, "from the + great proportion of the males, they are constantly fighting and + tearing each other on the spawning-beds."[388] This + disproportion, no doubt, can be accounted for in part, but + whether wholly is very doubtful, by the males ascending the + rivers before the females. Mr. F. Buckland remarks in regard to + trout, that "it is a curious fact that the males preponderate + very largely in number over the females. It _invariably_ + happens that when the first rush of fish is made to the net, + there will be at least seven or eight males to one female found + captive. I cannot quite account for this; either the males are + more numerous than the females, or the latter seek safety by + concealment rather than flight." He then adds, that by + carefully searching the banks, sufficient females for obtaining + ova can be found.[389] Mr. H. Lee informs me that out of 212 + trout, taken for this purpose in Lord Portsmouth's park, 150 + were males and 62 females. + + With the Cyprinid the males likewise seem to be in excess; but + several members of this Family, viz., the carp, tench, bream + and minnow, appear regularly to follow the practice, rare in + the animal kingdom, of polyandry; for the female whilst + spawning is always attended by two males, one on each side, and + in the case of the bream by three or four males. This fact is + so well known, that it is always recommended to stock a pond + with two male tenches to one female, or at least with three + males to two females. With the minnow, an excellent observer + states, that on the spawning-beds the males are ten times as + numerous as the females; when a female comes amongst the males, + "she is immediately pressed closely by a male on each side; and + when they have been in that situation for a time, are + superseded by other two males."[390] + + + INSECTS. + + In this class, the Lepidoptera alone afford the means of + judging of the proportional numbers of the sexes; for they have + been collected with special care by many good observers, and + have been largely bred from the egg or caterpillar state. I had + hoped that some breeders of silk-moths might have kept an exact + record, but after writing to France and Italy, and consulting + various treatises, I cannot find that this has ever been done. + The general opinion appears to be that the sexes are nearly + equal, but in Italy as I hear from Professor Canestrini, many + breeders are convinced that the females are produced in excess. + The same naturalist, however, informs me, that in the two + yearly broods of the Ailanthus silk-moth (_Bombyx cynthia_), + the males greatly preponderate in the first, whilst in the + second the two sexes are nearly equal, or the females rather in + excess. + + In regard to Butterflies in a state of nature, several + observers have been much struck by the apparently enormous + preponderance of the males.[391] Thus Mr. Bates,[392] in + speaking of the species, no less than about a hundred in + number, which inhabit the Upper Amazons, says that the males + are much more numerous than the females, even in the proportion + of a hundred to one. In North America, Edwards, who had great + experience, estimates in the genus Papilio the males to the + females as four to one; and Mr. Walsh, who informed me of this + statement, says that with _P. turnus_ this is certainly the + case. In South Africa, Mr. R. Trimen found the males in excess + in 19 species;[393] and in one of these, which swarms in open + places, he estimated the number of males as fifty to one + female. With another species, in which the males are numerous + in certain localities, he collected during seven years only + five females. In the island of Bourbon, M. Maillard states that + the males of one species of Papilio are twenty times as + numerous as the females.[394] Mr. Trimen informs me that as far + as he has himself seen, or heard from others, it is rare for + the females of any butterfly to exceed in number the males; but + this is perhaps the case with three South African species. Mr. + Wallace[395] states that the females of _Ornithoptera + croesus_, in the Malay archipelago, are more common and more + easily caught than the males; but this is a rare butterfly. I + may here add, that in Hyperythra, a genus of moths, Guene + says, that from four to five females are sent in collections + from India for one male. + + When this subject of the proportional numbers of the sexes of + insects was brought before the Entomological Society,[396] it + was generally admitted that the males of most Lepidoptera, in + the adult or imago state, are caught in greater numbers than + the females; but this fact was attributed by various observers + to the more retiring habits of the females, and to the males + emerging earlier from the cocoon. This latter circumstance is + well known to occur with most Lepidoptera, as well as with + other insects. So that, as M. Personnat remarks, the males of + the domesticated _Bombyx yamamai_, are lost at the beginning of + the season, and the females at the end, from the want of + mates.[397] I cannot however persuade myself that these causes + suffice to explain the great excess of males in the cases, + above given, of butterflies which are extremely common in their + native countries. Mr. Stainton, who has paid such close + attention during many years to the smaller moths, informs me + that when he collected them in the imago state, he thought that + the males were ten times as numerous as the females, but that + since he has reared them on a large scale from the caterpillar + state, he is convinced that the females are the most numerous. + Several entomologists concur in this view. Mr. Doubleday, + however, and some others, take an opposite view, and are + convinced that they have reared from the egg and caterpillar + states a larger proportion of males than of females. + + Besides the more active habits of the males, their earlier + emergence from the cocoon, and their frequenting in some cases + more open stations, other causes may be assigned for an + apparent or real difference in the proportional numbers of the + sexes of Lepidoptera, when captured in the imago state, and + when reared from the egg or caterpillar state. It is believed + by many breeders in Italy, as I hear from Professor Canestrini, + that the female caterpillar of the silk-moth suffers more from + the recent disease than the male; and Dr. Staudinger informs me + that in rearing Lepidoptera more females die in the cocoon than + males. With many species the female caterpillar is larger than + the male, and a collector would naturally choose the finest + specimens, and thus unintentionally collect a larger number of + females. Three collectors have told me that this was their + practice; but Dr. Wallace is sure that most collectors take all + the specimens which they can find of the rarer kinds, which + alone are worth the trouble of rearing. Birds when surrounded + by caterpillars would probably devour the largest; and + Professor Canestrini informs me that in Italy some breeders + believe, though on insufficient evidence, that in the first + brood of the Ailanthus silk-moth, the wasps destroy a larger + number of the female than of the male caterpillars. Dr. Wallace + further remarks that female caterpillars, from being larger + than the males, require more time for their development and + consume more food and moisture; and thus they would be exposed + during a longer time to danger from ichneumons, birds, &c., and + in times of scarcity would perish in greater numbers. Hence it + appears quite possible that, in a state of nature, fewer female + Lepidoptera may reach maturity than males; and for our special + object we are concerned with the numbers at maturity, when the + sexes are ready to propagate their kind. + + The manner in which the males of certain moths congregate in + extraordinary numbers round a single female, apparently + indicates a great excess of males, though this fact may perhaps + be accounted for by the earlier emergence of the males from + their cocoons. Mr. Stainton informs me that from twelve to + twenty males may often be seen congregated round a female + _Elachista rufocinerea_. It is well known that if a virgin + _Lasiocampa quercus_ or _Saturnia carpini_ be exposed in a + cage, vast numbers of males collect round her, and if confined + in a room will even come down the chimney to her. Mr. + Doubleday believes that he has seen from fifty to a hundred + males of both these species attracted in the course of a single + day by a female under confinement. Mr. Trimen exposed in the + Isle of Wight a box in which a female of the Lasiocampa had + been confined on the previous day, and five males soon + endeavoured to gain admittance. M. Verreaux, in Australia, + having placed the female of a small Bombyx in a box in his + pocket, was followed by a crowd of males, so that about 200 + entered the house with him.[398] + + Mr. Doubleday has called my attention to Dr. Staudinger's[399] + list of Lepidoptera, which gives the prices of the males and + females of 300 species or well-marked varieties of + (Rhopalocera) butterflies. The prices for both sexes of the + very common species are of course the same; but with 114 of the + rarer species they differ; the males being in all cases, + excepting one, the cheapest. On an average of the prices of the + 113 species, the price of the male to that of the female is as + 100 to 149; and this apparently indicates that inversely the + males exceed the females in number in the same proportion. + About 2000 species or varieties of moths (Heterocera) are + catalogued, those with wingless females being here excluded on + account of the difference in habits of the two sexes: of these + 2000 species, 141 differ in price according to sex, the males + of 130 being cheaper, and the males of only 11 being dearer + than the females. The average price of the males of the 130 + species, to that of the females, is as 100 to 143. With respect + to the butterflies in this priced list, Mr. Doubleday thinks + (and no man in England has had more experience), that there is + nothing in the habits of the species which can account for the + difference in the prices of the two sexes, and that it can be + accounted for only by an excess in the numbers of the males. + But I am bound to add that Dr. Staudinger himself, as he + informs me, is of a different opinion. He thinks that the less + active habits of the females and the earlier emergence of the + males will account for his collectors securing a larger number + of males than of females, and consequently for the lower prices + of the former With respect to specimens reared from the + caterpillar-state, Dr. Staudinger believes, as previously + stated, that a greater number of females than of males die + under confinement in the cocoons. He adds that with certain + species one sex seems to preponderate over the other during + certain years. + + Of direct observations on the sexes of Lepidoptera, reared + either from eggs or caterpillars, I have received only the few + following cases:-- + + -------------------------------------------------------------------------------------------------------------------------------------- + Males. | Females. + The Rev. J. Hellins[400] of Exeter reared, during | + 1868, imagos of 73 species, which consisted of 153 | 137 + | + Mr. Albert Jones of Eltham reared, during 1868, | + imagos of 9 species, which, consisted of 159 | 126 + | + During 1869 he reared imagos from 4 species, | + consisting of 114 | 112 + | + Mr. Buckler of Emsworth, Hants, during 1869, | + reared imagos from 74 species, consisting of 180 | 169 + | + Dr. Wallace of Colchester reared from one brood | + of Bombyx cynthia 52 | 48 + | + Dr. Wallace raised, from cocoons of Bombyx | + Pernyi sent from China, during 1869 224 | 123 + | + Dr. Wallace raised, during 1868 and 1869, | + from two lots of cocoons of Bombyx yamamai 52 | 46 + --------------+------------ + Total 934 | 761 + ------------------------------------------------------------------------------------------------------------------+------------------ + + So that in these eight lots of cocoons and eggs, males were + produced in excess. Taken together the proportion of males is + as 122.7 to 100 females. But the numbers are hardly large + enough to be trustworthy. + + On the whole, from the above various sources of evidence, all + pointing to the same direction, I infer that with most species + of Lepidoptera, the males in the imago state generally exceed + the females in number, whatever the proportions may be at their + first emergence from the egg. + + With reference to the other Orders of insects, I have been able + to collect very little reliable information. With the + stag-beetle (_Lucanus cervus_) "the males appear to be much + more numerous than the females;" but when, as Cornelius + remarked during 1867, an unusual number of these beetles + appeared in one part of Germany, the females appeared to exceed + the males as six so one. With one of the Elaterid, the males + are said to be much more numerous than the females, and "two or + three are often found united with one female;"[401] so that here + polyandry seems to prevail. With Siagonium (Staphylinid), in + which the males are furnished with horns, "the females are far + more numerous than the opposite sex." Mr. Janson stated at the + Entomological Society that the females of the bark-feeding + _Tomicus villosus_ are so common as to be a plague, whilst the + males are so rare as to be hardly known. In other Orders, from + unknown causes, but apparently in some instances owing to + parthenogenesis, the males of certain species have never been + discovered or are excessively rare, as with several of the + Cynipid.[402] In all the gall-making Cynipid known to Mr. + Walsh, the females are four or five times as numerous as the + males; and so it is, as he informs me, with the gall-making + Cecidomyii (Diptera). With some common species of Saw-flies + (Tenthredin) Mr. F. Smith has reared hundreds of specimens + from larv of all sizes, but has never reared a single male: on + the other hand Curtis says,[403] that with certain species + (Athalia), bred by him, the males to the females were as six to + one; whilst exactly the reverse occurred with the mature + insects of the same species caught in the fields. With the + Neuroptera, Mr. Walsh states that in many, but by no means in + all, the species of the Odonatous groups (Ephemerina), there is + a great overplus of males: in the genus Hetrina, also, the + males are generally at least four times as numerous as the + females. In certain species in the genus Gomphus the males are + equally numerous, whilst in two other species, the females are + twice or thrice as numerous as the males. In some European + species of Psocus thousands of females may be collected without + a single male, whilst with other species of the same genus both + sexes are common.[404] In England, Mr. MacLachlan has captured + hundreds of the female _Apatania muliebris_, but has never seen + the male; and of _Boreus hyemalis_ only four or five males have + been here seen.[405] With most of these species (excepting, as + I have heard, with the Tenthredin) there is no reason to + suppose that the females are subject to parthenogenesis; and + thus we see how ignorant we are on the causes of the apparent + discrepancy in the proportional numbers of the two sexes. + + In the other Classes of the Articulata I have been able to + collect still less information. With Spiders, Mr. Blackwall, + who has carefully attended to this class during many years, + writes to me that the males from their more erratic habits are + more commonly seen, and therefore appear to be the more + numerous. This is actually the case with a few species; but he + mentions several species in six genera, in which the females + appear to be much more numerous than the males.[406] The small + size of the males in comparison with the females, which is + sometimes carried to an extreme degree, and their widely + different appearance, may account in some instances for their + rarity in collections.[407] + + Some of the lower Crustaceans are able to propagate their kind + asexually, and this will account for the extreme rarity of the + males. With some other forms (as with Tanais and Cypris) there + is reason to believe, as Fritz Mller informs me, that the male + is much shorter-lived than the female, which, supposing the two + sexes to be at first equal in number, would explain the + scarcity of the males. On the other hand this same naturalist + has invariably taken, on the shores of Brazil, far more males + than females of the Diastylid and of Cypridina; thus with a + species in the latter genus, 63 specimens caught the same day, + included 57 males; but he suggests that this preponderance may + be due to some unknown difference in the habits of the two + sexes. With one of the higher Brazilian crabs, namely a + Gelasimus, Fritz Mller found the males to be more numerous + than the females. The reverse seems to be the case, according + to the large experience of Mr. C. Spence Bate, with six common + British crabs, the names of which he has given me. + + +_On the Power of Natural Selection to regulate the proportional Numbers +of the Sexes, and General Fertility._--In some peculiar cases, an excess +in the number of one sex over the other might be a great advantage to a +species, as with the sterile females of social insects, or with those +animals in which more than one male is requisite to fertilise the +female, as with certain cirripedes and perhaps certain fishes. An +inequality between the sexes in these cases might have been acquired +through natural selection, but from their rarity they need not here be +further considered. In all ordinary cases an inequality would be no +advantage or disadvantage to certain individuals more than to others; +and therefore it could hardly have resulted from natural selection. We +must attribute the inequality to the direct action of those unknown +conditions, which with mankind lead to the males being born in a +somewhat larger excess in certain countries than in others, or which +cause the proportion between the sexes to differ slightly in legitimate +and illegitimate births. + +Let us now take the case of a species producing from the unknown causes +just alluded to, an excess of one sex--we will say of males--these being +superfluous and useless, or nearly useless. Could the sexes be equalised +through natural selection? We may feel sure, from all characters being +variable, that certain pairs would produce a somewhat less excess of +males over females than other pairs. The former, supposing the actual +number of the offspring to remain constant, would necessarily produce +more females, and would therefore be more productive. On the doctrine of +chances a greater number of the offspring of the more productive pairs +would survive; and these would inherit a tendency to procreate fewer +males and more females. Thus a tendency towards the equalisation of the +sexes would be brought about. But our supposed species would by this +process be rendered, as just remarked, more productive; and this would +in many cases be far from an advantage; for whenever the limit to the +numbers which exist, depends, not on destruction by enemies, but on the +amount of food, increased fertility will lead to severer competition and +to most of the survivors being badly fed. In this case, if the sexes +were equalised by an increase in the number of the females, a +simultaneous decrease in the total number of the offspring would be +beneficial, or even necessary, for the existence of the species; and +this, I believe, could be effected through natural selection in the +manner hereafter to be described. The same train of reasoning is +applicable in the above, as well as in the following case, if we assume +that females instead of males are produced in excess, for such females +from not uniting with males would be superfluous and useless. So it +would be with polygamous species, if we assume the excess of females to +be inordinately great. + +An excess of either sex, we will again say of the males, could, however, +apparently be eliminated through natural selection in another and +indirect manner, namely by an actual diminution of the males, without +any increase of the females, and consequently without any increase in +the productiveness of the species. From the variability of all +characters, we may feel assured that some pairs, inhabiting any +locality, would produce a rather smaller excess of superfluous males, +but an equal number of productive females. When the offspring from the +more and the less male-productive parents were all mingled together, +none would have any direct advantage over the others; but those that +produced few superfluous males would have one great indirect advantage, +namely that their ova or embryos would probably be larger and finer, or +their young better nurtured in the womb and afterwards. We see this +principle illustrated with plants; as those which bear a vast number of +seed produce small ones; whilst those which bear comparatively few +seeds, often produce large ones well-stocked with nutriment for the use +of the seedlings.[408] Hence the offspring of the parents which had +wasted least force in producing superfluous males would be the most +likely to survive, and would inherit the same tendency not to produce +superfluous males, whilst retaining their full fertility in the +production of females. So it would be with the converse case of the +female sex. Any slight excess, however, of either sex could hardly be +checked in so indirect a manner. Nor indeed has a considerable +inequality between the sexes been always prevented, as we have seen in +some of the cases given in the previous discussion. In these cases the +unknown causes which determine the sex of the embryo, and which under +certain conditions lead to the production of one sex in excess over the +other, have not been mastered by the survival of those varieties which +were subjected to the least waste of organised matter and force by the +production of superfluous individuals of either sex. Nevertheless we may +conclude that natural selection will always tend, though sometimes +inefficiently, to equalise the relative numbers of the two sexes. + +Having said this much on the equalisation of the sexes, it may be well +to add a few remarks on the regulation through natural selection of the +ordinary fertility of species. Mr. Herbert Spencer has shewn in an able +discussion[409] that with all organisms a ratio exists between what he +calls individuation and genesis; whence it follows that beings which +consume much matter or force in their growth, complicated structure or +activity, or which produce ova and embryos of large size, or which +expend much energy in nurturing their young, cannot be so productive as +beings of an opposite nature. Mr. Spencer further shews that minor +differences in fertility will be regulated through natural selection. +Thus the fertility of each species will tend to increase, from the more +fertile pairs producing a larger number of offspring, and these from +their mere number will have the best chance of surviving, and will +transmit their tendency to greater fertility. The only check to a +continued augmentation of fertility in each organism seems to be either +the expenditure of more power and the greater risks run by the parents +that produce a more numerous progeny, or the contingency of very +numerous eggs and young being produced of smaller size, or less +vigorous, or subsequently not so well nurtured. To strike a balance in +any case between the disadvantages which follow from the production of a +numerous progeny, and the advantages (such as the escape of at least +some individuals from various dangers) is quite beyond our power of +judgment. + +When an organism has once been rendered extremely fertile, how its +fertility can be reduced through natural selection is not so clear as +how this capacity was first acquired. Yet it is obvious that if +individuals of a species, from a decrease of their natural enemies, were +habitually reared in larger numbers than could be supported, all the +members would suffer. Nevertheless the offspring from the less fertile +parents would have no direct advantage over the offspring from the more +fertile parents, when all were mingled together in the same district. +All the individuals would mutually tend to starve each other. The +offspring indeed of the less fertile parents would lie under one great +disadvantage, for from the simple fact of being produced in smaller +numbers, they would be the most liable to extermination. Indirectly, +however, they would partake of one great advantage; for under the +supposed condition of severe competition, when all were pressed for +food, it is extremely probable that those individuals which from some +variation in their constitution produced fewer eggs or young, would +produce them of greater size or vigour; and the adults reared from such +eggs or young would manifestly have the best chance of surviving, and +would inherit a tendency towards lessened fertility. The parents, +moreover, which had to nourish or provide for fewer offspring would +themselves be exposed to a less severe strain in the struggle for +existence, and would have a better chance of surviving. By these steps, +and by no others as far as I can see, natural selection under the above +conditions of severe competition for food, would lead to the formation +of a new race less fertile, but better adapted for survival, than the +parent-race. + + + + +CHAPTER IX. + +SECONDARY SEXUAL CHARACTERS IN THE LOWER CLASSES OF THE ANIMAL KINGDOM. + + + These characters absent in the lowest classes--Brilliant + colours--Mollusca--Annelids--Crustacea, secondary sexual + characters strongly developed; dimorphism; colour; characters + not acquired before maturity--Spiders, sexual colours of; + stridulation by the males--Myriapoda. + + +In the lowest classes the two sexes are not rarely united in the same +individual, and therefore secondary sexual characters cannot be +developed. In many cases in which the two sexes are separate, both are +permanently attached to some support, and the one cannot search or +struggle for the other. Moreover it is almost certain that these animals +have too imperfect senses and much too low mental powers to feel mutual +rivalry, or to appreciate each other's beauty or other attractions. + +Hence in these classes, such as the Protozoa, Coelenterata, +Echinodermata, Scolecida, true secondary sexual characters do not occur; +and this fact agrees with the belief that such characters in the higher +classes have been acquired through sexual selection, which depends on +the will, desires, and choice of either sex. Nevertheless some few +apparent exceptions occur; thus, as I hear from Dr. Baird, the males of +certain Entozoa, or internal parasitic worms, differ slightly in colour +from the females; but we have no reason to suppose that such differences +have been augmented through sexual selection. + +Many of the lower animals, whether hermaphrodites or with the sexes +separate, are ornamented with the most brilliant tints, or are shaded +and striped in an elegant manner. This is the case with many corals and +sea-anemonies (Actini), with some jelly-fish (Medus, Porpita, &c.), +with some Planari, Ascidians, numerous Star-fishes, Echini, &c.; but we +may conclude from the reasons already indicated, namely the union of the +two sexes in some of these animals, the permanently affixed condition of +others, and the low mental powers of all, that such colours do not serve +as a sexual attraction, and have not been acquired through sexual +selection. With the higher animals the case is very different; for with +them when one sex is much more brilliantly or conspicuously coloured +than the other, and there is no difference in the habits of the two +sexes which will account for this difference, we have reason to believe +in the influence of sexual selection; and this belief is strongly +confirmed when the more ornamented individuals, which are almost always +the males, display their attractions before the other sex. We may also +extend this conclusion to both sexes, when coloured alike, if their +colours are plainly analogous to those of one sex alone in certain other +species of the same group. + +How, then, are we to account for the beautiful or even gorgeous colours +of many animals in the lowest classes? It appears very doubtful whether +such colours usually serve as a protection; but we are extremely liable +to err in regard to characters of all kinds in relation to protection, +as will be admitted by every one who has read Mr. Wallace's excellent +essay on this subject. It would not, for instance, at first occur to any +one that the perfect transparency of the Medus, or jelly-fishes, was of +the highest service to them as a protection; but when we are reminded +by Hckel that not only the medus but many floating mollusca, +crustaceans, and even small oceanic fishes partake of this same +glass-like structure, we can hardly doubt that they thus escape the +notice of pelagic birds and other enemies. + +Notwithstanding our ignorance how far colour in many cases serves as a +protection, the most probable view in regard to the splendid tints of +many of the lowest animals seems to be that their colours are the direct +result either of the chemical nature or the minute structure of their +tissues, independently of any benefit thus derived. Hardly any colour is +finer than that of arterial blood; but there is no reason to suppose +that the colour of the blood is in itself any advantage; and though it +adds to the beauty of the maiden's cheek, no one will pretend that it +has been acquired for this purpose. So again with many animals, +especially the lower ones, the bile is richly coloured; thus the extreme +beauty of the Eolid (naked sea-slugs) is chiefly due, as I am informed +by Mr. Hancock, to the biliary glands seen through the translucent +integuments; this beauty being probably of no service to these animals. +The tints of the decaying leaves in an American forest are described by +every one as gorgeous; yet no one supposes that these tints are of the +least advantage to the trees. Bearing in mind how many substances +closely analogous to natural organic compounds have been recently formed +by chemists, and which exhibit the most splendid colours, it would have +been a strange fact if substances similarly coloured had not often +originated, independently of any useful end being thus gained, in the +complex laboratory of living organisms. + + +_The sub-kingdom of the Mollusca._--Throughout this great division +(taken in its largest acceptation) of the animal kingdom, secondary +sexual characters, such as we are here considering, never, as far as I +can discover, occur. Nor could they be expected in the three lowest +classes, namely in the Ascidians, Polyzoa, and Brachiopods (constituting +the Molluscoida of Huxley), for most of these animals are permanently +affixed to a support or have their sexes united in the same individual. +In the Lamellibranchiata, or bivalve shells, hermaphroditism is not +rare. In the next higher class of the Gasteropoda, or marine univalve +shells, the sexes are either united or separate. But in this latter case +the males never possess special organs for finding, securing, or +charming the females, or for fighting with other males. The sole +external difference between the sexes consists, as I am informed by Mr. +Gwyn Jeffreys, in the shell sometimes differing a little in form; for +instance, the shell of the male periwinkle (_Littorina littorea_) is +narrower and has a more elongated spire than that of the female. But +differences of this nature, it may be presumed, are directly connected +with the act of reproduction or with the development of the ova. + +The Gasteropoda, though capable of locomotion and furnished with +imperfect eyes, do not appear to be endowed with sufficient mental +powers for the members of the same sex to struggle together in rivalry, +and thus to acquire secondary sexual characters. Nevertheless with the +pulmoniferous gasteropods, or land-snails, the pairing is preceded by +courtship; for these animals, though hermaphrodites, are compelled by +their structure to pair together. Agassiz remarks,[410] "Quiconque a eu +l'occasion d'observer les amours des limaons, ne saurait mettre en +doute la sduction dploye dans les mouvements et les allures qui +prparent et accomplissent le double embrassement de ces +hermaphrodites." These animals appear also susceptible of some degree of +permanent attachment: an accurate observer, Mr. Lonsdale, informs me +that he placed a pair of land-shells (_Helix pomatia_), one of which was +weakly, into a small and ill-provided garden. After a short time the +strong and healthy individual disappeared, and was traced by its track +of slime over a wall into an adjoining well-stocked garden. Mr. Lonsdale +concluded that it had deserted its sickly mate; but after an absence of +twenty-four hours it returned, and apparently communicated the result of +its successful exploration, for both then started along the same track +and disappeared over the wall. + +Even in the highest class of the Mollusca, namely the Cephalopoda or +cuttle-fishes, in which the sexes are separate, secondary sexual +characters of the kind which we are here considering, do not, as far as +I can discover, occur. This is a surprising circumstance, as these +animals possess highly-developed sense-organs and have considerable +mental powers, as will be admitted by every one who has watched their +artful endeavours to escape from an enemy.[411] Certain Cephalopoda, +however, are characterised by one extraordinary sexual character, +namely, that the male element collects within one of the arms or +tentacles, which is then cast off, and, clinging by its sucking-discs to +the female, lives for a time an independent life. So completely does the +cast-off arm resemble a separate animal, that it was described by Cuvier +as a parasitic worm under the name of Hectocotyle. But this marvellous +structure may be classed as a primary rather than as a secondary sexual +character. + +Although with the Mollusca sexual selection does not seem to have come +into play; yet many univalve and bivalve shells, such as volutes, cones, +scallops, &c., are beautifully coloured and shaped. The colours do not +appear in most cases to be of any use as a protection; they are probably +the direct result, as in the lowest classes, of the nature of the +tissues; the patterns and the sculpture of the shell depending on its +manner of growth. The amount of light seems to a certain extent to be +influential; for although, as repeatedly stated by Mr. Gwyn Jeffreys, +the shells of some species living at a profound depth are brightly +coloured, yet we generally see the lower surfaces and the parts covered +by the mantle less highly coloured than the upper and exposed +surfaces.[412] In some cases, as with shells living amongst corals or +brightly-tinted sea-weeds, the bright colours may serve as a protection. +But many of the nudibranch mollusca, or sea-slugs, are as beautifully +coloured as any shells, as may be seen in Messrs. Alder and Hancock's +magnificent work; and from information kindly given me by Mr. Hancock, +it is extremely doubtful whether these colours usually serve as a +protection. With some species this may be the case, as with one which +lives on the green leaves of alg, and is itself bright-green. But many +brightly-coloured, white or otherwise conspicuous species, do not seek +concealment; whilst again some equally conspicuous species, as well as +other dull-coloured kinds, live under stones and in dark recesses. So +that with these nudibranch molluscs, colour apparently does not stand in +any close relation to the nature of the places which they inhabit. + +These naked sea-slugs are hermaphrodites, yet they pair together, as do +land-snails, many of which have extremely pretty shells. It is +conceivable that two hermaphrodites, attracted by each others' greater +beauty, might unite and leave offspring which would inherit their +parents' greater beauty. But with such lowly-organised creatures this is +extremely improbable. Nor is it at all obvious how the offspring from +the more beautiful pairs of hermaphrodites would have any advantage, so +as to increase in numbers, over the offspring of the less beautiful, +unless indeed vigour and beauty generally coincided. We have not here a +number of males becoming mature before the females, and the more +beautiful ones selected by the more vigorous females. If, indeed, +brilliant colours were beneficial to an hermaphrodite animal in relation +to its general habits of life, the more brightly-tinted individuals +would succeed best and would increase in number; but this would be a +case of natural and not of sexual selection. + + +_Sub-kingdom of the Vermes or Annulosa_: Class, _Annelida (or +Sea-worms)_.--In this class, although the sexes (when separate) +sometimes differ from each other in characters of such importance that +they have been placed under distinct genera or even families, yet the +differences do not seem of the kind which can be safely attributed to +sexual selection. These animals, like those in the preceding classes, +apparently stand too low in the scale, for the individuals of either sex +to exert any choice in selecting a partner, or for the individuals of +the same sex to struggle together in rivalry. + + +_Sub-kingdom of the Arthropoda_: Class, _Crustacea_.--In this great +class we first meet with undoubted secondary sexual characters, often +developed in a remarkable manner. Unfortunately the habits of +crustaceans are very imperfectly known, and we cannot explain the uses +of many structures peculiar to one sex. With the lower parasitic species +the males are of small size, and they alone are furnished with perfect +swimming-legs, antenn and sense-organs; the females being destitute of +these organs, with their bodies often consisting of a mere distorted +mass. But these extraordinary differences between the two sexes are no +doubt related to their widely different habits of life, and consequently +do not concern us. In various crustaceans, belonging to distinct +families, the anterior antenn are furnished with peculiar thread-like +bodies, which are believed to act as smelling-organs, and these are much +more numerous in the males than in the females. As the males, without +any unusual development of their olfactory organs, would almost +certainly be able sooner or later to find the females, the increased +number of the smelling-threads has probably been acquired through sexual +selection, by the better provided males having been the most successful +in finding partners and in leaving offspring. Fritz Mller has described +a remarkable dimorphic species of Tanais, in which the male is +represented by two distinct forms, never graduating into each other. In +the one form the male is furnished with more numerous smelling-threads, +and in the other form with more powerful and more elongated chel or +pincers which serve to hold the female. Fritz Mller suggests that these +differences between the two male forms of the same species must have +originated in certain individuals having varied in the number of the +smelling-threads, whilst other individuals varied in the shape and size +of their chel; so that of the former, those which were best able to +find the female, and of the latter, those which were best able to hold +her when found, have left the greater number of progeny to inherit their +respective advantages.[413] + +[Illustration: Fig. 3. Labidocera Darwinii, (from Lubbock). + +_a._ Part of right-hand anterior antenna of male, forming a prehensile +organ. + +_b._ Posterior pair of thoracic legs of male. + +_c._ Ditto of female.] + +In some of the lower crustaceans, the right-hand anterior antenna of the +male differs greatly in structure from the left-hand one, the latter +resembling in its simple tapering joints the antenn of the female. In +the male the modified antenna is either swollen in the middle or +angularly bent, or converted (fig. 3) into an elegant, and sometimes +wonderfully complex, prehensile organ.[414] It serves, as I hear from +Sir J. Lubbock, to hold the female, and for this same purpose one of the +two posterior legs (_b_) on the same side of the body is converted into +a forceps. In another family the inferior or posterior antenn are +"curiously zigzagged" in the males alone. + +[Illustration: Fig. 4. Anterior part of body of Callianassa (from +Milne-Edwards), showing the unequal and differently-constructed right +and left-hand chel of the male. + +N.B.--The artist by mistake has reversed the drawing, and made the +left-hand chela the largest.] + +[Illustration: Fig. 5. Second leg of male Orchestia Tucuratinga (from +Fritz Mller).] + +[Illustration: Fig. 6. Ditto of female.] + +In the higher crustaceans the anterior legs form a pair of chel or +pincers, and these are generally larger in the male than in the female. +In many species the chel on the opposite sides of the body are of +unequal size, the right-hand one being, as I am informed by Mr. C. +Spence Bate, generally, though not invariably, the largest. This +inequality is often much greater in the male than in the female. The two +chel also often differ in structure (figs. 4, 5, and 6), the smaller one +resembling those of the female. What advantage is gained by their +inequality in size on the opposite sides of the body, and by the +inequality being much greater in the male than in the female; and why, +when they are of equal size, both are often much larger in the male than +in the female, is not known. The chel are sometimes of such length and +size that they cannot possibly be used, as I hear from Mr. Spence Bate, +for carrying food to the mouth. In the males of certain freshwater +prawns (Palmon) the right leg is actually longer than the whole +body.[415] It is probable that the great size of one leg with its chel +may aid the male in fighting with his rivals; but this use will not +account for their inequality in the female on the opposite sides +of the body. In Gelasimus, according to a statement quoted by +Milne-Edwards,[416] the male and female live in the same burrow, which +is worth notice, as shewing that they pair, and the male closes the +mouth of the burrow with one of its chel, which is enormously +developed; so that here it indirectly serves as a means of defence. +Their main use, however, probably is to seize and to secure the female, +and this in some instances, as with Gammarus, is known to be the case. +The sexes, however, of the common shore-crab (_Carcinus mnas_), as Mr. +Spence Bate informs me, unite directly after the female has moulted her +hard shell, and when she is so soft that she would be injured if seized +by the strong pincers of the male; but as she is caught and carried +about by the male previously to the act of moulting, she could then be +seized with impunity. + +Fritz Mller states that certain species of Melita are distinguished +from all other amphipods by the females having "the coxal lamell of the +penultimate pair of feet produced into hook-like processes, of which the +males lay hold with the hands of the first pair." The development of +these hook-like processes probably resulted from those females which +were the most securely held during the act of reproduction, having left +the largest number of offspring. Another Brazilian amphipod (_Orchestia +Darwinii_, fig. 7) is described by Fritz Mller, as presenting a case of +dimorphism, like that of Tanais; for there are two male forms, which +differ in the structure of their chel.[417] As chel of either shape +would certainly have sufficed to hold the female, for both are now used +for this purpose, the two male forms probably originated, by some having +varied in one manner and some in another; both forms having derived +certain special, but nearly equal advantages, from their differently +shaped organs. + +It is not known that male crustaceans fight together for the possession +of the females, but this is probable; for with most animals when the +male is larger than the female, he seems to have acquired his greater +size by having conquered during many generations other males. Now Mr. +Spence Bate informs me that in most of the crustacean orders, especially +in the highest or the Brachyura, the male is larger than the female; the +parasitic genera, however, in which the sexes follow different habits of +life, and most of the Entomostraca must be excepted. The chel of many +crustaceans are weapons well adapted for fighting. Thus a Devil-crab +(_Portunus puber_) was seen by a son of Mr. Bate fighting with a +_Carcinus mnas_, and the latter was soon thrown on its back, and had +every limb torn from its body. When several males of a Brazilian +Gelasimus, a species furnished with immense pincers, were placed +together by Fritz Mller in a glass vessel, they mutilated and killed +each other. Mr. Bate put a large male _Carcinus mnas_ into a pan of +water, inhabited by a female paired with a smaller male; the latter was +soon dispossessed, but, as Mr. Bate adds, "if they fought, the victory +was a bloodless one, for I saw no wounds." This same naturalist +separated a male sand-skipper (so common on our sea-shores), _Gammarus +marinus_, from its female, both of which were imprisoned in the same +vessel with many individuals of the same species. The female being thus +divorced joined her comrades. After an interval the male was again put +into the same vessel and he then, after swimming about for a time, +dashed into the crowd, and without any fighting at once took away his +wife. This fact shews that in the Amphipoda, an order low in the scale, +the males and females recognise each other, and are mutually attached. + +[Illustration: Fig. 7. Orchestia Darwinii (from Fritz Mller), showing +the differently-constructed chel of the two male forms.] + +The mental powers of the Crustacea are probably higher than might have +been expected. Any one who has tried to catch one of the shore-crabs, so +numerous on many tropical coasts, will have perceived how wary and alert +they are. There is a large crab (_Birgus latro_), found on coral +islands, which makes at the bottom of a deep burrow a thick bed of the +picked fibres of the cocoa-nut. It feeds on the fallen fruit of this +tree by tearing off the husk, fibre by fibre; and it always begins at +that end where the three eye-like depressions are situated. It then +breaks through one of these eyes by hammering with its heavy front +pincers, and turning round, extracts the albuminous core with its narrow +posterior pincers. But these actions are probably instinctive, so that +they would be performed as well by a young as by an old animal. The +following case, however, can hardly be so considered: a trustworthy +naturalist, Mr. Gardner,[418] whilst watching a shore-crab (Gelasimus) +making its burrow, threw some shells towards the hole. One rolled in, +and three other shells remained within a few inches of the mouth. In +about five minutes the crab brought out the shell which had fallen in, +and carried it away to the distance of a foot; it then saw the three +other shells lying near, and evidently thinking that they might likewise +roll in, carried them to the spot where it had laid the first. It would, +I think, be difficult to distinguish this act from one performed by man +by the aid of reason. + +With respect to colour which so often differs in the two sexes of +animals belonging to the higher classes, Mr. Spence Bate does not know +of any well-marked instances with our British crustaceans. In some +cases, however, the male and female differ slightly in tint, but Mr. +Bate thinks not more than may be accounted for by their different habits +of life, such as by the male wandering more about and being thus more +exposed to the light. In a curious Bornean crab, which inhabits sponges, +Mr. Bate could always distinguish the sexes by the male not having the +epidermis so much rubbed off. Dr. Power tried to distinguish by colour +the sexes of the species which inhabit the Mauritius, but always failed, +except with one species of Squilla, probably the _S. stylifera_, the +male of which is described as being "of a beautiful blueish-green," with +some of the appendages cherry-red, whilst the female is clouded with +brown and grey, "with the red about her much less vivid than in the +male."[419] In this case, we may suspect the agency of sexual selection. +With Saphirina (an oceanic genus of Entomostraca, and therefore low in +the scale) the males are furnished with minute shields or cell-like +bodies, which exhibit beautiful changing colours; these being absent in +the females, and in the case of one species in both sexes.[420] It +would, however, be extremely rash to conclude that these curious organs +serve merely to attract the females. In the female of a Brazilian +species of Gelasimus, the whole body, as I am informed by Fritz Mller, +is of a nearly uniform greyish-brown. In the male the posterior part of +the cephalo-thorax is pure white, with the anterior part of a rich +green, shading into dark brown; and it is remarkable that these colours +are liable to change in the course of a few minutes--the white becoming +dirty grey or even black, the green "losing much of its brilliancy." The +males apparently are much more numerous than the females. It deserves +especial notice that they do not acquire their bright colours until they +become mature. They differ also from the females in the larger size of +their chel. In some species of the genus, probably in all, the sexes +pair and inhabit the same burrow. They are also, as we have seen, highly +intelligent animals. From these various considerations it seems highly +probable that the male in this species has become gaily ornamented in +order to attract or excite the female. + +It has just been stated that the male Gelasimus does not acquire his +conspicuous colours until mature and nearly ready to breed. This seems +the general rule in the whole class with the many remarkable differences +in structure between the two sexes. We shall hereafter find the same law +prevailing throughout the great sub-kingdom of the Vertebrata, and in +all cases it is eminently distinctive of characters which have been +acquired through sexual selection. Fritz Mller[421] gives some +striking instances of this law; thus the male sand-hopper (Orchestia) +does not acquire his large claspers, which are very differently +constructed from those of the female, until nearly full-grown; whilst +young his claspers resemble those of the female. Thus, again, the male +Brachyscelus possesses, like all other amphipods, a pair of posterior +antenn; the female, and this is a most extraordinary circumstance, is +destitute of them, and so is the male as long as he remains immature. + + +Class, _Arachnida_ (Spiders).--The males are often darker, but sometimes +lighter than the females, as may be seen in Mr. Blackwall's magnificent +work.[422] In some species the sexes differ conspicuously from each +other in colour; thus the female of _Sparassus smaragdulus_ is +dullish-green; whilst the adult male has the abdomen of a fine yellow, +with three longitudinal stripes of rich red. In some species of Thomisus +the two sexes closely resemble each other; in others they differ much; +thus in _T. citreus_ the legs and body of the female are pale-yellow or +green, whilst the front legs of the male are reddish-brown: in _T. +floricolens_, the legs of the female are pale-green, those of the male +being ringed in a conspicuous manner with various tints. Numerous +analogous cases could be given in the genera Epeira, Nephila, +Philodromus, Theridion, Linyphia, &c. It is often difficult to say which +of the two sexes departs most from the ordinary coloration of the genus +to which the species belong; but Mr. Blackwall thinks that, as a +general rule, it is the male. Both sexes whilst young, as I am informed +by the same author, usually resemble each other; and both often undergo +great changes in colour during their successive moults before arriving +at maturity. In other cases the male alone appears to change colour. +Thus the male of the above-mentioned brightly-coloured Sparassus at +first resembles the female and acquires his peculiar tints only when +nearly adult. Spiders are possessed of acute senses, and exhibit much +intelligence. The females often shew, as is well known, the strongest +affection for their eggs, which they carry about enveloped in a silken +web. On the whole it appears probable that well-marked differences in +colour between the sexes have generally resulted from sexual selection, +either on the male or female side. But doubts may be entertained on this +head from the extreme variability in colour of some species, for +instance of _Theridion lineatum_, the sexes of which differ when adult; +this great variability indicates that their colours have not been +subjected to any form of selection. + +Mr. Blackwall does not remember to have seen the males of any species +fighting together for the possession of the female. Nor, judging from +analogy, is this probable; for the males are generally much smaller than +the females, sometimes to an extraordinary degree.[423] Had the males +been in the habit of fighting together, they would, it is probable, have +gradually acquired greater size and strength. Mr. Blackwall has +sometimes seen two or more males on the same web with a single female; +but their courtship is too tedious and prolonged an affair to be easily +observed. The male is extremely cautious in making his advances, as the +female carries her coyness to a dangerous pitch. De Geer saw a male that +"in the midst of his preparatory caresses was seized by the object of +his attentions, enveloped by her in a web and then devoured, a sight +which, as he adds, filled him with horror and indignation."[424] + +Westring has made the interesting discovery that the males of several +species of Theridion[425] have the power of making a stridulating sound +(like that made by many beetles and other insects, but feebler), whilst +the females are quite mute. The apparatus consists of a serrated ridge +at the base of the abdomen, against which the hard hinder part of the +thorax is rubbed; and of this structure not a trace could be detected in +the females. From the analogy of the Orthoptera and Homoptera, to be +described in the next chapter, we may feel almost sure that the +stridulation serves, as Westring remarks, either to call or to excite +the female; and this is the first case in the ascending scale of the +animal kingdom, known to me, of sounds emitted for this purpose. + + +Class, _Myriapoda_.--In neither of the two orders in this class, +including the millipedes and centipedes, can I find any well-marked +instances of sexual differences such as more particularly concern us. In +_Glomeris limbata_, however, and perhaps in some few other species, the +males differ slightly in colour from the females; but this Glomeris is a +highly variable species. In the males of the Diplopoda, the legs +belonging to one of the anterior segments of the body, or to the +posterior segment, are modified into prehensile hooks which serve to +secure the female. In some species of Iulus the tarsi of the male are +furnished with membranous suckers for the same purpose. It is a much +more unusual circumstance, as we shall see when we treat of Insects, +that it is the female in Lithobius which is furnished with prehensile +appendages at the extremity of the body for holding the male.[426] + + + + +CHAPTER X. + +SECONDARY SEXUAL CHARACTERS OF INSECTS. + + + Diversified structures possessed by the males for seizing the + females--Differences between the sexes, of which the meaning is + not understood--Difference in size between the + sexes--Thysanura--Diptera--Hemiptera--Homoptera, musical powers + possessed by the males alone--Orthoptera, musical instruments + of the males, much diversified in structure; pugnacity; + colours--Neuroptera, sexual differences in colour--Hymenoptera, + pugnacity and colours--Coleoptera, colours; furnished with + great horns, apparently as an ornament; battles; stridulating + organs generally common to both sexes. + + +In the immense class of insects the sexes sometimes differ in their +organs for locomotion, and often in their sense-organs, as in the +pectinated and beautifully plumose antenn of the males of many species. +In one of the Ephemer, namely Chloon, the male has great pillared +eyes, of which the female is entirely destitute.[427] The ocelli are +absent in the females of certain other insects, as in the Mutillid, +which are likewise destitute of wings. But we are chiefly concerned with +structures by which one male is enabled to conquer another, either in +battle or courtship, through his strength, pugnacity, ornaments, or +music. The innumerable contrivances, therefore, by which the male is +able to seize the female, may be briefly passed over. Besides the +complex structures at the apex of the abdomen, which ought perhaps to be +ranked as primary organs,[428] "it is astonishing," as Mr. B. D. +Walsh[429] has remarked, "how many different organs are worked in by +nature, for the seemingly insignificant object of enabling the male to +grasp the female firmly." The mandibles or jaws are sometimes used for +this purpose; thus the male _Corydalis cornutus_ (a neuropterous insect +in some degree allied to the Dragon-flies, &c.) has immense curved jaws, +many times longer than those of the female; and they are smooth instead +of being toothed, by which means he is enabled to seize her without +injury.[430] One of the stag-beetles of North America (_Lucanus +elaphus_) uses his jaws, which are much larger than those of the female, +for the same purpose, but probably likewise for fighting. In one of the +sand-wasps (_Ammophila_) the jaws in the two sexes are closely alike, +but are used for widely different purposes; the males, as Professor +Westwood observes, "are exceedingly ardent, seizing their partners round +the neck with their sickle-shaped jaws;"[431] whilst the females use +these organs for burrowing in sand-banks and making their nests. + +[Illustration: Fig. 8. Crabro cribrarius. Upper figure, male: lower +figure, female.] + +The tarsi of the front-legs are dilated in many male beetles, or are +furnished with broad cushions of hairs; and in many genera of +water-beetles they are armed with a round flat sucker, so that the male +may adhere to the slippery body of the female. It is a much more unusual +circumstance that the females of some water-beetles (Dytiscus) have +their elytra deeply grooved, and in _Acilius sulcatus_ thickly set with +hairs, as an aid to the male. The females of some other water-beetles +(Hydroporus) have their elytra punctured for the same object.[432] In +the male of _Crabro cribrarius_ (fig. 8.), it is the tibia which is +dilated into a broad horny plate, with minute membraneous dots, giving +to it a singular appearance like that of a riddle.[433] In the male of +Penthe (a genus of beetles) a few of the middle joints of the antenn +are dilated and furnished on the inferior surface with cushions of +hair, exactly like those on the tarsi of the Carabid, "and obviously +for the same end." In male dragon-flies, "the appendages at the tip of +the tail are modified in an almost infinite variety of curious patterns +to enable them to embrace the neck of the female." Lastly in the males +of many insects, the legs are furnished with peculiar spines, knobs or +spurs; or the whole leg is bowed or thickened, but this is by no means +invariably a sexual character; or one pair, or all three pairs are +elongated, sometimes to an extravagant length.[434] + +[Illustration: Fig. 9. Taphroderes distortus (much enlarged). Upper +figure, male; lower figure, female.] + +In all the orders, the sexes of many species present differences, of +which the meaning is not understood. One curious case is that of a +beetle (fig. 9), the male of which has the left mandible much enlarged; +so that the mouth is greatly distorted. In another Carabidous beetle, +the Eurygnathus,[435] we have the unique case, as far as known to Mr. +Wollaston, of the head of the female being much broader and larger, +though in a variable degree, than that of the male. Any number of such +cases could be given. They abound in the Lepidoptera: one of the most +extraordinary is that certain male butterflies have their fore-legs more +or less atrophied, with the tibi and tarsi reduced to mere rudimentary +knobs. The wings, also, in the two sexes often differ in neuration,[436] +and sometimes considerably in outline, as in the _Aricoris epitus_, +which was shown to me in the British Museum by Mr. A. Butler. The males +of certain South American butterflies have tufts of hair on the margins +of the wings, and horny excrescences on the discs of the posterior +pair.[437] In several British butterflies, the males alone, as shewn by +Mr. Wonfor, are in parts clothed with peculiar scales. + +The purpose of the luminosity in the female glow-worm is likewise not +understood; for it is very doubtful whether the primary use of the light +is to guide the male to the female. It is no serious objection to this +latter belief that the males emit a feeble light; for secondary sexual +characters proper to one sex are often developed in a slight degree in +the other sex. It is a more valid objection that the larv shine, and in +some species brilliantly: Fritz Mller informs me that the most luminous +insect which he ever beheld in Brazil, was the larva of some beetle. +Both sexes of certain luminous species of Elater emit light. Kirby and +Spence suspect that the phosphorescence serves to frighten and drive +away enemies. + + +_Difference in Size between the Sexes._--With insects of all kinds the +males are commonly smaller than the females;[438] and this difference +can often be detected even in the larval state. So considerable is the +difference between the male and female cocoons of the silk-moth +(_Bombyx mori_), that in France they are separated by a particular mode +of weighing.[439] In the lower classes of the animal kingdom, the +greater size of the females seems generally to depend on their +developing an enormous number of ova; and this may to a certain extent +hold good with insects. But Dr. Wallace has suggested a much more +probable explanation. He finds, after carefully attending to the +development of the caterpillars of _Bombyx cynthia_ and _yamamai_, and +especially of some dwarfed caterpillars reared from a second brood on +unnatural food, "that in proportion as the individual moth is finer, so +is the time required for its metamorphosis longer; and for this reason +the female, which is the larger and heavier insect, from having to carry +her numerous eggs, will be preceded by the male, which is smaller and +has less to mature."[440] Now as most insects are short-lived, and as +they are exposed to many dangers, it would manifestly be advantageous to +the female to be impregnated as soon as possible. This end would be +gained by the males being first matured in large numbers ready for the +advent of the females; and this again would naturally follow, as Mr. A. +E. Wallace has remarked,[441] through natural selection; for the smaller +males would be first matured, and thus would procreate a large number of +offspring which would inherit the reduced size of their male parents, +whilst the larger males from being matured later would leave fewer +offspring. + +There are, however, exceptions to the rule of male insects being smaller +than the females; and some of these exceptions are intelligible. Size +and strength would be an advantage to the males, which fight for the +possession of the female; and in these cases the males, as with the +stag-beetle (Lucanus), are larger than the females. There are, however, +other beetles which are not known to fight together, of which the males +exceed the females in size; and the meaning of this fact is not known; +but in some of these cases, as with the huge Dynastes and Megasoma, we +can at least see that there would be no necessity for the males to be +smaller than the females, in order to be matured before them, for these +beetles are not short-lived, and there would be ample time for the +pairing of the sexes. So, again, male dragon-flies (Libellulid) are +sometimes sensibly larger, and never smaller, than the females;[442] and +they do not, as Mr. MacLachlan believes, generally pair with the +females, until a week or fortnight has elapsed, and until they have +assumed their proper masculine colours. But the most curious case, +shewing on what complex and easily-overlooked relations, so trifling a +character as a difference in size between the sexes may depend, is that +of the aculeate Hymenoptera; for Mr. F. Smith informs me that throughout +nearly the whole of this large group the males, in accordance with the +general rule, are smaller than the females and emerge about a week +before them; but amongst the Bees, the males of _Apis mellifica_, +_Anthidium manicatum_ and _Anthophora acervorum_, and amongst the +Fossores, the males of the _Methoca ichneumonides_, are larger than the +females. The explanation of this anomaly is that a marriage-flight is +absolutely necessary with these species, and the males require great +strength and size in order to carry the females through the air. +Increased size has here been acquired in opposition to the usual +relation between size and the period of development, for the males, +though larger, emerge before the smaller females. + +We will now review the several Orders, selecting such facts as more +particularly concern us. The Lepidoptera (Butterflies and Moths) will be +retained for a separate chapter. + + +Order, _Thysanura_.--The members of this Order are lowly organised for +their class. They are wingless, dull-coloured, minute insects, with +ugly, almost misshapen heads and bodies. The sexes do not differ; but +they offer one interesting fact, by showing that the males pay sedulous +court to their females even low down in the animal scale. Sir J. +Lubbock[443] in describing the _Smynthurus luteus_, says: "it is very +amusing to see these little creatures coquetting together. The male, +which is much smaller than the female, runs round her, and they butt one +another, standing face to face, and moving backward and forward like two +playful lambs. Then the female pretends to run away and the male runs +after her with a queer appearance of anger, gets in front and stands +facing her again; then she turns coyly round, but he, quicker and more +active, scuttles round too, and seems to whip her with his antenn; then +for a bit they stand face to face, play with their antenn, and seem to +be all in all to one another." + + +Order, _Diptera_ (Flies).--The sexes differ little in colour. The +greatest difference, known to Mr. F. Walker, is in the genus Bibio, in +which the males are blackish or quite black, and the females obscure +brownish-orange. The genus Elaphomyia, discovered by Mr. Wallace[444] in +New Guinea, is highly remarkable, as the males are furnished with horns, +of which the females are quite destitute. The horns spring from beneath +the eyes, and curiously resemble those of stags, being either branched +or palmated. They equal in length the whole of the body in one of the +species. They might be thought to serve for fighting, but as in one +species they are of a beautiful pink colour, edged with black, with a +pale central stripe, and as these insects have altogether a very elegant +appearance, it is perhaps more probable that the horns serve as +ornaments. That the males of some Diptera fight together is certain; for +Prof. Westwood[445] has several times seen this with some species of +Tipula or Harry-long-legs. Many observers believe that when gnats +(Culicid) dance in the air in a body, alternately rising and falling, +the males are courting the females. The mental faculties of the Diptera +are probably fairly well developed, for their nervous system is more +highly developed than in most other Orders of insects.[446] + + +Order, _Hemiptera_ (Field-Bugs).--Mr. J. W. Douglas, who has +particularly attended to the British species, has kindly given me an +account of their sexual differences. The males of some species are +furnished with wings, whilst the females are wingless; the sexes differ +in the form of the body and elytra; in the second joints of their +antenn and in their tarsi; but as the signification of these +differences is quite unknown, they may be here passed over. The females +are generally larger and more robust than the males. With British, and, +as far as Mr. Douglas knows, with exotic species, the sexes do not +commonly differ much in colour; but in about six British species the +male is considerably darker than the female, and in about four other +species the female is darker than the male. Both sexes of some species +are beautifully marked with vermilion and black. It is doubtful whether +these colours serve as a protection. If in any species the males had +differed from the females in an analogous manner, we might have been +justified in attributing such conspicuous colours to sexual selection +with transference to both sexes. + +Some species of Reduvid make a stridulating noise; and, in the case of +_Pirates stridulus_, this is said[447] to be effected by the movement of +the neck within the pro-thoracic cavity. According to Westring, +_Reduvius personatus_ also stridulates. But I have not been able to +learn any particulars about these insects; nor have I any reason to +suppose that they differ sexually in this respect. + + +Order, _Homoptera._--Every one who has wandered in a tropical forest +must have been astonished at the din made by the male Cicad. The +females are mute; as the Grecian poet Xenarchus says, "Happy the Cicadas +live, since they all have voiceless wives." The noise thus made could be +plainly heard on board the "Beagle," when anchored at a quarter of a +mile from the shore of Brazil; and Captain Hancock says it can be heard +at the distance of a mile. The Greeks formerly kept, and the Chinese now +keep, these insects in cages for the sake of their song, so that it +must be pleasing to the ears of some men.[448] The Cicadid usually sing +during the day; whilst the Fulgorid appear to be night-songsters. The +sound, according to Landois,[449] who has recently studied the subject, +is produced by the vibration of the lips of the spiracles, which are set +into motion by a current of air emitted from the trache. It is +increased by a wonderfully complex resounding apparatus, consisting of +two cavities covered by scales. Hence the sound may truly be called a +voice. In the female the musical apparatus is present, but very much +less developed than in the male, and is never used for producing sound. + +With respect to the object of the music, Dr. Hartman in speaking of the +_Cicada septemdecim_ of the United States, says,[450] "the drums are now +(June 6th and 7th, 1851) heard in all directions. This I believe to be +the marital summons from the males. Standing in thick chestnut sprouts +about as high as my head, where hundreds were around me, I observed the +females coming around the drumming males." He adds, "this season (Aug. +1868) a dwarf pear-tree in my garden produced about fifty larv of _Cic. +pruinosa_; and I several times noticed the females to alight near a male +while he was uttering his clanging notes." Fritz Mller writes to me +from S. Brazil that he has often listened to a musical contest between +two or three males of a Cicada, having a particularly loud voice, and +seated at a considerable distance from each other. As soon as the first +had finished his song, a second immediately began; and after he had +concluded, another began, and so on. As there is so much rivalry between +the males, it is probable that the females not only discover them by the +sounds emitted, but that, like female birds, they are excited or allured +by the male with the most attractive voice. + +I have not found any well-marked cases of ornamental differences between +the sexes of the Homoptera. Mr. Douglas informs me that there are three +British species, in which the male is black or marked with black bands, +whilst the females are pale-coloured or obscure. + + +Order, _Orthoptera_.--The males in the three saltatorial families +belonging to this Order are remarkable for their musical powers, namely +the Achetid or crickets, the Locustid for which there is no exact +equivalent name in English, and the Acridiid or grasshoppers. The +stridulation produced by some of the Locustid is so loud that it can be +heard during the night at the distance of a mile;[451] and that made by +certain species is not unmusical even to the human ear, so that the +Indians on the Amazons keep them in wicker cages. All observers agree +that the sounds serve either to call or excite the mute females. But it +has been noticed[452] that the male migratory locust of Russia (one of +the Acridiid) whilst coupled with the female, stridulates from anger or +jealousy when approached by another male. The house-cricket when +surprised at night uses its voice to warn its fellows.[453] In North +America the Katy-did (_Platyphyllum concavum_, one of the Locustid) is +described[454] as mounting on the upper branches of a tree, and in the +evening beginning "his noisy babble, while rival notes issue from the +neighbouring trees, and the groves resound with the call of +_Katy-did-she-did_, the live-long night." Mr. Bates, in speaking of the +European field-cricket (one of the Achetid), says, "the male has been +observed to place itself in the evening at the entrance of its burrow, +and stridulate until a female approaches, when the louder notes are +succeeded by a more subdued tone, whilst the successful musician +caresses with his antenn the mate he has won."[455] Dr. Scudder was +able to excite one of these insects to answer him, by rubbing on a file +with a quill.[456] In both sexes a remarkable auditory apparatus has +been discovered by Von Siebold, situated in the front legs.[457] + +[Illustration: Fig. 10. Gryllus campestris (from Landois). + +Right-hand figure, under side of part of the wing-nervure, much +magnified, showing the teeth, _st_. + +Left-hand figure, upper surface of wing-cover, with the projecting, +smooth nervure, _r._, across which the teeth (_st_) are scraped.] + +In the three Families the sounds are differently produced. In the males +Of the Achetid both wing-covers have the same structure; and this in +the field-cricket (_Gryllus campestris_, fig. 10) consists, as +described by Landois,[458] of from 131 to 138 sharp, transverse ridges +or teeth (_st_) on the under side of one of the nervures of the +wing-cover. This toothed nervure is rapidly scraped across a projecting, +smooth, hard nervure (_r_) on the upper surface of the opposite wing. +First one wing is rubbed over the other, and then the movement is +reversed. Both wings are raised a little at the same time, so as to +increase the resonance. In some species the wing-covers of the males are +furnished at the base with a talc-like plate.[459] I have here given a +drawing (fig. 11) of the teeth on the under side of the nervure of +another species of Gryllus, viz. _G. domesticus_. + +[Illustration: Fig. 11. Teeth of Nervure of Gryllus domesticus (from +Landois).] + +In the Locustid the opposite wing-covers differ in structure (fig. 12), +and cannot, as in the last family, be indifferently used in a reversed +manner. The left wing, which acts as the bow of the fiddle, lies over +the right wing which serves as the fiddle itself. One of the nervures +(_a_) on the under surface of the former is finely serrated, and is +scraped across the prominent nervures on the upper surface of the +opposite or right wing. In our British _Phasgonura viridissima_ it +appeared to me that the serrated nervure is rubbed against the rounded +hind corner of the opposite wing, the edge of which is thickened, +coloured brown, and very sharp. In the right wing, but not in the left, +there is a little plate, as transparent as talc, surrounded by nervures, +and called the speculum. In _Ephippiger vitium_, a member of this same +family, we have a curious subordinate modification; for the wing-covers +are greatly reduced in size, but "the posterior part of the pro-thorax +is elevated into a kind of dome over the wing-covers, and which has +probably the effect of increasing the sound."[460] + +[Illustration: Fig. 12. Chlorocoelus Tanana (from Bates). _a_, _b_. +Lobes of opposite wing-covers.] + +We thus see that the musical apparatus is more differentiated or +specialised in the Locustid, which includes I believe the most powerful +performers in the Order, than in the Achetid, in which both wing-covers +have the same structure and the same function.[461] Landois, however, +detected in one of the Locustid, namely in Decticus, a short and narrow +row of small teeth, mere rudiments, on the inferior surface of the +right wing-cover, which underlies the other and is never used as the +bow. I observed the same rudimentary structure on the under side of the +right wing-cover in _Phasgonura viridissima_. Hence we may with +confidence infer that the Locustid are descended from a form, in which, +as in the existing Achetid, both wing-covers had serrated nervures on +the under surface, and could be indifferently used as the bow; but that +in the Locustid the two wing-covers gradually became differentiated and +perfected, on the principle of the division of labour, the one to act +exclusively as the bow and the other as the fiddle. By what steps the +more simple apparatus in the Achetid originated, we do not know, but it +is probable that the basal portions of the wing-covers overlapped each +other formerly as at present, and that the friction of the nervures +produced a grating sound, as I find is now the case with the wing-covers +of the females.[462] A grating sound thus occasionally and accidentally +made by the males, if it served them ever so little as a love-call to +the females, might readily have been intensified through sexual +selection by fitting variations in the roughness of the nervures having +been continually preserved. + +[Illustration: Fig. 13, Hind-leg of Stenobothrus pratorum: _r_, the +stridulating ridge; lower figure, the teeth, forming the ridge, much +magnified (from Landois).] + +In the last and third Family, namely the Acridiid or grasshoppers, the +stridulation is produced in a very different manner, and is not so +shrill, according to Dr. Scudder, as in the preceding Families. The +inner surface of the femur (fig. 13, _r_) is furnished with a +longitudinal row of minute, elegant, lancet-shaped, elastic teeth, from +85 to 93 in number;[463] and these are scraped across the sharp, +projecting nervures on the wing-covers, which, are thus made to vibrate +and resound. Harris[464] says that when one of the males begins to play, +he first "bends the shank of the hind-leg beneath, the thigh, where it +is lodged in a furrow designed to receive it, and then draws the leg +briskly up and down. He does not play both fiddles together, but +alternately first upon one and then on the other." In many species, the +base of the abdomen is hollowed out into a great cavity which is +believed to act as a resounding board. In Pneumora (fig. 14), a S. +African genus belonging to this same family, we meet with a new and +remarkable modification: in the males a small notched ridge projects +obliquely from each side of the abdomen, against which the hind femora +are rubbed.[465] As the male is furnished with wings, the female being +wingless, it is remarkable that the thighs are not rubbed in the usual +manner against the wing-covers; but this may perhaps be accounted for by +the unusually small size of the hind-legs. I have not been able to +examine the inner surface of the thighs, which, judging from analogy, +would be finely serrated. The species of Pneumora have been more +profoundly modified for the sake of stridulation than any other +orthopterous insect; for in the male the whole body has been converted +into a musical instrument, being distended with air, like a great +pellucid bladder, so as to increase the resonance. Mr. Trimen informs me +that at the Cape of Good Hope these insects make a wonderful noise +during the night There is one exception to the rule that the females in +these three Families are destitute of an efficient musical apparatus; +for both sexes of Ephippiger (Locustid) are said[466] to be thus +provided. This case may be compared with that of the reindeer, in which +species alone both sexes possess horns. Although the female orthoptera +are thus almost invariably mute, yet Landois[467] found rudiments of the +stridulating organs on the femora of the female Acridiid, and similar +rudiments on the under surface of the wing-covers of the female +Achetid; but he failed to find any rudiments in the females of +Decticus, one of the Locustid. In the Homoptera the mute females of +Cicada, have the proper musical apparatus in an undeveloped state; and +we shall hereafter meet in other divisions of the animal kingdom with +innumerable instances of structures proper to the male being present in +a rudimentary condition in the female. Such cases appear at first sight +to indicate that both sexes were primordially constructed in the same +manner, but that certain organs were subsequently lost by the females. +It is, however, a more probable view, as previously explained, that the +organs in question were acquired by the males and partially transferred +to the females. + +[Illustration: Fig. 14. Pneumora (from specimens in the British Museum). +Upper figure, male; lower figure, female.] + +Landois has observed another interesting fact, namely that in the +females of the Acridiid, the stridulating teeth on the femora remain +throughout life in the same condition in which they first appear in both +sexes during the larval state. In the males, on the other hand, they +become fully developed and acquire their perfect structure at the last +moult, when the insect is mature and ready to breed. + +From the facts now given, we see that the means by which the males +produce their sounds are extremely diversified in the Orthoptera, and +are altogether different from those employed by the Homoptera. But +throughout the animal kingdom we incessantly find the same object +gained by the most diversified means; this being due to the whole +organisation undergoing in the course of ages multifarious changes; and +as part after part varies, different variations are taken advantage of +for the same general purpose. The diversification of the means for +producing sound in the three families of the Orthoptera and in the +Homoptera, impresses the mind with the high importance of these +structures to the males, for the sake of calling or alluring the +females. We need feel no surprise at the amount of modification which +the Orthoptera have undergone in this respect, as we now know, from Dr. +Scudder's remarkable discovery,[468] that there has been more than ample +time. This naturalist has lately found a fossil insect in the Devonian +formation of New Brunswick, which is furnished with "the well-known +tympanum or stridulating apparatus of the male Locustid." This insect, +though in most respects related to the Neuroptera, appears to connect, +as is so often the case with very ancient forms, the two Orders of the +Neuroptera and Orthoptera which are now generally ranked as quite +distinct. + +I have but little more to say on the Orthoptera. Some of the species are +very pugnacious: when two male field-crickets (_Gryllus campestris_) are +confined together, they fight till one kills the other; and the species +of Mantis are described as manoeuvring with their sword-like +front-limbs, like hussars with their sabres. The Chinese keep these +insects in little bamboo cages and match them like game-cocks.[469] With +respect to colour, some exotic locusts are beautifully ornamented; the +posterior wings being marked with red, blue, and black; but as +throughout the Order the two sexes rarely differ much in colour, it is +doubtful whether they owe these bright tints to sexual selection. +Conspicuous colours may be of use to these insects as a protection, on +the principle to be explained in the next chapter, by giving notice to +their enemies that they are unpalatable. Thus it has been observed[470] +that an Indian brightly-coloured locust was invariably rejected when +offered to birds and lizards. Some cases, however, of sexual differences +in colour in this Order are known. The male of an American cricket[471] +is described as being as white as ivory, whilst the female varies from +almost white to greenish-yellow or dusky. Mr. Walsh informs me that the +adult male of _Spectrum femoratum_ (one of the Phasmid) "is of a +shining brownish-yellow colour; the adult female being of a dull, +opaque, cinereous-brown; the young of both sexes being green." Lastly, I +may mention that the male of one curious kind of cricket[472] is +furnished with "a long membranous appendage, which falls over the face +like a veil;" but whether this serves as an ornament is not known. + + +Order, _Neuroptera_.--Little need here be said, except in regard to +colour. In the Ephemerid the sexes often differ slightly in their +obscure tints;[473] but it is not probable that the males are thus +rendered attractive to the females. The Libellulid or dragon-flies are +ornamented with splendid green, blue, yellow, and vermilion metallic +tints; and the sexes often differ. Thus, the males of some of the +Agrionid, as Prof. Westwood remarks[474] "are of a rich blue with black +wings, whilst the females are fine green with colourless wings." But in +_Agrion Ramburii_ these colours are exactly reversed in the two +sexes.[475] In the extensive N. American genus of Hetrina, the males +alone have a beautiful carmine spot at the base of each wing. In _Anax +junius_ the basal part of the abdomen in the male is a vivid +ultra-marine blue, and in the female grass-green. In the allied genus +Gomphus, on the other hand, and in some other genera, the sexes differ +but little in colour. Throughout the animal kingdom, similar cases of +the sexes of closely-allied forms either differing greatly, or very +little, or not at all, are of frequent occurrence. Although with many +Libellulid there is so wide a difference in colour between the sexes, +it is often difficult to say which is the most brilliant; and the +ordinary coloration of the two sexes is exactly reversed, as we have +just seen, in one species of Agrion. It is not probable that their +colours in any case have been gained as a protection. As Mr. MacLachlan, +who has closely attended to this family, writes to me, dragon-flies--the +tyrants of the insect-world--are the least liable of any insect to be +attacked by birds or other enemies. He believes that their bright +colours serve as a sexual attraction. It deserves notice, as bearing on +this subject, that certain dragon-flies appear to be attracted by +particular colours: Mr. Patterson observed[476] that the species of +Agrionid, of which the males are blue, settled in numbers on the blue +float of a fishing line; whilst two other species were attracted by +shining white colours. + +It is an interesting fact, first observed by Schelver, that the males, +in several genera belonging to two sub-families, when they first emerge +from the pupal state are coloured exactly like the females; but that +their bodies in a short time assume a conspicuous milky-blue tint, owing +to the exudation of a kind of oil, soluble in ether and alcohol. Mr. +MacLachlan believes that in the male of _Libellula depressa_ this change +of colour does not occur until nearly a fortnight after the +metamorphosis, when the sexes are ready to pair. + +Certain species of Neurothemis present, according to Brauer[477] a +curious case of dimorphism, some of the females having their wings +netted in the usual manner; whilst other females have them "very richly +netted as in the males of the same species." Brauer "explains the +phenomenon on Darwinian principles by the supposition that the close +netting of the veins is a secondary sexual character in the males." This +latter character is generally developed in the males alone, but being, +like every other masculine character, latent in the female, is +occasionally developed in them. We have here an illustration of the +manner in which the two sexes of many animals have probably come to +resemble each other, namely by variations first appearing in the males, +being preserved in them, and then transmitted to and developed in the +females; but in this particular genus a complete transference is +occasionally and abruptly effected. Mr. MacLachlan informs me of another +case of dimorphism occurring in several species of Agrion in which a +certain number of individuals are found of an orange colour, and these +are invariably females. This is probably a case of reversion, for in +the true Libellul, when the sexes differ in colour, the females are +always orange or yellow, so that supposing Agrion to be descended from +some primordial form having the characteristic sexual colours of the +typical Libellul, it would not be surprising that a tendency to vary in +this manner should occur in the females alone. + +Although many dragon-flies are such large, powerful, and fierce insects, +the males have not been observed by Mr. MacLachlan to fight together, +except, as he believes, in the case of some of the smaller species of +Agrion. In another very distinct group in this Order, namely in the +Termites or white ants, both sexes at the time of swarming may be seen +running about, "the male after the female, sometimes two chasing one +female, and contending with great eagerness who shall win the +prize."[478] + + +Order, _Hymenoptera_.--That inimitable observer, M. Fabre,[479] in +describing the habits of Cerceris, a wasp-like insect, remarks that +"fights frequently ensue between the males for the possession of some +particular female, who sits an apparently unconcerned beholder of the +struggle for supremacy, and when the victory is decided, quietly flies +away in company with the conqueror." Westwood[480] says that the males +of one of the saw-flies (Tenthredin) "have been found fighting +together, with their mandibles locked." As M. Fabre speaks of the males +of Cerceris striving to obtain a particular female, it may be well to +bear in mind that insects belonging to this Order have the power of +recognising each other after long intervals of time, and are deeply +attached. For instance, Pierre Huber, whose accuracy no one doubts, +separated some ants, and when after an interval of four months they met +others which had formerly belonged to the same community, they mutually +recognised and caressed each other with their antenn. Had they been +strangers they would have fought together. Again, when two communities +engage in a battle, the ants on the same side in the general confusion +sometimes attack each other, but they soon perceive their mistake, and +the one ant soothes the other.[481] + +In this Order slight differences in colour, according to sex, are +common, but conspicuous differences are rare except in the family of +Bees; yet both sexes of certain groups are so brilliantly coloured--for +instance in Chrysis, in which vermilion and metallic greens +prevail--that we are tempted to attribute the result to sexual +selection. In the Ichneumonid, according to Mr. Walsh,[482] the males +are almost universally lighter coloured than the females. On the other +hand, in the Tenthredinid the males are generally darker than the +females. In the Siricid the sexes frequently differ; thus the male of +_Sirex juvencus_ is banded with orange, whilst the female is dark +purple; but it is difficult to say which sex is the most ornamented. In +_Tremex columb_ the female is much brighter coloured than the male. +With ants, as I am informed by Mr. F. Smith, the males of several +species are black, the females being testaceous. In the family of Bees, +especially in the solitary species, as I hear from the same +distinguished entomologist, the sexes often differ in colour. The males +are generally the brightest, and in Bombus as well as in Apathus, much +more variable in colour than the females. In _Anthophora retusa_ the +male is of a rich fulvous-brown, whilst the female is quite black: so +are the females of several species of Xylocopa, the males being bright +yellow. In an Australian bee (_Lestis bombylans_), the female is of an +extremely brilliant steel-blue, sometimes tinted with vivid green; the +male being of a bright brassy colour clothed with rich fulvous +pubescence. As in this group the females are provided with excellent +defensive weapons in their stings, it is not probable that they have +come to differ in colour from the males for the sake of protection. + +_Mutilla Europa_ emits a stridulating noise; and according to +Goureau[483] both sexes have this power. He attributes the sound to the +friction of the third and preceding abdominal segments; and I find that +these surfaces are marked with very fine concentric ridges, but so is +the projecting thoracic collar, on which the head articulates; and this +collar, when scratched with the point of a needle, emits the proper +sound. It is rather surprising that both sexes should have the power of +stridulating, as the male is winged and the female wingless. It is +notorious that Bees express certain emotions, as of anger, by the tone +of their humming, as do some dipterous insects; but I have not referred +to these sounds, as they are not known to be in any way connected with +the act of courtship. + + +Order, _Coleoptera_ (Beetles).--Many beetles are coloured so as to +resemble the surfaces which they habitually frequent. Other species are +ornamented with gorgeous metallic tints,--for instance, many Carabid, +which live on the ground and have the power of defending themselves by +an intensely acrid secretion,--the splendid diamond-beetles which are +protected by an extremely hard covering,--many species of Chrysomela, +such as _C. cerealis_, a large species beautifully striped with various +colours, and in Britain confined to the bare summit of Snowdon,--and a +host of other species. These splendid colours, which are often arranged +in stripes, spots, crosses and other elegant patterns, can hardly be +beneficial, as a protection, except in the case of some flower-feeding +species; and we cannot believe that they are purposeless. Hence the +suspicion arises, that they serve as a sexual attraction; but we have no +evidence on this head, for the sexes rarely differ in colour. Blind +beetles, which cannot of course behold each other's beauty, never +exhibit, as I hear from Mr. Waterhouse, jun., bright colours, though +they often have polished coats: but the explanation of their obscurity +may be that blind insects inhabit caves and other obscure stations. + +Some Longicorns, however, especially certain Prionid, offer an +exception to the common rule that the sexes of beetles do not differ in +colour. Most of these insects are large and splendidly coloured. The +males in the genus Pyrodes,[484] as I saw in Mr. Bates' collection, are +generally redder but rather duller than the females, the latter being +coloured of a more or less splendid golden green. On the other hand, in +one species the male is golden-green, the female being richly tinted +with red and purple. In the genus Esmeralda the sexes differ so greatly +in colour that they have been ranked as distinct species: in one species +both are of a beautiful shining green, but the male has a red thorax. On +the whole, as far as I could judge, the females of those Prionid, in +which the sexes differ, are coloured more richly than the males; and +this does not accord with the common rule in regard to colour when +acquired through sexual selection. + +[Illustration: Fig. 15. Chalcosoma atlas. Upper figure, male (reduced); +lower figure, female (nat. size).] + +[Illustration: Fig. 16. Copris isidis. (Left-hand figures, males.)] + +[Illustration: Fig. 17. Phanus faunus.] + +[Illustration: Fig. 18. Dipelicus cantori.] + +[Illustration: Fig. 19. Onthophagus rangifer, enlarged.] + +A most remarkable distinction between the sexes of many beetles is +presented by the great horns which rise from the head, thorax, or +clypeus of the males; and in some few cases from the under surface of +the body. These horns, in the great family of the Lamellicorns, resemble +those of various quadrupeds, such as stags, rhinoceroses, &c., and are +wonderful both from their size and diversified shapes. Instead of +describing them, I have given figures of the males and females of some +of the more remarkable forms. (Figs. 15 to 19.) The females generally +exhibit rudiments of the horns in the form of small knobs or ridges; but +some are destitute of even a rudiment. On the other hand, the horns are +nearly as well developed in the female as in the male of _Phanus +lancifer_; and only a little less well developed in the females of some +other species of the same genus and of Copris. In the several +subdivisions of the family, the differences in structure of the horns do +not run parallel, as I am informed by Mr. Bates, with their more +important and characteristic differences; thus within the same natural +section of the genus Onthophagus, there are species which have either a +single cephalic horn, or two distinct horns. + +In almost all cases, the horns are remarkable from their excessive +variability; so that a graduated series can be formed, from the most +highly developed males to others so degenerate that they can barely be +distinguished from the females. Mr. Walsh[485] found that in _Phanus +carnifex_ the horns were thrice as long in some males as in others. Mr. +Bates, after examining above a hundred males of _Onthophagus rangifer_ +(fig. 19), thought that he had at last discovered a species in which +the horns did not vary; but further research proved the contrary. + +The extraordinary size of the horns, and their widely different +structure in closely-allied forms, indicate that they have been formed +for some important purpose; but their excessive variability in the males +of the same species leads to the inference that this purpose cannot be +of a definite nature. The horns do not show marks of friction, as if +used for any ordinary work. Some authors suppose[486] that as the males +wander much more than the females, they require horns as a defence +against their enemies; but in many cases the horns do not seem well +adapted for defence, as they are not sharp. The most obvious conjecture +is that they are used by the males for fighting together; but they have +never been observed to fight; nor could Mr. Bates, after a careful +examination of numerous species, find any sufficient evidence in their +mutilated or broken condition of their having been thus used. If the +males had been habitual fighters, their size would probably have been +increased through sexual selection, so as to have exceeded that of the +female; but Mr. Bates, after comparing the two sexes in above a hundred +species of the Coprid, does not find in well-developed individuals any +marked difference in this respect. There is, moreover, one beetle, +belonging to the same great division of the Lamellicorns, namely +Lethrus, the males of which are known to fight, but they are not +provided with horns, though their mandibles are much larger than those +of the female. + +The conclusion, which best agrees with the fact of the horns having been +so immensely yet not fixedly developed,--as shewn by their extreme +variability in the same species and by their extreme diversity in +closely-allied species--is that they have been acquired as ornaments. +This view will at first appear extremely improbable; but we shall +hereafter find with many animals, standing much higher in the scale, +namely fishes, amphibians, reptiles and birds, that various kinds of +crests, knobs, horns and combs have been developed apparently for this +sole purpose. + +[Illustration: Fig. 20. Onitis furcifer, male, viewed from beneath.] + +[Illustration: Fig. 21. Left-hand figure, male of Onitis furcifer, +viewed laterally. Right-hand figure, female. _a._ Rudiment of cephalic +horn. _b._ Trace of thoracic horn or crest.] + +The males of _Onitis furcifer_ (fig. 20) are furnished with singular +projections on their anterior femora, and with a great fork or pair of +horns on the lower surface of the thorax. This situation seems extremely +ill adapted for the display of these projections, and they may be of +some real service; but no use can at present be assigned to them. It is +a highly remarkable fact, that although the males do not exhibit even a +trace of horns on the upper surface of the body, yet in the females a +rudiment of a single horn on the head (fig. 21, _a_), and of a crest +(_b_) on the thorax, are plainly visible. That the slight thoracic crest +in the female is a rudiment of a projection proper to the male, though +entirely absent in the male of this particular species, is clear: for +the female of _Bubas bison_ (a form which comes next to _Onitis_) has a +similar slight crest on the thorax, and the male has in the same +situation a great projection. So again there can be no doubt that the +little point (_a_) on the head of the female _Onitis furcifer_, as well +of the females of two or three allied species, is a rudimentary +representative of the cephalic horn, which is common to the males of so +many lamellicorn beetles, as in Phanus, fig. 17. The males indeed of +some unnamed beetles in the British Museum, which are believed actually +to belong to the genus Onitis, are furnished with a similar horn. The +remarkable nature of this case will be best perceived by an +illustration: the Ruminant quadrupeds run parallel with the lamellicorn +beetles, in some females possessing horns as large as those of the male, +in others having them much smaller, or existing as mere rudiments +(though this is as rare with ruminants as it is common with +Lamellicorns), or in having none at all. Now if a new species of deer or +sheep were discovered with the female bearing distinct rudiments of +horns, whilst the head of the male was absolutely smooth, we should have +a case like that of _Onitis furcifer_. + +In this case the old belief of rudiments having been created to complete +the scheme of nature is so far from holding good, that all ordinary +rules are completely broken through. The view which seems the most +probable is that some early progenitor of Onitis acquired, like other +Lamellicorns, horns on the head and thorax, and then transferred them, +in a rudimentary condition, as with so many existing species, to the +female, by whom they have ever since been retained. The subsequent loss +of the horns by the male may have resulted through the principle of +compensation from the development of the projections on the lower +surface, whilst the female has not been thus affected, as she is not +furnished with these projections, and consequently has retained the +rudiments of the horns on the upper surface. Although this view is +supported by the case of Bledius immediately to be given, yet the +projections on the lower surface differ greatly in structure and +development in the males of the several species of Onitis, and are even +rudimentary in some; nevertheless the upper surface in all these species +is quite destitute of horns. As secondary sexual characters are so +eminently variable, it is possible that the projections on the lower +surface may have been first acquired by some progenitor of Onitis and +produced their effect through compensation, and then have been in +certain cases almost completely lost. + +[Illustration: Fig. 22. Bledius taurus, magnified. Left-hand figure, +male; right-hand figure, female.] + +All the cases hitherto given refer to the Lamellicorns, but the males of +some few other beetles, belonging to two widely distinct groups, namely, +the Curculionid and Staphylinid, are furnished with horns,--in the +former on the lower surface of the body,[487] in the latter on the upper +surface of the head and thorax. In the Staphylinid the horns of the +males in the same species are extraordinarily variable, just as we have +seen with the Lamellicorns. In Siagonium we have a case of dimorphism, +for the males can be divided into two sets, differing greatly in the +size of their bodies, and in the development of their horns, without any +intermediate gradations. In a species of Bledius (fig. 22), also +belonging to the Staphylinid, male specimens can be found in the same +locality, as Professor Westwood states, "in which the central horn of +the thorax is very large, but the horns of the head quite rudimental; +and others, in which the thoracic horn is much shorter, whilst the +protuberances on the head are long."[488] Here, then, we apparently have +an instance of compensation of growth, which throws light on the curious +case just given of the loss of the upper horns by the males of _Onitis +furcifer_. + + +_Law of Battle._--Some male beetles, which seem ill fitted for fighting, +nevertheless engage in conflicts for the possession of the females. Mr. +Wallace[489] saw two males of _Leptorhynchus angustatus_, a linear +beetle with a much elongated rostrum, "fighting for a female, who stood +close by busy at her boring. They pushed at each other with their +rostra, and clawed and thumped, apparently in the greatest rage." The +smaller male, however, "soon ran away, acknowledging himself +vanquished." In some few cases the males are well adapted for fighting, +by possessing great toothed mandibles, much larger than those of the +females. This is the case with the common stag-beetle (_Lucanus +cervus_), the males of which emerge from the pupal state about a week +before the other sex, so that several may often be seen pursuing the +same female. At this period they engage in fierce conflicts. When Mr. A. +H. Davis[490] enclosed two males with one female in a box, the larger +male severely pinched the smaller one, until he resigned his +pretensions. A friend informs me that when a boy he often put the males +together to see them fight, and he noticed that they were much bolder +and fiercer than the females, as is well known to be the case with the +higher animals. The males would seize hold of his finger, if held in +front, but not so the females. With many of the Lucanid, as well as +with the above-mentioned Leptorhynchus, the males are larger and more +powerful insects than the females. The two sexes of _Lethrus cephalotes_ +(one of the Lamellicorns) inhabit the same burrow; and the male has +larger mandibles than the female. If, during the breeding-season, a +strange male attempts to enter the burrow, he is attacked; the female +does not remain passive, but closes the mouth of the burrow, and +encourages her mate by continually pushing him on from behind. The +action does not cease until the aggressor is killed or runs away.[491] +The two sexes of another lamellicorn beetle, the _Ateuchus cicatricosus_ +live in pairs, and seem much attached to each other; the male excites +the female to roll the balls of dung in which the ova are deposited; and +if she is removed, he becomes much agitated. If the male is removed, the +female ceases all work, and as M. Brulerie[492] believes, would remain +on the spot until she died. + +The great mandibles of the male Lucanid are extremely variable both in +size and structure, and in this respect resemble the horns on the head +and thorax of many male Lamellicorns and Staphylinid. A perfect series +can be formed from the best-provided to the worst-provided or degenerate +males. Although the mandibles of the common stag-beetle, and probably of +many other species, are used as efficient weapons for fighting, it is +doubtful whether their great size can thus be accounted for. We have +seen that with the _Lucanus elaphus_ of N. America they are used for +seizing the female. As they are so conspicuous and so elegantly +branched, the suspicion has sometimes crossed my mind that they may be +serviceable to the males as an ornament, in the same manner as the horns +on the head and thorax of the various above described species. The male +_Chiasognathus grantii_ of S. Chile--a splendid beetle belonging to the +same family--has enormously-developed mandibles (fig. 23); he is bold +and pugnacious; when threatened on any side he faces round, opening his +great jaws, and at the same time stridulating loudly; but the mandibles +were not strong enough to pinch my finger so as to cause actual pain. + +[Illustration: Fig. 23. Chiasognathus grantii, reduced. Upper figure, +male; lower figure, female.] + +Sexual selection, which implies the possession of considerable +perceptive powers and of strong passions, seems to have been more +effective with the Lamellicorns than with any other family of the +Coleoptera or beetles. With some species the males are provided with +weapons for fighting; some live in pairs and show mutual affection; +many have the power of stridulating when excited; many are furnished +with the most extraordinary horns, apparently for the sake of ornament; +some which are diurnal in their habits are gorgeously coloured; and, +lastly, several of the largest beetles in the world belong to this +family, which was placed by Linnus and Fabricius at the head of the +Order of the Coleoptera.[493] + + +_Stridulating organs._--Beetles belonging to many and widely distinct +families possess these organs. The sound can sometimes be heard at the +distance of several feet or even yards,[494] but is not comparable with +that produced by the Orthoptera. The part which may be called the rasp +generally consists of a narrow slightly-raised surface, crossed by very +fine, parallel ribs, sometimes so fine as to cause iridescent colours, +and having a very elegant appearance under the microscope. In some +cases, for instance, with Typhus, it could be plainly seen that +extremely minute, bristly, scale-like prominences, which cover the whole +surrounding surface in approximately parallel lines, give rise to the +ribs of the rasp by becoming confluent and straight, and at the same +time more prominent and smooth. A hard ridge on any adjoining part of +the body, which in some cases is specially modified for the purpose, +serves as the scraper for the rasp. The scraper is rapidly moved across +the rasp, or conversely the rasp across the scraper. + +[Illustration: Fig. 24. Necrophorus (from Landois). _r._ The two rasps. +Left-hand figure, part of the rasp highly magnified.] + +These organs are situated in widely different positions. In the +carrion-beetles (Necrophorus) two parallel rasps (_r_, fig. 24) stand on +the dorsal surface of the fifth abdominal segment, each rasp being +crossed, as described by Landois,[495] by from 126 to 140 fine ribs. +These ribs are scraped by the posterior margins of the elytra, a small +portion of which projects beyond the general outline. In many +Criocerid, and in _Clythra 4-punctata_ (one of the Chrysomelid), and +in some Tenebrionid, &c.,[496] the rasp is seated on the dorsal apex of +the abdomen, on the pygidium or pro-pygidium, and is scraped as above by +the elytra. In Heterocerus, which belongs to another family, the rasps +are placed on the sides of the first abdominal segment, and are scraped +by ridges on the femora.[497] In certain Curculionid and Carabid,[498] +the parts are completely reversed in position, for the rasps are seated +on the inferior surface of the elytra, near their apices, or along their +outer margins, and the edges of the abdominal segments serve as the +scrapers. In _Pelobius hermanni_ (one of Dytiscid or water-beetles) a +strong ridge runs parallel and near to the sutural margin of the elytra, +and is crossed by ribs, coarse in the middle part, but becoming +gradually finer at both ends, especially at the upper end; when this +insect is held under water or in the air, a stridulating noise is +produced by scraping the extreme horny margin of the abdomen against the +rasp. In a great number of long-horned beetles (Longicornia) the organs +are altogether differently situated, the rasp being on the meso-thorax, +which is rubbed against the pro-thorax; Landois counted 238 very fine +ribs on the rasp of _Cerambyx heros_. + +[Illustration: Fig. 25. Hind-leg of Geotrupes stercorarius (from +Landois). _r._ Rasp. _c._ Coxa. _f._ Femur. _t._ Tibia. _tr._ Tarsi.] + +Many Lamellicorns have the power of stridulating, and the organs differ +greatly in position. Some species stridulate very loudly, so that when +Mr. F. Smith caught a _Trox sabulosus_, a gamekeeper who stood by +thought that he had caught a mouse; but I failed to discover the proper +organs in this beetle. In Geotrupes and Typhus a narrow ridge runs +obliquely across (_r_, fig. 25) the coxa of each hind-leg, having in +_G. stercorarius_ 84 ribs, which are scraped by a specially-projecting +part of one of the abdominal segments. In the nearly allied _Copris +lunaris_, an excessively narrow fine rasp runs along the sutural margin +of the elytra, with another short rasp near the basal outer margin; but +in some other Coprini the rasp is seated, according to Leconte,[499] on +the dorsal surface of the abdomen. In Oryctes it is seated on the +pro-pygidium, and in some other Dynastini, according to the same +entomologist, on the under surface of the elytra. Lastly, Westring +states that in _Omaloplia brunnea_ the rasp is placed on the +pro-sternum, and the scraper on the meta-sternum, the parts thus +occupying the under surface of the body, instead of the upper surface as +in the Longicorns. + +We thus see that the stridulating organs in the different coleopterous +families are wonderfully diversified in position, but not much in +structure. Within the same family some species are provided with these +organs, and some are quite destitute of them. This diversity is +intelligible, if we suppose that originally various species made a +shuffling or hissing noise by the rubbing together of the hard and rough +parts of their bodies which were in contact; and that from the noise +thus produced being in some way useful, the rough surfaces were +gradually developed into regular stridulating organs. Some beetles as +they move, now produce, either intentionally or unintentionally, a +shuffling noise, without possessing any proper organs for the purpose. +Mr. Wallace informs me that the _Euchirus longimanus_ (a Lamellicorn, +with the anterior legs wonderfully elongated in the male) "makes, whilst +moving, a low hissing sound by the protrusion and contraction of the +abdomen; and when seized it produces a grating sound by rubbing its +hind-legs against the edges of the elytra." The hissing sound is clearly +due to a narrow rasp running along the sutural margin of each elytron; +and I could likewise make the grating sound by rubbing the shagreened +surface of the femur against the granulated margin of the corresponding +elytron; but I could not here detect any proper rasp; nor is it likely +that I could have overlooked it in so large an insect. After examining +Cychrus and reading what Westring has written in his two papers about +this beetle, it seems very doubtful whether it possesses any true rasp, +though it has the power of emitting a sound. + +From the analogy of the Orthoptera and Homoptera, I expected to find +that the stridulating organs in the Coleoptera differed according to +sex; but Landois, who has carefully examined several species, observed +no such difference; nor did Westring; nor did Mr. G. R. Crotch in +preparing the numerous specimens which he had the kindness to send me +for examination. Any slight sexual difference, however, would be +difficult to detect, on account of the great variability of these +organs. Thus in the first pair of the _Necrophorus humator_ and of the +_Pelobius_ which I examined, the rasp was considerably larger in the +male than in the female; but not so with succeeding specimens. In +_Geotrupes stercorarius_ the rasp appeared to me thicker, opaquer, and +more prominent in three males than in the same number of females; +consequently my son, Mr. F. Darwin, in order to discover whether the +sexes differed in their power of stridulating, collected 57 living +specimens, which he separated into two lots, according as they made, +when held in the same manner, a greater or lesser noise. He then +examined their sexes, but found that the males were very nearly in the +same proportion to the females in both lots. Mr. F. Smith has kept alive +numerous specimens of _Mononychus pseudacori_ (Curculionid), and is +satisfied that both sexes stridulate, and apparently in an equal degree. + +Nevertheless the power of stridulating is certainly a sexual character +in some few Coleoptera. Mr. Crotch has discovered that the males alone +of two species of Heliopathes (Tenebrionid) possess stridulating +organs. I examined five males of _H. gibbus_, and in all these there was +a well-developed rasp, partially divided into two, on the dorsal surface +of the terminal abdominal segment; whilst in the same number of females +there was not even a rudiment of the rasp, the membrane of this segment +being transparent and much thinner than in the male. In _H. +cribratostriatus_ the male has a similar rasp, excepting that it is not +partially divided into two portions, and the female is completely +destitute of this organ; but in addition the male has on the apical +margins of the elytra, on each side of the suture, three or four short +longitudinal ridges, which are crossed by extremely fine ribs, parallel +to and resembling those on the abdominal rasp; whether these ridges +serve as an independent rasp, or as a scraper for the abdominal rasp, I +could not decide: the female exhibits no trace of this latter structure. + +Again, in three species of the Lamellicorn genus Oryctes, we have a +nearly parallel case. In the females of _O. gryphus_ and _nasicornis_ +the ribs on the rasp of the pro-pygidium are less continuous and less +distinct than in the males; but the chief difference is that the whole +upper surface of this segment, when held in the proper light, is seen to +be clothed with hairs, which are absent or are represented by +excessively fine down in the males. It should be noticed that in all +Coleoptera the effective part of the rasp is destitute of hairs. In _O. +senegalensis_ the difference between the sexes is more strongly marked, +and this is best seen when the proper segment is cleaned and viewed as a +transparent object. In the female the whole surface is covered with +little separate crests, bearing spines; whilst in the male these crests +become, in proceeding towards the apex, more and more confluent, +regular, and naked; so that three-fourths of the segment is covered with +extremely fine parallel ribs, which are quite absent in the female. In +the females, however, of all three species of Oryctes, when the abdomen +of a softened specimen is pushed backwards and forwards, a slight +grating or stridulating sound can be produced. + +In the case of the Heliopathes and Oryctes there can hardly be a doubt +that the males stridulate in order to call or to excite the females; but +with most beetles the stridulation apparently serves both sexes as a +mutual call. This view is not rendered improbable from beetles +stridulating under various emotions; we know that birds use their voices +for many purposes besides singing to their mates. The great +Chiasognathus stridulates in anger or defiance; many species do the same +from distress or fear, when held so that they cannot escape; Messrs. +Wollaston and Crotch were able, by striking the hollow stems of trees in +the Canary Islands, to discover the presence of beetles belonging to the +genus Acalles by their stridulation. Lastly the male Ateuchus +stridulates to encourage the female in her work, and from distress when +she is removed.[500] Some naturalists believe that beetles make this +noise to frighten away their enemies; but I cannot think that the +quadrupeds and birds which are able to devour the larger beetles with +their extremely hard coats, would be frightened by so slight a grating +sound. The belief that the stridulation serves as a sexual call is +supported by the fact that death-ticks (_Anobium tessellatum_) are well +known to answer each other's ticking, or, as I have myself observed, a +tapping noise artificially made; and Mr. Doubleday informs me that he +has twice or thrice observed a female ticking,[501] and in the course of +an hour or two has found her united with a male, and on one occasion +surrounded by several males. Finally, it seems probable that the two +sexes of many kinds of beetles were at first enabled to find each other +by the slight shuffling noise produced by the rubbing together of the +adjoining parts of their hard bodies; and that as the males or females +which made the greatest noise succeeded best in finding partners, the +rugosities on various parts of their bodies were gradually developed by +means of sexual selection into true stridulating organs. + + + + +CHAPTER XI. + +INSECTS, _continued_.--ORDER LEPIDOPTERA. + + + Courtship of butterflies--Battles--Ticking noise--Colours + common to both sexes, or more brilliant in the + males--Examples--Not due to the direct action of the conditions + of life--Colours adapted for protection--Colours of + moths--Display--Perceptive powers of the + Lepidoptera--Variability--Causes of the difference in colour + between the males and females--Mimickry, female butterflies + more brilliantly coloured than the males--Bright colours of + caterpillars--Summary and concluding remarks on the secondary + sexual characters of insects--Birds and insects compared. + + +In this great Order the most interesting point for us is the difference +in colour between the sexes of the same species, and between the +distinct species of the same genus. Nearly the whole of the following +chapter will be devoted to this subject; but I will first make a few +remarks on one or two other points. Several males may often be seen +pursuing and crowding round the same female. Their courtship appears to +be a prolonged affair, for I have frequently watched one or more males +pirouetting round a female until I became tired, without seeing the end +of the courtship. Although butterflies are such weak and fragile +creatures, they are pugnacious, and an Emperor butterfly[502] has been +captured with the tips of its wings broken from a conflict with another +male. Mr. Collingwood in speaking of the frequent battles between the +butterflies of Borneo says, "They whirl round each other with the +greatest rapidity, and appear to be incited by the greatest ferocity." +One case is known of a butterfly, namely the _Ageronia feronia_, which +makes a noise like that produced by a toothed wheel passing under a +spring catch, and which could be heard at the distance of several yards. +At Rio de Janeiro this sound was noticed by me, only when two were +chasing each other in an irregular course, so that it is probably made +during the courtship of the sexes; but I neglected to attend to this +point.[503] + +Every one has admired the extreme beauty of many butterflies and of some +moths; and we are led to ask, how has this beauty been acquired? Have +their colours and diversified patterns simply resulted from the direct +action of the physical conditions to which these insects have been +exposed, without any benefit being thus derived? Or have successive +variations been accumulated and determined either as a protection or for +some unknown purpose, or that one sex might be rendered attractive to +the other? And, again, what is the meaning of the colours being widely +different in the males and females of certain species, and alike in the +two sexes of other species? Before attempting to answer these questions +a body of facts must be given. + +With most of our English butterflies, both those which are beautiful, +such as the admiral, peacock, and painted lady (Vaness), and those +which are plain-coloured, such as the meadow-browns (Hipparchi), the +sexes are alike. This is also the case with the magnificent Heliconid +and Danaid of the tropics. But in certain other tropical groups, and +with some of our English butterflies, as the purple emperor, orange-tip, +&c. (_Apatura Iris_ and _Anthocharis cardamines_), the sexes differ +either greatly or slightly in colour. No language suffices to describe +the splendour of the males of some tropical species. Even within the +same genus we often find species presenting an extraordinary difference +between the sexes, whilst others have their sexes closely alike. Thus in +the South American genus Epicalia, Mr. Bates, to whom I am much indebted +for most of the following facts and for looking over this whole +discussion, informs me that he knows twelve species, the two sexes of +which haunt the same stations (and this is not always the case with +butterflies), and therefore cannot have been differently affected by +external conditions[504]. In nine of these species the males rank +amongst the most brilliant of all butterflies, and differ so greatly +from the comparatively plain females that they were formerly placed in +distinct genera. The females of these nine species resemble each other +in their general type of coloration, and likewise resemble both sexes in +several allied genera, found in various parts of the world. Hence in +accordance with the descent-theory we may infer that these nine species, +and probably all the others of the genus, are descended from an +ancestral form which was coloured in nearly the same manner. In the +tenth species the female still retains the same general colouring, but +the male resembles her, so that he is coloured in a much less gaudy and +contrasted manner than the males of the previous species. In the +eleventh and twelfth species, the females depart from the type of +colouring which is usual with their sex in this genus, for they are +gaily decorated in nearly the same manner as the males, but in a +somewhat less degree. Hence in these two species the bright colours of +the males seem to have been transferred to the females; whilst the male +of the tenth species has either retained or recovered the plain colours +of the female as well as of the parent-form of the genus; the two sexes +being thus rendered in both cases, though in an opposite manner, nearly +alike. In the allied genus Eubagis, both sexes of some of the species +are plain-coloured and nearly alike; whilst with the greater number the +males are decorated with beautiful metallic tints, in a diversified +manner, and differ much from their females. The females throughout the +genus retain the same general style of colouring, so that they commonly +resemble each other much more closely than they resemble their own +proper males. + +In the genus Papilio, all the species of the neas group are remarkable +for their conspicuous and strongly contrasted colours, and they +illustrate the frequent tendency to gradation in the amount of +difference between the sexes. In a few species, for instance in _P. +ascanius_, the males and females are alike; in others the males are a +little or very much more superbly coloured than the females. The genus +Junonia allied to our Vaness offers a nearly parallel case, for +although the sexes of most of the species resemble each other and are +destitute of rich colours, yet in certain species, as in _J. oenone_, +the male is rather more brightly coloured than the female, and in a few +(for instance _J. andremiaja_) the male is so different from the female +that he might be mistaken for an entirely distinct species. + +Another striking case was pointed out to me in the British museum by Mr. +A. Butler, namely one of the Tropical American Thecl, in which both +sexes are nearly alike and wonderfully splendid; in another, the male +is coloured in a similarly gorgeous manner, whilst the whole upper +surface of the female is of a dull uniform brown. Our common little +English blue butterflies of the genus Lycna, illustrate the various +differences in colour between the sexes, almost as well, though not in +so striking a manner, as the above exotic genera. In _Lycna agestis_ +both sexes have wings of a brown colour, bordered with small ocellated +orange spots, and are consequently alike. In _L. oegon_ the wings of +the male are of a fine blue, bordered with black; whilst the wings of +the female are brown, with a similar border, and closely resemble those +of _L. agestis_. Lastly, in _L. arion_ both sexes are of a blue colour +and nearly alike, though in the female the edges of the wings are rather +duskier, with the black spots plainer; and in a bright blue Indian +species both sexes are still more closely alike. + +I have given the foregoing cases in some detail in order to shew, in the +first place, that when the sexes of butterflies differ, the male as a +general rule is the most beautiful, and departs most from the usual type +of colouring of the group to which the species belongs. Hence in most +groups the females of the several species resemble each other much more +closely than do the males. In some exceptional cases, however, to which +I shall hereafter allude, the females are coloured more splendidly than +the males. In the second place these cases have been given to bring +clearly before the mind that within the same genus, the two sexes +frequently present every gradation from no difference in colour to so +great a difference that it was long before the two were placed by +entomologists in the same genus. In the third place, we have seen that +when the sexes nearly resemble each other, this apparently may be due +either to the male having transferred his colours to the female, or to +the male having retained, or perhaps recovered, the primordial colours +of the genus to which the species belongs. It also deserves notice that +in those groups in which the sexes present any difference of colour, the +females usually resemble the males to a certain extent, so that when the +males are beautiful to an extraordinary degree, the females almost +invariably exhibit some degree of beauty. From the numerous cases of +gradation in the amount of difference between the sexes, and from the +prevalence of the same general type of coloration throughout the whole +of the same group, we may conclude that the causes, whatever they may +be, which have determined the brilliant colouring of the males alone of +some species, and of both sexes in a more or less equal degree of other +species, have generally been the same. + +As so many gorgeous butterflies inhabit the tropics, it has often been +supposed that they owe their colours to the great heat and moisture of +these zones; but Mr. Bates[505] has shewn by the comparison of various +closely-allied groups of insects from the temperate and tropical +regions, that this view cannot be maintained; and the evidence becomes +conclusive when brilliantly-coloured males and plain-coloured females of +the same species inhabit the same district, feed on the same food, and +follow exactly the same habits of life. Even when the sexes resemble +each other, we can hardly believe that their brilliant and +beautifully-arranged colours are the purposeless result of the nature of +the tissues, and the action of the surrounding conditions. + +With animals of all kinds, whenever colour has been modified for some +special purpose, this has been, as far as we can judge, either for +protection or as an attraction between the sexes. With many species of +butterflies the upper surfaces of the wings are obscurely coloured, and +this in all probability leads to their escaping observation and danger. +But butterflies when at rest would be particularly liable to be attacked +by their enemies; and almost all the kinds when resting raise their +wings vertically over their backs, so that the lower sides alone are +exposed to view. Hence it is this side which in many cases is obviously +coloured so as to imitate the surfaces on which these insects commonly +rest. Dr. Rssler, I believe, first noticed the similarity of the closed +wings of certain Vaness and other butterflies to the bark of trees. +Many analogous and striking facts could be given. The most interesting +one is that recorded by Mr. Wallace[506] of a common Indian and Sumatran +butterfly (Kallima), which disappears like magic when it settles in a +bush; for it hides its head and antenn between its closed wings, and +these in form, colour, and veining cannot be distinguished from a +withered leaf together with the footstalk. In some other cases the lower +surfaces of the wings are brilliantly coloured, and yet are protective; +thus in _Thecla rubi_ the wings when closed are of an emerald green and +resemble the young leaves of the bramble, on which this butterfly in the +spring may often be seen seated. + +Although the obscure tints of the upper or under surface of many +butterflies no doubt serve to conceal them, yet we cannot possibly +extend this view to the brilliant and conspicuous colours of many kinds, +such as our admiral and peacock Vaness, our white cabbage-butterflies +(Pieris), or the great swallow-tail Papilio which haunts the open +fens--for these butterflies are thus rendered visible to every living +creature. With these species both sexes are alike; but in the common +brimstone butterfly (_Gonepteryx rhamni_), the male is of an intense +yellow, whilst the female is much paler; and in the orange-tip +(_Anthocharis cardamines_) the males alone have the bright orange tips +to their wings. In these cases the males and females are equally +conspicuous, and it is not credible that their difference in colour +stands in any relation to ordinary protection. Nevertheless it is +possible that the conspicuous colours of many species may be in an +indirect manner beneficial, as will hereafter be explained, by leading +their enemies at once to recognise them as unpalatable. Even in this +case it does not certainly follow that their bright colours and +beautiful patterns were acquired for this special purpose. In some other +remarkable cases, beauty has been gained for the sake of protection, +through the imitation of other beautiful species, which inhabit the same +district and enjoy an immunity from attack by being in some way +offensive to their enemies. + +The female of our orange-tip butterfly, above referred to, and of an +American species (_Anth. genutia_) probably shew us, as Mr. Walsh has +remarked to me, the primordial colours of the parent-species of the +genus; for both sexes of four or five widely-distributed species are +coloured in nearly the same manner. We may infer here, as in several +previous cases, that it is the males of _Anth. cardamines_ and _genutia_ +which have departed from the usual type of colouring of their genus. In +the _Anth. sara_ from California, the orange-tips have become partially +developed in the female; for her wings are tipped with reddish-orange, +but paler than in the male, and slightly different in some other +respects. In an allied Indian form, the _Iphias glaucippe_, the +orange-tips are fully developed in both sexes. In this Iphias the under +surface of the wings marvellously resembles, as pointed out to me by Mr. +A. Butler, a pale-coloured leaf; and in our English orange-tip, the +under surface resembles the flower-head of the wild parsley, on which it +may be seen going to rest at night.[507] The same reasoning power which +compels us to believe that the lower surfaces have here been coloured +for the sake of protection, leads us to deny that the wings have been +tipped, especially when this character is confined to the males, with +bright orange for the same purpose. + + +Turning now to Moths: most of these rest motionless with their wings +depressed during the whole or greater part of the day; and the upper +surfaces of their wings are often shaded and coloured in an admirable +manner, as Mr. Wallace has remarked, for escaping detection. With most +of the Bombycid and Noctuid,[508] when at rest, the front-wings +overlap and conceal the hind-wings; so that the latter might be brightly +coloured without much risk; and they are thus coloured in many species +of both families. During the act of flight, moths would often be able to +escape from their enemies; nevertheless, as the hind-wings are then +fully exposed to view, their bright colours must generally have been +acquired at the cost of some little risk. But the following fact shews +us how cautious we ought to be in drawing conclusions on this head. The +common yellow under-wings (Triphaena) often fly about during the day or +early evening, and are then conspicuous from the colour of their +hind-wings. It would naturally be thought that this would be a source of +danger; but Mr. J. Jenner Weir believes that it actually serves them as +a means of escape, for birds strike at these brightly coloured and +fragile surfaces, instead of at the body. For instance, Mr. Weir turned +into his aviary a vigorous specimen of _Triphaena pronuba_, which was +instantly pursued by a robin; but the bird's attention being caught by +the coloured wings, the moth was not captured until after about fifty +attempts, and small portions of the wings were repeatedly broken off. He +tried the same experiment, in the open air, with a _T. fimbria_ and +swallow; but the large size of this moth probably interfered with its +capture.[509] We are thus reminded of a statement made by Mr. +Wallace,[510] namely, that in the Brazilian forests and Malayan islands, +many common and highly-decorated butterflies are weak flyers, though +furnished with a broad expanse of wings; and they "are often captured +with pierced and broken wings, as if they had been seized by birds, from +which they had escaped: if the wings had been much smaller in proportion +to the body, it seems probable that the insect would more frequently +have been struck or pierced in a vital part, and thus the increased +expanse of the wings may have been indirectly beneficial." + + +_Display._--The bright colours of butterflies and of some moths are +specially arranged for display, whether or not they serve in addition as +a protection. Bright colours would not be visible during the night; and +there can be no doubt that moths, taken as a body, are much less gaily +decorated than butterflies, all of which are diurnal in their habits. +But the moths in certain families, such as the Zygnid, various +Sphingid, Uraniid, some Arctiid and Saturniid, fly about during the +day or early evening, and many of these are extremely beautiful, being +far more brightly coloured than the strictly nocturnal kinds. A few +exceptional cases, however, of brightly-coloured nocturnal species have +been recorded.[511] + +There is evidence of another kind in regard to display. Butterflies, as +before remarked, elevate their wings when at rest, and whilst basking in +the sunshine often alternately raise and depress them, thus exposing to +full view both surfaces; and although the lower surface is often +coloured in an obscure manner as a protection, yet in many species it is +as highly coloured as the upper surface, and sometimes in a very +different manner. In some tropical species the lower surface is even +more brilliantly coloured than the upper.[512] In one English +fritillary, the _Argynnis aglaia_, the lower surface alone is ornamented +with shining silver discs. Nevertheless, as a general rule, the upper +surface, which is probably the most fully exposed, is coloured more +brightly and in a more diversified manner than the lower. Hence the +lower surface generally affords to entomologists the most useful +character for detecting the affinities of the various species. + +Now if we turn to the enormous group of moths, which do not habitually +expose to full view the under surface of their wings, this side is very +rarely, as I hear from Mr. Stainton, coloured more brightly than the +upper side, or even with equal brightness. Some exceptions to the rule, +either real or apparent, must be noticed, as that of Hypopyra, specified +by Mr. Wormald.[513] Mr. R. Trimen informs me that in Guene's great +work, three moths are figured, in which the under surface is much the +most brilliant. For instance, in the Australian Gastrophora the upper +surface of the fore-wing is pale greyish-ochreous, while the lower +surface is magnificently ornamented by an ocellus of cobalt-blue, placed +in the midst of a black mark, surrounded by orange-yellow, and this by +bluish-white. But the habits of these three moths are unknown; so that +no explanation can be given of their unusual style of colouring. Mr. +Trimen also informs me that the lower surface of the wings in certain +other Geometr[514] and quadrifid Noctu are either more variegated or +more brightly-coloured than the upper surface; but some of these species +have the habit of "holding their wings quite erect over their backs, +retaining them in this position for a considerable time," and thus +exposing to view the under surface. Other species when settled on the +ground or herbage have the habit of now and then suddenly and slightly +lifting up their wings. Hence the lower surface of the wings being more +brightly-coloured than the upper surface in certain moths is not so +anomalous a circumstance as it at first appears. The Saturniid include +some of the most beautiful of all moths, their wings being decorated, as +in our British Emperor moth, with fine ocelli; and Mr. T. W. Wood[515] +observes that they resemble butterflies in some of their movements; "for +instance, in the gentle waving up and down of the wings, as if for +display, which is more characteristic of diurnal than of nocturnal +Lepidoptera." + +It is a singular fact that no British moths, nor as far as I can +discover hardly any foreign species, which are brilliantly coloured, +differ much in colour according to sex; though this is the case with +many brilliant butterflies. The male, however, of one American moth, the +_Saturnia Io_, is described as having its fore-wings deep yellow, +curiously marked with purplish-red spots; whilst the wings of the female +are purple-brown, marked with grey lines.[516] The British moths which +differ sexually in colour are all brown, or various tints of dull +yellow, or nearly white. In several species the males are much darker +than the females,[517] and these belong to groups which generally fly +about during the afternoon. On the other hand, in many genera, as Mr. +Stainton informs me, the males have the hind-wings whiter than those of +the female--of which, fact _Agrotis exclamationis_ offers a good +instance. The males are thus rendered more conspicuous than the females, +whilst flying about in the dusk. In the Ghost Moth (_Hepialus humuli_) +the difference is more strongly marked; the males being white and the +females yellow with darker markings. It is difficult to conjecture what +the meaning can be of these differences between the sexes in the shades +of darkness or lightness; but we can hardly suppose that they are the +result of mere variability with sexually-limited inheritance, +independently of any benefit thus derived. + +From the foregoing statements it is impossible to admit that the +brilliant colours of butterflies and of some few moths, have commonly +been acquired for the sake of protection. We have seen that their +colours and elegant patterns are arranged and exhibited as if for +display. Hence I am led to suppose that the females generally prefer, or +are most excited by the more brilliant males; for on any other +supposition the males would be ornamented, as far as we can see, for no +purpose. We know that ants and certain lamellicorn beetles are capable +of feeling an attachment for each other, and that ants recognise their +fellows after an interval of several months. Hence there is no abstract +improbability in the Lepidoptera, which probably stand nearly or quite +as high in the scale as these insects, having sufficient mental capacity +to admire bright colours. They certainly discover flowers by colour, +and, as I have elsewhere shewn, the plants which are fertilised +exclusively by the wind never have a conspicuously-coloured corolla. The +Humming-bird Sphinx may often be seen to swoop down from a distance on a +bunch of flowers in the midst of green foliage; and I have been assured +by a friend, that these moths repeatedly visited flowers painted on the +walls of a room in the South of France. The common white butterfly, as I +hear from Mr. Doubleday, often flies down to a bit of paper on the +ground, no doubt mistaking it for one of its own species. Mr. +Collingwood[518] in speaking of the difficulty of collecting certain +butterflies in the Malay Archipelago, states that "a dead specimen +pinned upon a conspicuous twig will often arrest an insect of the same +species in its headlong flight, and bring it down within easy reach of +the net, especially if it be of the opposite sex." + +The courtship of butterflies is a prolonged affair. The males sometimes +fight together in rivalry; and many may be seen pursuing or crowding +round the same female. If, then, the females do not prefer one male to +another, the pairing must be left to mere chance, and this does not +appear to me a probable event. If, on the other hand, the females +habitually, or even occasionally, prefer the more beautiful males, the +colours of the latter will have been rendered brighter by degrees, and +will have been transmitted to both sexes or to one sex, according to +which law of inheritance prevailed. The process of sexual selection will +have been much facilitated, if the conclusions arrived at from various +kinds of evidence in the supplement to the ninth chapter can be trusted; +namely that the males of many Lepidoptera, at least in the imago state, +greatly exceed in number the females. + +Some facts, however, are opposed to the belief that female butterflies +prefer the more beautiful males; thus, as I have been assured by several +observers, fresh females may frequently be seen paired with battered, +faded or dingy males; but this is a circumstance which could hardly +fail often to follow from the males emerging from their cocoons earlier +than the females. With moths of the family of the Bombycid, the sexes +pair immediately after assuming the imago state; for they cannot feed, +owing to the rudimentary condition of their mouths. The females, as +several entomologists have remarked to me, lie in an almost torpid +state, and appear not to evince the least choice in regard to their +partners, This is the case with the common silk-moth (_B. mori_), as I +have been told by some continental and English breeders. Dr. Wallace, +who has had such immense experience in breeding _Bombyx cynthia_, is +convinced that the females evince no choice or preference. He has kept +above 300 of these moths living together, and has often found the most +vigorous females mated with stunted males. The reverse apparently seldom +occurs; for, as he believes, the more vigorous males pass over the +weakly females, being attracted by those endowed with most vitality. +Although we have been indirectly induced to believe that the females of +many species prefer the more beautiful males, I have no reason to +suspect, either with moths or butterflies, that the males are attracted +by the beauty of the females. If the more beautiful females had been +continually preferred, it is almost certain, from the colours of +butterflies being so frequently transmitted to one sex alone, that the +females would often have been rendered more beautiful than their male +partners. But this does not occur except in a few instances; and these +can be explained, as we shall presently see, on the principle of +mimickry and protection. + + +As sexual selection primarily depends on variability, a few words must +be added on this subject. In respect to colour there is no difficulty, +as any number of highly variable Lepidoptera could be named. One good +instance will suffice. Mr. Bates shewed me a whole series of specimens +of _Papilio sesostris_ and _children_; in the latter the males varied +much in the extent of the beautifully enamelled green patch on the +fore-wings, and in the size of the white mark, as well as of the +splendid crimson stripe on the hind-wings; so that there was a great +contrast between the most and least gaudy males. The male of _Papilio +sesostris_, though a beautiful insect, is much less so than _P. +children_. It likewise varies a little in the size of the green patch +on the fore-wings, and in the occasional appearance of a small crimson +stripe on the hind-wings, borrowed, as it would seem, from its own +female; for the females of this and of many other species in the neas +group possess this crimson stripe. Hence between the brightest specimens +of _P. sesostris_ and the least bright of _P. children_, there was but +a small interval; and it was evident that as far as mere variability is +concerned, there would be no difficulty in permanently increasing by +means of selection the beauty of either species. The variability is here +almost confined to the male sex; but Mr. Wallace and Mr. Bates have +shewn[519] that the females of some other species are extremely +variable, the males being nearly constant. As I have before mentioned +the Ghost Moth (_Hepialus humuli_) as one of the best instances in +Britain of a difference in colour between the sexes of moths, it may be +worth adding[520] that in the Shetland Islands, males are frequently +found which closely resemble the females. In a future chapter I shall +have occasion to shew that the beautiful eye-like spots or ocelli, so +common on the wings of many Lepidoptera, are eminently variable. + + +On the whole, although many serious objections may be urged, it seems +probable that most of the species of Lepidoptera which are brilliantly +coloured, owe their colours to sexual selection, excepting in certain +cases, presently to be mentioned, in which conspicuous colours are +beneficial as a protection. From the ardour of the male throughout the +animal kingdom, he is generally willing to accept any female; and it is +the female which usually exerts a choice. Hence if sexual selection has +here acted, the male, when the sexes differ, ought to be the most +brilliantly coloured; and this undoubtedly is the ordinary rule. When +the sexes are brilliantly coloured and resemble each other, the +characters acquired by the males appear to have been transmitted to both +sexes. But will this explanation of the similarity and dissimilarity in +colour between the sexes suffice? + +The males and females of the same species of butterfly are known[521] in +several cases to inhabit different stations, the former commonly basking +in the sunshine, the latter haunting gloomy forests. It is therefore +possible that different conditions of life may have acted directly on +the two sexes; but this is not probable,[522] as in the adult state they +are exposed during a very short period to different conditions; and the +larv of both are exposed to the same conditions. Mr. Wallace believes +that the less brilliant colours of the female have been specially +gained in all or almost all cases for the sake of protection. On the +contrary it seems to me more probable that the males alone, in the large +majority of cases, have acquired their bright colours through sexual +selection, the females having been but little modified. Consequently the +females of distinct but allied species ought to resemble each other much +more closely than do the males of the same species; and this is the +general rule. The females thus approximately show us the primordial +colouring of the parent-species of the group to which they belong. They +have, however, almost always been modified to a certain extent by some +of the successive steps of variation, through the accumulation of which +the males were rendered beautiful, having been transferred to them. The +males and females of allied though distinct species will also generally +have been exposed during their prolonged larval state to different +conditions, and may have been thus indirectly affected; though with the +males any slight change of colour thus caused will often have been +completely masked by the brilliant tints gained through sexual +selection. When we treat of Birds, I shall have to discuss the whole +question whether the differences in colour between the males and females +have been in part specially gained by the latter as a protection; so +that I will here only give unavoidable details. + +In all cases when the more common form of equal inheritance by both +sexes has prevailed, the selection of bright-coloured males would tend +to make the females bright-coloured; and the selection of dull-coloured +females would tend to make the males dull. If both processes were +carried on simultaneously, they would tend to neutralise each other. As +far as I can see, it would be extremely difficult to change through +selection the one form of inheritance into the other. But by the +selection of successive variations, which were from the first sexually +limited in their transmission, there would not be the slightest +difficulty in giving bright colours to the males alone, and at the same +time or subsequently, dull colours to the females alone. In this latter +manner female butterflies and moths may, as I fully admit, have been +rendered inconspicuous for the sake of protection, and widely different +from their males. + +Mr. Wallace[523] has argued with much force in favour of his view that +when the sexes differ, the female has been specially modified for the +sake of protection; and that this has been effected by one form of +inheritance, namely, the transmission of characters to both sexes, +having been changed through the agency of natural selection into the +other form, namely, transmission to one sex. I was at first strongly +inclined to accept this view; but the more I have studied the various +classes throughout the animal kingdom, the less probable it has +appeared. Mr. Wallace urges that both sexes of the _Heliconid_, +_Danaid_, _Acroeid_ are equally brilliant because both are protected +from the attacks of birds and other enemies, by their offensive odour; +but that in other groups, which do not possess this immunity, the +females have been rendered inconspicuous, from having more need of +protection than the males. This supposed difference in the "need of +protection by the two sexes" is rather deceptive, and requires some +discussion. It is obvious that brightly-coloured individuals, whether +males or females, would equally attract, and obscurely-coloured +individuals equally escape, the attention of their enemies. But we are +concerned with the effects of the destruction or preservation of certain +individuals of either sex, on the character of the race. With insects, +after the male has fertilised the female, and after the latter has laid +her eggs, the greater or less immunity from danger of either sex could +not possibly have any effect on the offspring. Before the sexes have +performed their proper functions, if they existed in equal numbers and +if they strictly paired (all other circumstances being the same), the +preservation of the males and females would be equally important for the +existence of the species and for the character of the offspring. But +with most animals, as is known to be the case with the domestic +silk-moth, the male can fertilise two or three females; so that the +destruction of the males would not be so injurious to the species as +that of the females. On the other hand, Dr. Wallace believes that with +moths the progeny from a second or third fertilisation is apt to be +weakly, and therefore would not have so good chance of surviving. When +the males exist in much greater numbers than the females, no doubt many +males might be destroyed with impunity to the species; but I cannot see +that the results of ordinary selection for the sake of protection would +be influenced by the sexes existing in unequal numbers; for the same +proportion of the more conspicuous individuals, whether males or +females, would probably be destroyed. If indeed the males presented a +greater range of variation in colour, the result would be different; but +we need not here follow out such complex details. On the whole I cannot +perceive that an inequality in the numbers of the two sexes would +influence in any marked manner the effects of ordinary selection on the +character of the offspring. + +Female Lepidoptera require, as Mr. Wallace insists, some days to +deposit their fertilised ova and to search for a proper place; during +this period (whilst the life of the male was of no importance) the +brighter-coloured females would be exposed to danger and would be liable +to be destroyed. The duller-coloured females on the other hand would +survive, and thus would influence, it might be thought, in a marked +manner the character of the species,--either of both sexes or of one +sex, according to which form of inheritance prevailed. But it must not +be forgotten that the males emerge from the cocoon-state some days +before the females, and during this period, whilst the unborn females +were safe, the brighter-coloured males would be exposed to danger; so +that ultimately both sexes would probably be exposed during a nearly +equal length of time to danger, and the elimination of conspicuous +colours would not be much more effective in the one than the other sex. + +It is a more important consideration that female Lepidoptera, as Mr. +Wallace remarks, and as is known to every collector, are generally +slower flyers than the males. Consequently the latter, if exposed to +greater danger from being conspicuously coloured, might be able to +escape from their enemies, whilst the similarly-coloured females would +be destroyed; and thus the females would have the most influence in +modifying the colour of their progeny. + +There is one other consideration: bright colours, as far as sexual +selection is concerned, are commonly of no service to the females; so +that if the latter varied in brightness, and the variations were +sexually limited in their transmission, it would depend on mere chance +whether the females had their bright colours increased; and this would +tend throughout the Order to diminish the number of species with +brightly-coloured females in comparison with the species having +brightly-coloured males. On the other hand, as bright colours are +supposed to be highly serviceable to the males in their love-struggles, +the brighter males (as we shall see in the chapter on Birds) although +exposed to rather greater danger, would on an average procreate a +greater number of offspring than the duller males. In this case, if the +variations were limited in their transmission to the male sex, the males +alone would be rendered more brilliantly coloured; but if the variations +were not thus limited, the preservation and augmentation of such +variations would depend on whether more evil was caused to the species +by the females being rendered conspicuous, than good to the males by +certain individuals being successful over their rivals. + +As there can hardly be a doubt that both sexes of many butterflies and +moths have been rendered dull-coloured for the sake of protection, so +it may have been with the females alone of some species in which +successive variations towards dullness first appeared in the female sex +and were from the first limited in their transmission to the same sex. +If not thus limited, both sexes would become dull-coloured. We shall +immediately see, when we treat of mimickry, that the females alone of +certain butterflies have been rendered extremely beautiful for the sake +of protection, without any of the successive protective variations +having been transferred to the male, to whom they could not possibly +have been in the least degree injurious, and therefore could not have +been eliminated through natural selection. Whether in each particular +species, in which the sexes differ in colour, it is the female which has +been specially modified for the sake of protection; or whether it is the +male which has been specially modified for the sake of sexual +attraction, the female having retained her primordial colouring only +slightly changed through the agencies before alluded to; or whether +again both sexes have been modified, the female for protection and the +male for sexual attraction, can only be definitely decided when we know +the life-history of each species. + +Without distinct evidence, I am unwilling to admit that a double process +of selection has long been going on with a multitude of species,--the +males having been rendered more brilliant by beating their rivals; and +the females more dull-coloured by having escaped from their enemies. We +may take as an instance the common brimstone butterfly (Gonepteryx), +which appears early in the spring before any other kind. The male of +this species is of a far more intense yellow than the female, though she +is almost equally conspicuous; and in this case it does not seem +probable that she specially acquired her pale tints as a protection, +though it is probable that the male acquired his bright colours as a +sexual attraction. The female of _Anthocharis cardamines_ does not +possess the beautiful orange tips to her wings with which the male is +ornamented; consequently she closely resembles the white butterflies +(Pieris) so common in our gardens; but we have no evidence that this +resemblance is beneficial. On the contrary, as she resembles both sexes +of several species of the same genus inhabiting various quarters of the +world, it is more probable that she has simply retained to a large +extent her primordial colours. + +Various facts support the conclusion that with the greater number of +brilliantly-coloured Lepidoptera, it is the male which has been +modified; the two sexes having come to differ from each other, or to +resemble each other, according to which form of inheritance has +prevailed. Inheritance is governed by so many unknown laws or +conditions, that they seem to us to be most capricious in their +action;[524] and we can so far understand how it is that with +closely-allied species the sexes of some differ to an astonishing +degree, whilst the sexes of others are identical in colour. As the +successive steps in the process of variation are necessarily all +transmitted through the female, a greater or less number of such steps +might readily become developed in her; and thus we can understand the +frequent gradations from an extreme difference to no difference at all +between the sexes of the species within the same group. These cases of +gradation are much too common to favour the supposition that we here see +females actually undergoing the process of transition and losing their +brightness for the sake of protection; for we have every reason to +conclude that at any one time the greater number of species are in a +fixed condition. With respect to the differences between the females of +the species in the same genus or family, we can perceive that they +depend, at least in part, on the females partaking of the colours of +their respective males. This is well illustrated in those groups in +which the males are ornamented to an extraordinary degree; for the +females in these groups generally partake to a certain extent of the +splendour of their male partners. Lastly, we continually find, as +already remarked, that the females of almost all the species in the same +genus, or even family, resemble each other much more closely in colour +than do the males; and this indicates that the males have undergone a +greater amount of modification than the females. + + +_Mimickry._--This principle was first made clear in an admirable paper +by Mr. Bates,[525] who thus threw a flood of light on many obscure +problems. It had previously been observed that certain butterflies in S. +America belonging to quite distinct families, resembled the Heliconid +so closely in every stripe and shade of colour that they could not be +distinguished except by an experienced entomologist. As the Heliconid +are coloured in their usual manner, whilst the others depart from the +usual colouring of the groups to which they belong, it is clear that the +latter are the imitators, and the Heliconid the imitated. Mr. Bates +further observed that the imitating species are comparatively rare, +whilst the imitated swarm in large numbers; the two sets living mingled +together. From the fact of the Heliconid being conspicuous and +beautiful insects, yet so numerous in individuals and species, he +concluded that they must be protected from the attacks of birds by some +secretion or odour; and this hypothesis has now been confirmed by a +considerable body of curious evidence.[526] From these considerations +Mr. Bates inferred that the butterflies which imitate the protected +species had acquired their present marvellously deceptive appearance, +through variation and natural selection, in order to be mistaken for the +protected kinds and thus to escape being devoured. No explanation is +here attempted of the brilliant colours of the imitated, but only of the +imitating butterflies. We must account for the colours of the former in +the same general manner, as in the cases previously discussed in this +chapter. Since the publication of Mr. Bates' paper, similar and equally +striking facts have been observed by Mr. Wallace[527] in the Malayan +region, and by Mr. Trimen in South Africa. + +As some writers[528] have felt much difficulty in understanding how the +first steps in the process of mimickry could have been effected through +natural selection, it may be well to remark that the process probably +has never commenced with forms widely dissimilar in colour. But with two +species moderately like each other, the closest resemblance if +beneficial to either form could readily be thus gained; and if the +imitated form was subsequently and gradually modified through sexual +selection or any other means, the imitating form would be led along the +same track, and thus be modified to almost any extent, so that it might +ultimately assume an appearance or colouring wholly unlike that of the +other members of the group to which it belonged. As extremely slight +variations in colour would not in many cases suffice to render a species +so like another protected species as to lead to its preservation, it +should be remembered that many species of Lepidoptera are liable to +considerable and abrupt variations in colour. A few instances have been +given in this chapter; but under this point of view Mr. Bates' original +paper on mimickry, as well as Mr. Wallace's papers, should be consulted. + +In the foregoing cases both sexes of the imitating species +resemble the imitated; but occasionally the female alone mocks a +brilliantly-coloured and protected species inhabiting the same district. +Consequently the female differs in colour from her own male, and, which +is a rare and anomalous circumstance, is the more brightly-coloured of +the two. In all the few species of Pierid, in which the female is more +conspicuously coloured than the male, she imitates, as I am informed by +Mr. Wallace, some protected species inhabiting the same region. The +female of _Diadema anomala_ is rich purple-brown with almost the whole +surface glossed with satiny blue, and she closely imitates the +_Euploea midamus_, "one of the commonest butterflies of the East;" +whilst the male is bronzy or olive-brown, with only a slight blue gloss +on the outer parts of the wings.[529] Both sexes of this Diadema and of +_D. bolina_ follow the same habits of life, so that the differences in +colour between the sexes cannot be accounted for by exposure to +different conditions;[530] even if this explanation were admissible in +other instances.[531] + +The above cases of female butterflies which are more brightly-coloured +than the males, shew us, firstly, that variations have arisen in a state +of nature in the female sex, and have been transmitted exclusively, or +almost exclusively, to the same sex; and, secondly, that this form of +inheritance has not been determined through natural selection. For if we +assume that the females, before they became brightly coloured in +imitation of some protected kind, were exposed during each season for a +longer period to danger than the males; or if we assume that they could +not escape so swiftly from their enemies, we can understand how they +alone might originally have acquired through natural selection and +sexually-limited inheritance their present protective colours. But +except on the principle of these variations having been transmitted +exclusively to the female offspring, we cannot understand why the males +should have remained dull-coloured; for it would surely not have been in +any way injurious to each individual male to have partaken by +inheritance of the protective colours of the female, and thus to have +had a better chance of escaping destruction. In a group in which +brilliant colours are so common as with butterflies, it cannot be +supposed that the males have been kept dull-coloured through sexual +selection by the females rejecting the individuals which were rendered +as beautiful as themselves. We may, therefore, conclude that in these +cases inheritance by one sex is not due to the modification through +natural selection of a tendency to equal inheritance by both sexes. + +It may be well here to give an analogous case in another Order, of +characters acquired only by the female, though not in the least +injurious, as far as we can judge, to the male. Amongst the Phasmid, or +spectre-insects, Mr. Wallace states that "it is often the females alone +that so strikingly resemble leaves, while the males show only a rude +approximation." Now, whatever may be the habits of these insects, it is +highly improbable that it could be disadvantageous to the males to +escape detection by resembling leaves.[532] Hence we may conclude that +the females alone in this latter as in the previous cases originally +varied in certain characters; these characters having been preserved and +augmented through ordinary selection for the sake of protection and from +the first transmitted to the female offspring alone. + + +_Bright Colours of Caterpillars._--Whilst reflecting on the beauty of +many butterflies, it occurred to me that some caterpillars were +splendidly coloured, and as sexual selection could not possibly have +here acted, it appeared rash to attribute the beauty of the mature +insect to this agency, unless the bright colours of their larv could +be in some manner explained. In the first place it may be observed that +the colours of caterpillars do not stand in any close correlation with +those of the mature insect. Secondly, their bright colours do not serve +in any ordinary manner as a protection. As an instance of this, Mr. +Bates informs me that the most conspicuous caterpillar which he ever +beheld (that of a Sphinx) lived on the large green leaves of a tree on +the open llanos of South America; it was about four inches in length, +transversely banded with black and yellow, and with its head, legs, and +tail of a bright red. Hence it caught the eye of any man who passed by +at the distance of many yards, and no doubt of every passing bird. + +I then applied to Mr. Wallace, who has an innate genius for solving +difficulties. After some consideration he replied: "Most caterpillars +require protection, as may be inferred from some kinds being furnished +with spines or irritating hairs, and from many being coloured green like +the leaves on which they feed, or curiously like the twigs of the trees +on which they live." I may add as another instance of protection, that +there is a caterpillar of a moth, as I am informed by Mr. J. Mansel +Weale, which lives on the mimosas in South Africa, and fabricates for +itself a case, quite undistinguishable from the surrounding thorns. From +such considerations Mr. Wallace thought it probable that +conspicuously-coloured caterpillars were protected by having a nauseous +taste; but as their skin is extremely tender, and as their intestines +readily protrude from a wound, a slight peck from the beak of a bird +would be as fatal to them as if they had been devoured. Hence, as Mr. +Wallace remarks, "distastefulness alone would be insufficient to protect +a caterpillar unless some outward sign indicated to its would-be +destroyer that its prey was a disgusting morsel." Under these +circumstances it would be highly advantageous to a caterpillar to be +instantaneously and certainly recognised as unpalatable by all birds and +other animals. Thus the most gaudy colours would be serviceable, and +might have been gained by variation and the survival of the most +easily-recognised individuals. + +This hypothesis appears at first sight very bold; but when it was +brought before the Entomological Society[533] it was supported by +various statements; and Mr. J. Jenner Weir, who keeps a large number of +birds in an aviary, has made, as he informs me, numerous trials, and +finds no exception to the rule, that all caterpillars of nocturnal and +retiring habits with smooth skins, all of a green colour, and all which +imitate twigs, are greedily devoured by his birds. The hairy and spinose +kinds are invariably rejected, as were four conspicuously-coloured +species. When the birds rejected a caterpillar, they plainly shewed, by +shaking their heads and cleansing their beaks, that they were disgusted +by the taste.[534] Three conspicuous kinds of caterpillars and moths +were also given by Mr. A. Butler to some lizards and frogs, and were +rejected; though other kinds were eagerly eaten. Thus the probable truth +of Mr. Wallace's view is confirmed, namely, that certain caterpillars +have been made conspicuous for their own good, so as to be easily +recognised by their enemies, on nearly the same principle that certain +poisons are coloured by druggists for the good of man. This view will, +it is probable, be hereafter extended to many animals, which are +coloured in a conspicuous manner. + + +_Summary and Concluding Remarks on Insects._--Looking back to the +several Orders, we have seen that the sexes often differ in various +characters, the meaning of which is not understood. The sexes, also, +often differ in their organs of sense or locomotion, so that the males +may quickly discover or reach the females, and still oftener in the +males possessing diversified contrivances for retaining the females when +found. But we are not here much concerned with sexual differences of +these kinds. + +In almost all the Orders, the males of some species, even of weak and +delicate kinds, are known to be highly pugnacious; and some few are +furnished with special weapons for fighting with their rivals. But the +law of battle does not prevail nearly so widely with insects as with the +higher animals. Hence probably it is that the males have not often been +rendered larger and stronger than the females. On the contrary they are +usually smaller, in order that they may be developed within a shorter +time, so as to be ready in large numbers for the emergence of the +females. + +In two families of the Homoptera the males alone possess, in an +efficient state, organs which may be called vocal; and in three families +of the Orthoptera the males alone possess stridulating organs. In both +cases these organs are incessantly used during the breeding-season, not +only for calling the females, but for charming or exciting them in +rivalry with other males. No one who admits the agency of natural +selection, will dispute that these musical instruments have been +acquired through sexual selection. In four other Orders the members of +one sex, or more commonly of both sexes, are provided with organs for +producing various sounds, which apparently serve merely as call-notes. +Even when both sexes are thus provided, the individuals which were able +to make the loudest or most continuous noise would gain partners before +those which were less noisy, so that their organs have probably been +gained through sexual selection. It is instructive to reflect on the +wonderful diversity of the means for producing sound, possessed by the +males alone or by both sexes in no less than six Orders, and which were +possessed by at least one insect at an extremely remote geological +epoch. We thus learn how effectual sexual selection has been in leading +to modifications of structure, which sometimes, as with the Homoptera, +are of an important nature. + +From the reasons assigned in the last chapter, it is probable that the +great horns of the males of many lamellicorn, and some other beetles, +have been acquired as ornaments. So perhaps it may be with certain other +peculiarities confined to the male sex. From the small size of insects, +we are apt to undervalue their appearance. If we could imagine a male +Chalcosoma (fig. 15) with its polished, bronzed coat of mail, and vast +complex horns, magnified to the size of a horse or even of a dog, it +would be one of the most imposing animals in the world. + +The colouring of insects is a complex and obscure subject. When the male +differs slightly from the female, and neither are brilliantly coloured, +it is probable that the two sexes have varied in a slightly different +manner, with the variations transmitted to the same sex, without any +benefit having been thus derived or evil suffered. When the male is +brilliantly coloured and differs conspicuously from the female, as with +some dragon-flies and many butterflies, it is probable that he alone has +been modified, and that he owes his colours to sexual selection; whilst +the female has retained a primordial or very ancient type of colouring, +slightly modified by the agencies before explained, and has therefore +not been rendered obscure, at least in most cases, for the sake of +protection. But the female alone has sometimes been coloured +brilliantly so as to imitate other protected species inhabiting the same +district. When the sexes resemble each other and both are obscurely +coloured, there is no doubt that they have been in a multitude of cases +coloured for the sake of protection. So it is in some instances when +both are brightly coloured, causing them to resemble surrounding objects +such as flowers, or other protected species, or indirectly by giving +notice to their enemies that they are of an unpalatable nature. In many +other cases in which the sexes resemble each other and are brilliantly +coloured, especially when the colours are arranged for display, we may +conclude that they have been gained by the male sex as an attraction, +and have been transferred to both sexes. We are more especially led to +this conclusion whenever the same type of coloration prevails throughout +a group, and we find that the males of some species differ widely in +colour from the females, whilst both sexes of other species are quite +alike, with intermediate gradations connecting these extreme states. + +In the same manner as bright colours have often been partially +transferred from the males to the females, so it has been with the +extraordinary horns of many lamellicorn and some other beetles. So, +again, the vocal or instrumental organs proper to the males of the +Homoptera and Orthoptera have generally been transferred in a +rudimentary, or even in a nearly perfect condition to the females; yet +not sufficiently perfect to be used for producing sound. It is also an +interesting fact, as bearing on sexual selection, that the stridulating +organs of certain male Orthoptera are not fully developed until the last +moult; and that the colours of certain male dragon-flies are not fully +developed until some little time after their emergence from the pupal +state, and when they are ready to breed. + +Sexual selection implies that the more attractive individuals are +preferred by the opposite sex; and as with insects, when the sexes +differ, it is the male which, with rare exceptions, is the most +ornamented and departs most from the type to which the species +belongs;--and as it is the male which searches eagerly for the female, +we must suppose that the females habitually or occasionally prefer the +more beautiful males, and that these have thus acquired their beauty. +That in most or all the orders the females have the power of rejecting +any particular male, we may safely infer from the many singular +contrivances possessed by the males, such as great jaws, adhesive +cushions, spines, elongated legs, &c., for seizing the female; for these +contrivances shew that there is some difficulty in the act. In the case +of unions between distinct species, of which many instances have been +recorded, the female must have been a consenting party. Judging from +what we know of the perceptive powers and affections of various insects, +there is no antecedent improbability in sexual selection having come +largely into action; but we have as yet no direct evidence on this head, +and some facts are opposed to the belief. Nevertheless, when we see many +males pursuing the same female, we can hardly believe that the pairing +is left to blind chance--that the female exerts no choice, and is not +influenced by the gorgeous colours or other ornaments, with which the +male alone is decorated. + +If we admit that the females of the Homoptera and Orthoptera appreciate +the musical tones emitted by their male partners, and that the various +instruments for this purpose have been perfected through sexual +selection, there is little improbability in the females of other insects +appreciating beauty in form or colour, and consequently in such +characters having been thus gained by the males. But from the +circumstance of colour being so variable, and from its having been so +often modified for the sake of protection, it is extremely difficult to +decide in how large a proportion of cases sexual selection has come into +play. This is more especially difficult in those Orders, such as the +Orthoptera, Hymenoptera, and Coleoptera, in which the two sexes rarely +differ much in colour; for we are thus cut off from our best evidence of +some relation between the reproduction of the species and colour. With +the Coleoptera, however, as before remarked, it is in the great +lamellicorn group, placed by some authors at the head of the Order, and +in which we sometimes see a mutual attachment between the sexes, that we +find the males of some species possessing weapons for sexual strife, +others furnished with wonderful horns, many with stridulating organs, +and others ornamented with splendid metallic tints. Hence it seems +probable that all these characters have been gained through the same +means, namely sexual selection. + +When we treat of Birds, we shall see that they present in their +secondary sexual characters the closest analogy with insects. Thus, many +male birds are highly pugnacious, and some are furnished with special +weapons for fighting with their rivals. They possess organs which are +used during the breeding-season for producing vocal and instrumental +music. They are frequently ornamented with combs, horns, wattles and +plumes of the most diversified kinds, and are decorated with beautiful +colours, all evidently for the sake of display. We shall find that, as +with insects, both sexes, in certain groups, are equally beautiful, and +are equally provided with ornaments which are usually confined to the +male sex. In other groups both sexes are equally plain-coloured and +unornamented. Lastly, in some few anomalous cases, the females are more +beautiful than the males. We shall often find, in the same group of +birds, every gradation from no difference between the sexes, to an +extreme difference. In the latter case we shall see that the females, +like female insects, often possess more or less plain traces of the +characters which properly belong to the males. The analogy, indeed, in +all these respects between birds and insects, is curiously close. +Whatever explanation applies to the one class probably applies to the +other; and this explanation, as we shall hereafter attempt to shew, is +almost certainly sexual selection. + + + FOOTNOTES: + + [1] As the works of the first-named authors are so well known, + I need not give the titles; but as those of the latter are less + well known in England, I will give them:--'Sechs Vorlesungen + ber die Darwin'sche Theorie:' zweite Auflage, 1868, von Dr. L. + Bchner; translated into French under the title 'Confrences + sur la Thorie Darwinienne,' 1869. 'Der Mensch, im Lichte der + Darwin'sche Lehre,' 1865, von Dr. F. Rolle. I will not attempt + to give references to all the authors who have taken the same + side of the question. Thus G. Canestrini has published + ('Annuario della Soc. d. Nat.,' Modena, 1867, p. 81) a very + curious paper on rudimentary characters, as bearing on the + origin of man. Another work has (1869) been published by Dr. + Barrago Francesco, bearing in Italian the title of "Man, made + in the image of God, was also made in the image of the ape." + + [2] Prof. Hckel is the sole author who, since the publication + of the 'Origin,' has discussed, in his various works, in a very + able manner, the subject of sexual selection, and has seen its + full importance. + + [3] 'Grosshirnwindungen des Menschen,' 1868, s. 96. + + [4] 'Le. sur la Phys.' 1866, p. 890, as quoted by M. Dally, + 'L'Ordre des Primates et le Transformisme,' 1868, p. 29. + + [5] 'Naturgeschichte der Sugethiere von Paraguay,' 1830, s. + 50. + + [6] Brehm, 'Thierleben,' B. i. 1864, s. 75, 86. On the Ateles, + s. 105. For other analogous statements, see s. 25, 107. + + [7] With respect to insects see Dr. Laycock 'On a General Law + of Vital Periodicity,' British Association, 1842. Dr. + Macculloch, 'Silliman's North American Journal of Science,' + vol. xvii. p. 305, has seen a dog suffering from tertian ague. + + [8] I have given the evidence on this head in my 'Variation of + Animals and Plants under Domestication,' vol. ii. p. 15. + + [9] "Mares e diversis generibus Quadrumanorum sine dubio + dignoscunt feminas humanas a maribus. Primum, credo, odoratu, + postea aspectu. Mr. Youatt, qui diu in Hortis Zoologicis + (Bestiariis) medicus animalium erat, vir in rebus observandis + cautus et sagax, hoc mihi certissime probavit, et curatores + ejusdem loci et alii e ministris confirmaverunt. Sir Andrew + Smith et Brehm notabant idem in Cynocephalo. Illustrissimus + Cuvier etiam narrat multa de hac re qu ut opinor nihil turpius + potest indicari inter omnia hominibus et Quadrumanis communia. + Narrat enim Cynocephalum quendam in furorem incidere aspectu + feminarum aliquarum, sed nequaquam accendi tanto furore ab + omnibus. Semper eligebat juniores, et dignoscebat in turba, et + advocabat voce gestuque." + + [10] This remark is made with respect to Cynocephalus and the + anthropomorphous apes by Geoffroy Saint-Hilaire and F. Cuvier, + 'Hist. Nat. des Mammifres,' tom. i. 1824. + + [11] Huxley, 'Man's Place in Nature,' 1863, p. 34. + + [12] 'Man's Place in Nature,' 1863, p. 67. + + [13] The human embryo (upper fig.) is from Ecker, 'Icones + Phys.,' 1851-1859, tab. xxx. fig. 2. This embryo was ten lines + in length, so that the drawing is much magnified. The embryo of + the dog is from Bischoff, 'Entwicklungsgeschichte des + Hunde-Eies,' 1845, tab. xi. fig. 42 B. This drawing is five + times magnified, the embryo being 25 days old. The internal + viscera have been omitted, and the uterine appendages in both + drawings removed. I was directed to these figures by Prof. + Huxley, from whose work, 'Man's Place in Nature.' the idea of + giving them was taken. Hckel has also given analogous drawings + in his 'Schpfungsgeschichte.' + + [14] Prof. Wyman in 'Proc. of American Acad. of Sciences,' vol. + iv. 1860, p. 17. + + [15] Owen, 'Anatomy of Vertebrates,' vol. i. p. 533. + + [16] 'Die Grosshirnwindungen des Menschen,' 1868, s. 95. + + [17] 'Anatomy of Vertebrates,' vol. ii. p. 553. + + [18] 'Proc. Soc. Nat. Hist.' Boston, 1863, vol. ix. p. 185. + + [19] 'Man's Place in Nature,' p. 65. + + [20] I had written a rough copy of this chapter before reading + a valuable paper, "Caratteri rudimentali in ordine all'origine + del uomo" ('Annuario della Soc. d. Nat.,' Modena, 1867, p. 81), + by G. Canestrini, to which paper I am considerably indebted. + Hckel has given admirable discussions on this whole subject, + under the title of Dysteleology, in his 'Generelle Morphologie' + and 'Schpfungsgeschichte.' + + [21] Some good criticisms on this subject have been given by + Messrs. Murie and Mivart, in 'Transact. Zoolog. Soc.' 1869, + vol. vii. p. 92. + + [22] 'Variation of Animals and Plants under Domestication,' + vol. ii. pp. 317 and 397. See also 'Origin of Species,' 5th + edit. p. 535. + + [23] For instance M. Richard ('Annales des Sciences Nat.' 3rd + series, Zoolog. 1852, tom. xviii. p. 13) describes and figures + rudiments of what he calls the "muscle pdieux de la main," + which he says is sometimes "infiniment petit." Another muscle, + called "le tibial postrieur," is generally quite absent in the + hand, but appears from time to time in a more or less + rudimentary condition. + + [24] Prof. W. Turner, 'Proc. Royal Soc. Edinburgh,' 1866-67, p. + 65. + + [25] Canestrini quotes Hyrt. ('Annuario della Soc. dei + Naturalisti,' Modena, 1867, p. 97) to the same effect. + + [26] 'The Diseases of the Ear,' by J. Toynbee, F.R.S., 1860, p. + 12. + + [27] See also some remarks, and the drawings of the ears of the + Lemuroidea, in Messrs. Murie and Mivart's excellent paper in + 'Transact. Zoolog. Soc.' vol. vii. 1869, pp. 6 and 90. + + [28] Mller's 'Elements of Physiology,' Eng. translat., 1842, + vol. ii. p. 1117. Owen, 'Anatomy of Vertebrates,' vol. iii. p. + 260; ibid. on the Walrus, 'Proc. Zoolog. Soc.' November 8th, + 1854. See also R. Knox, 'Great Artists and Anatomists,' p. 106. + This rudiment apparently is somewhat larger in Negroes and + Australians than in Europeans, see Carl Vogt, 'Lectures on + Man,' Eng. translat. p. 129. + + [29] 'The Physiology and Pathology of Mind,' 2nd edit. 1868, p. + 134. + + [30] Eschricht, Ueber die Richtung der Haare am menschlichen + Krper 'Mllers Archiv fr Anat. und Phys.' 1837, s. 47. I + shall often have to refer to this very curious paper. + + [31] Paget, 'Lectures on Surgical Pathology,' 1853, vol. i. p. + 71. + + [32] Eschricht, ibid. s. 40, 47. + + [33] Dr. Webb, 'Teeth in Man and the Anthropoid Apes,' as + quoted by Dr. C. Carter Blake in 'Anthropological Review,' + July, 1867, p. 299. + + [34] Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 320, 321, + and 325. + + [35] 'On the Primitive Form of the Skull,' Eng. translat. in + 'Anthropological Review,' Oct. 1868, p. 426. + + [36] Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 416, 434, + 441. + + [37] 'Annuario della Soc. d. Nat.' Modena, 1867, p. 94. + + [38] M. C. Martins ("De l'Unit Organique," in 'Revue des Deux + Mondes,' June 15, 1862, p. 16), and Hckel ('Generelle + Morphologie,' B. ii. s. 278), have both remarked on the + singular fact of this rudiment sometimes causing death. + + [39] 'The Lancet,' Jan. 24, 1863, p. 83. Dr. Knox, 'Great + Artists and Anatomists,' p. 63. See also an important memoir on + this process by Dr. Grube, in the 'Bulletin de l'Acad. Imp. de + St. Ptersbourg,' tom. xii. 1867, p. 448. + + [40] "On the Caves of Gibraltar," 'Transact. Internat. Congress + of Prehist. Arch.' Third Session, 1869, p. 54. + + [41] Quatrefages has lately collected the evidence on this + subject. 'Revue des Cours Scientifiques,' 1867-1868, p. 625. + + [42] Owen, 'On the Nature of Limbs,' 1849, p. 114. + + [43] Leuckart, in Todd's 'Cyclop. of Anat.' 1849-52, vol. iv. + p. 1415. In man this organ is only from three to six lines in + length, but, like so many other rudimentary parts, it is + variable in development as well as in other characters. + + [44] See, on this subject, Owen, 'Anatomy of Vertebrates,' vol. + iii. pp. 675, 676, 706. + + [45] See the evidence on these points, as given by Lubbock, + 'Prehistoric Times,' p. 354, &c. + + [46] 'L'Instinct chez les Insectes.' 'Revue des Deux Mondes,' + Feb. 1870, p. 690. + + [47] 'The American Beaver and his Works,' 1868. + + [48] 'The Principles of Psychology,' 2nd edit. 1870, pp. + 418-443. + + [49] 'Contributions to the Theory of Natural Selection,' 1870, + p. 212 + + [50] 'Recherches sur les Moeurs des Fourmis,' 1810, p. 173. + + [51] All the following statements, given on the authority of + these two naturalists, are taken from Rengger's 'Naturges. der + Sugethiere von Paraguay,' 1830, s. 41-57, and from Brehm's + 'Thierleben,' B. i. s. 10-87. + + [52] 'Bridgewater Treatise,' p. 263. + + [53] W. C. L. Martin, 'Nat. Hist. of Mammalia,' 1841, p. 405. + + [54] Quoted by Vogt, 'Mmoire sur les Microcphales,' 1867, p. + 168. + + [55] 'The Variation of Animals and Plants under Domestication,' + vol. i. p. 27. + + [56] 'Les Moeurs des Fourmis,' 1810, p. 150. + + [57] Quoted in Dr. Maudsley's 'Physiology and Pathology of + Mind,' 1868, pp. 19, 220. + + [58] Dr. Jerdon, 'Birds of India,' vol. i. 1862, p. xxi. + + [59] Mr. L. H. Morgan's work on 'The American Beaver,' 1868, + offers a good illustration of this remark. I cannot, however, + avoid thinking that he goes too far in underrating the power of + Instinct. + + [60] 'The Moor and the Loch,' p. 45. Col. Hutchinson on 'Dog + Breaking,' 1850, p. 46. + + [61] 'Personal Narrative,' Eng. translat., vol. iii. p. 106. + + [62] Quoted by Sir C. Lyell, 'Antiquity of Man,' p. 497. + + [63] 'Journal of Researches during the Voyage of the "Beagle,"' + 1845, p. 398. 'Origin of Species,' 5th edit. p. 260. + + [64] 'Lettres Phil. sur l'Intelligence des Animaux,' nouvelle + edit. 1802, p. 86. + + [65] See the evidence on this head in chap. i. vol. i. 'On the + Variation of Animals and Plants under Domestication.' + + [66] 'Proc. Zoolog. Soc.' 1864, p. 186. + + [67] Savage and Wyman in 'Boston Journal of Nat. Hist.' vol. + iv. 1843-44, p. 383. + + [68] 'Sugethiere von Paraguay,' 1830, s. 51-56. + + [69] 'Thierleben,' B. i. s. 79, 82. + + [70] 'The Malay Archipelago,' vol. i. 1869, p. 87. + + [71] 'Primeval Man,' 1869, pp. 145, 147. + + [72] 'Prehistoric Times,' 1865, p. 473, &c. + + [73] Quoted in 'Anthropological Review,' 1864, p. 158. + + [74] Rengger, ibid. s. 45. + + [75] See my 'Variation of Animals and Plants under + Domestication,' vol. i. p. 27. + + [76] See a discussion on this subject in Mr. E. B. Tylor's very + interesting work, 'Researches into the Early History of + Mankind,' 1865, chaps. ii. to iv. + + [77] Hon. Daines Barrington in 'Philosoph. Transactions,' 1773, + p. 262. See also Dureau de la Malle, in 'Ann. des Sc. Nat.' 3rd + series, Zoolog. tom. x. p. 119. + + [78] 'On the Origin of Language,' by H. Wedgwood, 1866. + 'Chapters on Language,' by the Rev. F. W. Farrar, 1865. These + works are most interesting. See also 'De la Phys. et de + Parole,' par Albert Lemoine, 1865, p. 190. The work on this + subject, by the late Prof. Aug. Schleicher, has been translated + by Dr. Bikkers into English, under the title of 'Darwinism + tested by the Science of Language,' 1869. + + [79] Vogt, 'Mmoire sur les Microcphales,' 1867, p. 169. With + respect to savages, I have given some facts in my 'Journal of + Researches,' &c., 1845, p. 206. + + [80] See clear evidence on this head in the two works so often + quoted, by Brehm and Rengger. + + [81] See remarks on this head by Dr. Maudsley, 'The Physiology + and Pathology of Mind,' 2nd edit. 1868, p. 199. + + [82] Many curious cases have been recorded. See, for instance, + 'Inquiries Concerning the Intellectual Powers,' by Dr. + Abercrombie, 1838, p. 150. + + [83] 'The Variation of Animals and Plants under Domestication,' + vol. ii. p. 6. + + [84] See some good remarks to this effect by Dr. Maudsley, 'The + Physiology and Pathology of Mind,' 1808, p. 199. + + [85] Macgillivray, 'Hist. of British Birds,' vol. ii. 1839, p. + 29. An excellent observer, Mr. Blackwall, remarks that the + magpie learns to pronounce single words, and even short + sentences, more readily than almost any other British bird; + yet, as he adds, after long and closely investigating its + habits, he has never known it, in a state of nature, display + any unusual capacity for imitation. 'Researches in Zoology,' + 1834, p. 158. + + [86] See the very interesting parallelism between the + development of species and languages, given by Sir C. Lyell in + 'The Geolog. Evidences of the Antiquity of Man,' 1863, chap. + xxiii. + + [87] See remarks to this effect by the Rev. F. W. Farrar, in an + interesting article, entitled "Philology and Darwinism" in + 'Nature,' March 24th, 1870, p. 528. + + [88] 'Nature,' Jan. 6th, 1870, p. 257. + + [89] Quoted by C. S. Wake, 'Chapters on Man,' 1868, p. 101. + + [90] Buckland, 'Bridgewater Treatise,' p. 411. + + [91] See some good remarks on the simplification of languages, + by Sir J. Lubbock, 'Origin of Civilisation,' 1870, p. 278. + + [92] 'Confrences sur la Thorie Darwinienne,' French + translat., 1869, p. 132. + + [93] The Rev. Dr. J. M'Cann, 'Anti-Darwinism,' 1869, p. 13. + + [94] 'The Spectator,' Dec. 4th, 1869, p. 1430. + + [95] See an excellent article on this subject by the Rev. F. W. + Farrar, in the 'Anthropological Review,' Aug. 1864, p. ccxvii. + For further facts see Sir J. Lubbock, 'Prehistoric Times,' 2nd + edit. 1869. p. 564; and especially the chapters on Religion in + his 'Origin of Civilisation,' 1870. + + [96] The Worship of Animals and Plants, in the 'Fortnightly + Review,' Oct. 1, 1869, p. 422. + + [97] Tylor, 'Early History of Mankind,' 1865, p. 6. See also + the three striking chapters on the Development of Religion, in + Lubbock's 'Origin of Civilisation,' 1870. In a like manner Mr. + Herbert Spencer, in his ingenious essay in the 'Fortnightly + Review' (May 1st, 1870, p. 535), accounts for the earliest + forms of religious belief throughout the world, by man being + led through dreams, shadows, and other causes, to look at + himself as a double essence, corporeal and spiritual. As the + spiritual being is supposed to exist after death and to be + powerful, it is propitiated by various gifts and ceremonies, + and its aid invoked. He then further shews that names or + nicknames given from some animal or other object to the early + progenitors or founders of a tribe, are supposed after a long + interval to represent the real progenitor of the tribe; and + such animal or object is then naturally believed still to exist + as a spirit, is held sacred, and worshipped as a god. + Nevertheless I cannot but suspect that there is a still earlier + and ruder stage, when anything which manifests power or + movement is thought to be endowed with some form of life, and + with mental faculties analogous to our own. + + [98] See an able article on the Psychical Elements of Religion, + by Mr. L. Owen Pike, in 'Anthropolog. Review,' April, 1870, p. + lxiii. + + [99] 'Religion, Moral, &c., der Darwin'schen Art-Lehre,' 1869, + s. 53. + + [100] 'Prehistoric Times,' 2nd edit. p. 571. In this work (at + p. 553) there will be found an excellent account of the many + strange and capricious customs of savages. + + [101] See, for instance, on this subject, Quatrefages, 'Unit + de l'Espce Humaine,' 1861, p. 21, &c. + + [102] 'Dissertation on Ethical Philosophy,' 1837, p. 231, &c. + + [103] 'Metaphysics of Ethics,' translated by J. W. Semple, + Edinburgh, 1836, p. 136. + + [104] Mr. Bain gives a list ('Mental and Moral Science,' 1868, + p. 543-725) of twenty-six British authors who have written on + this subject, and whose names are familiar to every reader; to + these, Mr. Bain's own name, and those of Mr. Lecky, Mr. + Shadworth Hodgson, and Sir J. Lubbock, as well as of others, + may be added. + + [105] Sir B. Brodie, after observing that man is a social + animal ('Psychological Enquiries,' 1854, p. 192), asks the + pregnant question, "ought not this to settle the disputed + question as to the existence of a moral sense?" Similar ideas + have probably occurred to many persons, as they did long ago to + Marcus Aurelius. Mr. J. S. Mill speaks, in his celebrated work, + 'Utilitarianism,' (1864, p. 46), of the social feelings as a + "powerful natural sentiment," and as "the natural basis of + sentiment for utilitarian morality;" but on the previous page + he says, "if, as is my own belief, the moral feelings are not + innate, but acquired, they are not for that reason less + natural." It is with hesitation that I venture to differ from + so profound a thinker, but it can hardly be disputed that the + social feelings are instinctive or innate in the lower animals; + and why should they not be so in man? Mr. Bain (see, for + instance, 'The Emotions and the Will,' 1865, p. 481) and others + believe that the moral sense is acquired by each individual + during his lifetime. On the general theory of evolution this is + at least extremely improbable. + + [106] 'Die Darwin'sche Theorie,' s. 101. + + [107] Mr. R. Brown in 'Proc. Zoolog. Soc.' 1868, p. 409. + + [108] Brehm, 'Thierleben,' B. i. 1864, s. 52, 79. For the case + of the monkeys extracting thorns from each other, see s. 54. + With respect to the Hamadryas turning over stones, the fact is + given (s. 76) on the evidence of Alvarez, whose observations + Brehm thinks quite trustworthy. For the cases of the old male + baboons attacking the dogs, see s. 79; and with respect to the + eagle, s. 56. + + [109] 'Annals and Mag. of Nat. Hist.' November, 1868, p. 382. + + [110] Sir J. Lubbock, 'Prehistoric Times,' 2nd edit. p. 446. + + [111] As quoted by Mr. L. H. Morgan, 'The American Beaver,' + 1868, p. 272. Capt. Stansbury also gives an interesting account + of the manner in which a very young pelican, carried away by a + strong stream, was guided and encouraged in its attempts to + reach the shore by half a dozen old birds. + + [112] As Mr. Bain states, "effective aid to a sufferer springs + from sympathy proper:" 'Mental and Moral Science,' 1868, p. + 245. + + [113] 'Thierleben,' B. i. s. 85. + + [114] 'De l'Espce et de la Class.' 1869, p. 97. + + [115] 'Der Darwin'schen Art-Lehre,' 1869, s. 54. + + [116] Brehm, 'Thierleben,' B. i. s. 76. + + [117] See the first and striking chapter in Adam Smith's + 'Theory of Moral Sentiments.' Also Mr. Bain's 'Mental and Moral + Science,' 1868, p. 244, and 275-282. Mr. Bain states, that + "sympathy is, indirectly, a source of pleasure to the + sympathiser;" and he accounts for this through reciprocity. He + remarks that "the person benefited, or others in his stead, may + make up, by sympathy and good offices returned, for all the + sacrifice." But if, as appears to be the case, sympathy is + strictly an instinct, its exercise would give direct pleasure, + in the same manner as the exercise, as before remarked, of + almost every other instinct. + + [118] This fact, the Rev. L. Jenyns states (see his edition of + 'White's Nat. Hist. of Selborne,' 1853, p. 204) was first + recorded by the illustrious Jenner, in 'Phil. Transact.' 1824, + and has since been confirmed by several observers, especially + by Mr. Blackwall. This latter careful observer examined, late + in the autumn, during two years, thirty-six nests; he found + that twelve contained young dead birds, five contained eggs on + the point of being hatched, and three eggs not nearly hatched. + Many birds not yet old enough for a prolonged flight are + likewise deserted and left behind. See Blackwall, 'Researches + in Zoology,' 1834, pp. 108, 118. For some additional evidence, + although this is not wanted, see Leroy, 'Lettres Phil.' 1802, + p. 217. + + [119] Hume remarks ('An Enquiry Concerning the Principles of + Morals,' edit. of 1751, p. 132), "there seems a necessity for + confessing that the happiness and misery of others are not + spectacles altogether indifferent to us, but that the view of + the former ... communicates a secret joy; the appearance of the + latter ... throws a melancholy damp over the imagination." + + [120] 'Mental and Moral Science,' 1868, p. 254. + + [121] I have given one such case, namely of three Patagonian + Indians who preferred being shot, one after the other, to + betraying the plans of their companions in war ('Journal of + Researches,' 1845, p. 103). + + [122] Dr. Prosper Despine, in his 'Psychologie Naturelle,' 1868 + (tom. i. p. 243; tom ii. p. 169) gives many curious cases of + the worst criminals, who apparently have been entirely + destitute of conscience. + + [123] See an able article in the 'North British Review,' 1867, + p. 395. See also Mr. W. Bagehot's articles on the Importance of + Obedience and Coherence to Primitive Man, in the 'Fortnightly + Review,' 1867, p. 529, and 1868, p. 457, &c. + + [124] The fullest account which I have met with is by Dr. + Gerland, in his 'Ueber das Aussterben der Naturvlker,' 1868; + but I shall have to recur to the subject of infanticide in a + future chapter. + + [125] See the very interesting discussion on Suicide in Lecky's + 'History of European Morals,' vol. i. 1869, p. 223. + + [126] See, for instance, Mr. Hamilton's account of the Kaffirs, + 'Anthropological Review,' 1870, p. xv. + + [127] Mr. M'Lennan has given 'Primitive Marriage,' 1865, p. + 176, a good collection of facts on this head. + + [128] Lecky, 'History of European Morals,' vol. i. 1869, p. + 109. + + [129] 'Embassy to China,' vol. ii. p. 348. + + [130] See on this subject copious evidence in Chap. vii. of Sir + J. Lubbock, 'Origin of Civilisation,' 1870. + + [131] For instance Lecky, 'Hist. European Morals,' vol. i. p. + 124. + + [132] This term is used in an able article in the 'Westminster + Review,' Oct. 1869, p. 498. For the Greatest Happiness + principle, see J. S. Mill, 'Utilitarianism,' p. 17. + + [133] Good instances are given by Mr. Wallace in 'Scientific + Opinion,' Sept. 15, 1869; and more fully in his 'Contributions + to the Theory of Natural Selection,' 1870, p. 353. + + [134] Tennyson, 'Idylls of the King,' p. 244. + + [135] 'The Thoughts of the Emperor M. Aurelius Antoninus,' Eng. + translat., 2nd edit., 1869, p. 112. Marcus Aurelius was born + A.D. 121. + + [136] Letter to Mr. Mill in Bain's 'Mental and Moral Science,' + 1868, p. 722. + + [137] A writer in the 'North British Review' (July, 1869, p. + 531), well capable of forming a sound judgment, expresses + himself strongly to this effect. Mr. Lecky ('Hist. of Morals,' + vol. i. p. 143) seems to a certain extent to coincide. + + [138] See his remarkable work on 'Hereditary Genius,' 1869, p. + 349. The Duke of Argyll ('Primeval Man,' 1869, p. 188) has some + good remarks on the contest in man's nature between right and + wrong. + + [139] 'The Thoughts of Marcus Aurelius,' &c., p. 139. + + [140] 'Investigations in Military and Anthropolog. Statistics + of American Soldiers,' by B. A. Gould, 1869, p. 256. + + [141] With respect to the "Cranial forms of the American + aborigines," see Dr. Aitken Meigs in 'Proc. Acad. Nat. Sci.' + Philadelphia, May, 1866. On the Australians, see Huxley, in + Lyell's 'Antiquity of Man,' 1863, p. 87. On the Sandwich + Islanders, Prof. J. Wyman, 'Observations on Crania,' Boston, + 1868, p. 18. + + [142] 'Anatomy of the Arteries,' by R. Quain. + + [143] 'Transact. Royal Soc.' Edinburgh, vol. xxiv. p. 175, 189. + + [144] 'Proc. Royal Soc.' 1867, p. 544; also 1868, p. 483, 524. + There is a previous paper, 1866, p. 229. + + [145] 'Proc. R. Irish Academy,' vol. x. 1868, p. 141. + + [146] 'Act. Acad.,' St. Petersburg, 1778, part ii. p. 217. + + [147] Brehm, 'Thierleben,' B. i. s. 58, 87. Rengger, + 'Sugethiere von Paraguay,' s. 57. + + [148] 'Variation of Animals and Plants under Domestication,' + vol. ii. chap. xii. + + [149] 'Hereditary Genius: an Inquiry into its Laws and + Consequences,' 1869. + + [150] Mr. Bates remarks ('The Naturalist on the Amazons,' 1863, + vol. ii. p. 159), with respect to the Indians of the same S. + American tribe, "no two of them were at all similar in the + shape of the head; one man had an oval visage with fine + features, and another was quite Mongolian in breadth and + prominence of cheek, spread of nostrils, and obliquity of + eyes." + + [151] Blumenbach, 'Treatises on Anthropolog.' Eng. translat., + 1865, p. 205. + + [152] Godron, 'De l'Espce,' 1859, tom. ii. livre 3. + Quatrefages, 'Unit de l'Espce Humaine,' 1861. Also Lectures + on Anthropology, given in the 'Revue des Cours Scientifiques,' + 1866-1868. + + [153] 'Hist. Gen. et Part. des Anomalies de l'Organisation,' in + three volumes, tom. i. 1832. + + [154] I have fully discussed these laws in my 'Variation of + Animals and Plants under Domestication,' vol. ii. chap. xxii. + and xxiii. M. J. P. Durand has lately 1868; published a + valuable essay 'De l'Influence des Milieux, &c.' He lays much + stress on the nature of the soil. + + [155] 'Investigations in Military and Anthrop. Statistics,' &c. + 1869, by B. A. Gould, p. 93, 107, 126, 131, 134. + + [156] For the Polynesians, see Prichard's 'Physical Hist. of + Mankind,' vol. v. 1847, p. 145, 283. Also Godron, 'De + l'Espce,' tom. ii. p. 289. There is also a remarkable + difference in appearance between the closely-allied Hindoos + inhabiting the Upper Ganges and Bengal; see Elphinstone's + 'History of India,' vol. i. p. 324. + + [157] 'Memoirs, Anthropolog. Soc.' vol. iii. 1867-69, p. 561, + 565, 567. + + [158] Dr. Brakenridge, 'Theory of Diathesis,' 'Medical Times,' + June 19 and July 17, 1869. + + [159] I have given authorities for these several statements in + my 'Variation of Animals under Domestication,' vol. ii. p. + 297-300. Dr. Jaeger, "Ueber das Lngenwachsthum der Knochen," + 'Jenaischen Zeitschrift,' B. v. Heft i. + + [160] 'Investigations,' &c. By B. A. Gould, 1869, p. 288. + + [161] 'Sugethiere von Paraguay,' 1830, s. 4. + + [162] 'History of Greenland,' Eng. translat. 1767, vol. i. p. + 230. + + [163] 'Intermarriage.' By Alex. Walker, 1838. p. 377. + + [164] 'The Variation of Animals under Domestication,' vol. i. + p. 173. + + [165] 'Principles of Biology,' vol. i. p. 455. + + [166] Paget, 'Lectures on Surgical Pathology,' vol. i. 1853, p. + 209. + + [167] 'The Variation of Animals under Domestication,' vol. ii. + p. 8. + + [168] 'Sugethiere von Paraguay,' s. 8, 10. I have had good + opportunities for observing the extraordinary power of eyesight + in the Fuegians.' See also Lawrence ('Lectures on Physiology,' + &c., 1822, p. 404) on this same subject. M. Giraud-Teulon has + recently collected ('Revue des Cours Scientifiques,' 1870, p. + 625) a large and valuable body of evidence proving that the + cause of short-sight, "_C'est le travail assidu, de prs._" + + [169] Prichard, 'Phys. Hist. of Mankind,' on the authority of + Blumenbach, vol. i. 1851, p. 311; for the statement by Pallas, + vol. iv. 1844, p. 407. + + [170] Quoted by Prichard, 'Researches into the Phys. Hist. of + Mankind,' vol. v. p. 463. + + [171] Mr. Forbes' valuable paper is now published in the + 'Journal of the Ethnological Soc. of London,' new series, vol. + ii. 1870, p. 193. + + [172] Dr. Wilckens ('Landwirthschaft. Wochenblatt,' No. 10, + 1869) has lately published an interesting essay shewing how + domestic animals, which live in mountainous regions, have their + frames modified. + + [173] 'Mmoire sur les Microcphales,' 1867, p. 50, 125, 169, + 171, 184-198. + + [174] See Dr. A. Farre's well-known article in the 'Cyclop. of + Anat. and Phys.' vol. v. 1859, p. 642. Owen 'Anatomy of + Vertebrates,' vol. iii. 1868, p. 687. Prof. Turner in + 'Edinburgh Medical Journal,' Feb. 1865. + + [175] 'Annuario della Soc. dei Naturalisti in Modena,' 1867, p. + 83. Prof. Canestrini gives extracts on this subject from + various authorities. Laurillard remarks, that as he has found a + complete similarity in the form, proportions, and connexion of + the two malar bones in several human subjects and in certain + apes, he cannot consider this disposition of the parts as + simply accidental. + + [176] A whole series of cases is given by Isid. Geoffroy + St.-Hilaire, 'Hist. des Anomalies,' tom. iii. p. 437. + + [177] In my 'Variation of Animals under Domestication' (vol. + ii. p. 57) I attributed the not very rare cases of + supernumerary mamm in women to reversion. I was led to this as + a _probable_ conclusion, by the additional mamm being + generally placed symmetrically on the breast, and more + especially from one case, in which a single efficient mamma + occurred in the inguinal region of a woman, the daughter of + another woman with supernumerary mamm. But Prof. Preyer ('Der + Kampf um das Dasein,' 1869, s. 45) states that _mamm erratic_ + have been known to occur in other situations, even on the back; + so that the force of my argument is greatly weakened or perhaps + quite destroyed. + + With much hesitation I, in the same work (vol. ii. p. 12), + attributed the frequent cases of polydactylism in men to + reversion. I was partly led to this through Prof. Owen's + statement, that some of the Ichthyopterygia possess more than + five digits, and therefore, as I supposed, had retained a + primordial condition; but after reading Prof. Gegenbaur's paper + ('Jenaischen Zeitschrift,' B. v. Heft 3, s. 341), who is the + highest authority in Europe on such a point, and who disputes + Owen's conclusion, I see that it is extremely doubtful whether + supernumerary digits can thus be accounted for. It was the fact + that such digits not only frequently occur and are strongly + inherited, but have the power of regrowth after amputation, + like the normal digits of the lower vertebrata, that chiefly + led me to the above conclusion. This extraordinary fact of + their regrowth remains inexplicable, if the belief in reversion + to some extremely remote progenitor must be rejected. I cannot, + however, follow Prof. Gegenbaur in supposing that additional + digits could not reappear through reversion, without at the + same time other parts of the skeleton being simultaneously and + similarly modified; for single characters often reappear + through reversion. + + [178] 'Anatomy of Vertebrates,' vol. iii. 1868, p. 323. + + [179] 'Generelle Morphologie,' 1866, B. ii. s. clv. + + [180] Carl Vogt's 'Lectures on Man,' Eng. translat. 1864, p. + 151. + + [181] C. Carter Blake, on a jaw from La Naulette, 'Anthropolog. + Review,' 1867, p. 295. Schaaffhausen, ibid. 1868, p. 426. + + [182] 'The Anatomy of Expression,' 1844, p. 110, 131. + + [183] Quoted by Prof. Canestrini in the 'Annuario,' &c., 1867, + p. 90. + + [184] These papers deserve careful study by any one who desires + to learn how frequently our muscles vary, and in varying come + to resemble those of the Quadrumana. The following references + relate to the few points touched on in my text: Proc. Royal + Soc. vol. xiv. 1865, p. 379-384; vol. xv. 1866, p. 241, 242; + vol. xv. 1867, p. 544; vol. xvi. 1868, p. 524. I may here add + that Dr. Murie and Mr. St. George Mivart have shewn in their + Memoir on the Lemuroidea ('Transact. Zoolog. Soc.' vol. vii. + 1869, p. 96), how extraordinarily variable some of the muscles + are in these animals, the lowest members of the Primates. + Gradations, also, in the muscles leading to structures found in + animals still lower in the scale, are numerous in the + Lemuroidea. + + [185] Prof. Macalister in 'Proc. R. Irish Academy,' vol. x. + 1868, p. 124. + + [186] Prof. Macalister (ibid. p. 121) has tabulated his + observations, and finds that muscular abnormalities are most + frequent in the fore-arms, secondly in the face, thirdly in the + foot, &c. + + [187] The Rev. Dr. Haughton, after giving ('Proc. R. Irish + Academy,' June 27, 1864, p. 715) a remarkable case of variation + in the human _flexor pollicis longus_, adds, "This remarkable + example shews that man may sometimes possess the arrangement of + tendons of thumb and fingers characteristic of the macaque; but + whether such a case should be regarded as a macaque passing + upwards into a man, or a man passing downwards into a macaque, + or as a congenital freak of nature, I cannot undertake to say." + It is satisfactory to hear so capable an anatomist, and so + embittered an opponent of evolutionism, admitting even the + possibility of either of his first propositions. Prof. + Macalister has also described ('Proc. R. Irish Acad.' vol. x. + 1864, p. 138) variations in the _flexor pollicis longus_, + remarkable from their relations to the same muscle in the + Quadrumana. + + [188] The authorities for these several statements are given in + my 'Variation of Animals under Domestication,' vol. ii. p. + 320-335. + + [189] This whole subject has been discussed in chap. xxiii. + vol. ii. of my 'Variation of Animals and Plants under + Domestication.' + + [190] See the ever memorable 'Essay on the Principle of + Population,' by the Rev. T. Malthus, vol. i. 1826, p. 6, 517. + + [191] 'Variation of Animals and Plants under Domestication,' + vol. ii. p. 111-113, 163. + + [192] Mr. Sedgwick, 'British and Foreign Medico-Chirurg. + Review,' July, 1863, p. 170. + + [193] 'The Annals of Rural Bengal,' by W. W. Hunter, 1868, p. + 259. + + [194] 'Primitive Marriage,' 1865. + + [195] See some good remarks to this effect by W. Stanley + Jevons, "A Deduction from Darwin's Theory," 'Nature,' 1869, p. + 231. + + [196] Latham, 'Man and his Migrations,' 1851, p. 135. + + [197] Messrs. Murie and Mivart in their "Anatomy of the + Lemuroidea" ('Transact. Zoolog. Soc.' vol. vii. 1869, p. 96-98) + say, "some muscles are so irregular in their distribution that + they cannot be well classed in any of the above groups." These + muscles differ even on the opposite sides of the same + individual. + + [198] 'Quarterly Review,' April, 1869, p. 392. This subject is + more fully discussed in Mr. Wallace's 'Contributions to the + Theory of Natural Selection,' 1870, in which all the essays + referred to in this work are republished. The 'Essay on Man' + has been ably criticised by Prof. Claparde, one of the most + distinguished zoologists in Europe, in an article published in + the 'Bibliothque Universelle,' June, 1870. The remark quoted + in my text will surprise every one who has read Mr. Wallace's + celebrated paper on 'The Origin of Human Races deduced from the + Theory of Natural Selection,' originally published in the + 'Anthropological Review,' May, 1864, p. clviii. I cannot here + resist quoting a most just remark by Sir J. Lubbock + ('Prehistoric Times,' 1865, p. 479) in reference to this paper, + namely, that Mr. Wallace, "with characteristic unselfishness, + ascribes it (_i.e._ the idea of natural selection) unreservedly + to Mr. Darwin, although, as is well known, he struck out the + idea independently, and published it, though not with the same + elaboration, at the same time." + + [199] Quoted by Mr. Lawson Tait in his "Law of Natural + Selection,"--'Dublin Quarterly Journal of Medical Science,' + Feb. 1869. Dr. Keller is likewise quoted to the same effect. + + [200] Owen, 'Anatomy of Vertebrates,' vol. iii. p. 71. + + [201] 'Quarterly Review,' April, 1869, p. 392. + + [202] In _Hylobates syndactylus_, as the name expresses, two of + the digits regularly cohere; and this, as Mr. Blyth informs me, + is occasionally the case with the digits of _H. agilis_, _lar_, + and _leuciscus_. + + [203] Brehm, 'Thierleben,' B. i. s. 80. + + [204] "The Hand, its mechanism," &c. 'Bridgewater Treatise,' + 1833, p. 38. + + [205] Hckel has an excellent discussion on the steps by which + man became a biped: 'Natrliche Schpfungsgeschichte,' 1868, s. + 507. Dr. Bchner ('Confrences sur la Thorie Darwinienne,' + 1869, p. 135) has given good cases of the use of the foot as a + prehensile organ by man; also on the manner of progression of + the higher apes to which I allude in the following paragraph: + see also Owen ('Anatomy of Vertebrates,' vol. iii. p. 71) on + this latter subject. + + [206] "On the Primitive Form of the Skull," translated in + 'Anthropological Review,' Oct. 1868, p. 428. Owen ('Anatomy of + Vertebrates,' vol. ii. 1866, p. 551) on the mastoid processes + in the higher apes. + + [207] 'Die Grenzen der Thierwelt, eine Betrachtung zu Darwin's + Lehre,' 1868, s. 51. + + [208] Dujardin, 'Annales des Sc. Nat.' 3rd series, Zoolog. tom. + xiv. 1850, p. 203. See also Mr. Lowne, 'Anatomy and Phys. of + the _Musca vomitoria_,' 1870, p. 14. My son, Mr. F. Darwin, + dissected for me the cerebral ganglia of the _Formica rufa_. + + [209] 'Philosophical Transactions,' 1869, p. 513. + + [210] Quoted in C. Vogt's 'Lectures on Man,' Eng. translat. + 1864, p. 88, 90. Prichard, 'Phys. Hist. of Mankind,' vol. i. + 1838, p. 305. + + [211] 'Comptes Rendus des Sances,' &c. June 1, 1868. + + [212] 'The Variation of Animals and Plants under + Domestication,' vol. ii. p. 124-129. + + [213] Schaaffhausen gives from Blumenbach and Busch, the cases + of the spasms and cicatrix, in 'Anthropolog. Review,' Oct. + 1868, p. 420. Dr. Jarrold ('Anthropologia,' 1808, p. 115, 116) + adduces from Camper and from his own observations, cases of the + modification of the skull from the head being fixed in an + unnatural position. He believes that certain trades, such as + that of a shoemaker, by causing the head to be habitually held + forward, makes the forehead more rounded and prominent. + + [214] 'Variation of Animals,' &c., vol. i. p. 117 on the + elongation of the skull; p. 119, on the effect of the lopping + of one ear. + + [215] Quoted by Schaaffhausen, in 'Anthropolog. Review,' Oct. + 1868, p. 419. + + [216] Owen, 'Anatomy of Vertebrates,' vol. iii. p. 619. + + [217] Isidore Geoffroy St.-Hilaire remarks ('Hist. Nat. + Gnrale,' tom. ii. 1859, p. 215-217) on the head of man being + covered with long hair; also on the upper surfaces of monkeys + and of other mammals being more thickly clothed than the lower + surfaces. This has likewise been observed by various authors. + Prof. P. Gervais ('Hist. Nat. des Mammifres,' tom. i. 1854, p. + 28), however, states that in the Gorilla the hair is thinner on + the back, where it is partly rubbed off, than on the lower + surface. + + [218] Mr. St. George Mivart, 'Proc. Zoolog. Soc.' 1865, p. 562, + 583. Dr. J. E. Gray, 'Cat. Brit. Mus.: Skeletons.' Owen, + 'Anatomy of Vertebrates,' vol. ii. p. 517. Isidore Geoffroy, + 'Hist. Nat. Gn.' tom. ii. p. 244. + + [219] 'The Variation of Animals and Plants under + Domestication,' vol. ii. p. 280, 282. + + [220] 'Primeval Man,' 1869, p. 66. + + [221] 'Anthropological Review,' May, 1864, p. clviii. + + [222] After a time the members or tribes which are absorbed + into another tribe assume, as Mr. Maine remarks ('Ancient Law,' + 1861, p. 131), that they are the co-descendants of the same + ancestors. + + [223] Morlot, 'Soc. Vaud. Sc. Nat.' 1860, p. 294. + + [224] I have given instances in my 'Variation of Animals under + Domestication,' vol. ii. p. 196. + + [225] See a remarkable series of articles on Physics and + Politics in the 'Fortnightly Review,' Nov. 1867; April 1, 1868; + July 1, 1869. + + [226] 'Origin of Civilisation,' 1870, p. 265. + + [227] Mr. Wallace gives cases in his 'Contributions to the + Theory of Natural Selection,' 1870, p. 354. + + [228] 'Ancient Law,' 1861, p. 22. For Mr. Bagehot's remarks, + 'Fortnightly Review,' April 1, 1868, p. 452. + + [229] 'The Variation of Animals and Plants under + Domestication,' vol. i. p. 309. + + [230] 'Fraser's Magazine,' Sept. 1868, p. 353. This article + seems to have struck many persons, and has given rise to two + remarkable essays and a rejoinder in the 'Spectator,' Oct. 3rd + and 17th 1868. It has also been discussed in the 'Q. Journal of + Science,' 1869, p. 152, and by Mr. Lawson Tait in the 'Dublin + Q. Journal of Medical Science,' Feb. 1869, and by Mr. E. Ray + Lankester in his 'Comparative Longevity,' 1870, p. 128. Similar + views appeared previously in the 'Australasian,' July 13, 1867. + I have borrowed ideas from several of these writers. + + [231] For Mr. Wallace, see 'Anthropolog. Review,' as before + cited. Mr. Galton in 'Macmillan's Magazine,' Aug. 1865, p. 318; + also his great work, 'Hereditary Genius,' 1870. + + [232] 'Hereditary Genius,' 1870, p. 132-140. + + [233] See the fifth and sixth columns, compiled from good + authorities, in the table given in Mr. E. R. Lankester's + 'Comparative Longevity,' 1870, p. 115. + + [234] 'Hereditary Genius,' 1870, p. 330. + + [235] 'Origin of Species' (fifth edition, 1869), p. 104. + + [236] 'Hereditary Genius,' 1870, p. 347. + + [237] E. Ray Lankester, 'Comparative Longevity,' 1870, p. 115. + The table of the intemperate is from Nelson's 'Vital + Statistics.' In regard to profligacy, see Dr. Farr, "Influence + of Marriage on Mortality," 'Nat. Assoc. for the Promotion of + Social Science,' 1858. + + [238] 'Fraser's Magazine,' Sept. 1868, p. 353. 'Macmillan's + Magazine,' Aug. 1865, p. 318. The Rev. F. W. Farrar ('Fraser's + Mag.,' Aug. 1870, p. 264) takes a different view. + + [239] "On the Laws of the Fertility of Women," in 'Transact. + Royal Soc.' Edinburgh, vol. xxiv. p. 287. See, also, Mr. + Galton, 'Hereditary Genius,' p. 352-357, for observations to + the above effect. + + [240] 'Tenth Annual Report of Births, Deaths, &c., in + Scotland,' 1867, p. xxix. + + [241] These quotations are taken from our highest authority on + such questions, namely, Dr. Farr, in his paper "On the + Influence of Marriage on the Mortality of the French People," + read before the Nat. Assoc. for the Promotion of Social + Science, 1858. + + [242] Dr. Farr, ibid. The quotations given below are extracted + from the same striking paper. + + [243] I have taken the mean of the quinquennial means, given in + 'The Tenth Annual Report of Births, Deaths, &c., in Scotland,' + 1867. The quotation from Dr. Stark is copied from an article in + the 'Daily News,' Oct. 17th, 1868, which Dr. Farr considers + very carefully written. + + [244] See the ingenious and original argument on this subject + by Mr. Galton, 'Hereditary Genius,' p. 340-342. + + [245] Mr. Greg, 'Fraser's Magazine,' Sept. 1868, p. 357. + + [246] 'Hereditary Genius,' 1870, p. 357-359. The Rev. F. H. + Farrar ('Fraser's Mag.' , Aug. 1870, p. 257) advances arguments + on the other side. Sir C. Lyell had already ('Principles of + Geology,' vol. ii. 1868, p. 489) called attention, in a + striking passage, to the evil influence of the Holy Inquisition + in having lowered, through selection, the general standard of + intelligence in Europe. + + [247] Mr. Galton, 'Macmillan's Magazine,' August, 1865, p. 325. + See, also, 'Nature,' "On Darwinism and National Life," Dec. + 1869, p. 184. + + [248] 'Last Winter in the United States,' 1868, p. 29. + + [249] 'On the Origin of Civilisation,' 'Proc. Ethnological + Soc.' Nov. 26, 1867. + + [250] 'Primeval Man,' 1869. + + [251] 'Royal Institution of Great Britain,' March 15, 1867. + Also, 'Researches into the Early History of Mankind,' 1865. + + [252] 'Primitive Marriage,' 1865. See, likewise, an excellent + article, evidently by the same author, in the 'North British + Review,' July, 1869. Also, Mr. L. H. Morgan, "A Conjectural + Solution of the Origin of the Class. System of Relationship," + in 'Proc. American Acad. of Sciences,' vol. vii. Feb. 1868. + Prof. Schaaffhausen ('Anthropolog. Review,' Oct. 1869, p. 373) + remarks on "the vestiges of human sacrifices found both in + Homer and the Old Testament." + + [253] Sir J. Lubbock, 'Prehistoric Times,' 2nd edit. 1869, + chap. xv. and xvi. _et passim_. + + [254] Dr. F. Mller has made some good remarks to this effect + in the 'Reise der Novara: Anthropolog. Theil,' Abtheil. iii. + 1868, s. 127. + + [255] Isidore Geoffroy St.-Hilaire gives a detailed account of + the position assigned to man by various naturalists in their + classifications: 'Hist. Nat. Gn.' tom. ii. 1859, p. 170-189. + + [256] See the very interesting article, "L'Instinct chez les + Insectes," by M. George Pouchet, 'Revue des Deux Mondes,' Feb. + 1870, p. 682. + + [257] Westwood, 'Modern Class. of Insects,' vol. ii. 1840, p. + 87. + + [258] 'Proc. Zoolog. Soc.' 1869, p. 4. + + [259] 'Evidence as to Man's Place in Nature,' 1863, p. 70, _et + passim_. + + [260] Isid. Geoffroy, 'Hist. Nat. Gn.' tom. ii. 1859, p. 217. + + [261] "Ueber die Richtung der Haare," &c., Mller's 'Archiv fr + Anat. und Phys.' 1837, s. 51. + + [262] On the hair in Hylobates, see 'Nat. Hist. of Mammals,' by + C. L. Martin, 1841, p. 415. Also, Isid. Geoffroy on the + American monkeys and other kinds, 'Hist. Nat. Gn.' vol. ii. + 1859, p. 216, 243. Eschricht, ibid., s. 46, 55, 61. Owen, 'Anat. + of Vertebrates,' vol. iii. p. 619. Wallace, 'Contributions to + the Theory of Natural Selection,' 1870. p. 344. + + [263] 'Origin of Species,' 5th edit. 1869, p. 194. 'The + Variation of Animals and Plants under Domestication,' vol. ii. + 1868, p. 348. + + [264] 'An Introduction to the Classification of Animals,' 1869, + p. 99. + + [265] This is nearly the same classification as that + provisionally adopted by Mr. St. George Mivart ('Transact. + Philosoph. Soc.' 1867, p. 300), who, after separating the + Lemurid, divides the remainder of the Primates into the + Hominid, the Simiad answering to the Catarhines, the Cebid, + and the Hapalid,--these two latter groups answering to the + Platyrhines. + + [266] 'Transact. Zoolog. Soc.' vol. vi. 1867, p. 214. + + [267] Mr. St. G. Mivart, 'Transact. Phil. Soc.' 1867, p. 410. + + [268] Messrs. Murie and Mivart on the Lemuroidea. 'Transact. + Zoolog. Soc.' vol. vii. 1869, p. 5. + + [269] Hckel has come to this same conclusion. See 'Ueber die + Entstehung des Menschengeschlechts,' in Virchow's 'Sammlung. + gemein. wissen. Vortrge,' 1868, s. 61. Also his 'Natrliche + Schpfungsgeschichte,' 1868, in which he gives in detail his + views on the genealogy of man. + + [270] 'Anthropological Review,' April, 1867, p. 236. + + [271] 'Elements of Geology,' 1865, p. 583-585. 'Antiquity of + Man', 1863; p. 145. + + [272] 'Man's Place in Nature,' p. 105. + + [273] Elaborate tables are given in his 'Generelle Morphologie' + (B. ii. s. cliii. and s. 425); and with more especial reference + to man in his 'Natrliche Schpfungsgeschichte,' 1868. Prof. + Huxley, in reviewing this latter work ('The Academy,' 1869, p. + 42) says, that he considers the phylum or lines of descent of + the Vertebrata to be admirably discussed by Hckel, although he + differs on some points. He expresses, also, his high estimate + of the value of the general tenor and spirit of the whole work. + + [274] 'Palontology,' 1860, p. 199. + + [275] I had the satisfaction of seeing, at the Falkland + Islands, in April, 1833, and therefore some years before any + other naturalist, the locomotive larv of a compound Ascidian, + closely allied to, but apparently generically distinct from, + Synoicum. The tail was about five times as long as the oblong + head, and terminated in a very fine filament. It was plainly + divided, as sketched by me under a simple microscope, by + transverse opaque partitions, which I presume represent the + great cells figured by Kowalevsky. At an early stage of + development the tail was closely coiled round the head of the + larva. + + [276] 'Mmoires de l'Acad. des Sciences de St. Ptersbourg,' + tom. x. No. 15, 1866. + + [277] This is the conclusion of one of the highest authorities + in comparative anatomy, namely, Prof. Gegenbaur: 'Grundzge der + vergleich. Anat.' 1870, s. 876. The result has been arrived at + chiefly from the study of the Amphibia; but it appears from the + researches of Waldeyer (as quoted in Humphry's 'Journal of + Anat. and Phys.' 1869, p. 161), that the sexual organs of even + "the higher vertebrata are, in their early condition, + hermaphrodite." Similar views have long been held by some + authors, though until recently not well based. + + [278] The male Thylacinus offers the best instance. Owen, + 'Anatomy of Vertebrates,' vol. iii. p. 771. + + [279] Serranus is well known often to be in an hermaphrodite + condition; but Dr. Gnther informs me that he is convinced that + this is not its normal state. Descent from an ancient + androgynous prototype would, however, naturally favour and + explain, to a certain extent, the recurrence of this condition + in these fishes. + + [280] Mr. Lockwood believes (as quoted in 'Quart. Journal of + Science,' April, 1868, p. 269), from what he has observed of + the development of Hippocampus, that the walls of the abdominal + pouch of the male in some way afford nourishment. On male + fishes hatching the ova in their mouths, see a very interesting + paper by Prof. Wyman, in 'Proc. Boston Soc. of Nat. Hist.' + Sept. 15, 1857; also Prof. Turner, in 'Journal of Anat. and + Phys.' Nov. 1, 1866, p. 78. Dr. Gnther has likewise described + similar cases. + + [281] All vital functions tend to run their course in fixed and + recurrent periods, and with tidal animals the periods would + probably be lunar; for such animals must have been left dry or + covered deep with water,--supplied with copious food or + stinted,--during endless generations, at regular lunar + intervals. If then the Vertebrata are descended from an animal + allied to the existing tidal Ascidians, the mysterious fact, + that with the higher and now terrestrial Vertebrata, not to + mention other classes, many normal and abnormal vital processes + run their course according to lunar periods, is rendered + intelligible. A recurrent period, if approximately of the right + duration, when once gained, would not, as far as we can judge, + be liable to be changed; consequently it might be thus + transmitted during almost any number of generations. This + conclusion, if it could be proved sound, would be curious; for + we should then see that the period of gestation in each mammal, + and the hatching of each bird's eggs, and many other vital + processes, still betrayed the primordial birthplace of these + animals. + + [282] 'History of India,' 1841, vol. i. p. 323. Father Ripa + makes exactly the same remark with respect to the Chinese. + + [283] A vast number of measurements of Whites, Blacks, and + Indians, are given in the 'Investigations in the Military and + Anthropolog. Statistics of American Soldiers,' by B. A. Gould, + 1869, p. 298-358; on the capacity of the lungs, p. 471. See + also the numerous and valuable tables, by Dr. Weisbach, from + the observations of Dr. Scherzer and Dr. Schwarz, in the 'Reise + der Novara: Anthropolog. Theil,' 1867. + + [284] See, for instance, Mr. Marshall's account of the brain of + a Bush-woman, in 'Phil. Transact.' 1864, p. 519. + + [285] Wallace, 'The Malay Archipelago,' vol. ii. 1869, p. 178. + + [286] With respect to the figures in the famous Egyptian caves + of Abou-Simbel, M. Pouchet says ('The Plurality of the Human + Races,' Eng. translat. 1864, p. 50), that he was far from + finding recognisable representations of the dozen or more + nations which some authors believe that they can recognise. + Even some of the most strongly-marked races cannot be + identified with that degree of unanimity which might have been + expected from what has been written on the subject. Thus + Messrs. Nott and Gliddon ('Types of Mankind,' p. 148) state + that Rameses II., or the Great, has features superbly European; + whereas Knox, another firm believer in the specific distinction + of the races of man ('Races of Man,' 1850, p. 201), speaking of + young Memnon (the same person with Rameses II., as I am + informed by Mr. Birch) insists in the strongest manner that he + is identical in character with the Jews of Antwerp. Again, + whilst looking in the British Museum with two competent judges, + officers of the establishment, at the statue of Amunoph III., + we agreed that he had a strongly negro cast of features; but + Messrs. Nott and Gliddon (ibid. p. 146, fig. 53) describe him + as "a hybrid, but not of negro intermixture." + + [287] As quoted by Nott and Gliddon, 'Types of Mankind,' 1854, + p. 439. They give also corroborative evidence; but C. Vogt + thinks that the subject requires further investigation. + + [288] "Diversity of Origin of the Human Races," in the + 'Christian Examiner,' July, 1850. + + [289] 'Transact. B. Soc. of Edinburgh,' vol. xxii. 1861, p. + 567. + + [290] 'On the Phenomena of Hybridity in the Genus Homo,' Eng. + translat. 1864. + + [291] See the interesting letter by Mr. T. A. Murray, in the + 'Anthropolog. Review,' April, 1868, p. liii. In this letter + Count Strzelecki's statement, that Australian women who have + borne children to a white man are afterwards sterile with their + own race, is disproved. M. A. de Quatrefages has also collected + ('Revue des Cours Scientifiques,' March, 1869, p. 239) much + evidence that Australians and Europeans are not sterile when + crossed. + + [292] 'An Examination of Prof. Agassiz's Sketch of the Nat. + Provinces of the Animal World,' Charleston, 1855, p. 44. + + [293] 'Military and Anthropolog. Statistics of American + Soldiers,' by B. A. Gould, 1869, p. 319. + + [294] 'The Variation of Animals and Plants under + Domestication,' vol. ii. p. 109. I may here remind the reader + that the sterility of species when crossed is not a + specially-acquired quality; but, like the incapacity of certain + trees to be grafted together, is incidental on other acquired + differences. The nature of these differences is unknown, but + they relate more especially to the reproductive system, and + much less to external structure or to ordinary differences in + constitution. One important element in the sterility of crossed + species apparently lies in one or both having been long + habituated to fixed conditions; for we know that changed + conditions have a special influence on the reproductive system, + and we have good reason to believe (as before remarked) that + the fluctuating conditions of domestication tend to eliminate + that sterility which is so general with species in a natural + state when crossed. It has elsewhere been shewn by me (ibid. + vol. ii. p. 185, and 'Origin of Species,' 5th edit. p. 317) + that the sterility of crossed species has not been acquired + through natural selection: we can see that when two forms have + already been rendered very sterile, it is scarcely possible + that their sterility should be augmented by the preservation or + survival of the more and more sterile individuals; for as the + sterility increases fewer and fewer offspring will be produced + from which to breed, and at last only single individuals will + be produced, at the rarest intervals. But there is even a + higher grade of sterility than this. Both Grtner and Klreuter + have proved that in genera of plants including numerous + species, a series can be formed from species which when crossed + yield fewer and fewer seeds, to species which never produce a + single seed, but yet are affected by the pollen of the other + species, for the germen swells. It is here manifestly + impossible to select the more sterile individuals, which have + already ceased to yield seeds; so that the acme of sterility, + when the germen alone is affected, cannot be gained through + selection. This acme, and no doubt the other grades of + sterility, are the incidental results of certain unknown + differences in the constitution of the reproductive system of + the species which are crossed. + + [295] 'The Variation of Animals,' &c., vol. ii. p. 92. + + [296] M. de Quatrefages has given ('Anthropolog. Review,' Jan. + 1869, p. 22) an interesting account of the success and energy + of the Paulistas in Brazil, who are a much crossed race of + Portuguese and Indians, with a mixture of the blood of other + races. + + [297] For instance with the aborigines of America and + Australia. Prof. Huxley says ('Transact. Internat. Congress of + Prehist. Arch.' 1868. p. 105) that the skulls of many South + Germans and Swiss are "as short and as broad as those of the + Tartars," &c. + + [298] See a good discussion on this subject in Waitz, + 'Introduct. to Anthropology,' Eng. translat. 1863, p. 198-208, + 227. I have taken some of the above statements from H. Tuttle's + 'Origin and Antiquity of Physical Man,' Boston, 1866, p. 35. + + [299] Prof. Ngeli has carefully described several striking + cases in his 'Botanische Mittheilungen,' B. ii. 1866, s. + 294-369. Prof. Asa Gray has made analogous remarks on some + intermediate forms in the Composit of N. America. + + [300] 'Origin of Species,' 5th edit. p. 68. + + [301] See Prof. Huxley to this effect in the 'Fortnightly + Review,' 1865, p. 275. + + [302] 'Lectures on Man,' Eng. translat. 1864, p. 468. + + [303] 'Die Racen des Schweines,' 1860, s. 46. 'Vorstudien fr + Geschichte, &c., Schweineschdel,' 1864, s. 104. With respect + to cattle, see M. de Quatrefages, 'Unit de l'Espce Humaine,' + 1861, p. 119. + + [304] Tylor's 'Early History of Mankind,' 1865; for the + evidence with respect to gesture-language, see p. 54. Lubbock's + 'Prehistoric Times,' 2nd edit. 1869. + + [305] 'The Primitive Inhabitants of Scandinavia,' Eng. + translat. edited by Sir J. Lubbock, 1868, p. 104. + + [306] Hodder M. Westropp, on Cromlechs, &c., 'Journal of + Ethnological Soc.' as given in 'Scientific Opinion,' June 2nd. + 1869, p. 3. + + [307] 'Journal of Researches: Voyage of the "Beagle,"' p. 46. + + [308] 'Prehistoric Times,' 1869, p. 574. + + [309] Translation in 'Anthropological Review,' Oct. 1868, p. + 431. + + [310] 'Transact. Internat. Congress of Prehistoric Arch.' 1868, + p. 172-175. See also Broca (translation) in 'Anthropological + Review,' Oct. 1868, p. 410. + + [311] Dr. Gerland, 'Ueber das Aussterben der Naturvlker,' + 1868, s. 82. + + [312] Gerland (ibid. s. 12) gives facts in support of this + statement. + + [313] See remarks to this effect in Sir H. Holland's 'Medical + Notes and Reflections,' 1839, p. 390. + + [314] I have collected ('Journal of Researches, Voyage of the + "Beagle,"' p. 435) a good many cases bearing on this subject: + see also Gerland, ibid. s. 8. Poeppig speaks of the "breath of + civilisation as poisonous to savages." + + [315] Sproat, 'Scenes and Studies of Savage Life,' 1868, p. + 284. + + [316] Bagehot, "Physics and Politics," 'Fortnightly Review,' + April 1, 1868, p. 455. + + [317] "On Anthropology," translation, 'Anthropolog. Review,' + Jan. 1868, p. 38. + + [318] 'The Annals of Rural Bengal,' 1868, p. 134. + + [319] 'The Variation of Animals and Plants under + Domestication,' vol. ii. p. 95. + + [320] Pallas, 'Act. Acad. St. Petersburgh,' 1780, part ii. p. + 69. He was followed by Rudolphi, in his 'Beytrge zur + Anthropologie,' 1812. An excellent summary of the evidence is + given by Godron, 'De l'Espce,' 1859, vol. ii. p. 246, &c. + + [321] Sir Andrew Smith, as quoted by Knox, 'Races of Man,' + 1850, p. 473. + + [322] See De Quatrefages on this head, 'Revue des Cours + Scientifiques,' Oct. 17, 1868, p. 731. + + [323] Livingstone's 'Travels and Researches in S. Africa,' + 1857, p. 338, 329. D'Orbigny, as quoted by Godron, 'De + l'Espce,' vol. ii. p. 266. + + [324] See a paper read before the Royal Soc. in 1813, and + published in his Essays in 1818. I have given an account of Dr. + Wells' views in the Historical Sketch (p. xvi) to my 'Origin of + Species.' Various cases of colour correlated with + constitutional peculiarities are given in my 'Variation of + Animals under Domestication,' vol. ii. p. 227, 335. + + [325] See, for instance, Nott and Gliddon, 'Types of Mankind,' + p. 68. + + [326] Major Tulloch, in a paper read before the Statistical + Society, April 20th, 1840, and given in the 'Athenum,' 1840, + p. 353. + + [327] 'The Plurality of the Human Race' (translat.), 1864, p. + 60. + + [328] Quatrefages, 'Unit de l'Espce Humaine,' 1861, p. 205. + Waitz, 'Introduct. to Anthropology,' translat. vol. i. 1863, p. + 124. Livingstone gives analogous cases in his 'Travels.' + + [329] In the spring of 1862 I obtained permission from the + Director-General of the Medical department of the Army, to + transmit to the surgeons of the various regiments on foreign + service a blank table, with the following appended remarks, but + I have received no returns. "As several well-marked cases have + been recorded with our domestic animals of a relation between + the colour of the dermal appendages and the constitution; and + it being notorious that there is some limited degree of + relation between the colour of the races of man and the climate + inhabited by them; the following investigation seems worth + consideration. Namely, whether there is any relation in + Europeans between the colour of their hair, and their liability + to the diseases of tropical countries. If the surgeons of the + several regiments, when stationed in unhealthy tropical + districts, would be so good as first to count, as a standard of + comparison, how many men, in the force whence the sick are + drawn, have dark and light-coloured hair, and hair of + intermediate or doubtful tints; and if a similar account were + kept by the same medical gentlemen, of all the men who suffered + from malarious and yellow fevers, or from dysentery, it would + soon be apparent, after some thousand cases had been tabulated, + whether there exists any relation between the colour of the + hair and constitutional liability to tropical diseases. Perhaps + no such relation would be discovered, but the investigation is + well worth making. In case any positive result were obtained, + it might be of some practical use in selecting men for any + particular service. Theoretically the result would be of high + interest, as indicating one means by which a race of men + inhabiting from a remote period an unhealthy tropical climate, + might have become dark-coloured by the better preservation of + dark-haired or dark-complexioned individuals during a long + succession of generations." + + [330] 'Anthropological Review,' Jan. 1866, p. xxi. + + [331] See, for instance, Quatrefages ('Revue des Cours + Scientifiques,' Oct. 10, 1868, p. 724) on the effects of + residence in Abyssinia and Arabia, and other analogous cases. + Dr. Rolle ('Der Mensch, seine Abstammung,' &c., 1865, s. 99) + states, on the authority of Khanikof, that the greater number + of German families settled in Georgia, have acquired in the + course of two generations dark hair and eyes. Mr. D. Forbes + informs me that the Quichuas in the Andes vary greatly in + colour, according to the position of the valleys inhabited by + them. + + [332] Harlan, 'Medical Researches,' p. 532. Quatrefages ('Unit + de l'Espce Humaine,' 1861, p. 128) has collected much evidence + on this head. + + [333] See Prof. Schaaffhausen, translat. in 'Anthropological + Review,' Oct. 1868, p. 429. + + [334] Mr. Catlin states ('N. American Indians,' 3rd edit. 1842, + vol. i. p. 49) that in the whole tribe of the Mandans, about + one in ten or twelve of the members of all ages and both sexes + have bright silvery grey hair, which is hereditary. Now this + hair is as coarse and harsh as that of a horse's mane, whilst + the hair of other colours is fine and soft. + + [335] On the odour of the skin, Godron, 'Sur l'Espce,' tom. + ii. p. 217. On the pores in the skin, Dr. Wilckens, 'Die + Aufgaben der landwirth. Zootechnik,' 1869, s. 7. + + [336] Westwood, 'Modern Class. of Insects,' vol. ii. 1810, p. + 541. In regard to the statement about Tanais, mentioned below, + I am indebted to Fritz Mller. + + [337] Kirby and Spence, 'Introduction to Entomology,' vol. iii. + 1826, p. 309. + + [338] Even with those of plants in which the sexes are + separate, the male flowers are generally mature before the + female. Many hermaphrodite plants are, as first shewn by C. K. + Sprengel, dichogamous; that is, their male and female organs + are not ready at the same time, so that they cannot be + self-fertilised. Now with such plants the pollen is generally + mature in the same flower before the stigma, though there are + some exceptional species in which the female organs are mature + before the male. + + [339] I have received information, hereafter to be given, to + this effect with respect to poultry. Even with birds, such as + pigeons, which pair for life, the female, as I hear from Mr. + Jenner Weir, will desert her mate if he is injured or grows + weak. + + [340] On the Gorilla, Savage and Wyman, 'Boston Journal of Nat. + Hist.' vol. v. 1845-47, p. 423. On Cynocephalus, Brehm, + 'Illust. Thierleben,' B. i. 1864, s. 77. On Mycetes, Rengger, + 'Naturgesch.: Sugethiere von Paraguay,' 1830, s. 14, 20. On + Cebus, Brehm, ibid. s. 108. + + [341] Pallas, 'Spicilegia Zoolog.' Fasc. xii. 1777, p. 29. Sir + Andrew Smith, 'Illustrations of the Zoology of S. Africa,' + 1849, pl. 29, on the Kobus. Owen, in his 'Anatomy of + Vertebrates' (vol. iii. 1868, p. 633) gives a table + incidentally showing which species of Antelopes pair and which + are gregarious. + + [342] Dr. Campbell, in 'Proc. Zoolog. Soc.' 1869, p. 138. See + also an interesting paper, by Lieut. Johnstone, in 'Proc. + Asiatic Soc. of Bengal,' May, 1868. + + [343] 'The Ibis,' vol. iii. 1861, p. 133, on the Progne + Widow-bird. See also on the Vidua axillaris, ibid. vol. ii. + 1860, p. 211. On the polygamy of the Capercailzie and Great + Bustard, see L. Lloyd, 'Game Birds of Sweden,' 1867, p. 19, and + 182. Montagu and Selby speak of the Black Grouse as polygamous + and of the Red Grouse as monogamous. + + [344] The Rev. E. S. Dixon, however, speaks positively + ('Ornamental Poultry,' 1848, p. 76) about the eggs of the + guinea-fowl being infertile when more than one female is kept + with the same male. + + [345] Noel Humphreys, 'River Gardens,' 1857. + + [346] Kirby and Spence, 'Introduction to Entomology,' vol. iii. + 1826, p. 342. + + [347] One parasitic Hymenopterous insect (Westwood, 'Modern + Class. of Insects,' vol. ii, p. 160) forms an exception to the + rule, as the male has rudimentary wings, and never quits the + cell in which it is born, whilst the female has well-developed + wings. Audouin believes that the females are impregnated by the + males which are born in the same cells with them; but it is + much more probable that the females visit other cells, and thus + avoid close inter-breeding. We shall hereafter meet with a few + exceptional cases, in various classes, in which the female, + instead of the male, is the seeker and wooer. + + [348] 'Essays and Observations,' edited by Owen, vol. i. 1861, + p. 194. + + [349] Prof. Sachs ('Lehrbuch der Botanik,' 1870, s. 633) in + speaking of the male and female reproductive cells, remarks, + "verhlt sich die eine bei der Vereinigung activ, ... die + andere erscheint bei der Vereinigung passiv." + + [350] 'Reise der Novara: Anthropolog. Theil,' 1867, s. 216-269. + The results were calculated by Dr. Weisbach from measurements + made by Drs. K. Scherzer and Schwarz. On the greater + variability of the males of domesticated animals, see my + 'Variation of Animals and Plants under Domestication,' vol. ii. + 1868, p. 75. + + [351] 'Proceedings Royal Soc.' vol. xvi. July, 1868, p. 519 and + 524. + + [352] 'Proc. Royal Irish Academy,' vol. x. 1868, p. 123. + + [353] 'Massachusetts Medical Soc.' vol. ii. No. 3, 1808, p. 9. + + [354] 'The Variation of Animals and Plants under + Domestication,' vol. ii. 1868, p. 75. In the last chapter but + one, the provisional hypothesis of pangenesis, above alluded + to, is fully explained. + + [355] These facts are given on the high authority of a great + breeder, Mr. Teebay, in Tegetmeier's 'Poultry Book,' 1868, p. + 158. On the characters of chickens of different breeds, and on + the breeds of the pigeon, alluded to in the above paragraph, + see 'Variation of Animals,' &c., vol. i. p. 160, 249; vol. ii. + p. 77. + + [356] 'Nov species Quadrupedum e Glirium ordine,' 1778, p. 7. + On the transmission of colour by the horse, see 'Variation of + Animals, &c. under Domestication,' vol. i. p. 21. Also vol. ii. + p. 71, for a general discussion on Inheritance as limited by + Sex. + + [357] Dr. Chapuis, 'Le Pigeon Voyageur Belge,' 1865, p. 87. + Boitard et Corbi, 'Les Pigeons de Volire,' &c., 1824, p. 173. + + [358] References are given in my 'Variation of Animals under + Domestication,' vol. ii. p. 72. + + [359] I am much obliged to Mr. Cupples for having made + enquiries for me in regard to the Roebuck and Red Deer of + Scotland from Mr. Robertson, the experienced head-forester to + the Marquis of Breadalbane. In regard to Fallow-deer, I am + obliged to Mr. Eyton and others for information. For the + _Cervus alces_ of N. America, see 'Land and Water,' 1868, p. + 221 and 254; and for the _C. Virginianus_ and _strongyloceros_ + of the same continent, see J. D. Caton, in 'Ottawa Acad. of + Nat. Sc.' 1868, p. 13. For _Cervus Eldi_ of Pegu, see Lieut. + Beavan, 'Proc. Zoolog. Soc.' 1867, p. 762. + + [360] _Antilocapra Americana._ Owen, 'Anatomy of Vertebrates,' + vol. iii. p. 627. + + [361] I have been assured that the horns of the sheep in North + Wales can always be felt, and are sometimes even an inch in + length, at birth. With cattle Youatt says ('Cattle,' 1834, p. + 277) that the prominence of the frontal bone penetrates the + cutis at birth, and that the horny matter is soon formed over + it. + + [362] I am greatly indebted to Prof. Victor Carus for having + made inquiries for me, from the highest authorities, with + respect to the merino sheep of Saxony. On the Guinea coast of + Africa there is a breed of sheep in which, as with merinos, the + rams alone bear horns; and Mr. Winwood Reade informs me that in + the one case observed, a young ram born on Feb. 10th first + showed horns on March 6th, so that in this instance the + development of the horns occurred at a later period of life, + conformably with our rule, than in the Welsh sheep, in which + both sexes are horned. + + [363] In the common peacock (_Pavo cristatus_) the male alone + possesses spurs, whilst both sexes of the Java peacock (_P. + muticus_) offer the unusual case of being furnished with spurs. + Hence I fully expected that in the latter species they would + have been developed earlier in life than in the common peacock; + but M. Hegt of Amsterdam informs me, that with young birds of + the previous year, belonging to both species, compared on April + 23rd, 1869, there was no difference in the development of the + spurs. The spurs, however, were as yet represented merely by + slight knobs or elevations. I presume that I should have been + informed if any difference in the rate of development had + subsequently been observed. + + [364] In some other species of the Duck Family the speculum in + the two sexes differs in a greater degree; but I have not been + able to discover whether its full development occurs later in + life in the males of such species, than in the male of the + common duck, as ought to be the case according to our rule. + With the allied _Mergus cucullatus_ we have, however, a case of + this kind: the two sexes differ conspicuously in general + plumage, and to a considerable degree in the speculum, which is + pure white in the male and greyish-white in the female. Now the + young males at first resemble, in all respects, the female, and + have a greyish-white speculum, but this becomes pure white at + an earlier age than that at which the adult male acquires his + other more strongly-marked sexual differences in plumage: see + Audubon, 'Ornithological Biography,' vol. iii. 1835, p. + 249-250. + + [365] 'Das Ganze der Taubenzucht,' 1837, s. 21, 24. For the + case of the streaked pigeons, see Dr. Chapuis, 'Le Pigeon + Voyageur Belge.' 1865, p. 87. + + [366] For full particulars and references on all these points + respecting the several breeds of the Fowl, see 'Variation of + Animals and Plants under Domestication,' vol. i. p. 250, 256. + In regard to the higher animals, the sexual differences which + have arisen under domestication are described in the same work + under the head of each species. + + [367] 'Twenty-ninth Annual Report of the Registrar-General for + 1866.' In this report (p. xii) a special decennial table is + given. + + [368] For Norway and Russia, see abstract of Prof. Faye's + researches, in 'British and Foreign Medico-Chirurg. Review,' + April, 1867, p. 343, 345. For France, the 'Annuaire pour l'An + 1867.' p. 213. + + [369] In regard to the Jews, see M. Thury, 'La Loi de + Production des Sexes,' 1863, p. 25. + + [370] Babbage, 'Edinburgh Journal of Science,' 1829, vol. i. p. + 88; also p. 90, on still-born children. On illegitimate + children in England, see 'Report of Registrar-General for + 1866,' p. xv. + + [371] 'British and Foreign Medico-Chirurg. Review,' April, + 1867, p. 343. Dr. Stark also remarks ('Tenth Annual Report of + Births, Deaths, &c., in Scotland,' 1867, p. xxviii) that "These + examples may suffice to shew that, at almost every stage of + life, the males in Scotland have a greater liability to death + and a higher death-rate than the females. The fact, however, of + this peculiarity being most strongly developed at that + infantile period of life when the dress, food, and general + treatment of both sexes are alike, seems to prove that the + higher male death-rate is an impressed, natural, and + constitutional peculiarity due to sex alone." + + [372] With the savage Guaranys of Paraguay, according to the + accurate Azara ('Voyages dans l'Amrique mrid.' tom. ii. 1809, + p. 60, 179), the women in proportion to the men are as 14 to + 13. + + [373] Leuckart in Wagner, 'Handwrterbuch der Phys.' B. iv. + 1853, s. 774. + + [374] Anthropological Review, April, 1870, p. cviii. + + [375] During the last eleven years a record has been kept of + the number of mares which have proved barren or prematurely + slipped their foals; and it deserves notice, as shewing how + infertile these highly-nurtured and rather closely-interbred + animals have become, that not far from one-third of the mares + failed to produce living foals. Thus during 1866, 809 male + colts and 816 female colts were born, and 743 mares failed to + produce offspring. During 1867, 836 males and 902 females were + born, and 794 mares failed. + + [376] I am much indebted to Mr. Cupples for having procured for + me the above returns from Scotland, as well as some of the + following returns on cattle. Mr. R. Elliot, of Laighwood, first + called my attention to the premature deaths of the males,--a + statement subsequently confirmed by Mr. Aitchison and others. + To this latter gentleman, and to Mr. Payan, I owe my thanks for + the larger returns on sheep. + + [377] Bell, 'History of British Quadrupeds,' p. 100. + + [378] 'Illustrations of the Zoology of S. Africa,' 1849, pl. + 29. + + [379] Brehm ('Illust. Thierleben,' B. iv. s. 990) comes to the + same conclusion. + + [380] On the authority of L. Lloyd, 'Game Birds of Sweden,' + 1867, p. 12, 132. + + [381] 'Nat. Hist. of Selbourne,' letter xxix. edit. of 1825, + vol. i. p. 139. + + [382] Mr. Jenner Weir received similar information, on making + enquiries during the following year. To shew the number of + chaffinches caught, I may mention that in 1869 there was a + match between two experts; and one man caught in a day 62, and + another 40, male chaffinches. The greatest number ever caught + by one man in a single day was 70. + + [383] 'Ibis,' vol. ii. p. 260, as quoted in Gould's + 'Trochilid,' 1861, p. 52. For the foregoing proportions, I am + indebted to Mr. Salvin for a table of his results. + + [384] 'Ibis,' 1860, p. 137; and 1867, p. 369. + + [385] 'Ibis,' 1862, p. 137. + + [386] Leuckart quotes Bloch (Wagner, 'Handwrterbuch der Phys.' + B. iv. 1853, s. 775), that with fish there are twice as many + males as females. + + [387] Quoted in the 'Farmer,' March 18, 1869, p. 369. + + [388] 'The Stormontfield Piscicultural Experiments,' 1866, p. + 23. The 'Field' newspaper, June 29th, 1867. + + [389] 'Land and Water,' 1868, p. 41. + + [390] Yarrell, 'Hist. British Fishes,' vol. i. 1836, p. 307; on + the _Cyprinus carpio_, p. 331; on the _Tinca vulgaris_, p. 331; + on the _Abramis brama_, p. 336. See, for the minnow (_Leuciscus + phoxinus_), 'Loudon's Mag. of Nat. Hist.' vol. v. 1832, p. 682. + + [391] Leuckart quotes Meinecke (Wagner, 'Handwrterbuch der + Phys.' B. iv. 1853, s. 775) that with Butterflies the males are + three or four times as numerous as the females. + + [392] 'The Naturalist on the Amazons,' vol. ii. 1863, p. 228, + 347. + + [393] Four of these cases are given by Mr. Trimen in his + 'Rhopalocera Afric Australis.' + + [394] Quoted by Trimen, 'Transact. Ent. Soc.' vol. v. part iv. + 1866, p. 330. + + [395] 'Transact. Linn. Soc.' vol. xxv. p. 37. + + [396] 'Proc. Entomolog. Soc.' Feb. 17th, 1868. + + [397] Quoted by Dr. Wallace in 'Proc. Ent. Soc.' 3rd series, + vol. v. 1867, p. 487. + + [398] Blanchard, 'Metamorphoses, Moeurs des Insectes,' 1868, p. + 225-226. + + [399] 'Lepidopteren-Doubblettren Liste,' Berlin, No. x. 1866. + + [400] This naturalist has been so kind as to send me some + results from former years, in which the females seemed to + preponderate; but so many of the figures were estimates, that I + found it impossible to tabulate them. + + [401] Gnther's 'Record of Zoological Literature,' 1867, p. + 260. On the excess of female Lucanus, ibid. p. 250. On the + males of Lucanus in England, Westwood, 'Modern Class. of + Insects,' vol. i. p. 187. On the Siagonium, ibid. p. 172. + + [402] Walsh, in 'The American Entomologist,' vol. i. 1869, p. + 103. F. Smith, 'Record of Zoological Literature,' 1867, p. 328. + + [403] 'Farm Insects,' p. 45-46. + + [404] 'Observations on N. American Neuroptera,' by H. Hagen and + B. D. Walsh, 'Proc. Ent. Soc. Philadelphia,' Oct. 1863, p. 168, + 223, 239. + + [405] 'Proc. Ent. Soc. London,' Feb. 17, 1868. + + [406] Another great authority in this class, Prof. Thorell of + Upsala ('On European Spiders,' 1869-70, part i. p. 205) speaks + as if female spiders were generally commoner than the males. + + [407] See, on this subject, Mr. Pickard-Cambridge, as quoted in + 'Quarterly Journal of Science,' 1868, p. 429. + + [408] I have often been struck with the fact, that in several + species of Primula the seeds in the capsules which contained + only a few were very much larger than the numerous seeds in the + more productive capsules. + + [409] 'Principles of Biology,' vol. ii. 1867, chaps. ii.-xi. + + [410] 'De l'Espce et de la Class.' &c., 1869, p. 106. + + [411] See, for instance, the account which I have given in my + 'Journal of Researches,' 1845, p. 7. + + [412] I have given ('Geolog. Observations on Volcanic Islands,' + 1844, p. 53) a curious instance of the influence of light on + the colours of a frondescent incrustation, deposited by the + surf on the coast-rocks of Ascension, and formed by the + solution of triturated sea-shells. + + [413] 'Facts and Arguments for Darwin,' English translat. 1869, + p. 20. See the previous discussion on the olfactory threads. + Sars has described a somewhat analogous case (as quoted in + 'Nature,' 1870, p. 455) in a Norwegian crustacean, the + _Pontoporeia affinis_. + + [414] See Sir J. Lubbock in 'Annals. and Mag. of Nat. Hist.' + vol. xi. 1853, pl. i. and x.; and vol. xii. (1853) pl. vii. See + also Lubbock in 'Transact. Ent. Soc.' vol. iv. new series, + 1856-1858, p. 8. With respect to the zigzagged antenn + mentioned below, see Fritz Mller, 'Facts and Arguments for + Darwin' 1869, p. 40, foot-note. + + [415] See a paper by Mr. C. Spence Bate, with figures, in + 'Proc. Zoolog. Soc.' 1868, p. 363; and on the nomenclature of + the genus, ibid. p. 585. I am greatly indebted to Mr. Spence + Bate for nearly all the above statements with respect to the + chel of the higher crustaceans. + + [416] 'Hist. Nat. des Crust.' tom. ii. 1837, p. 50. + + [417] Fritz Mller, 'Facts and Arguments for Darwin,' 1869, p. + 25-28. + + [418] 'Travels in the Interior of Brazil,' 1846, p. 111. I have + given, in my 'Journal of Researches,' p. 463, an account of the + habits of the Birgos. + + [419] Mr. Ch. Fraser, in 'Proc. Zoolog. Soc.' 1869, p. 3. I am + indebted to Mr. Bate for the statement from Dr. Power. + + [420] Claus, 'Die freilebenden Copepoden,' 1863, s. 35. + + [421] 'Facts and Arguments,' &c., p. 79. + + [422] 'A History of the Spiders of Great Britain,' 1861-64. For + the following facts, see p. 102, 77, 88. + + [423] Aug. Vinson ('Aranides des Iles de la Runion,' pl. vi. + figs. 1 and 2) gives a good instance of the small size of the + male in _Epeira nigra_. In this species, as I may add, the male + is testaceous and the female black with legs banded with red. + Other even more striking cases of inequality in size between + the sexes have been recorded ('Quarterly Journal of Science,' + 1868, July, p. 429); but I have not seen the original accounts. + + [424] Kirby and Spence, 'Introduction to Entomology,' vol. i. + 1818, p. 280. + + [425] Theridion (Asagena, Sund.) serratipes, 4-punctatum et + guttatum; see Westring, in Kroyer, 'Naturhist. Tidskrift,' vol. + iv. 1842-1843, p. 349; and vol. ii. 1846-1849, p. 342. See, + also, for other species, 'Arane Svecic,' p. 184. + + [426] Walckenaer et P. Gervais, 'Hist. Nat. des Insectes: + Aptres,' tom. iv. 1847, p. 17, 19, 68. + + [427] Sir J. Lubbock, 'Transact. Linnean Soc.' vol. xxv. 1866, + p. 484. With respect to the Mutillid see Westwood, 'Modern + Class. of Insects,' vol. ii. p. 213. + + [428] These organs in the male often differ in closely-allied + species, and afford excellent specific characters. But their + importance, under a functional point of view, as Mr. E. + MacLachlan has remarked to me, has probably been overrated. It + has been suggested, that slight differences in these organs + would suffice to prevent the intercrossing of well-marked + varieties or incipient species, and would thus aid in their + development. That this can hardly be the case, we may infer + from the many recorded cases (see for instance, Bronn, + 'Geschichte der Natur,' B. ii. 1843, s. 164; and Westwood, + 'Transact. Ent. Soc.' vol. iii. 1842, p. 195) of distinct + species having been observed in union. Mr. MacLachlan informs + me (vide 'Stett. Ent. Zeitung,' 1867, s. 155) that when several + species of Phryganid, which present strongly-pronounced + differences of this kind, were confined together by Dr. Aug. + Meyer, _they coupled_, and one pair produced fertile ova. + + [429] 'The Practical Entomologist,' Philadelphia, vol. ii. May, + 1867, p. 88. + + [430] Mr. Walsh, ibid. p. 107. + + [431] 'Modern Classification of Insects,' vol. ii. 1840, p. + 206, 205. Mr. Walsh, who called my attention to this double use + of the jaws, says that he has repeatedly observed this fact. + + [432] We have here a curious and inexplicable case of + dimorphism, for some of the females of four European species of + Dytiscus, and of certain species of Hydroporus, have their + elytra smooth; and no intermediate gradations between sulcated + or punctured and quite smooth elytra have been observed. See + Dr. H. Schaum, as quoted in the 'Zoologist,' vol. v.-vi. + 1847-48, p. 1896. Also Kirby and Spence, 'Introduction to + Entomology,' vol. iii. 1826, p. 305. + + [433] Westwood, 'Modern Class.' vol. ii. p. 193. The following + statement about Penthe, and others in inverted commas, are + taken from Mr. Walsh, 'Practical Entomologist,' Philadelphia, + vol. ii. p. 88. + + [434] Kirby and Spence, 'Introduct.' &c., vol. iii. p. 332-336. + + [435] 'Insecta Maderensia,' 1854, p. 20. + + [436] E. Doubleday, 'Annals and Mag. of Nat. Hist.' vol. i. + 1848, p. 379. I may add that the wings in certain Hymenoptera + (see Shuckard, 'Fossorial Hymenop.' 1837, p. 39-43) differ in + neuration according to sex. + + [437] H. W. Bates, in 'Journal of Proc. Linn. Soc.' vol. vi. + 1862, p. 74. Mr. Wonfor's observations are quoted in 'Popular + Science Review,' 1868, p. 343. + + [438] Kirby and Spence, 'Introduction to Entomology,' vol. iii. + p. 299. + + [439] Robinet, 'Vers Soie,' 1848, p. 207. + + [440] 'Transact. Ent. Soc.' 3rd series, vol. v. p. 486. + + [441] 'Journal of Proc. Ent. Soc.' Feb. 4th, 1867, p. lxxi. + + [442] For this and other statements on the size of the sexes, + see Kirby and Spence, ibid. vol. iii. p. 300; on the duration + of life in insects, see p. 344. + + [443] 'Transact. Linnean Soc.' vol. xxvi. 1868, p. 296. + + [444] 'The Malay Archipelago,' vol. ii. 1869, p. 313. + + [445] 'Modern Classification of Insects,' vol. ii. 1840, p. + 526. + + [446] See Mr. B. T. Lowne's very interesting work, 'On the + Anatomy of the Blow-Fly, Musca vomitoria,' 1870, p. 14. + + [447] Westwood, 'Modern Class. of Insects,' vol. ii. p. 473. + + [448] These particulars are taken from Westwood's 'Modern + Class. of Insects,' vol. ii. 1840, p. 422. See, also, on the + Fulgorid, Kirby and Spence, 'Introduct.' vol. ii. p. 401. + + [449] 'Zeitschrift fr wissenschaft. Zoolog.' B. xvii. 1867, s. + 152-158. + + [450] I am indebted to Mr. Walsh for having sent me this + extract from a 'Journal of the Doings of Cicada septemdecim,' + by Dr. Hartman. + + [451] L. Guilding, 'Transact. Linn. Soc.' vol. xv. p. 154. + + [452] Kppen, as quoted in the 'Zoological Record,' for 1867, + p. 460. + + [453] Gilbert White, 'Nat. Hist. of Selborne,' vol. ii. 1825, + p. 262. + + [454] Harris, 'Insects of New England,' 1842, p. 128. + + [455] 'The Naturalist on the Amazons,' vol. i. 1863, p. 252. + Mr. Bates gives a very interesting discussion on the gradations + in the musical apparatus of the three families. See also + Westwood, 'Modern Class.' vol. ii. p. 445 and 453. + + [456] 'Proc. Boston Soc. of Nat. Hist.' vol. xi. April, 1868. + + [457] 'Nouveau Manuel d'Anat. Comp.' (French translat.), tom. + i. 1850 p. 567. + + [458] 'Zeitschrift fr wissenschaft. Zoolog.' B. xvii. 1867, s. + 117. + + [459] Westwood, 'Modern Class. of Insects,' vol. i. p. 440. + + [460] Westwood, 'Modern Class. of Insects,' vol. i. p. 453. + + [461] Landois, ibid. s. 121, 122. + + [462] Mr. Walsh also informs me that he has noticed that the + female of the _Platyphyllum concavum_, "when captured makes a + feeble grating noise by shuffling her wing-covers together." + + [463] Landois, ibid. s. 113. + + [464] 'Insects of New England,' 1842, p. 133. + + [465] Westwood, 'Modern Classification,' vol. i. p. 462. + + [466] Westwood, ibid. vol. i. p. 453. + + [467] Landois, ibid. s. 115, 116, 120, 122. + + [468] 'Transact. Ent. Soc.' 3rd series, vol. ii. ('Journal of + Proceedings, p. 117.) + + [469] Westwood, 'Modern Class. of Insects,' vol. i. p. 427; for + crickets, p. 445. + + [470] Mr. Ch. Horne, in 'Proc. Ent. Soc.' May 3, 1869, p. xii. + + [471] The _Oecanthus nivalis_, Harris, 'Insects of New + England,' 1842, p. 124. + + [472] Platyblemnus: Westwood, 'Modern. Class.' vol. i. p. 447. + + [473] B. D. Walsh, the Pseudo-neuroptera of Illinois, in 'Proc. + Ent. Soc. of Philadelphia,' 1862, p. 361. + + [474] 'Modern Class.' vol. ii. p. 37. + + [475] Walsh, ibid. p. 381. I am indebted to this naturalist for + the following facts on Hetrina, Anax, and Gomphus. + + [476] 'Transact. Ent. Soc' vol. i. 1836, p. lxxxi. + + [477] See abstract in the 'Zoological Record' for 1867, p. 450. + + [478] Kirby and Spence, 'Introduct. to Entomology,' vol. ii. + 1818, p. 35. + + [479] See an interesting article, "The Writings of Fabre," in + 'Nat. Hist. Review,' April, 1862, p. 122. + + [480] 'Journal of Proc. of Entomolog. Soc.' Sept. 7th, 1863, + p. 169. + + [481] P. Huber, 'Recherches sur les Moeurs des Fourmis,' 1810, + p. 150, 165. + + [482] 'Proc. Entomolog. Soc. of Philadelphia,' 1866, p. + 238-239. + + [483] Quoted by Westwood, 'Modern Class. of Insects,' vol. ii. + p. 214. + + [484] Pyrodes pulcherrimus, in which the sexes differ + conspicuously, has been described by Mr. Bates in 'Transact. + Ent. Soc.' 1869, p. 50. I will specify the few other cases in + which I have heard of a difference in colour between the sexes + of beetles. Kirby and Spence ('Introduct. to Entomology,' vol. + iii. p. 301) mention a Cantharis, Meloe, Rhagium, and the + _Leptura testacea_; the male of the latter being testaceous, + with a black thorax, and the female of a dull red all over. + These two latter beetles belong to the Order of Longicorns. + Messrs. R. Trimen and Waterhouse, junr., inform me of two + Lamellicorns, viz., a Peritrichia and Trichius, the male of the + latter being more obscurely coloured than the female. In + _Tillus elongatus_ the male is black, and the female always, as + it is believed, of a dark blue colour with a red thorax. The + male, also, of _Orsodacna atra_, as I hear from Mr. Walsh, is + black, the female (the so-called _O. ruficollis_) having a + rufous thorax. + + [485] 'Proc. Entomolog. Soc. of Philadelphia,' 1864, p. 228. + + [486] Kirby and Spence, 'Introduct. Entomolog.' vol. iii. p. + 300. + + [487] Kirby and Spence, ibid. vol. iii. p. 329. + + [488] 'Modern Classification of Insects,' vol. i. p. 172. On + the same page there is an account of Siagonium. In the British + Museum I noticed one male specimen of Siagonium in an + intermediate condition, so that the dimorphism is not strict. + + [489] 'The Malay Archipelago,' vol. ii. 1869, p. 276. + + [490] 'Entomological Magazine,' vol. i. 1833, p. 82. See also + on the conflicts of this species, Kirby and Spence, ibid. vol. + iii. p. 314; and Westwood, ibid. vol. i. p. 187. + + [491] Quoted from Fischer, in 'Dict. Class. d'Hist. Nat.' tom. + x. p. 324. + + [492] 'Ann. Soc. Entomolog. France,' 1866, as quoted in + 'Journal of Travel,' by A. Murray, 1868, p. 135. + + [493] Westwood, 'Modern Class.' vol. i. p. 184. + + [494] Wollaston, On certain musical Curculionid, 'Annals and + Mag. of Nat. Hist.' vol. vi. 1860, p. 14. + + [495] 'Zeitschrift fr wiss. Zoolog.' B. xvii. 1867, s. 127. + + [496] I am greatly indebted to Mr. G. R. Crotch for having sent + me numerous prepared specimens of various beetles belonging to + these three families and others, as well as for valuable + information of all kinds. He believes that the power of + stridulation in the Clythra has not been previously observed. I + am also much indebted to Mr. E. W. Janson, for information and + specimens. I may add that my son, Mr. F. Darwin, finds that + _Dermestes murinus_ stridulates, but he searched in vain for + the apparatus. Scolytus has lately been described by Mr. Algen + as a stridulator, in the 'Edinburgh Monthly Magazine,' 1869, + Nov., p. 130. + + [497] Schidte, translated in 'Annals and Mag. of Nat. Hist.' + vol. xx. 1867, p. 37. + + [498] Westring has described (Kroyer, 'Naturhist. Tidskrift,' + B. ii. 1848-49, p. 334) the stridulating organs in these two, + as well as in other families. In the Carabid I have examined + _Elaphrus uliginosus_ and _Blethisa multipunctata_, sent to me + by Mr. Crotch. In Blethisa the transverse ridges on the + furrowed border of the abdominal segment do not come into play, + as far as I could judge, in scraping the rasps on the elytra. + + [499] I am indebted to Mr. Walsh, of Illinois, for having sent + me extracts from Leconte's 'Introduction to Entomology,' p. + 101, 143. + + [500] M. P. de la Brulerie, as quoted in 'Journal of Travel,' + A. Murray, vol. i. 1868, p. 135. + + [501] Mr. Doubleday informs me that "the noise is produced by + the insect raising itself on its legs as high as it can, and + then sinking its thorax five or six times, in rapid succession, + against the substance upon which it is sitting." For references + on this subject see Landois, 'Zeitschrift fr wissen. Zoolog.' + B. xvii. s. 131. Olivier says (as quoted by Kirby and Spence, + 'Introduct.' vol. ii. p. 395) that the female of _Pimelia + striata_ produces a rather loud sound by striking her abdomen + against any hard substance, "and that the male, obedient to + this call, soon attends her and they pair." + + [502] Apatura Iris: 'The Entomologist's Weekly Intelligencer,' + 1859, p. 139. For the Bornean Butterflies see C. Collingwood, + 'Rambles of a Naturalist,' 1868, p. 183. + + [503] See my 'Journal of Researches,' 1845, p. 33. Mr. + Doubleday has detected ('Proc. Ent. Soc.' March 3rd, 1845, p. + 123) a peculiar membranous sac at the base of the front wings, + which is probably connected with the production of the sound. + + [504] See also Mr. Bates' paper in 'Proc. Ent. Soc. of + Philadelphia,' 1865, p. 206. Also Mr. Wallace on the same + subject, in regard to Diadema, in 'Transact. Entomolog. Soc. of + London,' 1869, p. 278. + + [505] 'The Naturalist on the Amazons,' vol. i. 1863, p. 19. + + [506] See the interesting article in the 'Westminster Review,' + July, 1867, p. 10. A woodcut of the Kallima is given by Mr. + Wallace in Hardwicke's 'Science Gossip,' Sept. 1867, p. 196. + + [507] See the interesting observations by Mr. T. W. Wood, 'The + Student,' Sept. 1868, p. 81. + + [508] Mr. Wallace in 'Hardwicke's Science Gossip,' Sept. 1867, + p. 193. + + [509] See also, on this subject, Mr. Weir's paper in 'Transact. + Ent. Soc.' 1869, p. 23. + + [510] 'Westminster Review,' July, 1867, p. 16. + + [511] For instance, Lithosia; but Prof. Westwood ('Modern + Class. of Insects,' vol. ii. p. 390) seems surprised at this + case. On the relative colours of diurnal and nocturnal + Lepidoptera, see ibid. p. 333 and 392; also Harris, 'Treatise + on the Insects of New England,' 1842, p. 315. + + [512] Such differences between the upper and lower surfaces of + the wings of several species of Papilio, may be seen in the + beautiful plates to Mr. Wallace's Memoir on the Papilionid of + the Malayan Region, in 'Transact. Linn. Soc.' vol. xxv. part i. + 1865. + + [513] 'Proc. Ent. Soc.' March 2nd, 1868. + + [514] See also an account of the S. American genus Erateina + (one of the Geometr) in 'Transact. Ent. Soc.' new series, vol. + v. pl. xv. and xvi. + + [515] 'Proc. Ent. Soc. of London,' July 6, 1868, p. xxvii. + + [516] Harris, 'Treatise,' &c., edited by Flint, 1862, p. 395. + + [517] For instance, I observe in my son's cabinet that the + males are darker than the females in the _Lasiocampa quercus_, + _Odonestis potatoria_, _Hypogymna dispar_, _Dasychira + pudibunda_, and _Cycnia mendica_. In this latter species the + difference in colour between the two sexes is strongly marked; + and Mr. Wallace informs me that we here have, as he believes, + an instance of protective mimickry confined to one sex, as will + hereafter be more fully explained. The white female of the + Cycnia resembles the very common _Spilosoma menthrasti_, both + sexes of which are white; and Mr. Stainton observed that this + latter moth was rejected with utter disgust by a whole brood of + young turkeys, which were fond of eating other moths; so that + if the Cycnia was commonly mistaken by British birds for the + Spilosoma, it would escape being devoured, and its white + deceptive colour would thus be highly beneficial. + + [518] 'Rambles of a Naturalist in the Chinese Seas,' 1868, p. + 182. + + [519] Wallace on the Papilionid of the Malayan Region, in + 'Transact. Linn. Soc.' vol. xxv. 1865, p. 8, 36. A striking case + of a rare variety, strictly intermediate between two other + well-marked female varieties, is given by Mr. Wallace. See also + Mr. Bates, in 'Proc. Entomolog. Soc.' Nov. 19th, 1866, p. xl. + + [520] Mr. R. MacLachlan, 'Transact. Ent. Soc.' vol. ii. part + 6th, 3rd series, 1866, p. 459. + + [521] H. W. Bates, 'The Naturalist on the Amazons,' vol. ii. + 1863, p. 228. A. R. Wallace, in 'Transact. Linn. Soc.' vol. xxv. + 1865, p. 10. + + [522] On this whole subject see 'The Variation of Animals and + Plants under Domestication,' vol. ii. 1868, chap. xxiii. + + [523] A. R. Wallace, in 'The Journal of Travel,' vol. i. 1868, + p. 88. 'Westminster Review,' July, 1857, p. 37. See also + Messrs. Wallace and Bates in 'Proc. Ent. Soc.' Nov. 19th, 1866, + p. xxxix. + + [524] 'The Variation of Animals and Plants under + Domestication,' vol. ii. chap. xii. p. 17. + + [525] 'Transact. Linn. Soc.' vol. xxiii. 1862, p. 495. + + [526] 'Proc. Ent. Soc.' Dec. 3rd, 1866, p. xlv. + + [527] 'Transact. Linn. Soc.' vol. xxv. 1865, p. 1; also + 'Transact. Ent. Soc.' vol. iv. (3rd series), 1867, p. 301. + + [528] See an ingenious article entitled, "Difficulties of the + Theory of Natural Selection," in the 'Month,' 1869. The writer + strangely supposes that I attribute the variations in colour of + the Lepidoptera, by which certain species belonging to distinct + families have come to resemble others, to reversion to a common + progenitor; but there is no more reason to attribute these + variations to reversion than in the case of any ordinary + variation. + + [529] Wallace, "Notes on Eastern Butterflies," 'Transact. Ent. + Soc.' 1869, p. 287. + + [530] Wallace, in 'Westminster Review,' July, 1867, p. 37; and + in 'Journal of Travel and Nat. Hist.' vol. i. 1868, p. 88. + + [531] See remarks by Messrs. Bates and Wallace, in 'Proc. Ent. + Soc.' Nov. 19, 1866, p. xxxix. + + [532] See Mr. Wallace in 'Westminster Review,' July, 1867, p. + 11 and 37. The male of no butterfly, as Mr. Wallace informs me, + is known to differ in colour, as a protection, from the female; + and he asks me how I can explain this fact on the principle + that one sex alone has varied and has transmitted its + variations exclusively to the same sex, without the aid of + selection to check the variations being inherited by the other + sex. No doubt if it could be shewn that the females of very + many species had been rendered beautiful through protective + mimickry, but that this has never occurred with the males, it + would be a serious difficulty. But the number of cases as yet + known hardly suffices for a fair judgment. We can see that the + males, from having the power of flying more swiftly, and thus + escaping danger, would not be so likely as the females to have + had their colours modified for the sake of protection; but this + would not in the least have interfered with their receiving + protective colours through inheritance from the females. In the + second place, it is probable that sexual selection would + actually tend to prevent a beautiful male from becoming + obscure, for the less brilliant individuals would be less + attractive to the females. Supposing that the beauty of the + male of any species had been mainly acquired through sexual + selection, yet if this beauty likewise served as a protection, + the acquisition would have been aided by natural selection. But + it would be quite beyond our power to distinguish between the + two processes of sexual and ordinary selection. Hence it is not + likely that we should be able to adduce cases of the males + having been rendered brilliant exclusively through protective + mimickry, though this is comparatively easy with the females, + which have rarely or never been rendered beautiful, as far as + we can judge, for the sake of sexual attraction, although they + have often received beauty through inheritance from their male + parents. + + [533] 'Proc. Entomolog. Soc.' Dec. 3rd, 1866, p. xlv., and + March 4th, 1867, p. lxxx. + + [534] See Mr. J. Jenner Weir's paper on insects and + insectivorous birds, in 'Transact. Ent. Soc.' 1869, p. 21; also + Mr. Butler's paper, ibid. p. 27. + + +END OF VOL. I. + +LONDON: + +PRINTED BY WILLIAM CLOWES AND SONS, DUKE STREET, STAMFORD STREET, AND +CHARING CROSS. + + +50A, ALBEMARLE STREET, LONDON, +_January, 1871_. + + +MR. MURRAY'S + +LIST OF POPULAR WORKS. + + +_One Shilling._ + +THE PRINCIPAL SPEECHES AND ADDRESSES OF H.R.H. 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By HERMAN MELVILLE. + +OMOO: A Narrative of Adventures in the South Seas; a Sequel to 'Typee.' +By HERMAN MELVILLE. + +THE BIBLE IN THE HOLY LAND; being Extracts from Dean Stanley's 'Sinai +and Palestine.' With Woodcuts. + +AN ESSAY ON ENGLISH POETRY. With Short Notices of the British Poets. By +THOMAS CAMPBELL. + +HISTORICAL AND CRITICAL ESSAYS. By LORD MAHON. + +LIFE OF LORD CLIVE. By REV. G. R. GLEIG. + +LIFE OF SIR THOMAS MUNRO. With Selections from his Correspondence. By +REV. G. R. GLEIG. + +THE RISE OF OUR INDIAN EMPIRE: being a History of British India from its +Origin till the Peace of 1783. By LORD MAHON. + +A NARRATIVE OF THE SIEGE OF KARS, and of the Six Months' Resistance by +the Turkish Garrison under General Williams. With Travels and Adventures +in Armenia, &c. By HUMPHREY SANDWITH, M.D. + +SKETCHES OF PERSIA. By SIR JOHN MALCOLM. + +THE WILD SPORTS AND NATURAL HISTORY OF THE HIGHLANDS. By CHARLES ST. +JOHN. + +GATHERINGS FROM SPAIN. 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You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: The Descent of Man and Selection in Relation to Sex, Vol. I (1st edition) + +Author: Charles Darwin + +Release Date: January 15, 2011 [EBook #34967] + +Language: English + +Character set encoding: UTF-8 + +*** START OF THIS PROJECT GUTENBERG EBOOK THE DESCENT OF MAN AND *** + + + + +Produced by StevenGibbs, Turgut Dincer and the Online +Distributed Proofreading Team at http://www.pgdp.net + + + + + + +</pre> + + +<h4>THE</h4> +<h1>DESCENT OF MAN,</h1> + +<h5>AND</h5> + +<h2>SELECTION IN RELATION TO SEX.</h2> + +<h3><span class="smcap">By</span> CHARLES DARWIN, M.A., F.R.S., &c.</h3> + +<h4>IN TWO VOLUMES.—<span class="smcap">Vol. I.</span></h4> + +<h6>WITH ILLUSTRATIONS.</h6> + +<h4>LONDON:<br /> + +JOHN MURRAY, ALBEMARLE STREET.<br /> + +1871.</h4> + +<h6>[<i>The right of Translation is reserved.</i>]</h6> + +<h4>BY THE SAME AUTHOR.</h4> + +<p class="indent">ON THE ORIGIN OF SPECIES BY MEANS OF NATURAL +SELECTION; or, <span class="smcap">The Preservation of Favoured Races in the Struggle +for Life</span>; Fifth Edition (<i>Tenth Thousand</i>), with Additions and Corrections. +1869. <span class="smcap">Murray</span>.</p> + +<p class="indent">THE VARIATION OF ANIMALS AND PLANTS UNDER +DOMESTICATION. In two vols. With Illustrations. 1868. <span class="smcap">Murray</span>.</p> + +<p class="indent">ON THE VARIOUS CONTRIVANCES <span class="smcap">by which</span> BRITISH +AND FOREIGN ORCHIDS ARE FERTILISED BY INSECTS; and on the +<span class="smcap">Good Effects of Crossing</span>. With numerous Woodcuts. <span class="smcap">Murray</span>.</p> + +<p class="indent">A NATURALIST‘S VOYAGE ROUND THE WORLD; or, +<span class="smcap">A Journal of Researches into the Natural History and Geology of the +Countries</span> visited during the voyage of H.M.S. ‘Beagle,’ under the command of +Captain <span class="smcap">FitzRoy</span>, R.N. <i>Eleventh Thousand</i>. <span class="smcap">Murray</span>.</p> + +<p class="indent">ON THE STRUCTURE AND DISTRIBUTION OF CORAL +REEFS. <span class="smcap"> Smith, Elder, & Co.</span></p> + +<p class="indent">GEOLOGICAL OBSERVATIONS ON VOLCANIC ISLANDS. +<span class="smcap">Smith, Elder, & Co.</span></p> + +<p class="indent">GEOLOGICAL OBSERVATIONS ON SOUTH AMERICA. +<span class="smcap">Smith, Elder & Co.</span></p> + +<p class="indent">A MONOGRAPH OF THE CIRRIPEDIA. With numerous +Illustrations. 2 vols. 8vo. <span class="smcap">Hardwicke</span>.</p> + +<p class="indent">ON THE MOVEMENTS AND HABITS OF CLIMBING +PLANTS. With Woodcuts. <span class="smcap">Williams</span> & <span class="smcap">Norgate</span>.</p> + +<hr /> + +<h6>LONDON: PRINTED BY WILLIAM CLOWES AND SONS, STAMFORD STREET, + +AND CHARING CROSS.</h6> + +<p><span class='pagenum'><a name="Page_v" id="Page_v">v</a></span></p> +<table width="100%" summary="contents" border="0"> + +<tr> +<td class="center" colspan="2"><h2>CONTENTS.</h2></td> +</tr> +<tr> +<td class="center" colspan="2"><hr /></td> +</tr> +<tr> +<td class="left85"><span class="smcap">Introduction</span></td> +<td class="right15">Page <a href="#Page_1">1-5</a></td> +</tr> +<tr> +<td class="center" colspan="2"><h3>PART I.</h3></td> +</tr> +<tr> +<td class="center2" colspan="2"><h4>ON THE DESCENT OF MAN</h4></td> +</tr> +<tr> +<td class="center" colspan="2">CHAPTER I.</td> +</tr> +<tr> +<td class="center2" colspan="2"><span class="smcap">The Evidence of the Descent of man from some +Lower form.</span></td> +</tr> +<tr> +<td class="left85">Nature of the evidence bearing on the origin of man—Homologous +structures in man and the lower animals—Miscellaneous points +of correspondence—Development—Rudimentary structures, +muscles, sense-organs, hair, bones, reproductive organs, &c.—The +bearing of these three great classes of facts on the origin of +man </td> +<td class="right15"><a href="#Page_9">9-33</a></td> +</tr> +<tr> +<td class="center" colspan="2">CHAPTER II.</td> +</tr> +<tr> +<td class="center2" colspan="2"><span class="smcap">Comparison of the Mental Powers of Man and the +Lower Animals</span>.</td> +</tr> +<tr> +<td class="left85">The difference in mental power between the highest ape and the +lowest savage, immense—Certain instincts in common—The +emotions—Curiosity—Imitation—Attention—Memory—Imagination—Reason—Progressive +improvement—Tools and +weapons used by animals—Language—Self-consciousness—Sense +of beauty—Belief in God, spiritual agencies, superstitions</td> +<td class="right15"><a href="#Page_34">34-69</a></td> +</tr> +<tr> +<td class="center" colspan="2">CHAPTER III.</td> +</tr> +<tr> +<td class="center2" colspan="2"><span class="smcap">Comparison of the Mental Powers of Man and the +Lower Animals</span>—<i>continued</i>.</td> +</tr> +<tr> +<td class="left85">The difference in mental power between the highest ape and the +lowest savage, immense—Certain instincts in common—The +emotions—Curiosity—Imitation—Attention—Memory—Imagination—Reason—Progressive +improvement—Tools and +weapons used by animals—Language—Self-consciousness—Sense +of beauty—Belief in God, spiritual agencies, superstitions</td> +<td class="right15"><a href="#Page_70">70-106</a></td> +</tr> + +<tr> +<td class="center" colspan="2">CHAPTER IV.</td> +</tr> +<tr> +<td class="center2" colspan="2"><span class="smcap">On the Manner of Development of Man from some +Lower Form.</span></td> +</tr> +<tr> +<td class="left85">Variability of body and mind in man—Inheritance—Causes of +variability—Laws of variation the same in man as in the lower +animals—Direct action of the conditions of life—Effects of the +increased use and disuse of parts—Arrested development—Reversion—Correlated +variation—Rate of increase—Checks to +increase—Natural selection—Man the most dominant animal +in the world—Importance of his corporeal structure—The +causes which have led to his becoming erect—Consequent changes +of structure—Decrease in size of the canine teeth—Increased +size and altered shape of the skull—Nakedness—Absence of +a tail—Defenceless condition of man</td> +<td class="right15"><a href="#Page_107">107-157</a></td> +</tr> + +<tr> +<td class="center" colspan="2">CHAPTER V.</td> +</tr> +<tr> +<td class="center2" colspan="2"><span class="smcap">On the Development of the Intellectual and Moral +Faculties during Primeval and Civilised Times.</span></td> +</tr> +<tr> +<td class="left85">The advancement of the intellectual powers through natural selection—Importance +of imitation—Social and moral faculties—Their +development within the limits of the same tribe—Natural +selection as affecting civilised nations—Evidence that civilised +nations were once barbarous</td> +<td class="right15"><a href="#Page_158">158-184</a></td> +</tr> +<tr> +<td class="center" colspan="2">CHAPTER VI.</td> +</tr> +<tr> +<td class="center2" colspan="2"><span class="smcap">On the Affinities and Genealogy of Man.</span></td> +</tr> +<tr> +<td class="left85">Position of man in the animal series—The natural system genealogical—Adaptive +characters of slight value—Various small +points of resemblance between man and the Quadrumana—Rank +of man in the natural system—Birthplace and antiquity +<span class='pagenum'><a name="Page_vii" id="Page_vii">vii</a></span> +of man—Absence of fossil connecting-links—Lower stages in +the genealogy of man, as inferred, firstly from his affinities and +secondly from his structure—Early androgynous condition of +the Vertebrata—Conclusion</td> +<td class="right15"><a href="#Page_195">185-213</a></td> +</tr> + +<tr> +<td class="center" colspan="2">CHAPTER VII.</td> +</tr> +<tr> +<td class="center2" colspan="2"><span class="smcap">On the Races of Man.</span></td> +</tr> +<tr> +<td class="left85">The nature and value of specific characters—Application to the races +of man—Arguments in favour of, and opposed to, ranking the +so-called races of man as distinct species—Sub-species—Monogenists +and polygenists—Convergence of character—Numerous +points of resemblance in body and mind between the most distinct +races of man—The state of man when he first spread over the +earth—Each race not descended from a single pair—The extinction +of races—The formation of races—The effects of crossing—Slight +influence of the direct action of the conditions of life—Slight +or no influence of natural selection—Sexual selection.</td> +<td class="right15"><a href="#Page_214">214-250</a></td> +</tr> +<tr> +<td class="center2" colspan="2"><hr /></td> +</tr> +</table> + +<table width="100%" summary="contents"> +<tr> +<td class="center" colspan="2"><h3>PART II.<br /><br /><small>SEXUAL SELECTION.</small></h3></td> +</tr> +<tr> +<td class="center" colspan="2">CHAPTER VIII.</td> +</tr> +<tr> +<td class="center2" colspan="2"><span class="smcap">Principles of Sexual Selection.</span></td> +</tr> +<tr> +<td class="left85">Secondary sexual characters—Sexual selection—Manner of action—Excess +of males—Polygamy—The male alone generally +modified through sexual selection—Eagerness of the male—Variability +of the male—Choice exerted by the female—Sexual +compared with natural selection—Inheritance at corresponding +periods of life, at corresponding seasons of the year, and as limited +by sex—Relations between the several forms of inheritance—Causes +why one sex and the young are not modified through +sexual selection—Supplement on the proportional numbers of +the two sexes throughout the animal kingdom—On the limitation +of the numbers of the two sexes through natural selection</td> +<td class="right15"><a href="#Page_253">253-320</a></td> +</tr> + +<tr> +<td class="center" colspan="2">CHAPTER IX.</td> +</tr> +<tr> +<td class="center2" colspan="2"><span class="smcap">Secondary Sexual Characters in the Lower Classes of +the Animal Kingdom.</span></td> +</tr> +<tr> +<td class="left85"><span class='pagenum'><a name="Page_viii" id="Page_viii">viii</a></span> +These characters absent in the lowest classes—Brilliant colours—Mollusca—Annelids—Crustacea, +secondary sexual characters +strongly developed; dimorphism; colour; characters not acquired +before maturity—Spiders, sexual colours of; stridulation by the +males—Myriapoda</td> +<td class="right15"><a href="#Page_321">321-340</a></td> +</tr> + +<tr> +<td class="center" colspan="2">CHAPTER X.</td> +</tr> +<tr> +<td class="center2" colspan="2"><span class="smcap">Secondary Sexual Characters of Insects.</span></td> +</tr> +<tr> +<td class="left85">Diversified structures possessed by the males for seizing the females—Differences +between the sexes, of which the meaning is not +understood—Difference in size between the sexes—Thysanura—Diptera—Hemiptera—Homoptera, +musical powers possessed +by the males alone—Orthoptera, musical instruments of the +males, much diversified in structure; pugnacity; colours—Neuroptera, +sexual differences in colour—Hymenoptera, pugnacity +and colours—Coleoptera, colours; furnished with great +horns, apparently as an ornament; battles; stridulating organs +generally common to both sexes</td> +<td class="right15"><a href="#Page_341">341-385</a></td> +</tr> +<tr> +<td class="center" colspan="2">CHAPTER XI.</td> +</tr> +<tr> +<td class="center2" colspan="2"><span class="smcap">Insects</span>, <i>continued</i>.—<span class="smcap">Order Lepidoptera.</span> +</td></tr> +<tr> +<td class="left85">Courtship of butterflies—Battles—Ticking noise—Colours common +to both sexes, or more brilliant in the males—Examples—Not +due to the direct action of the conditions of life—Colours +adapted for protection—Colours of moths—Display—Perceptive +powers of the Lepidoptera—Variability—Causes of the +difference in colour between the males and females—Mimickry, +female butterflies more brilliantly coloured than the males—Bright +colours of caterpillars—Summary and concluding remarks +on the secondary sexual characters of insects—Birds +and insects compared</td> +<td class="right15"><a href="#Page_386">386-423</a></td> +</tr> +</table> + +<hr /> + +<p><span class='pagenum'><a name="Page_1" id="Page_1">1</a></span></p> + +<h1>THE DESCENT OF MAN;</h1> + +<h6>AND ON</h6> + +<h3>SELECTION IN RELATION TO SEX.</h3> + +<h4>INTRODUCTION.</h4> + +<p>The nature of the following work will be best understood +by a brief account of how it came to be written. +During many years I collected notes on the origin or +descent of man, without any intention of publishing on +the subject, but rather with the determination not to +publish, as I thought that I should thus only add to +the prejudices against my views. It seemed to me sufficient +to indicate, in the first edition of my ‘Origin of +Species,’ that by this work “light would be thrown on +the origin of man and his history;” and this implies +that man must be included with other organic beings in +any general conclusion respecting his manner of appearance +on this earth. Now the case wears a wholly different +aspect. When a naturalist like Carl Vogt ventures +to say in his address as President of the National +Institution of Geneva (1869), “personne, en Europe +au moins, n’ose plus soutenir la création indépendante +et de toutes pièces, des espèces,” it is manifest that at +least a large number of naturalists must admit that +species are the modified descendants of other species;<span class='pagenum'><a name="Page_2" id="Page_2">2</a></span> +and this especially holds good with the younger and +rising naturalists. The greater number accept the +agency of natural selection; though some urge, whether +with justice the future must decide, that I have greatly +overrated its importance. Of the older and honoured +chiefs in natural science, many unfortunately are still +opposed to evolution in every form.</p> + +<p>In consequence of the views now adopted by most +naturalists, and which will ultimately, as in every other +case, be followed by other men, I have been led to put +together my notes, so as to see how far the general +conclusions arrived at in my former works were applicable +to man. This seemed all the more desirable +as I had never deliberately applied these views to a +species taken singly. When we confine our attention +to any one form, we are deprived of the weighty arguments +derived from the nature of the affinities which +connect together whole groups of organisms—their geographical +distribution in past and present times, and +their geological succession. The homological structure, +embryological development, and rudimentary organs of +a species, whether it be man or any other animal, to +which our attention may be directed, remain to be considered; +but these great classes of facts afford, as it +appears to me, ample and conclusive evidence in favour +of the principle of gradual evolution. The strong support +derived from the other arguments should, however, +always be kept before the mind.</p> + +<p>The sole object of this work is to consider, firstly, +whether man, like every other species, is descended +from some pre-existing form; secondly, the manner of<span class='pagenum'><a name="Page_3" id="Page_3">3</a></span> +his development; and thirdly, the value of the differences +between the so-called races of man. As I shall +confine myself to these points, it will not be necessary +to describe in detail the differences between the several +races—an enormous subject which has been fully discussed +in many valuable works. The high antiquity of +man has recently been demonstrated by the labours +of a host of eminent men, beginning with M. Boucher +de Perthes; and this is the indispensable basis for +understanding his origin. I shall, therefore, take this +conclusion for granted, and may refer my readers to +the admirable treatises of Sir Charles Lyell, Sir John +Lubbock, and others. Nor shall I have occasion to do +more than to allude to the amount of difference between +man and the anthropomorphous apes; for Prof. Huxley, +in the opinion of most competent judges, has conclusively +shewn that in every single visible character man +differs less from the higher apes than these do from the +lower members of the same order of Primates.</p> + +<p>This work contains hardly any original facts in +regard to man; but as the conclusions at which I +arrived, after drawing up a rough draft, appeared to +me interesting, I thought that they might interest +others. It has often and confidently been asserted, that +man’s origin can never be known: but ignorance more +frequently begets confidence than does knowledge: it is +those who know little, and not those who know much, +who so positively assert that this or that problem will +never be solved by science. The conclusion that man is +the co-descendant with other species of some ancient, +lower, and extinct form, is not in any degree new. La<span class='pagenum'><a name="Page_4" id="Page_4">4</a></span>marck +long ago came to this conclusion, which has lately +been maintained by several eminent naturalists and +philosophers; for instance by Wallace, Huxley, Lyell, +Vogt, Lubbock, Büchner, Rolle, &c.,<a name="FNanchor_1" id="FNanchor_1"></a><a href="#Footnote_1" class="fnanchor">1</a> and especially by +Häckel. This last naturalist, besides his great work, +'Generelle Morphologie ‘(1866), has recently (1868, +with a second edit. in 1870), published his ‘Natürliche +Schöpfungsgeschichte, ‘in which he fully discusses the +genealogy of man. If this work had appeared before +my essay had been written, I should probably never +have completed it. Almost all the conclusions at which +I have arrived I find confirmed by this naturalist, whose +knowledge on many points is much fuller than mine. +Wherever I have added any fact or view from Prof. +Häckel’s writings, I give his authority in the text, other +statements I leave as they originally stood in my manuscript, +occasionally giving in the foot-notes references +to his works, as a confirmation of the more doubtful or +interesting points.</p> + +<p>During many years it has seemed to me highly probable +that sexual selection has played an important +part in differentiating the races of man; but in my +<span class='pagenum'><a name="Page_5" id="Page_5">5<br />6</a></span>‘Origin of Species’ (first edition, p. 199) I contented +myself by merely alluding to this belief. When I came +to apply this view to man, I found it indispensable to +treat the whole subject in full detail.<a name="FNanchor_2" id="FNanchor_2"></a><a href="#Footnote_2" class="fnanchor">2</a> Consequently +the second part of the present work, treating of sexual +selection, has extended to an inordinate length, compared +with the first part; but this could not be +avoided.</p> + +<p>I had intended adding to the present volumes an +essay on the expression of the various emotions by man +and the lower animals. My attention was called to this +subject many years ago by Sir Charles Bell’s admirable +work. This illustrious anatomist maintains that man +is endowed with certain muscles solely for the sake +of expressing his emotions. As this view is obviously +opposed to the belief that man is descended from some +other and lower form, it was necessary for me to consider +it. I likewise wished to ascertain how far the emotions +are expressed in the same manner by the different races +of man. But owing to the length of the present work, +I have thought it better to reserve my essay, which is +partially completed, for separate publication.</p> +<hr /> +<p><span class='pagenum'><a name="Page_7" id="Page_7">7<br />8</a></span></p> + +<h4><span class="smcap">Part I.</span></h4> + +<h3>THE DESCENT OR ORIGIN OF MAN.</h3> +<hr /> +<p><span class='pagenum'><a name="Page_9" id="Page_9">9</a></span></p> + +<h3><span class="smcap">Part I.</span>—THE DESCENT OF MAN.</h3> + +<hr /> +<h4>CHAPTER I.</h4> + +<h4><span class="smcap">The Evidence of the Descent of Man from some +Lower Form.</span></h4> + +<div class="blockquot"><p>Nature of the evidence bearing on the origin of man—Homologous +structures in man and the lower animals—Miscellaneous points +of correspondence—Development—Rudimentary structures, +muscles, sense-organs, hair, bones, reproductive organs, &c.—The +bearing of these three great classes of facts on the origin +of man.</p></div> + +<p>He who wishes to decide whether man is the modified +descendant of some pre-existing form, would probably +first enquire whether man varies, however slightly, in +bodily structure and in mental faculties; and if so, +whether the variations are transmitted to his offspring +in accordance with the laws which prevail with the lower +animals; such as that of the transmission of characters +to the same age or sex. Again, are the variations the result, +as far as our ignorance permits us to judge, of the +same general causes, and are they governed by the same +general laws, as in the case of other organisms; for instance +by correlation, the inherited effects of use and +disuse, &c.? Is man subject to similar malconformations, +the result of arrested development, of reduplication of +parts, &c., and does he display in any of his anomalies +reversion to some former and ancient type of structure? +It might also naturally be enquired whether man, like +so many other animals, has given rise to varieties and +sub-races, differing but slightly from each other, or to<span class='pagenum'><a name="Page_10" id="Page_10">10</a></span> +races differing so much that they must be classed as +doubtful species? How are such races distributed over +the world; and how, when crossed, do they react on +each other, both in the first and succeeding generations? +And so with many other points.</p> + +<p class="tb">The enquirer would next come to the important point, +whether man tends to increase at so rapid a rate, as to +lead to occasional severe struggles for existence, and +consequently to beneficial variations, whether in body +or mind, being preserved, and injurious ones eliminated. +Do the races or species of men, whichever term may be +applied, encroach on and replace each other, so that +some finally become extinct? We shall see that all +these questions, as indeed is obvious in respect to most +of them, must be answered in the affirmative, in the +same manner as with the lower animals. But the +several considerations just referred to may be conveniently +deferred for a time; and we will first see how +far the bodily structure of man shows traces, more or +less plain, of his descent from some lower form. In the +two succeeding chapters the mental powers of man, in +comparison with those of the lower animals, will be considered.</p> + +<p><i>The Bodily Structure of Man.</i>—It is notorious that +man is constructed on the same general type or model +with other mammals. All the bones in his skeleton +can be compared with corresponding bones in a monkey, +bat, or seal. So it is with his muscles, nerves, blood-vessels +and internal viscera. The brain, the most important +of all the organs, follows the same law, as shewn +by Huxley and other anatomists. Bischoff,<a name="FNanchor_3" id="FNanchor_3"></a><a href="#Footnote_3" class="fnanchor">3</a> who is a +hostile witness, admits that every chief fissure and fold +<span class='pagenum'><a name="Page_11" id="Page_11">11</a></span>in the brain of man has its analogy in that of the orang; +but he adds that at no period of development do their +brains perfectly agree; nor could this be expected, for +otherwise their mental powers would have been the same. +Vulpian<a name="FNanchor_4" id="FNanchor_4"></a><a href="#Footnote_4" class="fnanchor">4</a> remarks: “Les différences réelles qui existent +entre l’encéphale de l’homme et celui des singes supérieurs, +sont bien minimes. Il ne faut pas se faire +d’illusions à cet égard. L’homme est bien plus près +des singes anthropomorphes par les caractères anatomiques +de son cerveau que ceux-ci ne le sont non-seulement +des autres mammifères, mais mêmes de +certains quadrumanes, des guenons et des macaques.” +But it would be superfluous here to give further details +on the correspondence between man and the higher +mammals in the structure of the brain and all other +parts of the body.</p> + +<p>It may, however, be worth while to specify a few +points, not directly or obviously connected with structure, +by which this correspondence or relationship is +well shewn.</p> + +<p>Man is liable to receive from the lower animals, and +to communicate to them, certain diseases as hydrophobia, +variola, the glanders, &c.; and this fact proves +the close similarity of their tissues and blood, both in +minute structure and composition, far more plainly than +does their comparison under the best microscope, or by +the aid of the best chemical analysis. Monkeys are +liable to many of the same non-contagious diseases as we +are; thus Rengger,<a name="FNanchor_5" id="FNanchor_5"></a><a href="#Footnote_5" class="fnanchor">5</a> who carefully observed for a long +time the <i>Cebus Azaræ</i> in its native land, found it liable +to catarrh, with the usual symptoms, and which when +<span class='pagenum'><a name="Page_12" id="Page_12">12</a></span>often recurrent led to consumption. These monkeys +suffered also from apoplexy, inflammation of the bowels, +and cataract in the eye. The younger ones when shedding +their milk-teeth often died from fever. Medicines +produced the same effect on them as on us. Many +kinds of monkeys have a strong taste for tea, coffee, and +spirituous liquors: they will also, as I have myself seen, +smoke tobacco with pleasure. Brehm asserts that the +natives of north-eastern Africa catch the wild baboons +by exposing vessels with strong beer, by which they are +made drunk. He has seen some of these animals, which +he kept in confinement, in this state; and he gives +a laughable account of their behaviour and strange +grimaces. On the following morning they were very +cross and dismal; they held their aching heads with +both hands and wore a most pitiable expression: when +beer or wine was offered them, they turned away with +disgust, but relished the juice of lemons.<a name="FNanchor_6" id="FNanchor_6"></a><a href="#Footnote_6" class="fnanchor">6</a> An American +monkey, an Ateles, after getting drunk on brandy, would +never touch it again, and thus was wiser than many +men. These trifling facts prove how similar the nerves +of taste must be in monkeys and man, and how similarly +their whole nervous system is affected.</p> + +<p>Man is infested with internal parasites, sometimes +causing fatal effects, and is plagued by external parasites, +all of which belong to the same genera or families +with those infesting other mammals. Man is subject like +other mammals, birds, and even insects, to that mysterious +law, which causes certain normal processes, such +as gestation, as well as the maturation and duration of +various diseases, to follow lunar periods.<a name="FNanchor_7" id="FNanchor_7"></a><a href="#Footnote_7" class="fnanchor">7</a> His wounds +<span class='pagenum'><a name="Page_13" id="Page_13">13</a></span>are repaired by the same process of healing; and the +stumps left after the amputation of his limbs occasionally +possess, especially during an early embryonic +period, some power of regeneration, as in the lowest +animals.<a name="FNanchor_8" id="FNanchor_8"></a><a href="#Footnote_8" class="fnanchor">8</a></p> + +<p class="tb">The whole process of that most important function, +the reproduction of the species, is strikingly the same +in all mammals, from the first act of courtship by the +male<a name="FNanchor_9" id="FNanchor_9"></a><a href="#Footnote_9" class="fnanchor">9</a> to the birth and nurturing of the young. Monkeys +are born in almost as helpless a condition as our +own infants; and in certain genera the young differ +fully as much in appearance from the adults, as do our +children from their full-grown parents.<a name="FNanchor_10" id="FNanchor_10"></a><a href="#Footnote_10" class="fnanchor">10</a> It has been +urged by some writers as an important distinction, that +with man the young arrive at maturity at a much later +age than with any other animal; but if we look to the +races of mankind which inhabit tropical countries the +difference is not great, for the orang is believed not to +be adult till the age of from ten to fifteen years.<a name="FNanchor_11" id="FNanchor_11"></a><a href="#Footnote_11" class="fnanchor">11</a> Man +<span class='pagenum'><a name="Page_14" id="Page_14">14</a></span>differs from woman in size, bodily strength, hairyness, +&c., as well as in mind, in the same manner as do the +two sexes of many mammals. It is, in short, scarcely +possible to exaggerate the close correspondence in general +structure, in the minute structure of the tissues, in +chemical composition and in constitution, between man +and the higher animals, especially the anthropomorphous +apes.</p> + +<p><i>Embryonic Development.</i>—Man is developed from an +ovule, about the 125th of an inch in diameter, which +differs in no respect from the ovules of other animals. +The embryo itself at a very early period can hardly be +distinguished from that of other members of the vertebrate +kingdom. At this period the arteries run in arch-like +branches, as if to carry the blood to branchiæ +which are not present in the higher vertebrata, though +the slits on the sides of the neck still remain (<i>f</i>, <i>g</i>, fig. 1), +marking their former position. At a somewhat later +period, when the extremities are developed, “the feet of +lizards and mammals,” as the illustrious Von Baer +remarks, “the wings and feet of birds, no less than the +hands and feet of man, all arise from the same fundamental +form.” It is, says Prof. Huxley,<a name="FNanchor_12" id="FNanchor_12"></a><a href="#Footnote_12" class="fnanchor">12</a> “quite in +the later stages of development that the young human +being presents marked differences from the young +ape, while the latter departs as much from the dog +in its developments, as the man does. Startling as +this last assertion may appear to be, it is demonstrably +true.”</p> + +<p>As some of my readers may never have seen a drawing +of an embryo, I have given one of man and another +of a dog, at about the same early stage of development, +<span class='pagenum'><a name="Page_15" id="Page_15">15</a></span>carefully copied from two works of undoubted accuracy.<a name="FNanchor_13" id="FNanchor_13"></a><a href="#Footnote_13" class="fnanchor">13</a></p> + +<div class="figcenter" style="width: 550px;"><a name="f1" id="f1"></a><img src="images/fig1.png" width="550" height="972" +alt="Upper figure human embryo, from Ecker. Lower figure that of a dog." title="" /> + +<p class="center2">Fig. 1. Upper figure human embryo, from Ecker. Lower figure that of a dog, +from Bischoff.</p></div> + +<table width="100%" summary="embryos" border="0"> +<tr> +<td class="left50"><i>a.</i> Fore-brain, cerebral hemispheres, &c.</td> +<td class="left25" colspan="2"><i>g.</i> Second visceral arch.</td> +</tr> +<tr> +<td class="left50"><i>b.</i> Mid-brain, corpora quadrigemina.</td> +<td class="left25" colspan="2">H. Vertebral columns and muscles in process of development.</td> +</tr> +<tr> +<td class="left50"><i>c.</i> Hind-brain, cerebellum, medulla oblongata.</td> +<td class="left25"><i>i.</i> Anterior</td> +<td class="left25m" rowspan="2">┐<br />│ extremities<br />┘</td> +</tr> +<tr> +<td class="left50"><i>d.</i> Eye.</td> +<td class="left25">K. Posterior</td> +</tr> +<tr> +<td class="left50"><i>e.</i> Ear.</td> +<td class="left25" colspan="2">L. Tail or os coccyx.</td> +</tr> +<tr> +<td class="left50"><i>f.</i> First visceral arch.</td> +<td class="left25" colspan="2"> </td> +</tr> +</table> + +<p><span class='pagenum'><a name="Page_16" id="Page_16">16</a></span></p> + +<p class="tb">After the foregoing statements made by such high +authorities, it would be superfluous on my part to give +a number of borrowed details, shewing that the embryo +of man closely resembles that of other mammals. It +may, however, be added that the human embryo likewise +resembles in various points of structure certain +low forms when adult. For instance, the heart +at first exists as a simple pulsating vessel; the excreta +are voided through a cloacal passage; and the os +coccyx projects like a true tail, “extending considerably +beyond the rudimentary legs.”<a name="FNanchor_14" id="FNanchor_14"></a><a href="#Footnote_14" class="fnanchor">14</a> In the embryos of +all air-breathing vertebrates, certain glands called the corpora +Wolffiana, correspond with and act like the kidneys +of mature fishes.<a name="FNanchor_15" id="FNanchor_15"></a><a href="#Footnote_15" class="fnanchor">15</a> Even at a later embryonic period, +some striking resemblances between man and the lower +animals may be observed. Bischoff says that the convolutions +of the brain in a human fœtus at the end of +the seventh month reach about the same stage of development +as in a baboon when adult.<a name="FNanchor_16" id="FNanchor_16"></a><a href="#Footnote_16" class="fnanchor">16</a> The great toe, as +Prof. Owen remarks,<a name="FNanchor_17" id="FNanchor_17"></a><a href="#Footnote_17" class="fnanchor">17</a> “which forms the fulcrum when +standing or walking, is perhaps the most characteristic +<span class='pagenum'><a name="Page_17" id="Page_17">17</a></span>peculiarity in the human structure;” but in an embryo, +about an inch in length, Prof. Wyman<a name="FNanchor_18" id="FNanchor_18"></a><a href="#Footnote_18" class="fnanchor">18</a> found +“that the great toe was shorter than the others, and, +instead of being parallel to them, projected at an +angle from the side of the foot, thus corresponding +with the permanent condition of this part in the +quadrumana.” I will conclude with a quotation from +Huxley,<a name="FNanchor_19" id="FNanchor_19"></a><a href="#Footnote_19" class="fnanchor">19</a> who after asking, does man originate in a +different way from a dog, bird, frog or fish? says, “the +reply is not doubtful for a moment; without question, +the mode of origin and the early stages of the development +of man are identical with those of the animals +immediately below him in the scale: without a doubt +in these respects, he is far nearer to apes, than the apes +are to the dog.”</p> + +<p><i>Rudiments.</i>—This subject, though not intrinsically +more important than the two last, will for several reasons +be here treated with more fullness.<a name="FNanchor_20" id="FNanchor_20"></a><a href="#Footnote_20" class="fnanchor">20</a> Not one of +the higher animals can be named which does not bear +some part in a rudimentary condition; and man forms +no exception to the rule. Rudimentary organs must +be distinguished from those that are nascent; though +in some cases the distinction is not easy. The former +are either absolutely useless, such as the mammæ of +male quadrupeds, or the incisor teeth of ruminants +which never cut through the gums; or they are of such +slight service to their present possessors, that we cannot +suppose that they were developed under the conditions +<span class='pagenum'><a name="Page_18" id="Page_18">18</a></span>which now exist. Organs in this latter state are not +strictly rudimentary, but they are tending in this direction. +Nascent organs, on the other hand, though not +fully developed, are of high service to their possessors, +and are capable of further development. Rudimentary +organs are eminently variable; and this is partly intelligible, +as they are useless or nearly useless, and +consequently are no longer subjected to natural selection. +They often become wholly suppressed. When +this occurs, they are nevertheless liable to occasional +reappearance through reversion; and this is a circumstance +well worthy of attention.</p> + +<p>Disuse at that period of life, when an organ is chiefly +used, and this is generally during maturity, together +with inheritance at a corresponding period of life, seem +to have been the chief agents in causing organs to become +rudimentary. The term “disuse” does not relate +merely to the lessened action of muscles, but includes +a diminished flow of blood to a part or organ, from +being subjected to fewer alternations of pressure, or +from becoming in any way less habitually active. Rudiments, +however, may occur in one sex of parts normally +present in the other sex; and such rudiments, as we +shall hereafter see, have often originated in a distinct +manner. In some cases organs have been reduced by +means of natural selection, from having become injurious +to the species under changed habits of life. The +process of reduction is probably often aided through the +two principles of compensation and economy of growth; +but the later stages of reduction, after disuse has done +all that can fairly be attributed to it, and when the saving +to be effected by the economy of growth would be very +small,<a name="FNanchor_21" id="FNanchor_21"></a><a href="#Footnote_21" class="fnanchor">21</a> are difficult to understand. The final and com<span class='pagenum'><a name="Page_19" id="Page_19">19</a></span>plete +suppression of a part, already useless and much +reduced in size, in which case neither compensation nor +economy can come into play, is perhaps intelligible by +the aid of the hypothesis of pangenesis, and apparently in +no other way. But as the whole subject of rudimentary +organs has been fully discussed and illustrated in my +former works,<a name="FNanchor_22" id="FNanchor_22"></a><a href="#Footnote_22" class="fnanchor">22</a> I need here say no more on this head.</p> + +<p>Rudiments of various muscles have been observed in +many parts of the human body;<a name="FNanchor_23" id="FNanchor_23"></a><a href="#Footnote_23" class="fnanchor">23</a> and not a few muscles, +which are regularly present in some of the lower animals +can occasionally be detected in man in a greatly +reduced condition. Every one must have noticed the +power which many animals, especially horses, possess +of moving or twitching their skin; and this is effected +by the panniculus carnosus. Remnants of this muscle +in an efficient state are found in various parts of our +bodies; for instance, on the forehead, by which the +eyebrows are raised. The <i>platysma myoides</i>, which is +well developed on the neck, belongs to this system, but +cannot be voluntarily brought into action. Prof. +Turner, of Edinburgh, has occasionally detected, as he +informs me, muscular fasciculi in five different situations, +namely in the axillæ, near the scapulæ, &c., all of +which must be referred to the system of the panniculus. +He has also shewn<a name="FNanchor_24" id="FNanchor_24"></a><a href="#Footnote_24" class="fnanchor">24</a> that the <i>musculus sternalis</i> or <i>sternalis +brutorum</i>, which is not an extension of the <i>rectus +abdominalis</i>, but is closely allied to the panniculus, oc<span class='pagenum'><a name="Page_20" id="Page_20">20</a></span>curred +in the proportion of about 3 per cent. in upwards +of 600 bodies: he adds, that this muscle affords “an +excellent illustration of the statement that occasional +and rudimentary structures are especially liable to +variation in arrangement.”</p> + +<p>Some few persons have the power of contracting the +superficial muscles on their scalps; and these muscles +are in a variable and partially rudimentary condition. +M. A. de Candolle has communicated to me a curious +instance of the long-continued persistence or inheritance +of this power, as well as of its unusual development. +He knows a family, in which one member, the present +head of a family, could, when a youth, pitch several +heavy books from his head by the movement of the +scalp alone; and he won wagers by performing this feat. +His father, uncle, grandfather, and all his three children +possess the same power to the same unusual degree. +This family became divided eight generations ago into +two branches; so that the head of the above-mentioned +branch is cousin in the seventh degree to the head of +the other branch. This distant cousin resides in another +part of France, and on being asked whether he possessed +the same faculty, immediately exhibited his power. +This case offers a good illustration how persistently an +absolutely useless faculty may be transmitted.</p> + +<p>The extrinsic muscles which serve to move the whole +external ear, and the intrinsic muscles which move the +different parts, all of which belong to the system of the +panniculus, are in a rudimentary condition in man; they +are also variable in development, or at least in function. +I have seen one man who could draw his ears forwards, +and another who could draw them backwards;<a name="FNanchor_25" id="FNanchor_25"></a><a href="#Footnote_25" class="fnanchor">25</a></p> +<p><span class='pagenum'><a name="Page_21" id="Page_21">21</a></span> +and from what one of these persons told me, it is probable +that most of us by often touching our ears and +thus directing our attention towards them, could by +repeated trials recover some power of movement. The +faculty of erecting the ears and of directing them to +different points of the compass, is no doubt of the +highest service to many animals, as they thus perceive +the point of danger; but I have never heard of a man +who possessed the least power of erecting his ears,—the +one movement which might be of use to him. The +whole external shell of the ear may be considered a +rudiment, together with the various folds and prominences +(helix and anti-helix, tragus and anti-tragus, &c.) +which in the lower animals strengthen and support the +ear when erect, without adding much to its weight. +Some authors, however, suppose that the cartilage of the +shell serves to transmit vibrations to the acoustic +nerve; but Mr. Toynbee,<a name="FNanchor_26" id="FNanchor_26"></a><a href="#Footnote_26" class="fnanchor">26</a> after collecting all the +known evidence on this head, concludes that the external +shell is of no distinct use. The ears of the chimpanzee +and orang are curiously like those of man, and I +am assured by the keepers in the Zoological Gardens +that these animals never move or erect them; so that +they are in an equally rudimentary condition, as far as +function is concerned, as in man. Why these animals, +as well as the progenitors of man, should have lost the +power of erecting their ears we cannot say. It may be, +though I am not quite satisfied with this view, that owing +to their arboreal habits and great strength they were +but little exposed to danger, and so during a lengthened +period moved their ears but little, and thus gradually +lost the power of moving them. This would be a +parallel case with that of those large and heavy birds, +<span class='pagenum'><a name="Page_22" id="Page_22">22</a></span>which from inhabiting oceanic islands have not been +exposed to the attacks of beasts of prey, and have consequently +lost the power of using their wings for flight.</p> + +<div class="figleft" style="width: 200px;"><a name="f2" id="f2"></a><img src="images/fig2.png" width="200" height="243" +alt="Human Ear, modelled +and drawn by Mr. Woolner." title="" /> + +<p class="indent2">Fig. 2. Human Ear, modelled +and drawn by Mr. Woolner.<br /> + <i>a.</i> The projecting point.</p></div> + +<p>The celebrated sculptor, Mr. Woolner, informs me of +one little peculiarity in the external ear, which he has +often observed both in men and women, and of which +he perceived the full signification. His attention was +first called to the subject whilst at work on his figure +of Puck, to which he had given pointed ears. He was +thus led to examine the ears of various monkeys, and +subsequently more carefully those of man. The peculiarity +consists in a little blunt point, projecting from +the inwardly folded margin, or helix. Mr. Woolner +made an exact model of one such case, and has sent +me the accompanying drawing. +(Fig. 2.) These points not only +project inwards, but often a little +outwards, so that they are visible +when the head is viewed from directly +in front or behind. They +are variable in size and somewhat +in position, standing either +a little higher or lower; and they +sometimes occur on one ear and +not on the other. Now the meaning +of these projections is not, +I think, doubtful; but it may +be thought that they offer too +trifling a character to be worth notice. This thought, +however, is as false as it is natural. Every character, +however slight, must be the result of some definite +cause; and if it occurs in many individuals deserves +consideration. The helix obviously consists of the extreme +margin of the ear folded inwards; and this folding +appears to be in some manner connected with the<span class='pagenum'><a name="Page_23" id="Page_23">23</a></span> +whole external ear being permanently pressed backwards. +In many monkeys, which do not stand high in +the order, as baboons and some species of macacus,<a name="FNanchor_27" id="FNanchor_27"></a><a href="#Footnote_27" class="fnanchor">27</a> the +upper portion of the ear is slightly pointed, and the +margin is not at all folded inwards; but if the margin +were to be thus folded, a slight point would necessarily +project inwards and probably a little outwards. This +could actually be observed in a specimen of the <i>Ateles +beelzebuth</i> in the Zoological Gardens; and we may safely +conclude that it is a similar structure—a vestige of +formerly pointed ears—which occasionally reappears in +man.</p> + +<p>The nictitating membrane, or third eyelid, with its +accessory muscles and other structures, is especially +well developed in birds, and is of much functional importance +to them, as it can be rapidly drawn across the +whole eye-ball. It is found in some reptiles and amphibians, +and in certain fishes, as in sharks. It is fairly +well developed in the two lower divisions of the mammalian +series, namely, in the monotremata and marsupials, +and in some few of the higher mammals, as in the +walrus. But in man, the quadrumana, and most other +mammals, it exists, as is admitted by all anatomists, as +a mere rudiment, called the semilunar fold.<a name="FNanchor_28" id="FNanchor_28"></a><a href="#Footnote_28" class="fnanchor">28</a></p> + +<p>The sense of smell is of the highest importance to +the greater number of mammals—to some, as the ruminants, +in warning them of danger; to others, as the +<span class='pagenum'><a name="Page_24" id="Page_24">24</a></span>carnivora, in finding their prey; to others, as the wild +boar, for both purposes combined. But the sense of +smell is of extremely slight service, if any, even to +savages, in whom it is generally more highly developed +than in the civilised races. It does not warn them of +danger, nor guide them to their food; nor does it prevent +the Esquimaux from sleeping in the most fetid +atmosphere, nor many savages from eating half-putrid +meat. Those who believe in the principle of gradual +evolution, will not readily admit that this sense in its +present state was originally acquired by man, as he +now exists. No doubt he inherits the power in <i>an</i> +enfeebled and so far rudimentary condition, from some +early progenitor, to whom it was highly serviceable +and by whom it was continually used. We can thus +perhaps understand how it is, as Dr. Maudsley has truly +remarked,<a name="FNanchor_29" id="FNanchor_29"></a><a href="#Footnote_29" class="fnanchor">29</a> that the sense of smell in man “is singularly +effective in recalling vividly the ideas and images +of forgotten scenes and places;” for we see in those +animals, which have this sense highly developed, such as +dogs and horses, that old recollections of persons and +places are strongly associated with their odour.</p> + +<p>Man differs conspicuously from all the other Primates +in being almost naked. But a few short straggling +hairs are found over the greater part of the body in +the male sex, and fine down on that of the female sex, +In individuals belonging to the same race these hairs +are highly variable, not only in abundance, but likewise +in position: thus the shoulders in some Europeans +are quite naked, whilst in others they bear thick tufts +of hair.<a name="FNanchor_30" id="FNanchor_30"></a><a href="#Footnote_30" class="fnanchor">30</a> There can be little doubt that the hairs +<span class='pagenum'><a name="Page_25" id="Page_25">25</a></span>thus scattered over the body are the rudiments of the +uniform hairy coat of the lower animals. This view +is rendered all the more probable, as it is known that +fine, short, and pale-coloured hairs on the limbs and +other parts of the body occasionally become developed +into “thickset, long, and rather coarse dark hairs,” +when abnormally nourished near old-standing inflamed +surfaces.<a name="FNanchor_31" id="FNanchor_31"></a><a href="#Footnote_31" class="fnanchor">31</a></p> + +<p>I am informed by Mr. Paget that persons belonging +to the same family often have a few hairs in their eyebrows +much longer than the others; so that this slight +peculiarity seems to be inherited. These hairs apparently +represent the vibrissæ, which are used as organs +of touch by many of the lower animals. In a young +chimpanzee I observed that a few upright, rather long, +hairs, projected above the eyes, where the true eyebrows, +if present, would have stood.</p> + +<p>The fine wool-like hair, or so-called lanugo, with +which the human fœtus during the sixth month is +thickly covered, offers a more curious case. It is first +developed, during the fifth month, on the eyebrows and +face, and especially round the mouth, where it is much +longer than that on the head. A moustache of this kind +was observed by Eschricht<a name="FNanchor_32" id="FNanchor_32"></a><a href="#Footnote_32" class="fnanchor">32</a> on a female fœtus; but this +is not so surprising a circumstance as it may at first appear, +for the two sexes generally resemble each other in +all external characters during an early period of growth. +The direction and arrangement of the hairs on all parts +of the fœtal body are the same as in the adult, but are +subject to much variability. The whole surface, including +even the forehead and ears, is thus thickly clothed; but +it is a significant fact that the palms of the hands and +<span class='pagenum'><a name="Page_26" id="Page_26">26</a></span>the soles of the feet are quite naked, like the inferior +surfaces of all four extremities in most of the lower +animals. As this can hardly be an accidental coincidence, +we must consider the woolly covering of the +fœtus to be the rudimental representative of the first +permanent coat of hair in those mammals which are +born hairy. This representation is much more complete, +in accordance with the usual law of embryological +development, than that afforded by the straggling hairs +on the body of the adult.</p> + +<p>It appears as if the posterior molar or wisdom-teeth +were tending to become rudimentary in the more civilised +races of man. These teeth are rather smaller +than the other molars, as is likewise the case with the +corresponding teeth in the chimpanzee and orang; and +they have only two separate fangs. They do not cut +through the gums till about the seventeenth year, and +I am assured by dentists that they are much more +liable to decay, and are earlier lost, than the other +teeth. It is also remarkable that they are much more +liable to vary both in structure and in the period of +their development than the other teeth.<a name="FNanchor_33" id="FNanchor_33"></a><a href="#Footnote_33" class="fnanchor">33</a> In the +Melanian races, on the other hand, the wisdom-teeth +are usually furnished with three separate fangs, and +are generally sound: they also differ from the other +molars in size less than in the Caucasian races.<a name="FNanchor_34" id="FNanchor_34"></a><a href="#Footnote_34" class="fnanchor">34</a> Prof. +Schaaffhausen accounts for this difference between the +races by “the posterior dental portion of the jaw being +always shortened” in those that are civilised,<a name="FNanchor_35" id="FNanchor_35"></a><a href="#Footnote_35" class="fnanchor">35</a> and this +shortening may, I presume, be safely attributed to civi<span class='pagenum'><a name="Page_27" id="Page_27">27</a></span>lised +men habitually feeding on soft, cooked food, and +thus using their jaws less. I am informed by Mr. Brace +that it is becoming quite a common practice in the United +States to remove some of the molar teeth of children, +as the jaw does not grow large enough for the perfect +development of the normal number.</p> + +<p>With respect to the alimentary canal I have met +with an account of only a single rudiment, namely the +vermiform appendage of the cæcum. The cæcum is +a branch or diverticulum of the intestine, ending in a +cul-de-sac, and it is extremely long in many of the +lower vegetable-feeding mammals. In the marsupial +koala it is actually more than thrice as long as the +whole body.<a name="FNanchor_36" id="FNanchor_36"></a><a href="#Footnote_36" class="fnanchor">36</a> It is sometimes produced into a long +gradually-tapering point, and is sometimes constricted +in parts. It appears as if, in consequence of changed +diet or habits, the cæcum had become much shortened +in various animals, the vermiform appendage being left +as a rudiment of the shortened part. That this appendage +is a rudiment, we may infer from its small +size, and from the evidence which Prof. Canestrini<a name="FNanchor_37" id="FNanchor_37"></a><a href="#Footnote_37" class="fnanchor">37</a> has +collected of its variability in man. It is occasionally +quite absent, or again is largely developed. The passage +is sometimes completely closed for half or two-thirds of +its length, with the terminal part consisting of a flattened +solid expansion. In the orang this appendage is +long and convoluted: in man it arises from the end of +the short cæcum, and is commonly from four to five +inches in length, being only about the third of an inch +in diameter. Not only is it useless, but it is sometimes +the cause of death, of which fact I have lately +heard two instances: this is due to small hard bodies, +<span class='pagenum'><a name="Page_28" id="Page_28">28</a></span>such as seeds, entering the passage and causing inflammation.<a name="FNanchor_38" id="FNanchor_38"></a><a href="#Footnote_38" class="fnanchor">38</a></p> + +<p>In the Quadrumana and some other orders of mammals, +especially in the Carnivora, there is a passage near +the lower end of the humerus, called the supra-condyloid +foramen, through which the great nerve of the fore limb +passes, and often the great artery. Now in the humerus +of man, as Dr. Struthers<a name="FNanchor_39" id="FNanchor_39"></a><a href="#Footnote_39" class="fnanchor">39</a> and others have shewn, there +is generally a trace of this passage, and it is sometimes +fairly well developed, being formed by a depending +hook-like process of bone, completed by a band of +ligament. When present the great nerve invariably +passes through it, and this clearly indicates that it is the +homologue and rudiment of the supra-condyloid foramen +of the lower animals. Prof. Turner estimates, as +he informs me, that it occurs in about one per cent. of +recent skeletons; but during ancient times it appears +to have been much more common. Mr. Busk<a name="FNanchor_40" id="FNanchor_40"></a><a href="#Footnote_40" class="fnanchor">40</a> has +collected the following evidence on this head: Prof. +Broca “noticed the perforation in four and a half per +cent. of the arm-bones collected in the ‘Cimetière du +Sud’ at Paris; and in the Grotto of Orrony, the contents +of which are referred to the Bronze period, as +many as eight humeri out of thirty-two were perforated; +but this extraordinary proportion, he thinks, +might be due to the cavern having been a sort of +<span class='pagenum'><a name="Page_29" id="Page_29">29</a></span> ‘family vault.’ Again, M. Dupont found 30 per cent. +of perforated bones in the caves of the Valley of the +Lesse, belonging to the Reindeer period; whilst M. +Leguay, in a sort of <i>dolmen</i> at Argenteuil, observed +twenty-five per cent. to be perforated; and M. Pruner-Bey +found twenty-six per cent. in the same condition +in bones from Vauréal. Nor should it be left unnoticed +that M. Pruner-Bey states that this condition is +common in Guanche skeletons.” The fact that ancient +races, in this and several other cases, more frequently +present structures which resemble those of the lower +animals than do the modern races, is interesting. One +chief cause seems to be that ancient races stand somewhat +nearer than modern races in the long line of +descent to their remote animal-like progenitors.</p> + +<p>The os coccyx in man, though functionless as a tail, +plainly represents this part in other vertebrate animals. +At an early embryonic period it is free, and, as we have +seen, projects beyond the lower extremities. In certain +rare and anomalous cases it has been known, according +to Isidore Geoffroy St.-Hilaire and others,<a name="FNanchor_41" id="FNanchor_41"></a><a href="#Footnote_41" class="fnanchor">41</a> to form a +small external rudiment of a tail. The os coccyx is +short, usually including only four vertebræ: and these +are in a rudimental condition, for they consist, with the +exception of the basal one, of the centrum alone.<a name="FNanchor_42" id="FNanchor_42"></a><a href="#Footnote_42" class="fnanchor">42</a> They +are furnished with some small muscles; one of which, as +I am informed by Prof. Turner, has been expressly +described by Theile as a rudimentary repetition of the +extensor of the tail, which is so largely developed in +many mammals.</p> + +<p>The spinal cord in man extends only as far downwards +as the last dorsal or first lumbar vertebra; but a +<span class='pagenum'><a name="Page_30" id="Page_30">30</a></span>thread-like structure (the <i>filum terminale</i>) runs down the +axis of the sacral part of the spinal canal, and even +along the back of the coccygeal bones. The upper +part of this filament, as Prof. Turner informs me, is +undoubtedly homologous with the spinal cord; but the +lower part apparently consists merely of the <i>pia mater</i>, +or vascular investing membrane. Even in this case the +os coccyx may be said to possess a vestige of so important +a structure as the spinal cord, though no longer +enclosed within a bony canal. The following fact, for +which I am also indebted to Prof. Turner, shews how +closely the os coccyx corresponds with the true tail in +the lower animals: Luschka has recently discovered at +the extremity of the coccygeal bones a very peculiar +convoluted body, which is continuous with the middle +sacral artery; and this discovery led Krause and Meyer +to examine the tail of a monkey (Macacus) and of a cat, +in both of which they found, though not at the extremity, +a similarly convoluted body.</p> + +<p class="tb">The reproductive system offers various rudimentary +structures; but these differ in one important respect +from the foregoing cases. We are not here concerned +with a vestige of a part which does not belong to the +species in an efficient state; but with a part which +is always present and efficient in the one sex, being +represented in the other by a mere rudiment. Nevertheless, +the occurrence of such rudiments is as difficult +to explain on the belief of the separate creation +of each species, as in the foregoing cases. Hereafter +I shall have to recur to these rudiments, and shall +shew that their presence generally depends merely on +inheritance; namely, on parts acquired by one sex +having been partially transmitted to the other. Here +I will only give some instances of such rudiments. It +is well known that in the males of all mammals, in<span class='pagenum'><a name="Page_31" id="Page_31">31</a></span>cluding +man, rudimentary mammæ exist. These in +several instances have become well developed, and have +yielded a copious supply of milk. Their essential identity +in the two sexes is likewise shewn by their occasional +sympathetic enlargement in both during an +attack of the measles. The <i>vesicula prostratica</i>, which +has been observed in many male mammals, is now universally +acknowledged to be the homologue of the +female uterus, together with the connected passage. It +is impossible to read Leuckart’s able description of this +organ, and his reasoning, without admitting the justness +of his conclusion. This is especially clear in the case of +those mammals in which the true female uterus bifurcates, +for in the males of these the vesicula likewise +bifurcates.<a name="FNanchor_43" id="FNanchor_43"></a><a href="#Footnote_43" class="fnanchor">43</a> Some additional rudimentary structures +belonging to the reproductive system might here have +been adduced.<a name="FNanchor_44" id="FNanchor_44"></a><a href="#Footnote_44" class="fnanchor">44</a></p> + +<p>The bearing of the three great classes of facts now +given is unmistakeable. But it would be superfluous here +fully to recapitulate the line of argument given in detail +in my ‘Origin of Species.’ The homological construction +of the whole frame in the members of the same class is +intelligible, if we admit their descent from a common +progenitor, together with their subsequent adaptation +to diversified conditions. On any other view the similarity +of pattern between the hand of a man or monkey, +the foot of a horse, the flipper of a seal, the wing of +a bat, &c., is utterly inexplicable. It is no scientific +<span class='pagenum'><a name="Page_32" id="Page_32">32</a></span>explanation to assert that they have all been formed on +the same ideal plan. With respect to development, +we can clearly understand, on the principle of variations +supervening at a rather late embryonic period, +and being inherited at a corresponding period, how it is +that the embryos of wonderfully different forms should +still retain, more or less perfectly, the structure of their +common progenitor. No other explanation has ever +been given of the marvellous fact that the embryo of a +man, dog, seal, bat, reptile, &c., can at first hardly be +distinguished from each other. In order to understand +the existence of rudimentary organs, we have only to +suppose that a former progenitor possessed the parts in +question in a perfect state, and that under changed +habits of life they became greatly reduced, either from +simple disuse, or through the natural selection of those +individuals which were least encumbered with a superfluous +part, aided by the other means previously indicated.</p> + +<p>Thus we can understand how it has come to pass +that man and all other vertebrate animals have been +constructed on the same general model, why they pass +through the same early stages of development, and why +they retain certain rudiments in common. Consequently +we ought frankly to admit their community of descent: +to take any other view, is to admit that our own structure +and that of all the animals around us, is a mere +snare laid to entrap our judgment. This conclusion +is greatly strengthened, if we look to the members of +the whole animal series, and consider the evidence derived +from their affinities or classification, their geographical +distribution and geological succession. It +is only our natural prejudice, and that arrogance +which made our forefathers declare that they were +descended from demi-gods, which leads us to demur to<span class='pagenum'><a name="Page_33" id="Page_33">33</a></span> +this conclusion. But the time will before long come +when it will be thought wonderful, that naturalists, who +were well acquainted with the comparative structure +and development of man and other mammals, should +have believed that each was the work of a separate act +of creation.</p> + +<hr /> +<p><span class='pagenum'><a name="Page_34" id="Page_34">34</a></span></p> +<h3>CHAPTER II.</h3> + +<h4><span class="smcap">Comparison of the Mental Powers of Man and the +Lower Animals.</span></h4> + +<div class="blockquot"><p>The difference in mental power between the highest ape and the +lowest savage, immense—Certain instincts in common—The +emotions—Curiosity—Imitation—Attention—Memory—Imagination—Reason—Progressive +improvement—Tools and +weapons used by animals—Language—Self-consciousness—Sense +of beauty—Belief in God, spiritual agencies, superstitions.</p></div> + +<p>We have seen in the last chapter that man bears in his +bodily structure clear traces of his descent from some +lower form; but it may be urged that, as man differs so +greatly in his mental power from all other animals, there +must be some error in this conclusion. No doubt the +difference in this respect is enormous, even if we compare +the mind of one of the lowest savages, who has no +words to express any number higher than four, and who +uses no abstract terms for the commonest objects or +affections,<a name="FNanchor_45" id="FNanchor_45"></a><a href="#Footnote_45" class="fnanchor">45</a> with that of the most highly organised ape. +The difference would, no doubt, still remain immense, +even if one of the higher apes had been improved or +civilised as much as a dog has been in comparison with +its parent-form, the wolf or jackal. The Fuegians rank +amongst the lowest barbarians; but I was continually +struck with surprise how closely the three natives on +board H.M.S. “Beagle,” who had lived some years in +England and could talk a little English, resembled us +in disposition and in most of our mental faculties. If no +<span class='pagenum'><a name="Page_35" id="Page_35">35</a></span>organic being excepting man had possessed any mental +power, or if his powers had been of a wholly different +nature from those of the lower animals, then we should +never have been able to convince ourselves that our +high faculties had been gradually developed. But it +can be clearly shewn that there is no fundamental +difference of this kind. We must also admit that +there is a much wider interval in mental power between +one of the lowest fishes, as a lamprey or lancelet, +and one of the higher apes, than between an ape and +man; yet this immense interval is filled up by numberless +gradations.</p> + +<p>Nor is the difference slight in moral disposition +between a barbarian, such as the man described by the +old navigator Byron, who dashed his child on the rocks +for dropping a basket of sea-urchins, and a Howard or +Clarkson; and in intellect, between a savage who does +not use any abstract terms, and a Newton or Shakspeare. +Differences of this kind between the highest men of the +highest races and the lowest savages, are connected by +the finest gradations. Therefore it is possible that they +might pass and be developed into each other.</p> + +<p>My object in this chapter is solely to shew that there +is no fundamental difference between man and the +higher mammals in their mental faculties. Each division +of the subject might have been extended into a +separate essay, but must here be treated briefly. As +no classification of the mental powers has been universally +accepted, I shall arrange my remarks in the order +most convenient for my purpose; and will select those +facts which have most struck me, with the hope that +they may produce some effect on the reader.</p> + +<p>With respect to animals very low in the scale, I shall +have to give some additional facts under Sexual Selection, +shewing that their mental powers are higher than<span class='pagenum'><a name="Page_36" id="Page_36">36</a></span> +might have been expected. The variability of the faculties +in the individuals of the same species is an important +point for us, and some few illustrations will here +be given. But it would be superfluous to enter into +many details on this head, for I have found on frequent +enquiry, that it is the unanimous opinion of all those +who have long attended to animals of many kinds, +including birds, that the individuals differ greatly in +every mental characteristic. In what manner the mental +powers were first developed in the lowest organisms, +is as hopeless an enquiry as how life first originated. +These are problems for the distant future, if they are +ever to be solved by man.</p> + +<p>As man possesses the same senses with the lower +animals, his fundamental intuitions must be the same. +Man has also some few instincts in common, as that of +self-preservation, sexual love, the love of the mother for +her new-born offspring, the power possessed by the +latter of sucking, and so forth. But man, perhaps, has +somewhat fewer instincts than those possessed by the +animals which come next to him in the series. The +orang in the Eastern islands, and the chimpanzee in +Africa, build platforms on which they sleep; and, as both +species follow the same habit, it might be argued that +this was due to instinct, but we cannot feel sure that it is +not the result of both animals having similar wants and +possessing similar powers of reasoning. These apes, as +we may assume, avoid the many poisonous fruits of the +tropics, and man has no such knowledge; but as our +domestic animals, when taken to foreign lands and when +first turned out in the spring, often eat poisonous herbs, +which they afterwards avoid, we cannot feel sure that +the apes do not learn from their own experience or +from that of their parents what fruits to select. It is +however certain, as we shall presently see, that apes have<span class='pagenum'><a name="Page_37" id="Page_37">37</a></span> +an instinctive dread of serpents, and probably of other +dangerous animals.</p> + +<p>The fewness and the comparative simplicity of the +instincts in the higher animals are remarkable in contrast +with those of the lower animals. Cuvier maintained +that instinct and intelligence stand in an inverse +ratio to each other; and some have thought that the +intellectual faculties of the higher animals have been +gradually developed from their instincts. But Pouchet, +in an interesting essay,<a name="FNanchor_46" id="FNanchor_46"></a><a href="#Footnote_46" class="fnanchor">46</a> has shewn that no such inverse +ratio really exists. Those insects which possess the most +wonderful instincts are certainly the most intelligent. +In the vertebrate series, the least intelligent members, +namely fishes and amphibians, do not possess complex +instincts; and amongst mammals the animal most remarkable +for its instincts, namely the beaver, is highly +intelligent, as will be admitted by every one who has +read Mr. Morgan’s excellent account of this animal.<a name="FNanchor_47" id="FNanchor_47"></a><a href="#Footnote_47" class="fnanchor">47</a></p> + +<p>Although the first dawnings of intelligence, according +to Mr. Herbert Spencer,<a name="FNanchor_48" id="FNanchor_48"></a><a href="#Footnote_48" class="fnanchor">48</a> have been developed +through the multiplication and co-ordination of reflex +actions, and although many of the simpler instincts +graduate into actions of this kind and can hardly be +distinguished from them, as in the case of young animals +sucking, yet the more complex instincts seem to have +originated independently of intelligence. I am, however, +far from wishing to deny that instinctive actions +may lose their fixed and untaught character, and be +replaced by others performed by the aid of the free will. +On the other hand, some intelligent actions—as when, +birds on oceanic islands first learn to avoid man—after +<span class='pagenum'><a name="Page_38" id="Page_38">38</a></span>being performed during many generations, become converted +into instincts and are inherited. They may then +be said to be degraded in character, for they are no +longer performed through reason or from experience. +But the greater number of the more complex instincts +appear to have been gained in a wholly different manner, +through the natural selection of variations of simpler +instinctive actions. Such variations appear to arise from +the same unknown causes acting on the cerebral organisation, +which induce slight variations or individual differences +in other parts of the body; and these variations, +owing to our ignorance, are often said to arise spontaneously. +We can, I think, come to no other conclusion +with respect to the origin of the more complex instincts, +when we reflect on the marvellous instincts of sterile +worker-ants and bees, which leave no offspring to inherit +the effects of experience and of modified habits.</p> + +<p>Although a high degree of intelligence is certainly +compatible with the existence of complex instincts, as +we see in the insects just named and in the beaver, it is +not improbable that they may to a certain extent interfere +with each other’s development. Little is known +about the functions of the brain, but we can perceive +that as the intellectual powers become highly developed, +the various parts of the brain must be connected by the +most intricate channels of intercommunication; and as +a consequence each separate part would perhaps tend to +become less well fitted to answer in a definite and uniform, +that is instinctive, manner to particular sensations +or associations.</p> + +<p>I have thought this digression worth giving, because +we may easily underrate the mental powers of the +higher animals, and especially of man, when we compare +their actions founded on the memory of past +events, on foresight, reason, and imagination, with<span class='pagenum'><a name="Page_39" id="Page_39">39</a></span> +exactly similar actions instinctively performed by the +lower animals; in this latter case the capacity of +performing such actions having been gained, step by +step, through the variability of the mental organs and +natural selection, without any conscious intelligence on +the part of the animal during each successive generation. +No doubt, as Mr. Wallace has argued,<a name="FNanchor_49" id="FNanchor_49"></a><a href="#Footnote_49" class="fnanchor">49</a> much of +the intelligent work done by man is due to imitation and +not to reason; but there is this great difference between +his actions and many of those performed by the lower +animals, namely, that man cannot, on his first trial, +make, for instance, a stone hatchet or a canoe, through +his power of imitation. He has to learn his work by +practice; a beaver, on the other hand, can make its dam +or canal, and a bird its nest, as well, or nearly as well, +the first time it tries, as when old and experienced.</p> + +<p>To return to our immediate subject: the lower +animals, like man, manifestly feel pleasure and pain, +happiness and misery. Happiness is never better exhibited +than by young animals, such as puppies, kittens, +lambs, &c., when playing together, like our own children. +Even insects play together, as has been described by +that excellent observer, P. Huber,<a name="FNanchor_50" id="FNanchor_50"></a><a href="#Footnote_50" class="fnanchor">50</a> who saw ants chasing +and pretending to bite each other, like so many puppies.</p> + +<p>The fact that the lower animals are excited by the +same emotions as ourselves is so well established, that +it will not be necessary to weary the reader by many +details. Terror acts in the same manner on them as on +us, causing the muscles to tremble, the heart to palpitate, +the sphincters to be relaxed, and the hair to +stand on end. Suspicion, the offspring of fear, is eminently +characteristic of most wild animals. Courage +<span class='pagenum'><a name="Page_40" id="Page_40">40</a></span>and timidity are extremely variable qualities in the +individuals of the same species, as is plainly seen in +our dogs. Some dogs and horses are ill-tempered and +easily turn sulky; others are good-tempered; and +these qualities are certainly inherited. Every one +knows how liable animals are to furious rage, and how +plainly they show it. Many anecdotes, probably true, +have been published on the long-delayed and artful +revenge of various animals. The accurate Rengger and +Brehm<a name="FNanchor_51" id="FNanchor_51"></a><a href="#Footnote_51" class="fnanchor">51</a> state that the American and African monkeys +which they kept tame, certainly revenged themselves. +The love of a dog for his master is notorious; in the +agony of death he has been known to caress his master, +and every one has heard of the dog suffering under +vivisection, who licked the hand of the operator; this +man, unless he had a heart of stone, must have felt +remorse to the last hour of his life. As Whewell<a name="FNanchor_52" id="FNanchor_52"></a><a href="#Footnote_52" class="fnanchor">52</a> has +remarked, “who that reads the touching instances of +maternal affection, related so often of the women of +all nations, and of the females of all animals, can +doubt that the principle of action is the same in the +two cases?”</p> + +<p>We see maternal affection exhibited in the most +trifling details; thus Rengger observed an American +monkey (a Cebus) carefully driving away the flies which +plagued her infant; and Duvaucel saw a Hylobates +washing the faces of her young ones in a stream. So +intense is the grief of female monkeys for the loss of +their young, that it invariably caused the death of certain +kinds kept under confinement by Brehm in N. +<span class='pagenum'><a name="Page_41" id="Page_41">41</a></span>Africa. Orphan-monkeys were always adopted and +carefully guarded by the other monkeys, both males +and females. One female baboon had so capacious a +heart that she not only adopted young monkeys of +other species, but stole young dogs and cats, which she +continually carried about. Her kindness, however, did +not go so far as to share her food with her adopted +offspring, at which Brehm was surprised, as his monkeys +always divided everything quite fairly with their own +young ones. An adopted kitten scratched the above-mentioned +affectionate baboon, who certainly had a +fine intellect, for she was much astonished at being +scratched, and immediately examined the kitten’s feet, +and without more ado bit off the claws. In the +Zoological Gardens, I heard from the keeper that +an old baboon (<i>C. chacma</i>) had adopted a Rhesus +monkey; but when a young drill and mandrill were +placed in the cage, she seemed to perceive that these +monkeys, though distinct species, were her nearer relatives, +for she at once rejected the Rhesus and adopted +both of them. The young Rhesus, as I saw, was greatly +discontented at being thus rejected, and it would, like +a naughty child, annoy and attack the young drill and +mandrill whenever it could do so with safety; this +conduct exciting great indignation in the old baboon. +Monkeys will also, according to Brehm, defend their +master when attacked by any one, as well as dogs to +whom they are attached, from the attacks of other +dogs. But we here trench on the subject of sympathy, +to which I shall recur. Some of Brehm’s monkeys +took much delight in teasing, in various ingenious +ways, a certain old dog whom they disliked, as well +as other animals.</p> + +<p>Most of the more complex emotions are common to +the higher animals and ourselves. Every one has seen<span class='pagenum'><a name="Page_42" id="Page_42">42</a></span> +how jealous a dog is of his master’s affection, if lavished +on any other creature; and I have observed the same fact +with monkeys. This shews that animals not only love, +but have the desire to be loved. Animals manifestly +feel emulation. They love approbation or praise; and +a dog carrying a basket for his master exhibits in a high +degree self-complacency or pride. There can, I think, +be no doubt that a dog feels shame, as distinct from +fear, and something very like modesty when begging +too often for food. A great dog scorns the snarling of +a little dog, and this may be called magnanimity. +Several observers have stated that monkeys certainly +dislike being laughed at; and they sometimes invent +imaginary offences. In the Zoological Gardens I saw a +baboon who always got into a furious rage when his +keeper took out a letter or book and read it aloud to +him; and his rage was so violent that, as I witnessed on +one occasion, he bit his own leg till the blood flowed.</p> + +<p>We will now turn to the more intellectual emotions +and faculties, which are very important, as forming the +basis for the development of the higher mental powers. +Animals manifestly enjoy excitement and suffer from +ennui, as may be seen with dogs, and, according to +Rengger, with monkeys. All animals feel Wonder, +and many exhibit Curiosity. They sometimes suffer +from this latter quality, as when the hunter plays antics +and thus attracts them; I have witnessed this with +deer, and so it is with the wary chamois, and with some +kinds of wild-ducks. Brehm gives a curious account of +the instinctive dread which his monkeys exhibited +towards snakes; but their curiosity was so great that +they could not desist from occasionally satiating their +horror in a most human fashion, by lifting up the lid +of the box in which the snakes were kept. I was so +much surprised at his account, that I took a stuffed and<span class='pagenum'><a name="Page_43" id="Page_43">43</a></span> +coiled-up snake into the monkey-house at the Zoological +Gardens, and the excitement thus caused was +one of the most curious spectacles which I ever beheld. +Three species of Cercopithecus were the most alarmed; +they dashed about their cages and uttered sharp signal-cries +of danger, which were understood by the other +monkeys. A few young monkeys and one old Anubis +baboon alone took no notice of the snake. I then +placed the stuffed specimen on the ground in one of +the larger compartments. After a time all the monkeys +collected round it in a large circle, and staring intently, +presented a most ludicrous appearance. They +became extremely nervous; so that when a wooden ball, +with which they were familiar as a plaything, was accidently +moved in the straw, under which it was partly +hidden, they all instantly started away. These monkeys +behaved very differently when a dead fish, a mouse, and +some other new objects were placed in their cages; for +though at first frightened, they soon approached, handled +and examined them. I then placed a live snake in a +paper bag, with the mouth loosely closed, in one of the +larger compartments. One of the monkeys immediately +approached, cautiously opened the bag a little, peeped +in, and instantly dashed away. Then I witnessed what +Brehm has described, for monkey after monkey, with +head raised high and turned on one side, could not +resist taking momentary peeps into the upright bag, +at the dreadful object lying quiet at the bottom. It +would almost appear as if monkeys had some notion of +zoological affinities, for those kept by Brehm exhibited +a strange, though mistaken, instinctive dread of innocent +lizards and frogs. An orang, also, has been known +to be much alarmed at the first sight of a turtle.<a name="FNanchor_53" id="FNanchor_53"></a><a href="#Footnote_53" class="fnanchor">53</a></p> +<p><span class='pagenum'><a name="Page_44" id="Page_44">44</a></span></p> +<p>The principle of <i>Imitation</i> is strong in man, and +especially in man in a barbarous state. Desor<a name="FNanchor_54" id="FNanchor_54"></a><a href="#Footnote_54" class="fnanchor">54</a> has +remarked that no animal voluntarily imitates an action +performed by man, until in the ascending scale we +come to monkeys, which are well-known to be ridiculous +mockers. Animals, however, sometimes imitate each +others’ actions: thus two species of wolves, which had +been reared by dogs, learned to bark, as does sometimes +the jackal,<a name="FNanchor_55" id="FNanchor_55"></a><a href="#Footnote_55" class="fnanchor">55</a> but whether this can be called voluntary +imitation is another question. From one account +which I have read, there is reason to believe that puppies +nursed by cats sometimes learn to lick their feet and +thus to clean their faces: it is at least certain, as I hear +from a perfectly trustworthy friend, that some dogs +behave in this manner. Birds imitate the songs of their +parents, and sometimes those of other birds; and parrots +are notorious imitators of any sound which they +often hear.</p> + +<p>Hardly any faculty is more important for the intellectual +progress of man than the power of <i>Attention</i>. +Animals clearly manifest this power, as when a cat +watches by a hole and prepares to spring on its prey. +Wild animals sometimes become so absorbed when thus +engaged, that they may be easily approached. Mr. +Bartlett has given me a curious proof how variable this +faculty is in monkeys. A man who trains monkeys to +act used to purchase common kinds from the Zoological +Society at the price of five pounds for each; but he +offered to give double the price, if he might keep three +or four of them for a few days, in order to select one. +When asked how he could possibly so soon learn whether +<span class='pagenum'><a name="Page_45" id="Page_45">45</a></span>a particular monkey would turn out a good actor, he +answered that it all depended on their power of attention. +If when he was talking and explaining anything +to a monkey, its attention was easily distracted, as by +a fly on the wall or other trifling object, the case was +hopeless. If he tried by punishment to make an inattentive +monkey act, it turned sulky. On the other +hand, a monkey which carefully attended to him could +always be trained.</p> + +<p>It is almost superfluous to state that animals have +excellent <i>Memories</i> for persons and places. A baboon +at the Cape of Good Hope, as I have been informed by +Sir Andrew Smith, recognised him with joy after an +absence of nine months. I had a dog who was savage +and averse to all strangers, and I purposely tried his +memory after an absence of five years and two days. I +went near the stable where he lived, and shouted to +him in my old manner; he showed no joy, but instantly +followed me out walking and obeyed me, +exactly as if I had parted with him only half-an-hour +before. A train of old associations, dormant during five +years, had thus been instantaneously awakened in his +mind. Even ants, as P. Huber<a name="FNanchor_56" id="FNanchor_56"></a><a href="#Footnote_56" class="fnanchor">56</a> has clearly shewn, +recognised their fellow-ants belonging to the same community +after a separation of four months. Animals +can certainly by some means judge of the intervals of +time between recurrent events.</p> + +<p>The <i>Imagination</i> is one of the highest prerogatives +of man. By this faculty he unites, independently of +the will, former images and ideas, and thus creates brilliant +and novel results. A poet, as Jean Paul Richter +remarks,<a name="FNanchor_57" id="FNanchor_57"></a><a href="#Footnote_57" class="fnanchor">57</a> “who must reflect whether he shall make a +<span class='pagenum'><a name="Page_46" id="Page_46">46</a></span>character say yes or no—to the devil with him; he is +only a stupid corpse.” Dreaming gives us the best +notion of this power; as Jean Paul again says, “The +dream is an involuntary art of poetry.” The value of +the products of our imagination depends of course on +the number, accuracy, and clearness of our impressions; +on our judgment and taste in selecting or rejecting the +involuntary combinations, and to a certain extent on +our power of voluntarily combining them. As dogs, +cats, horses, and probably all the higher animals, even +birds, as is stated on good authority,<a name="FNanchor_58" id="FNanchor_58"></a><a href="#Footnote_58" class="fnanchor">58</a> have vivid dreams, +and this is shewn by their movements and voice, we must +admit that they possess some power of imagination.</p> + +<p>Of all the faculties of the human mind, it will, I +presume, be admitted that <i>Reason</i> stands at the summit. +Few persons any longer dispute that animals possess +some power of reasoning. Animals may constantly be +seen to pause, deliberate, and resolve. It is a significant +fact, that the more the habits of any particular animal +are studied by a naturalist, the more he attributes to +reason and the less to unlearnt instincts.<a name="FNanchor_59" id="FNanchor_59"></a><a href="#Footnote_59" class="fnanchor">59</a> In future +chapters we shall see that some animals extremely low in +the scale apparently display a certain amount of reason. +No doubt it is often difficult to distinguish between the +power of reason and that of instinct. Thus Dr. Hayes, +in his work on ‘The Open Polar Sea,’ repeatedly remarks +that his dogs, instead of continuing to draw the +sledges in a compact body, diverged and separated when +they came to thin ice, so that their weight might be +more evenly distributed. This was often the first warn<span class='pagenum'><a name="Page_47" id="Page_47">47</a></span>ing +and notice which the travellers received that the ice +was becoming thin and dangerous. Now, did the dogs +act thus from the experience of each individual, or from +the example of the older and wiser dogs, or from an +inherited habit, that is from an instinct? This instinct +might possibly have arisen since the time, long ago, +when dogs were first employed by the natives in drawing +their sledges; or the Arctic wolves, the parent-stock +of the Esquimaux dog, may have acquired this instinct, +impelling them not to attack their prey in a close pack +when on thin ice. Questions of this kind are most +difficult to answer.</p> + +<p>So many facts have been recorded in various works +shewing that animals possess some degree of reason, +that I will here give only two or three instances, authenticated +by Rengger, and relating to American monkeys, +which stand low in their order. He states that when +he first gave eggs to his monkeys, they smashed them +and thus lost much of their contents; afterwards they +gently hit one end against some hard body, and picked +off the bits of shell with their fingers. After cutting +themselves only once with any sharp tool, they would +not touch it again, or would handle it with the greatest +care. Lumps of sugar were often given them wrapped +up in paper; and Rengger sometimes put a live wasp +in the paper, so that in hastily unfolding it they got +stung; after this had once happened, they always first +held the packet to their ears to detect any movement +within. Any one who is not convinced by such facts as +these, and by what he may observe with his own dogs, +that animals can reason, would not be convinced by +anything that I could add. Nevertheless I will give +one case with respect to dogs, as it rests on two distinct +observers, and can hardly depend on the modification +of any instinct.</p> + +<p><span class='pagenum'><a name="Page_48" id="Page_48">48</a></span>Mr. Colquhoun<a name="FNanchor_60" id="FNanchor_60"></a><a href="#Footnote_60" class="fnanchor">60</a> winged two wild-ducks, which fell +on the opposite side of a stream; his retriever tried to +bring over both at once, but could not succeed; she +then, though never before known to ruffle a feather, +deliberately killed one, brought over the other, and returned +for the dead bird. Col. Hutchinson relates that +two partridges were shot at once, one being killed, the +other wounded; the latter ran away, and was caught by +the retriever, who on her return came across the dead +bird; “she stopped, evidently greatly puzzled, and +after one or two trials, finding she could not take it up +without permitting the escape of the winged bird, she +considered a moment, then deliberately murdered it +by giving it a severe crunch, and afterwards brought +away both together. This was the only known instance +of her ever having wilfully injured any game.” +Here we have reason, though not quite perfect, for the +retriever might have brought the wounded bird first +and then returned for the dead one, as in the case of +the two wild-ducks.</p> + +<p class="tb">The muleteers in S. America say, “I will not give +you the mule whose step is easiest, but <i>la mas rational,</i>—the +one that reasons best;” and Humboldt<a name="FNanchor_61" id="FNanchor_61"></a><a href="#Footnote_61" class="fnanchor">61</a> adds, +“this popular expression, dictated by long experience, +combats the system of animated machines, better perhaps +than all the arguments of speculative philosophy.”</p> + +<p>It has, I think, now been shewn that man and the +higher animals, especially the Primates, have some few +instincts in common. All have the same senses, intuitions +and sensations—similar passions, affections, and +emotions, even the more complex ones; they feel +<span class='pagenum'><a name="Page_49" id="Page_49">49</a></span>wonder and curiosity; they possess the same faculties +of imitation, attention, memory, imagination, and reason, +though in very different degrees. Nevertheless many +authors have insisted that man is separated through his +mental faculties by an impassable barrier from all the +lower animals. I formerly made a collection of above +a score of such aphorisms, but they are not worth +giving, as their wide difference and number prove the +difficulty, if not the impossibility, of the attempt. It +has been asserted that man alone is capable of progressive +improvement; that he alone makes use of tools or +fire, domesticates other animals, possesses property, or +employs language; that no other animal is self-conscious, +comprehends itself, has the power of abstraction, +or possesses general ideas; that man alone has a sense +of beauty, is liable to caprice, has the feeling of gratitude, +mystery, &c.; believes in God, or is endowed with +a conscience. I will hazard a few remarks on the more +important and interesting of these points.</p> + +<p>Archbishop Sumner formerly maintained<a name="FNanchor_62" id="FNanchor_62"></a><a href="#Footnote_62" class="fnanchor">62</a> that man +alone is capable of progressive improvement. With +animals, looking first to the individual, every one who +has had any experience in setting traps knows that +young animals can be caught much more easily than +old ones; and they can be much more easily approached +by an enemy. Even with respect to old animals, it is +impossible to catch many in the same place and in the +same kind of trap, or to destroy them by the same kind +of poison; yet it is improbable that all should have +partaken of the poison, and impossible that all should +have been caught in the trap. They must learn caution +by seeing their brethren caught or poisoned. In North +America, where the fur-bearing animals have long been +<span class='pagenum'><a name="Page_50" id="Page_50">50</a></span>pursued, they exhibit, according to the unanimous testimony +of all observers, an almost incredible amount +of sagacity, caution, and cunning; but trapping has +been there so long carried on that inheritance may have +come into play.</p> + +<p>If we look to successive generations, or to the race, +there is no doubt that birds and other animals gradually +both acquire and lose caution in relation to man or +other enemies;<a name="FNanchor_63" id="FNanchor_63"></a><a href="#Footnote_63" class="fnanchor">63</a> and this caution is certainly in chief +part an inherited habit or instinct, but in part the result +of individual experience. A good observer, Leroy,<a name="FNanchor_64" id="FNanchor_64"></a><a href="#Footnote_64" class="fnanchor">64</a> +states that in districts where foxes are much hunted, +the young when they first leave their burrows are incontestably +much more wary than the old ones in districts +where they are not much disturbed.</p> + +<p>Our domestic dogs are descended from wolves and +jackals,<a name="FNanchor_65" id="FNanchor_65"></a><a href="#Footnote_65" class="fnanchor">65</a> and though they may not have gained in +cunning, and may have lost in waryness and suspicion, +yet they have progressed in certain moral qualities, +such as in affection, trust-worthiness, temper, and probably +in general intelligence. The common rat has +conquered and beaten several other species throughout +Europe, in parts of North America, New Zealand, +and recently in Formosa, as well as on the mainland of +China. Mr. Swinhoe,<a name="FNanchor_66" id="FNanchor_66"></a><a href="#Footnote_66" class="fnanchor">66</a> who describes these latter cases, +attributes the victory of the common rat over the large +<i>Mus coninga</i> to its superior cunning; and this latter +quality may be attributed to the habitual exercise of +all its faculties in avoiding extirpation by man, as well +<span class='pagenum'><a name="Page_51" id="Page_51">51</a></span>as to nearly all the less cunning or weak-minded rats +having been successively destroyed by him. To maintain, +independently of any direct evidence, that no +animal during the course of ages has progressed in +intellect or other mental faculties, is to beg the question +of the evolution of species. Hereafter we shall see that, +according to Lartet, existing mammals belonging to +several orders have larger brains than their ancient +tertiary prototypes.</p> + +<p>It has often been said that no animal uses any tool; +but the chimpanzee in a state of nature cracks a native +fruit, somewhat like a walnut, with a stone.<a name="FNanchor_67" id="FNanchor_67"></a><a href="#Footnote_67" class="fnanchor">67</a> Rengger<a name="FNanchor_68" id="FNanchor_68"></a><a href="#Footnote_68" class="fnanchor">68</a> +easily taught an American monkey thus to break open +hard palm-nuts, and afterwards of its own accord it +used stones to open other kinds of nuts, as well as +boxes. It thus also removed the soft rind of fruit that +had a disagreeable flavour. Another monkey was taught +to open the lid of a large box with a stick, and afterwards +it used the stick as a lever to move heavy bodies; +and I have myself seen a young orang put a stick into +a crevice, slip his hand to the other end, and use it in +the proper manner as a lever. In the cases just mentioned +stones and sticks were employed as implements; +but they are likewise used as weapons. Brehm<a name="FNanchor_69" id="FNanchor_69"></a><a href="#Footnote_69" class="fnanchor">69</a> states, +on the authority of the well-known traveller Schimper, +that in Abyssinia when the baboons belonging to one +species (<i>C. gelada</i>) descend in troops from the mountains +to plunder the fields, they sometimes encounter +troops of another species (<i>C. hamadryas</i>), and then a +fight ensues. The Geladas roll down great stones, which +the Hamadryas try to avoid, and then, both species, +<span class='pagenum'><a name="Page_52" id="Page_52">52</a></span>making a great uproar, rush furiously against each +other. Brehm, when accompanying the Duke of Coburg-Gotha, +aided in an attack with fire-arms on a troop of +baboons in the pass of Mensa in Abyssinia. The baboons +in return rolled so many stones down the mountain, +some as large as a man’s head, that the attackers had +to beat a hasty retreat; and the pass was actually for +a time closed against the caravan. It deserves notice +that these baboons thus acted in concert. Mr. Wallace<a name="FNanchor_70" id="FNanchor_70"></a><a href="#Footnote_70" class="fnanchor">70</a> +on three occasions saw female orangs, accompanied +by their young, “breaking off branches and +the great spiny fruit of the Durian tree, with every +appearance of rage; causing such a shower of missiles +as effectually kept us from approaching too near the +tree.”</p> + +<p>In the Zoological Gardens a monkey which had weak +teeth used to break open nuts with a stone; and I was +assured by the keepers that this animal, after using the +stone, hid it in the straw, and would not let any other +monkey touch it. Here, then, we have the idea of +property; but this idea is common to every dog with a +bone, and to most or all birds with their nests.</p> + +<p class="tb">The Duke of Argyll<a name="FNanchor_71" id="FNanchor_71"></a><a href="#Footnote_71" class="fnanchor">71</a> remarks, that the fashioning of +an implement for a special purpose is absolutely peculiar +to man; and he considers that this forms an immeasurable +gulf between him and the brutes. It is no doubt +a very important distinction, but there appears to me +much truth in Sir J. Lubbock’s suggestion,<a name="FNanchor_72" id="FNanchor_72"></a><a href="#Footnote_72" class="fnanchor">72</a> that when +primeval man first used flint-stones for any purpose, he +would have accidentally splintered them, and would +then have used the sharp fragments. From this step +it would be a small one to intentionally break the +<span class='pagenum'><a name="Page_53" id="Page_53">53</a></span>flints, and not a very wide step to rudely fashion them. +This latter advance, however, may have taken long +ages, if we may judge by the immense interval of time +which elapsed before the men of the neolithic period +took to grinding and polishing their stone tools. In +breaking the flints, as Sir J. Lubbock likewise remarks, +sparks would have been emitted, and in grinding them +heat would have been evolved: “thus the two usual +methods of obtaining fire may have originated.” The +nature of fire would have been known in the many +volcanic regions where lava occasionally flows through +forests. The anthropomorphous apes, guided probably +by instinct, build for themselves temporary platforms; +but as many instincts are largely controlled by reason, +the simpler ones, such as this of building a platform, +might readily pass into a voluntary and conscious act. +The orang is known to cover itself at night with the +leaves of the Pandanus; and Brehm states that one of +his baboons used to protect itself from the heat of the +sun by throwing a straw-mat over its head. In these +latter habits, we probably see the first steps towards +some of the simpler arts; namely rude architecture +and dress, as they arose amongst the early progenitors +of man.</p> + +<p><i>Language.</i>—This faculty has justly been considered as +one of the chief distinctions between man and the lower +animals. But man, as a highly competent judge, Archbishop +Whately remarks, “is not the only animal that +can make use of language to express what is passing in +his mind, and can understand, more or less, what is so +expressed by another.”<a name="FNanchor_73" id="FNanchor_73"></a><a href="#Footnote_73" class="fnanchor">73</a> In Paraguay the <i>Cebus Azaræ</i> +when excited utters at least six distinct sounds, which +<span class='pagenum'><a name="Page_54" id="Page_54">54</a></span>excite in other monkeys similar emotions.<a name="FNanchor_74" id="FNanchor_74"></a><a href="#Footnote_74" class="fnanchor">74</a> The movements +of the features and gestures of monkeys are understood +by us, and they partly understand ours, as +Rengger and others declare. It is a more remarkable +fact that the dog, since being domesticated, has +learnt to bark<a name="FNanchor_75" id="FNanchor_75"></a><a href="#Footnote_75" class="fnanchor">75</a> in at least four or five distinct tones. +Although barking is a new art, no doubt the wild species, +the parents of the dog, expressed their feelings +by cries of various kinds. With the domesticated +dog we have the bark of eagerness, as in the chase; +that of anger; the yelping or howling bark of despair, +as when shut up; that of joy, as when starting on a +walk with his master; and the very distinct one of +demand or supplication, as when wishing for a door or +window to be opened.</p> + +<p>Articulate language is, however, peculiar to man; +but he uses in common with the lower animals inarticulate +cries to express his meaning, aided by gestures +and the movements of the muscles of the face.<a name="FNanchor_76" id="FNanchor_76"></a><a href="#Footnote_76" class="fnanchor">76</a> This +especially holds good with the more simple and vivid +feelings, which are but little connected with our higher +intelligence. Our cries of pain, fear, surprise, anger, together +with their appropriate actions, and the murmur +of a mother to her beloved child, are more expressive +than any words. It is not the mere power of articulation +that distinguishes man from other animals, for as +every one knows, parrots can talk; but it is his large +power of connecting definite sounds with definite ideas; +and this obviously depends on the development of the +mental faculties.</p> + +<p><span class='pagenum'><a name="Page_55" id="Page_55">55</a></span></p><p>As Horne Tooke, one of the founders of the noble +science of philology, observes, language is an art, like +brewing or baking; but writing would have been a +much more appropriate simile. It certainly is not a +true instinct, as every language has to be learnt. It +differs, however, widely from all ordinary arts, for man +has an instinctive tendency to speak, as we see in the +babble of our young children; whilst no child has an +instinctive tendency to brew, bake, or write. Moreover, +no philologist now supposes that any language has +been deliberately invented; each has been slowly and +unconsciously developed by many steps. The sounds +uttered by birds offer in several respects the nearest +analogy to language, for all the members of the same +species utter the same instinctive cries expressive of +their emotions; and all the kinds that have the power +of singing exert this power instinctively; but the actual +song, and even the call-notes, are learnt from their +parents or foster-parents. These sounds, as Daines +Barrington<a name="FNanchor_77" id="FNanchor_77"></a><a href="#Footnote_77" class="fnanchor">77</a> has proved, “are no more innate than +language is in man.” The first attempts to sing +“may be compared to the imperfect endeavour in a +child to babble.” The young males continue practising, +or, as the bird-catchers say, recording, for ten +or eleven months. Their first essays show hardly a +rudiment of the future song; but as they grow older +we can perceive what they are aiming at; and at last +they are said “to sing their song round.” Nestlings +which have learnt the song of a distinct species, as +with the canary-birds educated in the Tyrol, teach and +transmit their new song to their offspring. The slight +natural differences of song in the same species inha<span class='pagenum'><a name="Page_56" id="Page_56">56</a></span>biting +different districts may be appositely compared, +as Barrington remarks, “to provincial dialects;” and +the songs of allied, though distinct species may be compared +with the languages of distinct races of man. I +have given the foregoing details to shew that an instinctive +tendency to acquire an art is not a peculiarity +confined to man.</p> + +<p>With respect to the origin of articulate language, +after having read on the one side the highly interesting +works of Mr. Hensleigh Wedgwood, the Rev. F. Farrar, +and Prof. Schleicher,<a name="FNanchor_78" id="FNanchor_78"></a><a href="#Footnote_78" class="fnanchor">78</a> and the celebrated lectures of +Prof. Max Müller on the other side, I cannot doubt that +language owes its origin to the imitation and modification, +aided by signs and gestures, of various +natural sounds, the voices of other animals, and man’s +own instinctive cries. When we treat of sexual selection +we shall see that primeval man, or rather some +early progenitor of man, probably used his voice largely, +as does one of the gibbon-apes at the present day, in +producing true musical cadences, that is in singing; +we may conclude from a widely-spread analogy that +this power would have been especially exerted during +the courtship of the sexes, serving to express various +emotions, as love, jealousy, triumph, and serving as a +challenge to their rivals. The imitation by articulate +sounds of musical cries might have given rise to +words expressive of various complex emotions. As +bearing on the subject of imitation, the strong tendency +in our nearest allies, the monkeys, in microcephalous +<span class='pagenum'><a name="Page_57" id="Page_57">57</a></span>idiots,<a name="FNanchor_79" id="FNanchor_79"></a><a href="#Footnote_79" class="fnanchor">79</a> and in the barbarous races of mankind, to imitate +whatever they hear deserves notice. As monkeys +certainly understand much that is said to them by man, +and as in a state of nature they utter signal-cries of +danger to their fellows,<a name="FNanchor_80" id="FNanchor_80"></a><a href="#Footnote_80" class="fnanchor">80</a> it does not appear altogether +incredible, that some unusually wise ape-like animal +should have thought of imitating the growl of a beast +of prey, so as to indicate to his fellow monkeys the +nature of the expected danger. And this would have +been a first step in the formation of a language.</p> + +<p>As the voice was used more and more, the vocal +organs would have been strengthened and perfected +through the principle of the inherited effects of use; +and this would have reacted on the power of speech. +But the relation between the continued use of language +and the development of the brain has no doubt been far +more important. The mental powers in some early progenitor +of man must have been more highly developed +than in any existing ape, before even the most imperfect +form of speech could have come into use; but we may +confidently believe that the continued use and advancement +of this power would have reacted on the mind by +enabling and encouraging it to carry on long trains of +thought. A long and complex train of thought can no +more be carried on without the aid of words, whether +spoken or silent, than a long calculation without the +use of figures or algebra. It appears, also, that even +ordinary trains of thought almost require some form of +language, for the dumb, deaf, and blind girl, Laura +Bridgman, was observed to use her fingers whilst dream<span class='pagenum'><a name="Page_58" id="Page_58">58</a></span>ing.<a name="FNanchor_81" id="FNanchor_81"></a><a href="#Footnote_81" class="fnanchor">81</a> +Nevertheless a long succession of vivid and connected +ideas, may pass through the mind without the +aid of any form of language, as we may infer from the +prolonged dreams of dogs. We have, also, seen that +retriever-dogs are able to reason to a certain extent; +and this they manifestly do without the aid of language. +The intimate connection between the brain, as it is +now developed in us, and the faculty of speech, is well +shewn by those curious cases of brain-disease, in which +speech is specially affected, as when the power to remember +substantives is lost, whilst other words can be +correctly used.<a name="FNanchor_82" id="FNanchor_82"></a><a href="#Footnote_82" class="fnanchor">82</a> There is no more improbability in +the effects of the continued use of the vocal and mental +organs being inherited, than in the case of handwriting, +which depends partly on the structure of the +hand and partly on the disposition of the mind; and +handwriting is certainly inherited.<a name="FNanchor_83" id="FNanchor_83"></a><a href="#Footnote_83" class="fnanchor">83</a></p> + +<p>Why the organs now used for speech should have +been originally perfected for this purpose, rather than +any other organs, it is not difficult to see. Ants have +considerable powers of intercommunication by means +of their antennæ, as shewn by Huber, who devotes a +whole chapter to their language. We might have used +our fingers as efficient instruments, for a person with +practice can report to a deaf man every word of a speech +rapidly delivered at a public meeting; but the loss of +our hands, whilst thus employed, would have been +a serious inconvenience. As all the higher mammals +possess vocal organs constructed on the same general +<span class='pagenum'><a name="Page_59" id="Page_59">59</a></span>plan with ours, and which are used as a means of communication, +it was obviously probable, if the power of communication +had to be improved, that these same organs +would have been still further developed; and this has +been effected by the aid of adjoining and well-adapted +parts, namely the tongue and lips.<a name="FNanchor_84" id="FNanchor_84"></a><a href="#Footnote_84" class="fnanchor">84</a> The fact of the +higher apes not using their vocal organs for speech, no +doubt depends on their intelligence not having been +sufficiently advanced. The possession by them of organs, +which with long-continued practice might have been +used for speech, although not thus used, is paralleled by +the case of many birds which possess organs fitted for +singing, though they never sing. Thus, the nightingale +and crow have vocal organs similarly constructed, these +being used by the former for diversified song, and by +the latter merely for croaking.<a name="FNanchor_85" id="FNanchor_85"></a><a href="#Footnote_85" class="fnanchor">85</a></p> + +<p>The formation of different languages and of distinct +species, and the proofs that both have been developed +through a gradual process, are curiously the same.<a name="FNanchor_86" id="FNanchor_86"></a><a href="#Footnote_86" class="fnanchor">86</a> +But we can trace the origin of many words further +back than in the case of species, for we can perceive +that they have arisen from the imitation of various +sounds, as in alliterative poetry. We find in distinct +languages striking homologies due to community of +descent, and analogies due to a similar process of +<span class='pagenum'><a name="Page_60" id="Page_60">60</a></span>formation. The manner in which certain letters or +sounds change when others change is very like correlated +growth. We have in both cases the reduplication +of parts, the effects of long-continued use, and so forth. +The frequent presence of rudiments, both in languages +and in species, is still more remarkable. The letter <i>m</i> +in the word <i>am</i>, means <i>I</i>; so that in the expression <i>I am</i>, +a superfluous and useless rudiment has been retained. +In the spelling also of words, letters often remain +as the rudiments of ancient forms of pronunciation. +Languages, like organic beings, can be classed in groups +under groups; and they can be classed either naturally +according to descent, or artificially by other characters. +Dominant languages and dialects spread widely and +lead to the gradual extinction of other tongues. A language, +like a species, when once extinct, never, as Sir +C. Lyell remarks, reappears. The same language never +has two birthplaces. Distinct languages may be crossed +or blended together.<a name="FNanchor_87" id="FNanchor_87"></a><a href="#Footnote_87" class="fnanchor">87</a> We see variability in every +tongue, and new words are continually cropping up; but +as there is a limit to the powers of the memory, single +words, like whole languages, gradually become extinct. +As Max Müller<a name="FNanchor_88" id="FNanchor_88"></a><a href="#Footnote_88" class="fnanchor">88</a> has well remarked:—“A struggle for +life is constantly going on amongst the words and grammatical +forms in each language. The better, the +shorter, the easier forms are constantly gaining the +upper hand, and they owe their success to their own +inherent virtue.” To these more important causes of +the survival of certain words, mere novelty may, I +think, be added; for there is in the mind of man a strong +love for slight changes in all things. The survival or +<span class='pagenum'><a name="Page_61" id="Page_61">61</a></span>preservation of certain favoured words in the struggle +for existence is natural selection.</p> + +<p>The perfectly regular and wonderfully complex construction +of the languages of many barbarous nations +has often been advanced as a proof, either of the divine +origin of these languages, or of the high art and former +civilisation of their founders. Thus F. von Schlegel +writes: “In those languages which appear to be at the +lowest grade of intellectual culture, we frequently observe +a very high and elaborate degree of art in their +grammatical structure. This is especially the case with +the Basque and the Lapponian, and many of the American +languages.”<a name="FNanchor_89" id="FNanchor_89"></a><a href="#Footnote_89" class="fnanchor">89</a> But it is assuredly an error to speak +of any language as an art in the sense of its having +been elaborately and methodically formed. Philologists +now admit that conjugations, declensions, &c., originally +existed as distinct words, since joined together; +and as such words express the most obvious relations +between objects and persons, it is not surprising that +they should have been used by the men of most races +during the earliest ages. With respect to perfection, +the following illustration will best shew how easily we +may err: a Crinoid sometimes consists of no less +than 150,000 pieces of shell,<a name="FNanchor_90" id="FNanchor_90"></a><a href="#Footnote_90" class="fnanchor">90</a> all arranged with perfect +symmetry in radiating lines; but a naturalist does +not consider an animal of this kind as more perfect +than a bilateral one with comparatively few parts, +and with none of these alike, excepting on the opposite +sides of the body. He justly considers the differentiation +and specialisation of organs as the test of perfection. +So with languages, the most symmetrical and +complex ought not to be ranked above irregular, abbre<span class='pagenum'><a name="Page_62" id="Page_62">62</a></span>viated, +and bastardised languages, which have borrowed +expressive words and useful forms of construction from +various conquering, or conquered, or immigrant races.</p> + +<p class="tb">From these few and imperfect remarks I conclude +that the extremely complex and regular construction of +many barbarous languages, is no proof that they owe +their origin to a special act of creation.<a name="FNanchor_91" id="FNanchor_91"></a><a href="#Footnote_91" class="fnanchor">91</a> Nor, as +we have seen, does the faculty of articulate speech in +itself offer any insuperable objection to the belief that +man has been developed from some lower form.</p> + +<p><i>Self-consciousness, Individuality, Abstraction, General +Ideas, &c.</i>—It would be useless to attempt discussing +these high faculties, which, according to several recent +writers, make the sole and complete distinction between +man and the brutes, for hardly two authors agree in their +definitions. Such faculties could not have been fully +developed in man until his mental powers had advanced +to a high standard, and this implies the use of a perfect +language. No one supposes that one of the lower animals +reflects whence he comes or whither he goes,—what +is death or what is life, and so forth. But can +we feel sure that an old dog with an excellent memory +and some power of imagination, as shewn by his +dreams, never reflects on his past pleasures in the +chase? and this would be a form of self-consciousness. +On the other hand, as Büchner<a name="FNanchor_92" id="FNanchor_92"></a><a href="#Footnote_92" class="fnanchor">92</a> has remarked, how +little can the hard-worked wife of a degraded Australian +savage, who uses hardly any abstract words and cannot +count above four, exert her self-consciousness, or reflect +on the nature of her own existence.</p> + +<p class="tb"><span class='pagenum'><a name="Page_63" id="Page_63">63</a></span>That animals retain their mental individuality is +unquestionable. When my voice awakened a train of +old associations in the mind of the above-mentioned +dog, he must have retained his mental individuality, +although every atom of his brain had probably undergone +change more than once during the interval of five +years. This dog might have brought forward the +argument lately advanced to crush all evolutionists, +and said, “I abide amid all mental moods and all +material changes.... The teaching that atoms leave +their impressions as legacies to other atoms falling +into the places they have vacated is contradictory of +the utterance of consciousness, and is therefore false; +but it is the teaching necessitated by evolutionism, +consequently the hypothesis is a false one.”<a name="FNanchor_93" id="FNanchor_93"></a><a href="#Footnote_93" class="fnanchor">93</a></p> + +<p><i>Sense of Beauty.</i>—This sense has been declared to +be peculiar to man. But when we behold male birds +elaborately displaying their plumes and splendid colours +before the females, whilst other birds not thus decorated +make no such display, it is impossible to doubt +that the females admire the beauty of their male partners. +As women everywhere deck themselves with +these plumes, the beauty of such ornaments cannot be +disputed. The Bower-birds by tastefully ornamenting +their playing-passages with gaily-coloured objects, as +do certain humming-birds their nests, offer additional +evidence that they possess a sense of beauty. So with +the song of birds, the sweet strains poured forth by the +males during the season of love are certainly admired +by the females, of which fact evidence will hereafter be +given. If female birds had been incapable of appreciating +the beautiful colours, the ornaments, and voices +<span class='pagenum'><a name="Page_64" id="Page_64">64</a></span>of their male partners, all the labour and anxiety exhibited +by them in displaying their charms before the +females would have been thrown away; and this it is +impossible to admit. Why certain bright colours and +certain sounds should excite pleasure, when in harmony, +cannot, I presume, be explained any more than why +certain flavours and scents are agreeable; but assuredly +the same colours and the same sounds are admired by +us and by many of the lower animals.</p> + +<p>The taste for the beautiful, at least as far as female +beauty is concerned, is not of a special nature in the +human mind; for it differs widely in the different +races of man, as will hereafter be shewn, and is not +quite the same even in the different nations of the +same race. Judging from the hideous ornaments and +the equally hideous music admired by most savages, it +might be urged that their æsthetic faculty was not so +highly developed as in certain animals, for instance, in +birds. Obviously no animal would be capable of admiring +such scenes as the heavens at night, a beautiful +landscape, or refined music; but such high tastes, depending +as they do on culture and complex associations, +are not enjoyed by barbarians or by uneducated +persons.</p> + +<p class="tb">Many of the faculties, which have been of inestimable +service to man for his progressive advancement, +such as the powers of the imagination, wonder, +curiosity, an undefined sense of beauty, a tendency +to imitation, and the love of excitement or novelty, +could not fail to have led to the most capricious +changes of customs and fashions. I have alluded to +this point, because a recent writer<a name="FNanchor_94" id="FNanchor_94"></a><a href="#Footnote_94" class="fnanchor">94</a> has oddly fixed +on Caprice “as one of the most remarkable and +<span class='pagenum'><a name="Page_65" id="Page_65">65</a></span>typical differences between savages and brutes.” But +not only can we perceive how it is that roan is capricious, +but the lower animals are, as we shall hereafter +see, capricious in their affections, aversions, and sense +of beauty. There is also good reason to suspect that +they love novelty, for its own sake.</p> + +<p><i>Belief in God—Religion.</i>—There is no evidence that +man was aboriginally endowed with the ennobling +belief in the existence of an Omnipotent God. On the +contrary there is ample evidence, derived not from hasty +travellers, but from men who have long resided with +savages, that numerous races have existed and still +exist, who have no idea of one or more gods, and who +have no words in their languages to express such an +idea.<a name="FNanchor_95" id="FNanchor_95"></a><a href="#Footnote_95" class="fnanchor">95</a> The question is of course wholly distinct from +that higher one, whether there exists a Creator and +Ruler of the universe; and this has been answered in +the affirmative by the highest intellects that have ever +lived.</p> + +<p>If, however, we include under the term “religion” the +belief in unseen or spiritual agencies, the case is wholly +different; for this belief seems to be almost universal +with the less civilised races. Nor is it difficult to +comprehend how it arose. As soon as the important +faculties of the imagination, wonder, and curiosity, +together with some power of reasoning, had become +partially developed, man would naturally have craved +to understand what was passing around him, and +have vaguely speculated on his own existence. As +<span class='pagenum'><a name="Page_66" id="Page_66">66</a></span>Mr. M’Lennan<a name="FNanchor_96" id="FNanchor_96"></a><a href="#Footnote_96" class="fnanchor">96</a> has remarked, “Some explanation of +the phenomena of life, a man must feign for himself; +and to judge from the universality of it, the simplest +hypothesis, and the first to occur to men, seems to have +been that natural phenomena are ascribable to the presence +in animals, plants, and things, and in the forces +of nature, of such spirits prompting to action as men +are conscious they themselves possess.” It is probable, +as Mr. Tylor has clearly shewn, that dreams may have +first given rise to the notion of spirits; for savages do +not readily distinguish between subjective and objective +impressions. When a savage dreams, the figures which +appear before him are believed to have come from a +distance and to stand over him; or “the soul of the +dreamer goes out on its travels, and comes home with +a remembrance of what it has seen.”<a name="FNanchor_97" id="FNanchor_97"></a><a href="#Footnote_97" class="fnanchor">97</a> But until the +above-named faculties of imagination, curiosity, reason, +&c., had been fairly well developed in the mind of man, +his dreams would not have led him to believe in spirits, +any more than in the case of a dog.</p> + +<p><span class='pagenum'><a name="Page_67" id="Page_67">67</a></span></p><p>The tendency in savages to imagine that natural +objects and agencies are animated by spiritual or living +essences, is perhaps illustrated by a little fact which I +once noticed: my dog, a full-grown and very sensible +animal, was lying on the lawn during a hot and still +day; but at a little distance a slight breeze occasionally +moved an open parasol, which would have been wholly +disregarded by the dog, had any one stood near it. As +it was, every time that the parasol slightly moved, the +dog growled fiercely and barked. He must, I think, +have reasoned to himself in a rapid and unconscious +manner, that movement without any apparent cause +indicated the presence of some strange living agent, and +no stranger had a right to be on his territory.</p> + +<p>The belief in spiritual agencies would easily pass into +the belief in the existence of one or more gods. For +savages would naturally attribute to spirits the same +passions, the same love of vengeance or simplest form +of justice, and the same affections which they themselves +experienced. The Fuegians appear to be in this respect +in an intermediate condition, for when the surgeon on +board the “Beagle” shot some young ducklings as +specimens, York Minster declared in the most solemn +manner, “Oh! Mr. Bynoe, much rain, much snow, blow +much;” and this was evidently a retributive punishment +for wasting human food. So again he related +how, when his brother killed a “wild man,” storms long +raged, much rain and snow fell. Yet we could never +discover that the Fuegians believed in what we should +call a God, or practised any religious rites; and Jemmy +Button, with justifiable pride, stoutly maintained that +there was no devil in his land. This latter assertion is +the more remarkable, as with savages the belief in +bad spirits is far more common than the belief in good +spirits.<span class='pagenum'><a name="Page_68" id="Page_68">68</a></span></p> + +<p>The feeling of religious devotion is a highly complex +one, consisting of love, complete submission to +an exalted and mysterious superior, a strong sense of +dependence,<a name="FNanchor_98" id="FNanchor_98"></a><a href="#Footnote_98" class="fnanchor">98</a> fear, reverence, gratitude, hope for the +future, and perhaps other elements. No being could +experience so complex an emotion until advanced in +his intellectual and moral faculties to at least a moderately +high level. Nevertheless we see some distant +approach to this state of mind, in the deep love of a +dog for his master, associated with complete submission, +some fear, and perhaps other feelings. The behaviour +of a dog when returning to his master after an absence, +and, as I may add, of a monkey to his beloved +keeper, is widely different from that towards their +fellows. In the latter case the transports of joy +appear to be somewhat less, and the sense of equality +is shewn in every action. Professor Braubach<a name="FNanchor_99" id="FNanchor_99"></a><a href="#Footnote_99" class="fnanchor">99</a> goes +so far as to maintain that a dog looks on his master as +on a god.</p> + +<p>The same high mental faculties which first led man +to believe in unseen spiritual agencies, then in fetishism, +polytheism, and ultimately in monotheism, would +infallibly lead him, as long as his reasoning powers +remained poorly developed, to various strange superstitions +and customs. Many of these are terrible to +think of—such as the sacrifice of human beings to a +blood-loving god; the trial of innocent persons by the +ordeal of poison or fire; witchcraft, &c.—yet it is well +occasionally to reflect on these superstitions, for they +shew us what an infinite debt of gratitude we owe to +the improvement of our reason, to science, and our +<span class='pagenum'><a name="Page_69" id="Page_69">69</a></span>accumulated knowledge.<a name="FNanchor_100" id="FNanchor_100"></a><a href="#Footnote_100" class="fnanchor">100</a> As Sir J. Lubbock has well +observed, “it is not too much to say that the horrible +dread of unknown evil hangs like a thick cloud over +savage life, and embitters every pleasure.” These +miserable and indirect consequences of our highest +faculties may be compared with the incidental and +occasional mistakes of the instincts of the lower animals.</p> + +<hr /> +<p><span class='pagenum'><a name="Page_70" id="Page_70">70</a></span></p> +<h3>CHAPTER III.</h3> + +<h4><span class="smcap">Comparison of the Mental Powers of Man and the +Lower Animals</span>—<i>continued</i>.</h4> + +<div class="blockquot"><p>The moral sense—Fundamental proposition—The qualities of social +animals—Origin of sociability—Struggle between opposed instincts—Man +a social animal—The more enduring social instincts +conquer other less persistent instincts—The social virtues alone +regarded by savages—The self-regarding virtues acquired at a +later stage of development—The importance of the judgment +of the members of the same community on conduct—Transmission +of moral tendencies—Summary.</p></div> + +<p>I fully subscribe to the judgment of those writers<a name="FNanchor_101" id="FNanchor_101"></a><a href="#Footnote_101" class="fnanchor">101</a> +who maintain that of all the differences between man +and the lower animals, the moral sense or conscience +is by far the most important. This sense, as Mackintosh<a name="FNanchor_102" id="FNanchor_102"></a><a href="#Footnote_102" class="fnanchor">102</a> +remarks, “has a rightful supremacy over every +other principle of human action;” it is summed up +in that short but imperious word <i>ought</i>, so full of high +significance. It is the most noble of all the attributes +of man, leading him without a moment’s hesitation +to risk his life for that of a fellow-creature; or after +due deliberation, impelled simply by the deep feeling +of right or duty, to sacrifice it in some great cause. +Immanuel Kant exclaims, “Duty! Wondrous thought, +that workest neither by fond insinuation, flattery, nor +by any threat, but merely by holding up thy naked +law in the soul, and so extorting for thyself always +<span class='pagenum'><a name="Page_71" id="Page_71">71</a></span>reverence, if not always obedience; before whom all +appetites are dumb, however secretly they rebel; +whence thy original?”<a name="FNanchor_103" id="FNanchor_103"></a><a href="#Footnote_103" class="fnanchor">103</a></p> + +<p>This great question has been discussed by many +writers<a name="FNanchor_104" id="FNanchor_104"></a><a href="#Footnote_104" class="fnanchor">104</a> of consummate ability; and my sole excuse +for touching on it is the impossibility of here passing +it over, and because, as far as I know, no one has approached +it exclusively from the side of natural history. +The investigation possesses, also, some independent interest, +as an attempt to see how far the study of the +lower animals can throw light on one of the highest +psychical faculties of man.</p> + +<p>The following proposition seems to me in a high +degree probable—namely, that any animal whatever, +endowed with well-marked social instincts,<a name="FNanchor_105" id="FNanchor_105"></a><a href="#Footnote_105" class="fnanchor">105</a> would inevitably +acquire a moral sense or conscience, as soon as +<span class='pagenum'><a name="Page_72" id="Page_72">72</a></span>its intellectual powers had become as well developed, or +nearly as well developed, as in man. For, <i>firstly</i>, the +social instincts lead an animal to take pleasure in +the society of its fellows, to feel a certain amount of +sympathy with them, and to perform various services +for them. The services may be of a definite and evidently +instinctive nature; or there may be only a wish +and readiness, as with most of the higher social animals, +to aid their fellows in certain general ways. But these +feelings and services are by no means extended to all +the individuals of the same species, only to those of +the same association. <i>Secondly</i>, as soon as the mental +faculties had become highly developed, images of all +past actions and motives would be incessantly passing +through the brain of each individual; and that feeling +of dissatisfaction which invariably results, as we shall +hereafter see, from any unsatisfied instinct, would arise, +as often as it was perceived that the enduring and +always present social instinct had yielded to some other +instinct, at the time stronger, but neither enduring in +its nature, nor leaving behind it a very vivid impression. +It is clear that many instinctive desires, such as +that of hunger, are in their nature of short duration; +and after being satisfied are not readily or vividly recalled. +<i>Thirdly</i>, after the power of language had been +acquired and the wishes of the members of the same +community could be distinctly expressed, the common +opinion how each member ought to act for the public +good, would naturally become to a large extent the guide +to action. But the social instincts would still give the +impulse to act for the good of the community, this impulse +being strengthened, directed, and sometimes even +deflected by public opinion, the power of which rests, as +we shall presently see, on instinctive sympathy. <i>Lastly</i>, +habit in the individual would ultimately play a very<span class='pagenum'><a name="Page_73" id="Page_73">73</a></span> +important part in guiding the conduct of each member; +for the social instincts and impulses, like all other instincts, +would be greatly strengthened by habit, as +would obedience to the wishes and judgment of the community. +These several subordinate propositions must +now be discussed; and some of them at considerable +length.</p> + +<p class="tb">It may be well first to premise that I do not wish to +maintain that any strictly social animal, if its intellectual +faculties were to become as active and as highly +developed as in man, would acquire exactly the same +moral sense as ours. In the same manner as various +animals have some sense of beauty, though they admire +widely different objects, so they might have a sense of +right and wrong, though led by it to follow widely different +lines of conduct. If, for instance, to take an extreme +case, men were reared under precisely the same +conditions as hive-bees, there can hardly be a doubt +that our unmarried females would, like the worker-bees, +think it a sacred duty to kill their brothers, and +mothers would strive to kill their fertile daughters; +and no one would think of interfering. Nevertheless +the bee, or any other social animal, would in our supposed +case gain, as it appears to me, some feeling of +right and wrong, or a conscience. For each individual +would have an inward sense of possessing certain +stronger or more enduring instincts, and others less +strong or enduring; so that there would often be a +struggle which impulse should be followed; and satisfaction +or dissatisfaction would be felt, as past impressions +were compared during their incessant passage +through the mind. In this case an inward monitor +would tell the animal that it would have been better to +have followed the one impulse rather than the other. +The one course ought to have been followed: the one<span class='pagenum'><a name="Page_74" id="Page_74">74</a></span> +would have been right and the other wrong; but to +these terms I shall have to recur.</p> + +<p><i>Sociability.</i>—Animals of many kinds are social; we +find even distinct species living together, as with some +American monkeys, and with the united flocks of rooks, +jackdaws, and starlings. Man shows the same feeling +in his strong love for the dog, which the dog returns +with interest. Every one must have noticed how miserable +horses, dogs, sheep, &c. are when separated from +their companions; and what affection at least the two +former kinds show on their reunion. It is curious to +speculate on the feelings of a dog, who will rest peacefully +for hours in a room with his master or any of the +family, without the least notice being taken of him; but +if left for a short time by himself, barks or howls dismally. +We will confine our attention to the higher +social animals, excluding insects, although these aid +each other in many important ways. The most common +service which the higher animals perform for each other, +is the warning each other of danger by means of the +united senses of all. Every sportsman knows, as Dr. +Jaeger remarks,<a name="FNanchor_106" id="FNanchor_106"></a><a href="#Footnote_106" class="fnanchor">106</a> how difficult it is to approach animals +in a herd or troop. Wild horses and cattle do not, I +believe, make any danger-signal; but the attitude of +any one who first discovers an enemy, warns the others. +Rabbits stamp loudly on the ground with their hind-feet +as a signal: sheep and chamois do the same, but with +their fore-feet, uttering likewise a whistle. Many birds +and some mammals post sentinels, which in the case of +seals are said<a name="FNanchor_107" id="FNanchor_107"></a><a href="#Footnote_107" class="fnanchor">107</a> generally to be the females. The leader +of a troop of monkeys acts as the sentinel, and utters +cries expressive both of danger and of safety.<a name="FNanchor_108" id="FNanchor_108"></a><a href="#Footnote_108" class="fnanchor">108</a> Social +<span class='pagenum'><a name="Page_75" id="Page_75">75</a></span>animals perform many little services for each other: +horses nibble, and cows lick each other, on any spot +which itches: monkeys search for each other’s external +parasites; and Brehm states that after a troop of the +<i>Cercopithecus griseo-viridis</i> has rushed through a thorny +brake, each monkey stretches itself on a branch, and +another monkey sitting by “conscientiously” examines +its fur and extracts every thorn or burr.</p> + +<p>Animals also render more important services to each +other: thus wolves and some other beasts of prey hunt +in packs, and aid each other in attacking their victims. +Pelicans fish in concert. The Hamadryas baboons turn +over stones to find insects, &c.; and when they come to +a large one, as many as can stand round, turn it over +together and share the booty. Social animals mutually +defend each other. The males of some ruminants come +to the front when there is danger and defend the herd +with their horns. I shall also in a future chapter give +cases of two young wild bulls attacking an old one in +concert, and of two stallions together trying to drive +away a third stallion from a troop of mares. Brehm +encountered in Abyssinia a great troop of baboons which +were crossing a valley: some had already ascended the +opposite mountain, and some were still in the valley: +the latter were attacked by the dogs, but the old males +immediately hurried down from the rocks, and with +mouths widely opened roared so fearfully, that the dogs +precipitately retreated. They were again encouraged +to the attack; but by this time all the baboons had reascended +the heights, excepting a young one, about six +<span class='pagenum'><a name="Page_76" id="Page_76">76</a></span>months old, who, loudly calling for aid, climbed on a +block of rock and was surrounded. Now one of the +largest males, a true hero, came down again from the +mountain, slowly went to the young one, coaxed him, +and triumphantly led him away—the dogs being too +much astonished to make an attack. I cannot resist +giving another scene which was witnessed by this same +naturalist; an eagle seized a young Cercopithecus, +which, by clinging to a branch, was not at once carried +off; it cried loudly for assistance, upon which the other +members of the troop with much uproar rushed to the +rescue, surrounded the eagle, and pulled out so many +feathers, that he no longer thought of his prey, but only +how to escape. This eagle, as Brehm remarks, assuredly +would never again attack a monkey in a troop.</p> + +<p>It is certain that associated animals have a feeling of +love for each other which is not felt by adult and non-social +animals. How far in most cases they actually +sympathise with each other’s pains and pleasures is +more doubtful, especially with respect to the latter. +Mr. Buxton, however, who had excellent means of +observation,<a name="FNanchor_109" id="FNanchor_109"></a><a href="#Footnote_109" class="fnanchor">109</a> states that his macaws, which lived free in +Norfolk, took “an extravagant interest” in a pair +with a nest, and whenever the female left it, she was +surrounded by a troop “screaming horrible acclamations +in her honour.” It is often difficult to judge +whether animals have any feeling for each other’s +sufferings. Who can say what cows feel, when they +surround and stare intently on a dying or dead +companion? That animals sometimes are far from +feeling any sympathy is too certain; for they will expel +a wounded animal from the herd, or gore or worry +it to death. This is almost the blackest fact in natural +<span class='pagenum'><a name="Page_77" id="Page_77">77</a></span>history, unless indeed the explanation which has been +suggested is true, that their instinct or reason leads them +to expel an injured companion, lest beasts of prey, +including man, should be tempted to follow the troop. +In this case their conduct is not much worse than that +of the North American Indians who leave their feeble +comrades to perish on the plains, or the Feegeans, who, +when their parents get old or fall ill, bury them alive.<a name="FNanchor_110" id="FNanchor_110"></a><a href="#Footnote_110" class="fnanchor">110</a></p> + +<p>Many animals, however, certainly sympathise with +each other’s distress or danger. This is the case even +with birds; Capt. Stansbury<a name="FNanchor_111" id="FNanchor_111"></a><a href="#Footnote_111" class="fnanchor">111</a> found on a salt lake in +Utah an old and completely blind pelican, which was +very fat, and must have been long and well fed by his +companions. Mr. Blyth, as he informs me, saw Indian +crows feeding two or three of their companions which +were blind; and I have heard of an analogous case +with the domestic cock. We may, if we choose, call +these actions instinctive; but such cases are much too +rare for the development of any special instinct.<a name="FNanchor_112" id="FNanchor_112"></a><a href="#Footnote_112" class="fnanchor">112</a> I +have myself seen a dog, who never passed a great +friend of his, a cat which lay sick in a basket, without +giving her a few licks with his tongue, the surest +sign of kind feeling in a dog.</p> + +<p>It must be called sympathy that leads a courageous +dog to fly at any one who strikes his master, as he +certainly will. I saw a person pretending to beat a +lady who had a very timid little dog on her lap, and +the trial had never before been made. The little crea<span class='pagenum'><a name="Page_78" id="Page_78">78</a></span>ture +instantly jumped away, but after the pretended +beating was over, it was really pathetic to see how perseveringly +he tried to lick his mistress’s face and comfort +her. Brehm<a name="FNanchor_113" id="FNanchor_113"></a><a href="#Footnote_113" class="fnanchor">113</a> states that when a baboon in confinement +was pursued to be punished, the others tried +to protect him. It must have been sympathy in the +cases above given which led the baboons and Cercopitheci +to defend their young comrades from the dogs +and the eagle. I will give only one other instance of +sympathetic and heroic conduct in a little American +monkey. Several years ago a keeper at the Zoological +Gardens, showed me some deep and scarcely healed +wounds on the nape of his neck, inflicted on him whilst +kneeling on the floor by a fierce baboon. The little +American monkey, who was a warm friend of this +keeper, lived in the same large compartment, and was +dreadfully afraid of the great baboon. Nevertheless, as +soon as he saw his friend the keeper in peril, he rushed +to the rescue, and by screams and bites so distracted the +baboon that the man was able to escape, after running +great risk, as the surgeon who attended him thought, +of his life.</p> + +<p>Besides love and sympathy, animals exhibit other +qualities which in us would be called moral; and I agree +with Agassiz<a name="FNanchor_114" id="FNanchor_114"></a><a href="#Footnote_114" class="fnanchor">114</a> that dogs possess something very like a +conscience. They certainly possess some power of self-command, +and this does not appear to be wholly the +result of fear. As Braubach<a name="FNanchor_115" id="FNanchor_115"></a><a href="#Footnote_115" class="fnanchor">115</a> remarks, a dog will +refrain from stealing food in the absence of his master. +Dogs have long been accepted as the very type of +fidelity and obedience. All animals living in a body +which defend each other or attack their enemies +<span class='pagenum'><a name="Page_79" id="Page_79">79</a></span>in concert, must be in some degree faithful to each +other; and those that follow a leader must be in +some degree obedient. When the baboons in Abyssinia<a name="FNanchor_116" id="FNanchor_116"></a><a href="#Footnote_116" class="fnanchor">116</a> +plunder a garden, they silently follow their +leader; and if an imprudent young animal makes a +noise, he receives a slap from the others to teach him +silence and obedience; but as soon as they are sure +that there is no danger, all show their joy by much +clamour.</p> + +<p>With respect to the impulse which leads certain +animals to associate together, and to aid each other in +many ways, we may infer that in most cases they are +impelled by the same sense of satisfaction or pleasure +which they experience in performing other instinctive +actions; or by the same sense of dissatisfaction, as in +other cases of prevented instinctive actions. We see +this in innumerable instances, and it is illustrated in +a striking manner by the acquired instincts of our +domesticated animals; thus a young shepherd-dog +delights in driving and running round a flock of sheep, +but not in worrying them; a young foxhound delights +in hunting a fox, whilst some other kinds of dogs +as I have witnessed, utterly disregard foxes. What a +strong feeling of inward satisfaction must impel a bird, +so full of activity, to brood day after day over her eggs. +Migratory birds are miserable if prevented from migrating, +and perhaps they enjoy starting on their long +flight. Some few instincts are determined solely by +painful feelings, as by fear, which leads to self-preservation, +or is specially directed against certain enemies. +No one, I presume, can analyse the sensations of +pleasure or pain. In many cases, however, it is probable +that instincts are persistently followed from the +<span class='pagenum'><a name="Page_80" id="Page_80">80</a></span>mere force of inheritance, without the stimulus of either +pleasure or pain. A young pointer, when it first scents +game, apparently cannot help pointing. A squirrel in +a cage who pats the nuts which it cannot eat, as if to +bury them in the ground, can hardly be thought to act +thus either from pleasure or pain. Hence the common +assumption that men must be impelled to every action +by experiencing some pleasure or pain may be erroneous. +Although a habit may be blindly and implicitly +followed, independently of any pleasure or pain felt at +the moment, yet if it be forcibly and abruptly checked, +a vague sense of dissatisfaction is generally experienced; +and this is especially true in regard to persons +of feeble intellect.</p> + +<p>It has often been assumed that animals were in the +first place rendered social, and that they feel as a consequence +uncomfortable when separated from each other, +and comfortable whilst together; but it is a more probable +view that these sensations were first developed, in +order that those animals which would profit by living +in society, should be induced to live together. In the +same manner as the sense of hunger and the pleasure of +eating were, no doubt, first acquired in order to induce +animals to eat. The feeling of pleasure from society +is probably an extension of the parental or filial affections; +and this extension may be in chief part attributed +to natural selection, but perhaps in part to mere habit. +For with those animals which were benefited by living +in close association, the individuals which took the +greatest pleasure in society would best escape various +dangers; whilst those that cared least for their comrades +and lived solitary would perish in greater numbers. +With respect to the origin of the parental and filial +affections, which apparently lie at the basis of the +social affections, it is hopeless to speculate; but we<span class='pagenum'><a name="Page_81" id="Page_81">81</a></span> +may infer that they have been to a large extent gained +through natural selection. So it has almost certainly +been with the unusual and opposite feeling of hatred +between the nearest relations, as with the worker-bees +which kill their brother-drones, and with the queen-bees +which kill their daughter-queens; the desire to destroy, +instead of loving, their nearest relations having been +here of service to the community.</p> + +<p>The all-important emotion of sympathy is distinct +from that of love. A mother may passionately love her +sleeping and passive infant, but she can then hardly be +said to feel sympathy for it. The love of a man for +his dog is distinct from sympathy, and so is that of a +dog for his master. Adam Smith formerly argued, as +has Mr. Bain recently, that the basis of sympathy lies +in our strong retentiveness of former states of pain or +pleasure. Hence, “the sight of another person enduring +hunger, cold, fatigue, revives in us some recollection +of these states, which are painful even in idea.” We +are thus impelled to relieve the sufferings of another, +in order that our own painful feelings may be at the +same time relieved. In like manner we are led to +participate in the pleasures of others.<a name="FNanchor_117" id="FNanchor_117"></a><a href="#Footnote_117" class="fnanchor">117</a> But I cannot +see how this view explains the fact that sympathy +is excited in an immeasurably stronger degree by a +beloved than by an indifferent person. The mere +<span class='pagenum'><a name="Page_82" id="Page_82">82</a></span>sight of suffering, independently of love, would suffice +to call up in us vivid recollections and associations. +Sympathy may at first have originated in the manner +above suggested; but it seems now to have become +an instinct, which is especially directed towards beloved +objects, in the same manner as fear with animals +is especially directed against certain enemies. As +sympathy is thus directed, the mutual love of the +members of the same community will extend its limits. +No doubt a tiger or lion feels sympathy for the sufferings +of its own young, but not for any other animal. +With strictly social animals the feeling will be more +or less extended to all the associated members, as we +know to be the case. With mankind selfishness, experience, +and imitation probably add, as Mr. Bain has +shewn, to the power of sympathy; for we are led +by the hope of receiving good in return to perform +acts of sympathetic kindness to others; and there can +be no doubt that the feeling of sympathy is much +strengthened by habit. In however complex a manner +this feeling may have originated, as it is one of high +importance to all those animals which aid and defend +each other, it will have been increased, through natural +selection; for those communities, which included the +greatest number of the most sympathetic members, +would flourish best and rear the greatest number of +offspring.</p> + +<p>In many cases it is impossible to decide whether +certain social instincts have been acquired through +natural selection, or are the indirect result of other +instincts and faculties, such as sympathy, reason, experience, +and a tendency to imitation; or again, whether +they are simply the result of long-continued habit. +So remarkable an instinct as the placing sentinels to +warn the community of danger, can hardly have been<span class='pagenum'><a name="Page_83" id="Page_83">83</a></span> +the indirect result of any other faculty; it must therefore +have been directly acquired. On the other hand, +the habit followed by the males of some social animals, +of defending the community and of attacking their +enemies or their prey in concert, may perhaps have +originated from mutual sympathy; but courage, and +in most cases strength, must have been previously +acquired, probably through natural selection.</p> + +<p>Of the various instincts and habits, some are much +stronger than others, that is, some either give more +pleasure in their performance and more distress in their +prevention than others; or, which is probably quite as +important, they are more persistently followed through +inheritance without exciting any special feeling of pleasure +or pain. We are ourselves conscious that some +habits are much more difficult to cure or change than +others. Hence a struggle may often be observed in +animals between different instincts, or between an +instinct and some habitual disposition; as when a dog +rushes after a hare, is rebuked, pauses, hesitates, pursues +again or returns ashamed to his master; or as between +the love of a female dog for her young puppies and for +her master, for she may be seen to slink away to them, +as if half ashamed of not accompanying her master. +But the most curious instance known to me of one +instinct conquering another, is the migratory instinct +conquering the maternal instinct. The former is wonderfully +strong; a confined bird will at the proper +season beat her breast against the wires of her cage, until +it is bare and bloody. It causes young salmon to leap +out of the fresh water, where they could still continue to +live, and thus unintentionally to commit suicide. Every +one knows how strong the maternal instinct is, leading +even timid birds to face great danger, though with +hesitation and in opposition to the instinct of self<span class='pagenum'><a name="Page_84" id="Page_84">84</a></span>preservation. +Nevertheless the migratory instinct is so +powerful that late in the autumn swallows and house-martins +frequently desert their tender young, leaving +them to perish miserably in their nests.<a name="FNanchor_118" id="FNanchor_118"></a><a href="#Footnote_118" class="fnanchor">118</a></p> + +<p class="tb">We can perceive that an instinctive impulse, if it be in +any way more beneficial to a species than some other or +opposed instinct, would be rendered the more potent of +the two through natural selection; for the individuals +which had it most strongly developed would survive in +larger numbers. Whether this is the case with the +migratory in comparison with the maternal instinct, +may well be doubted. The great persistence or steady +action of the former at certain seasons of the year +during the whole day, may give it for a time paramount +force.</p> + +<p><i>Man a social animal.</i>—Most persons admit that man +is a social being. We see this in his dislike of solitude, +and in his wish for society beyond that of his own +family. Solitary confinement is one of the severest +punishments which can be inflicted. Some authors suppose +that man primevally lived in single families; but +at the present day, though single families, or only two +or three together, roam the solitudes of some savage +lands, they are always, as far as I can discover, friendly +with other families inhabiting the same district. Such +families occasionally meet in council, and they unite +<span class='pagenum'><a name="Page_85" id="Page_85">85</a></span>for their common defence. It is no argument against +savage man being a social animal, that the tribes inhabiting +adjacent districts are almost always at war +with each other; for the social instincts never extend +to all the individuals of the same species. Judging +from the analogy of the greater number of the Quadrumana, +it is probable that the early ape-like progenitors +of man were likewise social; but this is not of +much importance for us. Although man, as he now +exists, has few special instincts, having lost any which +his early progenitors may have possessed, this is no +reason why he should not have retained from an extremely +remote period some degree of instinctive love +and sympathy for his fellows. We are indeed all conscious +that we do possess such sympathetic feelings;<a name="FNanchor_119" id="FNanchor_119"></a><a href="#Footnote_119" class="fnanchor">119</a> +but our consciousness does not tell us whether they are +instinctive, having originated long ago in the same +manner as with the lower animals, or whether they have +been acquired by each of us during our early years. +As man is a social animal, it is also probable that he +would inherit a tendency to be faithful to his comrades, +for this quality is common to most social animals. He +would in like manner possess some capacity for self-command, +and perhaps of obedience to the leader of +the community. He would from an inherited tendency +still be willing to defend, in concert with others, his +fellow-men, and would be ready to aid them in any +way which did not too greatly interfere with his own +welfare or his own strong desires.</p> + +<p class="tb">The social animals which stand at the bottom of the +<span class='pagenum'><a name="Page_86" id="Page_86">86</a></span>scale are guided almost exclusively, and those which +stand higher in the scale are largely guided, in the aid +which they give to the members of the same community, +by special instincts; but they are likewise in part impelled +by mutual love and sympathy, assisted apparently +by some amount of reason. Although man, as just +remarked, has no special instincts to tell him how to aid +his fellow-men, he still has the impulse, and with his +improved intellectual faculties would naturally be much +guided in this respect by reason and experience. Instinctive +sympathy would, also, cause him to value highly +the approbation of his fellow-men; for, as Mr. Bain has +clearly shewn,<a name="FNanchor_120" id="FNanchor_120"></a><a href="#Footnote_120" class="fnanchor">120</a> the love of praise and the strong feeling +of glory, and the still stronger horror of scorn and infamy, +“are due to the workings of sympathy.” Consequently +man would be greatly influenced by the wishes, +approbation, and blame of his fellow-men, as expressed +by their gestures and language. Thus the social instincts, +which must have been acquired by man in a +very rude state, and probably even by his early ape-like +progenitors, still give the impulse to many of his best +actions; but his actions are largely determined by the +expressed wishes and judgment of his fellow-men, and +unfortunately still oftener by his own strong, selfish +desires. But as the feelings of love and sympathy and +the power of self-command become strengthened by +habit, and as the power of reasoning becomes clearer so +that man can appreciate the justice of the judgments of +his fellow-men, he will feel himself impelled, independently +of any pleasure or pain felt at the moment, to +certain lines of conduct. He may then say, I am the +supreme judge of my own conduct, and in the words of +Kant, I will not in my own person violate the dignity +of humanity.</p> + +<p><span class='pagenum'><a name="Page_87" id="Page_87">87</a></span><i>The more enduring Social Instincts conquer the less +Persistent Instincts.</i>—We have, however, not as yet considered +the main point, on which the whole question of +the moral sense hinges. Why should a man feel that +he ought to obey one instinctive desire rather than +another? Why does he bitterly regret if he has yielded +to the strong sense of self-preservation, and has not +risked his life to save that of a fellow-creature; or why +does he regret having stolen food from severe hunger?</p> + +<p>It is evident in the first place, that with mankind the +instinctive impulses have different degrees of strength; +a young and timid mother urged by the maternal instinct +will, without a moment’s hesitation, run the +greatest danger for her infant, but not for a mere fellow-creature. +Many a man, or even boy, who never +before risked his life for another, but in whom courage +and sympathy were well developed, has, disregarding the +instinct of self-preservation, instantaneously plunged +into a torrent to save a drowning fellow-creature. In +this case man is impelled by the same instinctive motive, +which caused the heroic little American monkey, +formerly described, to attack the great and dreaded +baboon, to save his keeper. Such actions as the above +appear to be the simple result of the greater strength of +the social or maternal instincts than of any other instinct +or motive; for they are performed too instantaneously +for reflection, or for the sensation of pleasure or pain; +though if prevented distress would be caused.</p> + +<p>I am aware that some persons maintain that actions +performed impulsively, as in the above cases, do not +come under the dominion of the moral sense, and +cannot be called moral. They confine this term to +actions done deliberately, after a victory over opposing +desires, or to actions prompted by some lofty motive. +But it appears scarcely possible to draw any clear line<span class='pagenum'><a name="Page_88" id="Page_88">88</a></span> +of distinction of this kind; though the distinction may +be real. As far as exalted motives are concerned, many +instances have been recorded of barbarians, destitute of +any feeling of general benevolence towards mankind, +and not guided by any religious motive, who have deliberately +as prisoners sacrificed their lives,<a name="FNanchor_121" id="FNanchor_121"></a><a href="#Footnote_121" class="fnanchor">121</a> rather than +betray their comrades; and surely their conduct ought +to be considered as moral. As far as deliberation and +the victory over opposing motives are concerned, animals +may be seen doubting between opposed instincts, +as in rescuing their offspring or comrades from danger; +yet their actions, though done for the good of +others, are not called moral. Moreover, an action +repeatedly performed by us, will at last be done without +deliberation or hesitation, and can then hardly be +distinguished from an instinct; yet surely no one will +pretend that an action thus done ceases to be moral. +On the contrary, we all feel that an act cannot be +considered as perfect, or as performed in the most +noble manner, unless it be done impulsively, without +deliberation or effort, in the same manner as by a man +in whom the requisite qualities are innate. He +who is forced to overcome his fear or want of sympathy +before he acts, deserves, however, in one way +higher credit than the man whose innate disposition +leads him to a good act without effort. As we cannot +distinguish between motives, we rank all actions of a +certain class as moral, when they are performed by a +moral being. A moral being is one who is capable of +comparing his past and future actions or motives, and +of approving or disapproving of them. We have no +reason to suppose that any of the lower animals have +<span class='pagenum'><a name="Page_89" id="Page_89">89</a></span>this capacity; therefore when a monkey faces danger +to rescue its comrade, or takes charge of an orphan-monkey, +we do not call its conduct moral. But in the +case of man, who alone can with certainty be ranked as +a moral being, actions of a certain class are called moral, +whether performed deliberately after a struggle with +opposing motives, or from the effects of slowly-gained +habit, or impulsively through instinct.</p> + +<p>But to return to our more immediate subject; although +some instincts are more powerful than others, +thus leading to corresponding actions, yet it cannot +be maintained that the social instincts are ordinarily +stronger in man, or have become stronger through +long-continued habit, than the instincts, for instance, +of self-preservation, hunger, lust, vengeance, &c. Why +then does man regret, even though he may endeavour +to banish any such regret, that he has followed the +one natural impulse, rather than the other; and why +does he further feel that he ought to regret his conduct? +Man in this respect differs profoundly from the lower +animals. Nevertheless we can, I think, see with some +degree of clearness the reason of this difference.</p> + +<p>Man, from the activity of his mental faculties, cannot +avoid reflection: past impressions and images are incessantly +passing through his mind with distinctness. +Now with those animals which live permanently in a +body, the social instincts are ever present and persistent. +Such animals are always ready to utter the +danger-signal, to defend the community, and to give +aid to their fellows in accordance with their habits; +they feel at all times, without the stimulus of any +special passion or desire, some degree of love and +sympathy for them; they are unhappy if long separated +from them, and always happy to be in their company. +So it is with ourselves. A man who possessed no trace<span class='pagenum'><a name="Page_90" id="Page_90">90</a></span> +of such feelings would be an unnatural monster. On +the other hand, the desire to satisfy hunger, or any +passion, such as vengeance, is in its nature temporary, +and can for a time be fully satisfied. Nor is it easy, +perhaps hardly possible, to call up with complete vividness +the feeling, for instance, of hunger; nor indeed, as +has often been remarked, of any suffering. The instinct +of self-preservation is not felt except in the presence +of danger; and many a coward has thought himself +brave until he has met his enemy face to face. +The wish for another man’s property is perhaps as +persistent a desire as any that can be named; but even +in this case the satisfaction of actual possession is generally +a weaker feeling than the desire: many a thief, if +not an habitual one, after success has wondered why he +stole some article.</p> + +<p>Thus, as man cannot prevent old impressions continually +repassing through his mind, he will be compelled +to compare the weaker impressions of, for instance, +past hunger, or of vengeance satisfied or danger +avoided at the cost of other men, with the instinct of +sympathy and good-will to his fellows, which is still present +and ever in some degree active in his mind. He +will then feel in his imagination that a stronger instinct +has yielded to one which now seems comparatively +weak; and then that sense of dissatisfaction will inevitably +be felt with which man is endowed, like every +other animal, in order that his instincts may be obeyed. +The case before given, of the swallow, affords an illustration, +though of a reversed nature, of a temporary +though for the time strongly persistent instinct conquering +another instinct which is usually dominant over +all others. At the proper season these birds seem all +day long to be impressed with the desire to migrate; +their habits change; they become restless, are noisy,<span class='pagenum'><a name="Page_91" id="Page_91">91</a></span> +and congregate in flocks. Whilst the mother-bird is +feeding or brooding over her nestlings, the maternal +instinct is probably stronger than the migratory; but +the instinct which is more persistent gains the victory, +and at last, at a moment when her young ones are not +in sight, she takes flight and deserts them. When +arrived at the end of her long journey, and the migratory +instinct ceases to act, what an agony of remorse +each bird would feel, if, from being endowed with great +mental activity, she could not prevent the image continually +passing before her mind of her young ones +perishing in the bleak north from cold and hunger.</p> + +<p>At the moment of action, man will no doubt be apt +to follow the stronger impulse; and though this may +occasionally prompt him to the noblest deeds, it will +far more commonly lead him to gratify his own desires +at the expense of other men. But after their gratification, +when past and weaker impressions are contrasted +with the ever-enduring social instincts, retribution +will surely come. Man will then feel dissatisfied +with himself, and will resolve with more or less +force to act differently for the future. This is conscience; +for conscience looks backwards and judges past +actions, inducing that kind of dissatisfaction, which if +weak we call regret, and if severe remorse.</p> + +<p>These sensations are, no doubt, different from those +experienced when other instincts or desires are left +unsatisfied; but every unsatisfied instinct has its own +proper prompting sensation, as we recognise with hunger, +thirst, &c. Man thus prompted, will through long +habit acquire such perfect self-command, that his desires +and passions will at last instantly yield to his social +sympathies, and there will no longer be a struggle +between them. The still hungry, or the still revengeful +man will not think of stealing food, or of wreaking his<span class='pagenum'><a name="Page_92" id="Page_92">92</a></span> +vengeance. It is possible, or, as we shall hereafter +see, even probable, that the habit of self-command +may, like other habits, be inherited. Thus at last +man comes to feel, through acquired and perhaps inherited +habit, that it is best for him to obey his more +persistent instincts. The imperious word <i>ought</i> seems +merely to imply the consciousness of the existence of a +persistent instinct, either innate or partly acquired, +serving him as a guide, though liable to be disobeyed. +We hardly use the word <i>ought</i> in a metaphorical sense, +when we say hounds ought to hunt, pointers to point, +and retrievers to retrieve their game. If they fail thus +to act, they fail in their duty and act wrongly.</p> + +<p>If any desire or instinct, leading to an action opposed +to the good of others, still appears to a man, when recalled +to mind, as strong as, or stronger than, his social +instinct, he will feel no keen regret at having followed +it; but he will be conscious that if his conduct were +known to his fellows, it would meet with their disapprobation; +and few are so destitute of sympathy as not +to feel discomfort when this is realised. If he has no +such sympathy, and if his desires leading to bad actions +are at the time strong, and when recalled are not overmastered +by the persistent social instincts, then he is +essentially a bad man;<a name="FNanchor_122" id="FNanchor_122"></a><a href="#Footnote_122" class="fnanchor">122</a> and the sole restraining motive +left is the fear of punishment, and the conviction that +in the long run it would be best for his own selfish +interests to regard the good of others rather than his +own.</p> + +<p class="tb">It is obvious that every one may with an easy conscience +gratify his own desires, if they do not interfere +<span class='pagenum'><a name="Page_93" id="Page_93">93</a></span>with his social instincts, that is with the good of others; +but in order to be quite free from self-reproach, or at +least of anxiety, it is almost necessary for him to avoid +the disapprobation, whether reasonable or not, of his +fellow men. Nor must he break through the fixed habits +of his life, especially if these are supported by reason; +for if he does, he will assuredly feel dissatisfaction. +He must likewise avoid the reprobation of the one +God or gods, in whom according to his knowledge or +superstition he may believe; but in this case the additional +fear of divine punishment often supervenes.</p> + +<p><i>The strictly Social Virtues at first alone regarded.</i>—The +above view of the first origin and nature of the moral +sense, which tells us what we ought to do, and of +the conscience which reproves us if we disobey it, +accords well with what we see of the early and undeveloped +condition of this faculty in mankind. The +virtues which must be practised, at least generally, by +rude men, so that they may associate in a body, are +those which are still recognised as the most important. +But they are practised almost exclusively in relation to +the men of the same tribe; and their opposites are not +regarded as crimes in relation to the men of other tribes. +No tribe could hold together if murder, robbery, treachery, +&c., were common; consequently such crimes +within the limits of the same tribe “are branded +with everlasting infamy;”<a name="FNanchor_123" id="FNanchor_123"></a><a href="#Footnote_123" class="fnanchor">123</a> but excite no such sentiment +beyond these limits. A North-American Indian +is well pleased with himself, and is honoured by others, +when he scalps a man of another tribe; and a Dyak +<span class='pagenum'><a name="Page_94" id="Page_94">94</a></span>cuts off the head of an unoffending person and dries it +as a trophy. The murder of infants has prevailed on +the largest scale throughout the world,<a name="FNanchor_124" id="FNanchor_124"></a><a href="#Footnote_124" class="fnanchor">124</a> and has met +with no reproach; but infanticide, especially of females, +has been thought to be good for the tribe, or at least +not injurious. Suicide during former times was not +generally considered as a crime,<a name="FNanchor_125" id="FNanchor_125"></a><a href="#Footnote_125" class="fnanchor">125</a> but rather from the +courage displayed as an honourable act; and it is still +largely practised by some semi-civilised nations without +reproach, for the loss to a nation of a single individual +is not felt: whatever the explanation may be, suicide, +as I hear from Sir J. Lubbock, is rarely practised by the +lowest barbarians. It has been recorded that an Indian +Thug conscientiously regretted that he had not strangled +and robbed as many travellers as did his father before +him. In a rude state of civilisation the robbery of +strangers is, indeed, generally considered as honourable.</p> + +<p>The great sin of Slavery has been almost universal, and +slaves have often been treated in an infamous manner. +As barbarians do not regard the opinion of their women, +wives are commonly treated like slaves. Most savages +are utterly indifferent to the sufferings of strangers, or +even delight in witnessing them. It is well known that +the women and children of the North-American Indians +aided in torturing their enemies. Some savages take a +horrid pleasure in cruelty to animals,<a name="FNanchor_126" id="FNanchor_126"></a><a href="#Footnote_126" class="fnanchor">126</a> and humanity +with them is an unknown virtue. Nevertheless, feelings +of sympathy and kindness are common, especially +<span class='pagenum'><a name="Page_95" id="Page_95">95</a></span>during sickness, between the members of the same tribe, +and are sometimes extended beyond the limits of the +tribe. Mungo Park’s touching account of the kindness +of the negro women of the interior to him is well known. +Many instances could be given of the noble fidelity of +savages towards each other, but not to strangers; +common experience justifies the maxim of the Spaniard, +“Never, never trust an Indian.” There cannot be +fidelity without truth; and this fundamental virtue is +not rare between the members of the same tribe: thus +Mungo Park heard the negro women teaching their +young children to love the truth. This, again, is one of +the virtues which becomes so deeply rooted in the mind +that it is sometimes practised by savages even at a high +cost, towards strangers; but to lie to your enemy has +rarely been thought a sin, as the history of modern +diplomacy too plainly shews. As soon as a tribe has +a recognised leader, disobedience becomes a crime, and +even abject submission is looked at as a sacred virtue.</p> + +<p>As during rude times no man can be useful or faithful +to his tribe without courage, this quality has universally +been placed in the highest rank; and although, in +civilised countries, a good, yet timid, man may be far +more useful to the community than a brave one, we +cannot help instinctively honouring the latter above +a coward, however benevolent. Prudence, on the other +hand, which does not concern the welfare of others, +though a very useful virtue, has never been highly +esteemed. As no man can practise the virtues necessary +for the welfare of his tribe without self-sacrifice, self-command, +and the power of endurance, these qualities +have been at all times highly and most justly valued. +The American savage voluntarily submits without +a groan to the most horrid tortures to prove and +strengthen his fortitude and courage; and we cannot<span class='pagenum'><a name="Page_96" id="Page_96">96</a></span> +help admiring him, or even an Indian Fakir, who, from +a foolish religious motive, swings suspended by a hook +buried in his flesh.</p> + +<p>The other self-regarding virtues, which do not obviously, +though they may really, affect the welfare of +the tribe, have never been esteemed by savages, though +now highly appreciated by civilised nations. The +greatest intemperance with savages is no reproach. +Their utter licentiousness, not to mention unnatural +crimes, is something astounding.<a name="FNanchor_127" id="FNanchor_127"></a><a href="#Footnote_127" class="fnanchor">127</a> As soon, however, as +marriage, whether polygamous or monogamous, becomes +common, jealousy will lead to the inculcation of female +virtue; and this being honoured will tend to spread to the +unmarried females. How slowly it spreads to the male +sex we see at the present day. Chastity eminently +requires self-command; therefore it has been honoured +from a very early period in the moral history of civilised +man. As a consequence of this, the senseless practice +of celibacy has been ranked from a remote period as +a virtue.<a name="FNanchor_128" id="FNanchor_128"></a><a href="#Footnote_128" class="fnanchor">128</a> The hatred of indecency, which appears to +us so natural as to be thought innate, and which is so +valuable an aid to chastity, is a modern virtue, appertaining +exclusively, as Sir G. Staunton remarks,<a name="FNanchor_129" id="FNanchor_129"></a><a href="#Footnote_129" class="fnanchor">129</a> to +civilised life. This is shewn by the ancient religious +rites of various nations, by the drawings on the walls of +Pompeii, and by the practices of many savages.</p> + +<p>We have now seen that actions are regarded by +savages, and were probably so regarded by primeval +man, as good or bad, solely as they affect in an obvious +manner the welfare of the tribe,—not that of the +species, nor that of man as an individual member of the +<span class='pagenum'><a name="Page_97" id="Page_97">97</a></span>tribe. This conclusion agrees well with the belief that +the so-called moral sense is aboriginally derived from +the social instincts, for both relate at first exclusively to +the community. The chief causes of the low morality +of savages, as judged by our standard, are, firstly, the +confinement of sympathy to the same tribe. Secondly, +insufficient powers of reasoning, so that the bearing of +many virtues, especially of the self-regarding virtues, on +the general welfare of the tribe is not recognised. +Savages, for instance, fail to trace the multiplied evils +consequent on a want of temperance, chastity, &c. +And, thirdly, weak power of self-command; for this +power has not been strengthened through long-continued, +perhaps inherited, habit, instruction and religion.</p> + +<p class="tb">I have entered into the above details on the immorality +of savages,<a name="FNanchor_130" id="FNanchor_130"></a><a href="#Footnote_130" class="fnanchor">130</a> because some authors have recently +taken a high view of their moral nature, or have attributed +most of their crimes to mistaken benevolence.<a name="FNanchor_131" id="FNanchor_131"></a><a href="#Footnote_131" class="fnanchor">131</a> +These authors appear to rest their conclusion on savages +possessing, as they undoubtedly do possess, and often +in a high degree, those virtues which are serviceable, +or even necessary, for the existence of a tribal community.</p> + +<p><i>Concluding Remarks.</i>—Philosophers of the derivative<a name="FNanchor_132" id="FNanchor_132"></a><a href="#Footnote_132" class="fnanchor">132</a> +school of morals formerly assumed that the foundation +of morality lay in a form of Selfishness; but more +recently in the “Greatest Happiness principle.” According +to the view given above, the moral sense is +<span class='pagenum'><a name="Page_98" id="Page_98">98</a></span>fundamentally identical with the social instincts; and +in the case of the lower animals it would be absurd to +speak of these instincts as having been developed from +selfishness, or for the happiness of the community. +They have, however, certainly been developed for the +general good of the community. The term, general +good, may be defined as the means by which the greatest +possible number of individuals can be reared in +full vigour and health, with all their faculties perfect, +under the conditions to which they are exposed. As +the social instincts both of man and the lower animals +have no doubt been developed by the same steps, it +would be advisable, if found practicable, to use the +same definition in both cases, and to take as the test +of morality, the general good or welfare of the community, +rather than the general happiness; but this +definition would perhaps require some limitation on +account of political ethics.</p> + +<p>When a man risks his life to save that of a fellow-creature, +it seems more appropriate to say that he acts +for the general good or welfare, rather than for the +general happiness of mankind. No doubt the welfare +and the happiness of the individual usually coincide; +and a contented, happy tribe will flourish better than +one that is discontented and unhappy. We have seen +that at an early period in the history of man, the expressed +wishes of the community will have naturally +influenced to a large extent the conduct of each member; +and as all wish for happiness, the “greatest happiness +principle” will have become a most important +secondary guide and object; the social instincts, including +sympathy, always serving as the primary impulse +and guide. Thus the reproach of laying the foundation +of the most noble part of our nature in the base principle +of selfishness is removed; unless indeed the satis<span class='pagenum'><a name="Page_99" id="Page_99">99</a></span>faction +which every animal feels when it follows its +proper instincts, and the dissatisfaction felt when prevented, +be called selfish.</p> + +<p>The expression of the wishes and judgment of the +members of the same community, at first by oral and +afterwards by written language, serves, as just remarked, +as a most important secondary guide of +conduct, in aid of the social instincts, but sometimes +in opposition to them. This latter fact is well exemplified +by the <i>Law of Honour</i>, that is the law of the +opinion of our equals, and not of all our countrymen. +The breach of this law, even when the breach +is known to be strictly accordant with true morality, +has caused many a man more agony than a real +crime. We recognise the same influence in the burning +sense of shame which most of us have felt even +after the interval of years, when calling to mind some +accidental breach of a trifling though fixed rule of etiquette. +The judgment of the community will generally +be guided by some rude experience of what is best in +the long run for all the members; but this judgment +will not rarely err from ignorance and from weak powers +of reasoning. Hence the strangest customs and superstitions, +in complete opposition to the true welfare and +happiness of mankind, have become all-powerful throughout +the world. We see this in the horror felt by a +Hindoo who breaks his caste, in the shame of a Mahometan +woman who exposes her face, and in innumerable +other instances. It would be difficult to distinguish +between the remorse felt by a Hindoo who has eaten +unclean food, from that felt after committing a theft; +but the former would probably be the more severe.</p> + +<p>How so many absurd rules of conduct, as well as so +many absurd religious beliefs, have originated we do +not know; nor how it is that they have become, in all<span class='pagenum'><a name="Page_100" id="Page_100">100</a></span> +quarters of the world, so deeply impressed on the mind +of men; but it is worthy of remark that a belief constantly +inculcated during the early years of life, whilst +the brain is impressible, appears to acquire almost the +nature of an instinct; and the very essence of an instinct +is that it is followed independently of reason. +Neither can we say why certain admirable virtues, such +as the love of truth, are much more highly appreciated +by some savage tribes than by others;<a name="FNanchor_133" id="FNanchor_133"></a><a href="#Footnote_133" class="fnanchor">133</a> nor, +again, why similar differences prevail even amongst +civilised nations. Knowing how firmly fixed many +strange customs and superstitions have become, we +need feel no surprise that the self-regarding virtues +should now appear to us so natural, supported as they +are by reason, as to be thought innate, although they +were not valued by man in his early condition.</p> + +<p>Notwithstanding many sources of doubt, man can +generally and readily distinguish between the higher +and lower moral rules. The higher are founded on the +social instincts, and relate to the welfare of others. +They are supported by the approbation of our fellow-men +and by reason. The lower rules, though some of +them when implying self-sacrifice hardly deserve to be +called lower, relate chiefly to self, and owe their origin +to public opinion, when matured by experience and +cultivated; for they are not practised by rude tribes.</p> + +<p>As man advances in civilisation, and small tribes +are united into larger communities, the simplest reason +would tell each individual that he ought to extend his +social instincts and sympathies to all the members of +the same nation, though personally unknown to him. +This point being once reached, there is only an arti<span class='pagenum'><a name="Page_101" id="Page_101">101</a></span>ficial +barrier to prevent his sympathies extending to the +men of all nations and races. If, indeed, such men are +separated from him by great differences in appearance +or habits, experience unfortunately shews us how long +it is before we look at them as our fellow-creatures. +Sympathy beyond the confines of man, that is humanity +to the lower animals, seems to be one of the latest +moral acquisitions. It is apparently unfelt by savages, +except towards their pets. How little the old Romans +knew of it is shewn by their abhorrent gladiatorial +exhibitions. The very idea of humanity, as far as I +could observe, was new to most of the Gauchos of the +Pampas. This virtue, one of the noblest with which +man is endowed, seems to arise incidentally from our +sympathies becoming more tender and more widely diffused, +until they are extended to all sentient beings. +As soon as this virtue is honoured and practised by some +few men, it spreads through instruction and example to +the young, and eventually through public opinion.</p> + +<p>The highest stage in moral culture at which we can +arrive, is when we recognise that we ought to control +our thoughts, and “not even in inmost thought to think +again the sins that made the past so pleasant to us.”<a name="FNanchor_134" id="FNanchor_134"></a><a href="#Footnote_134" class="fnanchor">134</a> +Whatever makes any bad action familiar to the mind, +renders its performance by so much the easier. As +Marcus Aurelius long ago said, “Such as are thy habitual +thoughts, such also will be the character of thy +mind; for the soul is dyed by the thoughts.”<a name="FNanchor_135" id="FNanchor_135"></a><a href="#Footnote_135" class="fnanchor">135</a></p> + +<p>Our great philosopher, Herbert Spencer, has recently +explained his views on the moral sense. He says,<a name="FNanchor_136" id="FNanchor_136"></a><a href="#Footnote_136" class="fnanchor">136</a> “I +<span class='pagenum'><a name="Page_102" id="Page_102">102</a></span>believe that the experiences of utility organised and +consolidated through all past generations of the human +race, have been producing corresponding modifications, +which, by continued transmission and accumulation, +have become in us certain faculties of moral intuition—certain +emotions responding to right and wrong conduct, +which have no apparent basis in the individual +experiences of utility.” There is not the least inherent +improbability, as it seems to me, in virtuous tendencies +being more or less strongly inherited; for, not +to mention the various dispositions and habits transmitted +by many of our domestic animals, I have heard +of cases in which a desire to steal and a tendency to lie +appeared to run in families of the upper ranks; and +as stealing is so rare a crime in the wealthy classes, +we can hardly account by accidental coincidence for the +tendency occurring in two or three members of the +same family. If bad tendencies are transmitted, it is +probable that good ones are likewise transmitted. Excepting +through the principle of the transmission of +moral tendencies, we cannot understand the differences +believed to exist in this respect between the various +races of mankind. We have, however, as yet, hardly +sufficient evidence on this head.</p> + +<p class="tb">Even the partial transmission of virtuous tendencies +would be an immense assistance to the primary impulse +derived directly from the social instincts, and indirectly +from the approbation of our fellow-men. Admitting +for the moment that virtuous tendencies are inherited, +it appears probable, at least in such cases as chastity, +temperance, humanity to animals, &c., that they become +first impressed on the mental organisation through +habit, instruction, and example, continued during several +generations in the same family, and in a quite subordinate +degree, or not at all, by the individuals pos<span class='pagenum'><a name="Page_103" id="Page_103">103</a></span>sessing +such virtues, having succeeded best in the +struggle for life. My chief source of doubt with respect +to any such inheritance, is that senseless customs, superstitions, +and tastes, such as the horror of a Hindoo for +unclean food, ought on the same principle to be transmitted. +Although this in itself is perhaps not less probable +than that animals should acquire inherited tastes +for certain kinds of food or fear of certain foes, I have +not met with any evidence in support of the transmission +of superstitious customs or senseless habits.</p> + +<p>Finally, the social instincts which no doubt were +acquired by man, as by the lower animals, for the good +of the community, will from the first have given to him +some wish to aid his fellows, and some feeling of sympathy. +Such impulses will have served him at a very +early period as a rude rule of right and wrong. But as +man gradually advanced in intellectual power and was +enabled to trace the more remote consequences of his +actions; as he acquired sufficient knowledge to reject +baneful customs and superstitions; as he regarded +more and more not only the welfare but the happiness +of his fellow-men; as from habit, following on +beneficial experience, instruction, and example, his +sympathies became more tender and widely diffused, +so as to extend to the men of all races, to the imbecile, +the maimed, and other useless members of +society, and finally to the lower animals,—so would the +standard of his morality rise higher and higher. And +it is admitted by moralists of the derivative school and +by some intuitionists, that the standard of morality has +risen since an early period in the history of man.<a name="FNanchor_137" id="FNanchor_137"></a><a href="#Footnote_137" class="fnanchor">137</a></p> + +<p class="tb">As a struggle may sometimes be seen going on +<span class='pagenum'><a name="Page_104" id="Page_104">104</a></span>between the various instincts of the lower animals, it is +not surprising that there should be a struggle in man +between his social instincts, with their derived virtues, +and his lower, though at the moment, stronger impulses +or desires. This, as Mr. Galton<a name="FNanchor_138" id="FNanchor_138"></a><a href="#Footnote_138" class="fnanchor">138</a> has remarked, is all +the less surprising, as man has emerged from a state of +barbarism within a comparatively recent period. After +having yielded to some temptation we feel a sense of +dissatisfaction, analogous to that felt from other unsatisfied +instincts, called in this case conscience; for we +cannot prevent past images and impressions continually +passing through our minds, and these in their weakened +state we compare with the ever-present social instincts, +or with habits gained in early youth and strengthened +during our whole lives, perhaps inherited, so that they +are at last rendered almost as strong as instincts. +Looking to future generations, there is no cause to fear +that the social instincts will grow weaker, and we may +expect that virtuous habits will grow stronger, becoming +perhaps fixed by inheritance. In this case the struggle +between our higher and lower impulses will be less +severe, and virtue will be triumphant.</p> + +<p class="tb"><i>Summary of the two last Chapters.</i>—There can be no +doubt that the difference between the mind of the lowest +man and that of the highest animal is immense. An +anthropomorphous ape, if he could take a dispassionate +view of his own case, would admit that though he could +form an artful plan to plunder a garden—though he +could use stones for fighting or for breaking open nuts, +<span class='pagenum'><a name="Page_105" id="Page_105">105</a></span>yet that the thought of fashioning a stone into a tool +was quite beyond his scope. Still less, as he would +admit, could he follow out a train of metaphysical +reasoning, or solve a mathematical problem, or reflect +on God, or admire a grand natural scene. Some apes, +however, would probably declare that they could and +did admire the beauty of the coloured skin and fur of +their partners in marriage. They would admit, that +though they could make other apes understand by cries +some of their perceptions and simpler wants, the notion +of expressing definite ideas by definite sounds had +never crossed their minds. They might insist that they +were ready to aid their fellow-apes of the same troop in +many ways, to risk their lives for them, and to take +charge of their orphans; but they would be forced to +acknowledge that disinterested love for all living creatures, +the most noble attribute of man, was quite beyond +their comprehension.</p> + +<p>Nevertheless the difference in mind between man +and the higher animals, great as it is, is certainly one +of degree and not of kind. We have seen that the +senses and intuitions, the various emotions and faculties, +such as love, memory, attention, curiosity, imitation, +reason, &c., of which man boasts, may be found in an +incipient, or even sometimes in a well-developed condition, +in the lower animals. They are also capable of +some inherited improvement, as we see in the domestic +dog compared with the wolf or jackal. If it be maintained +that certain powers, such as self-consciousness, +abstraction, &c., are peculiar to man, it may well be +that these are the incidental results of other highly-advanced +intellectual faculties; and these again are +mainly the result of the continued use of a highly +developed language. At what age does the new-born +infant possess the power of abstraction, or become self<span class='pagenum'><a name="Page_106" id="Page_106">106</a></span>conscious +and reflect on its own existence? We cannot +answer; nor can we answer in regard to the ascending +organic scale. The half-art and half-instinct of language +still bears the stamp of its gradual evolution. +The ennobling belief in God is not universal with man; +and the belief in active spiritual agencies naturally follows +from his other mental powers. The moral sense +perhaps affords the best and highest distinction between +man and the lower animals; but I need not say anything +on this head, as I have so lately endeavoured +to shew that the social instincts,—the prime principle +of man’s moral constitution<a name="FNanchor_139" id="FNanchor_139"></a><a href="#Footnote_139" class="fnanchor">139</a>—with the aid of active +intellectual powers and the effects of habit, naturally +lead to the golden rule, “As ye would that men should +do to you, do ye to them likewise;” and this lies at +the foundation of morality.</p> + +<p>In a future chapter I shall make some few remarks +on the probable steps and means by which the several +mental and moral faculties of man have been gradually +evolved. That this at least is possible ought not +to be denied, when we daily see their development in +every infant; and when we may trace a perfect gradation +from the mind of an utter idiot, lower than that of +the lowest animal, to the mind of a Newton.</p> + +<hr /> +<p><span class='pagenum'><a name="Page_107" id="Page_107">107</a></span></p> + + +<h3>CHAPTER IV.</h3> + +<h4><span class="smcap">On the Manner of Development of Man from some +lower Form.</span></h4> + +<div class="blockquot"><p>Variability of body and mind in man—Inheritance—Causes of +variability—Laws of variation the same in man as in the lower +animals—Direct action of the conditions of life—Effects of the +increased use and disuse of parts—Arrested development—Reversion—Correlated +variation—Rate of increase—Checks to +increase—Natural selection—Man the most dominant animal +in the world—Importance of his corporeal structure—The +causes which have led to his becoming erect—Consequent changes +of structure—Decrease in size of the canine teeth—Increased +size and altered shape of the skull—Nakedness—Absence of +a tail—Defenceless condition of man.</p></div> + +<p>We have seen in the first chapter that the homological +structure of man, his embryological development and +the rudiments which he still retains, all declare in the +plainest manner that he is descended from some lower +form. The possession of exalted mental powers is no +insuperable objection to this conclusion. In order that +an ape-like creature should have been transformed into +man, it is necessary that this early form, as well as +many successive links, should all have varied in mind +and body. It is impossible to obtain direct evidence on +this head; but if it can be shewn that man now varies—that +his variations are induced by the same general +causes, and obey the same general laws, as in the case +of the lower animals—there can be little doubt that +the preceding intermediate links varied in a like +manner. The variations at each successive stage of +descent must, also, have been in some manner accumulated +and fixed.</p> + +<p><span class='pagenum'><a name="Page_108" id="Page_108">108</a></span>The facts and conclusions to be given in this chapter +relate almost exclusively to the probable means by +which the transformation of man has been effected, +as far as his bodily structure is concerned. The following +chapter will be devoted to the development of +his intellectual and moral faculties. But the present +discussion likewise bears on the origin of the different +races or species of mankind, whichever term may be +preferred.</p> + +<p>It is manifest that man is now subject to much +variability. No two individuals of the same race are +quite alike. We may compare millions of faces, and +each will be distinct. There is an equally great +amount of diversity in the proportions and dimensions +of the various parts of the body; the length of the legs +being one of the most variable points.<a name="FNanchor_140" id="FNanchor_140"></a><a href="#Footnote_140" class="fnanchor">140</a> Although in +some quarters of the world an elongated skull, and in +other quarters a short skull prevails, yet there is great +diversity of shape even within the limits of the same +race, as with the aborigines of America and South +Australia,—the latter a race “probably as pure and +homogeneous in blood, customs, and language as any +in existence”—and even with the inhabitants of so +confined an area as the Sandwich Islands.<a name="FNanchor_141" id="FNanchor_141"></a><a href="#Footnote_141" class="fnanchor">141</a> An eminent +dentist assures me that there is nearly as much +diversity in the teeth, as in the features. The chief +arteries so frequently run in abnormal courses, that it +has been found useful for surgical purposes to calculate +<span class='pagenum'><a name="Page_109" id="Page_109">109</a></span>from 12,000 corpses how often each course prevails.<a name="FNanchor_142" id="FNanchor_142"></a><a href="#Footnote_142" class="fnanchor">142</a> +The muscles are eminently variable: thus those of the +foot were found by Prof. Turner<a name="FNanchor_143" id="FNanchor_143"></a><a href="#Footnote_143" class="fnanchor">143</a> not to be strictly +alike in any two out of fifty bodies; and in some the +deviations were considerable. Prof. Turner adds that +the power of performing the appropriate movements +must have been modified in accordance with the several +deviations. Mr. J. Wood has recorded<a name="FNanchor_144" id="FNanchor_144"></a><a href="#Footnote_144" class="fnanchor">144</a> the occurrence +of 295 muscular variations in thirty-six subjects, and in +another set of the same number no less than 558 variations, +reckoning both sides of the body as one. In the +last set, not one body out of the thirty-six was “found +totally wanting in departures from the standard descriptions +of the muscular system given in anatomical +text-books.” A single body presented the extraordinary +number of twenty-five distinct abnormalities. The +same muscle sometimes varies in many ways: thus +Prof. Macalister describes<a name="FNanchor_145" id="FNanchor_145"></a><a href="#Footnote_145" class="fnanchor">145</a> no less than twenty distinct +variations in the <i>palmaris accessorius</i>.</p> + +<p>The famous old anatomist, Wolff,<a name="FNanchor_146" id="FNanchor_146"></a><a href="#Footnote_146" class="fnanchor">146</a> insists that the +internal viscera are more variable than the external +parts: <i>Nulla particula est quæ non aliter et aliter in +aliis se habeat hominibus.</i> He has even written a treatise +on the choice of typical examples of the viscera for +representation. A discussion on the beau-ideal of the +liver, lungs, kidneys, &c., as of the human face divine, +sounds strange in our ears.</p> + +<p>The variability or diversity of the mental faculties +in men of the same race, not to mention the greater +<span class='pagenum'><a name="Page_110" id="Page_110">110</a></span>differences between the men of distinct races, is so +notorious that not a word need here be said. So it +is with the lower animals, as has been illustrated by +a few examples in the last chapter. All who have had +charge of menageries admit this fact, and we see it +plainly in our dogs and other domestic animals. Brehm +especially insists that each individual monkey of those +which he kept under confinement in Africa had its own +peculiar disposition and temper: he mentions one baboon +remarkable for its high intelligence; and the keepers +in the Zoological Gardens pointed out to me a monkey, +belonging to the New World division, equally remarkable +for intelligence. Rengger, also, insists on the diversity +in the various mental characters of the monkeys +of the same species which he kept in Paraguay; and +this diversity, as he adds, is partly innate, and partly +the result of the manner in which they have been +treated or educated.<a name="FNanchor_147" id="FNanchor_147"></a><a href="#Footnote_147" class="fnanchor">147</a></p> + +<p>I have elsewhere<a name="FNanchor_148" id="FNanchor_148"></a><a href="#Footnote_148" class="fnanchor">148</a> so fully discussed the subject of +Inheritance that I need here add hardly anything. A +greater number of facts have been collected with respect +to the transmission of the most trifling, as well as of the +most important characters in man than in any of the +lower animals; though the facts are copious enough +with respect to the latter. So in regard to mental +qualities, their transmission is manifest in our dogs, +horses, and other domestic animals. Besides special +tastes and habits, general intelligence, courage, bad and +good temper, &c., are certainly transmitted. With man +we see similar facts in almost every family; and we +<span class='pagenum'><a name="Page_111" id="Page_111">111</a></span>now know through the admirable labours of Mr. Galton<a name="FNanchor_149" id="FNanchor_149"></a><a href="#Footnote_149" class="fnanchor">149</a> +that genius, which implies a wonderfully complex combination +of high faculties, tends to be inherited; and, +on the other hand, it is too certain that insanity and +deteriorated mental powers likewise run in the same +families.</p> + +<p>With respect to the causes of variability we are in +all cases very ignorant; but we can see that in man as +in the lower animals, they stand in some relation with +the conditions to which each species has been exposed +during several generations. Domesticated animals vary +more than those in a state of nature; and this is apparently +due to the diversified and changing nature of +their conditions. The different races of man resemble +in this respect domesticated animals, and so do the +individuals of the same race when inhabiting a very +wide area, like that of America. We see the influence +of diversified conditions in the more civilised nations, +the members of which belong to different grades of rank +and follow different occupations, presenting a greater +range of character than the members of barbarous +nations. But the uniformity of savages has often been +exaggerated, and in some cases can hardly be said +to exist.<a name="FNanchor_150" id="FNanchor_150"></a><a href="#Footnote_150" class="fnanchor">150</a> It is nevertheless an error to speak of man, +even if we look only to the conditions to which he +has been subjected, as “far more domesticated”<a name="FNanchor_151" id="FNanchor_151"></a><a href="#Footnote_151" class="fnanchor">151</a> than +<span class='pagenum'><a name="Page_112" id="Page_112">112</a></span>any other animal. Some savage races, such as the +Australians, are not exposed to more diversified conditions +than are many species which have very wide +ranges. In another and much more important respect, +man differs widely from any strictly domesticated +animal; for his breeding has not been controlled, +either through methodical or unconscious selection. No +race or body of men has been so completely subjugated +by other men, that certain individuals have been preserved +and thus unconsciously selected, from being in +some way more useful to their masters. Nor have +certain male and female individuals been intentionally +picked out and matched, except in the well-known +case of the Prussian grenadiers; and in this case man +obeyed, as might have been expected, the law of methodical +selection; for it is asserted that many tall men +were reared in the villages inhabited by the grenadiers +with their tall wives.</p> + +<p>If we consider all the races of man, as forming a single +species, his range is enormous; but some separate races, +as the Americans and Polynesians, have very wide +ranges. It is a well-known law that widely-ranging +species are much more variable than species with restricted +ranges; and the variability of man may with +more truth be compared with that of widely-ranging +species, than with that of domesticated animals.</p> + +<p class="tb">Not only does variability appear to be induced in +man and the lower animals by the same general causes, +but in both the same characters are affected in a closely +analogous manner. This has been proved in such full +detail by Godron and Quatrefages, that I need here +only refer to their works.<a name="FNanchor_152" id="FNanchor_152"></a><a href="#Footnote_152" class="fnanchor">152</a> Monstrosities, which gra<span class='pagenum'><a name="Page_113" id="Page_113">113</a></span>duate +into slight variations, are likewise so similar in +man and the lower animals, that the same classification +and the same terms can be used for both, as may +be seen in Isidore Geoffroy St.-Hilaire’s great work.<a name="FNanchor_153" id="FNanchor_153"></a><a href="#Footnote_153" class="fnanchor">153</a> +This is a necessary consequence of the same laws of +change prevailing throughout the animal kingdom. In +my work on the variation of domestic animals, I have +attempted to arrange in a rude fashion the laws of +variation under the following heads:—The direct and +definite action of changed conditions, as shewn by all or +nearly all the individuals of the same species varying +in the same manner under the same circumstances. +The effects of the long-continued use or disuse of +parts. The cohesion of homologous parts. The variability +of multiple parts. Compensation of growth; +but of this law I have found no good instances in the +case of man. The effects of the mechanical pressure of +one part on another; as of the pelvis on the cranium +of the infant in the womb. Arrests of development, +leading to the diminution or suppression of parts. The +reappearance of long-lost characters through reversion. +And lastly, correlated variation. All these so-called +laws apply equally to man and the lower animals; and +most of them even to plants. It would be superfluous +here to discuss all of them;<a name="FNanchor_154" id="FNanchor_154"></a><a href="#Footnote_154" class="fnanchor">154</a> but several are so important +for us, that they must be treated at considerable +length.</p> + +<p><i>The direct and definite action of changed conditions.</i>—This +is a most perplexing subject. It cannot be denied +<span class='pagenum'><a name="Page_114" id="Page_114">114</a></span>that changed conditions produce some effect, and occasionally +a considerable effect, on organisms of all kinds; +and it seems at first probable that if sufficient time +were allowed this would be the invariable result. But +I have failed to obtain clear evidence in favour of this +conclusion; and valid reasons may be urged on the +other side, at least as far as the innumerable structures +are concerned, which are adapted for special ends. +There can, however, be no doubt that changed conditions +induce an almost indefinite amount of fluctuating +variability, by which the whole organisation is rendered +in some degree plastic.</p> + +<p>In the United States, above 1,000,000 soldiers, who +served in the late war, were measured, and the States +in which they were born and reared recorded.<a name="FNanchor_155" id="FNanchor_155"></a><a href="#Footnote_155" class="fnanchor">155</a> From +this astonishing number of observations it is proved that +local influences of some kind act directly on stature; +and we further learn that “the State where the physical +growth has in great measure taken place, and the State +of birth, which indicates the ancestry, seem to exert +a marked influence on the stature.” For instance it +is established, “that residence in the Western States, +during the years of growth, tends to produce increase +of stature.” On the other hand, it is certain that with +sailors, their manner of life delays growth, as shewn “by +the great difference between the statures of soldiers and +sailors at the ages of 17 and 18 years.” Mr. B. A. Gould +endeavoured to ascertain the nature of the influences +which thus act on stature; but he arrived only at negative +results, namely, that they did not relate to climate, +the elevation of the land, soil, nor even “in any controlling +degree” to the abundance or need of the com<span class='pagenum'><a name="Page_115" id="Page_115">115</a></span>forts +of life. This latter conclusion is directly opposed +to that arrived at by Villermé from the statistics of the +height of the conscripts in different parts of France. +When we compare the differences in stature between the +Polynesian chiefs and the lower orders within the same +islands, or between the inhabitants of the fertile volcanic +and low barren coral islands of the same ocean,<a name="FNanchor_156" id="FNanchor_156"></a><a href="#Footnote_156" class="fnanchor">156</a> or +again between the Fuegians on the eastern and western +shores of their country, where the means of subsistence +are very different, it is scarcely possible to avoid the +conclusion that better food and greater comfort do influence +stature. But the preceding statements shew +how difficult it is to arrive at any precise result. Dr. +Beddoe has lately proved that, with the inhabitants of +Britain, residence in towns and certain occupations have +a deteriorating influence on height; and he infers that +the result is to a certain extent inherited, as is likewise +the case in the United States. Dr. Beddoe further +believes that wherever a “race attains its maximum of +physical development, it rises highest in energy and +moral vigour.”<a name="FNanchor_157" id="FNanchor_157"></a><a href="#Footnote_157" class="fnanchor">157</a></p> + +<p class="tb">Whether external conditions produce any other +direct effect on man is not known. It might have been +expected that differences of climate would have had a +marked influence, as the lungs and kidneys are brought +into fuller activity under a low temperature, and the +liver and skin under a high one.<a name="FNanchor_158" id="FNanchor_158"></a><a href="#Footnote_158" class="fnanchor">158</a> It was formerly +thought that the colour of the skin and the character +<span class='pagenum'><a name="Page_116" id="Page_116">116</a></span>of the hair were determined by light or heat; and +although it can hardly be denied that some effect is +thus produced, almost all observers now agree that the +effect has been very small, even after exposure during +many ages. But this subject will be more properly +discussed when we treat of the different races of mankind. +With our domestic animals there are grounds +for believing that cold and damp directly affect the +growth of the hair; but I have not met with any evidence +on this head in the case of man.</p> + +<p><i>Effects of the increased Use and Disuse of Parts.</i>—It +is well known that use strengthens the muscles in +the individual, and complete disuse, or the destruction +of the proper nerve, weakens them. When the eye +is destroyed the optic nerve often becomes atrophied. +When an artery is tied, the lateral channels increase +not only in diameter, but in the thickness and strength +of their coats. When one kidney ceases acting from +disease, the other increases in size and does double +work. Bones increase not only in thickness, but in +length, from carrying a greater weight.<a name="FNanchor_159" id="FNanchor_159"></a><a href="#Footnote_159" class="fnanchor">159</a> Different +occupations habitually followed lead to changed proportions +in various parts of the body. Thus it was +clearly ascertained by the United States Commission<a name="FNanchor_160" id="FNanchor_160"></a><a href="#Footnote_160" class="fnanchor">160</a> +that the legs of the sailors employed in the late war +were longer by 0.217 of an inch than those of the soldiers, +though the sailors were on an average shorter +men; whilst their arms were shorter by 1.09 of an inch, +and therefore out of proportion shorter in relation to +<span class='pagenum'><a name="Page_117" id="Page_117">117</a></span>their lesser height. This shortness of the arms is +apparently due to their greater use, and is an unexpected +result; but sailors chiefly use their arms in +pulling and not in supporting weights. The girth of +the neck and the depth of the instep are greater, whilst +the circumference of the chest, waist, and hips is less in +sailors than in soldiers.</p> + +<p>Whether the several foregoing modifications would +become hereditary, if the same habits of life were followed +during many generations, is not known, but is +probable. Rengger<a name="FNanchor_161" id="FNanchor_161"></a><a href="#Footnote_161" class="fnanchor">161</a> attributes the thin legs and thick +arms of the Payaguas Indians to successive generations +having passed nearly their whole lives in canoes, with +their lower extremities motionless. Other writers have +come to a similar conclusion in other analogous cases. +According to Cranz,<a name="FNanchor_162" id="FNanchor_162"></a><a href="#Footnote_162" class="fnanchor">162</a> who lived for a long time with the +Esquimaux, “the natives believe that ingenuity and +dexterity in seal-catching (their highest art and virtue) +is hereditary; there is really something in it, for the +son of a celebrated seal-catcher will distinguish himself +though he lost his father in childhood.” But in +this case it is mental aptitude, quite as much as bodily +structure, which appears to be inherited. It is asserted +that the hands of English labourers are at birth larger +than those of the gentry.<a name="FNanchor_163" id="FNanchor_163"></a><a href="#Footnote_163" class="fnanchor">163</a> From the correlation which +exists, at least in some cases,<a name="FNanchor_164" id="FNanchor_164"></a><a href="#Footnote_164" class="fnanchor">164</a> between the development +of the extremities and of the jaws, it is possible that +in those classes which do not labour much with their +hands and feet, the jaws would be reduced in size from +this cause. That they are generally smaller in refined +and civilised men than in hard-working men or savages, +<span class='pagenum'><a name="Page_118" id="Page_118">118</a></span>is certain. But with savages, as Mr. Herbert Spencer<a name="FNanchor_165" id="FNanchor_165"></a><a href="#Footnote_165" class="fnanchor">165</a> +has remarked, the greater use of the jaws in chewing +coarse, uncooked food, would act in a direct manner on +the masticatory muscles and on the bones to which +they are attached. In infants long before birth, the +skin on the soles of the feet is thicker than on any +other part of the body;<a name="FNanchor_166" id="FNanchor_166"></a><a href="#Footnote_166" class="fnanchor">166</a> and it can hardly be doubted +that this is due to the inherited effects of pressure +during a long series of generations.</p> + +<p>It is familiar to every one that watchmakers and +engravers are liable to become short-sighted, whilst +sailors and especially savages are generally long-sighted. +Short-sight and long-sight certainly tend to be inherited.<a name="FNanchor_167" id="FNanchor_167"></a><a href="#Footnote_167" class="fnanchor">167</a> +The inferiority of Europeans, in comparison +with savages, in eyesight and in the other +senses, is no doubt the accumulated and transmitted +effect of lessened use during many generations; for +Rengger<a name="FNanchor_168" id="FNanchor_168"></a><a href="#Footnote_168" class="fnanchor">168</a> states that he has repeatedly observed Europeans, +who had been brought up and spent their whole +lives with the wild Indians, who nevertheless did not +equal them in the sharpness of their senses. The same +naturalist observes that the cavities in the skull for +the reception of the several sense-organs are larger in +the American aborigines than in Europeans; and this +no doubt indicates a corresponding difference in the +dimensions of the organs themselves. Blumenbach has +also remarked on the large size of the nasal cavities +<span class='pagenum'><a name="Page_119" id="Page_119">119</a></span>in the skulls of the American aborigines, and connects +this fact with their remarkably acute power of smell. +The Mongolians of the plains of Northern Asia, according +to Pallas, have wonderfully perfect senses; and Prichard +believes that the great breadth of their skulls across +the zygomas follows from their highly-developed sense-organs.<a name="FNanchor_169" id="FNanchor_169"></a><a href="#Footnote_169" class="fnanchor">169</a></p> + +<p>The Quechua Indians inhabit the lofty plateaux of +Peru, and Alcide d’Orbigny states<a name="FNanchor_170" id="FNanchor_170"></a><a href="#Footnote_170" class="fnanchor">170</a> that from continually +breathing a highly rarefied atmosphere they +have acquired chests and lungs of extraordinary dimensions. +The cells, also, of the lungs are larger and more +numerous than in Europeans. These observations +have been doubted; but Mr. D. Forbes carefully +measured many Aymaras, an allied race, living at the +height of between ten and fifteen thousand feet; and +he informs me<a name="FNanchor_171" id="FNanchor_171"></a><a href="#Footnote_171" class="fnanchor">171</a> that they differ conspicuously from the +men of all other races seen by him, in the circumference +and length of their bodies. In his table of +measurements, the stature of each man is taken at +1000, and the other measurements are reduced to this +standard. It is here seen that the extended arms +of the Aymaras are shorter than those of Europeans, +and much shorter than those of Negroes. The legs are +likewise shorter, and they present this remarkable peculiarity, +that in every Aymara measured the femur is +actually shorter than the tibia. On an average the +length of the femur to that of the tibia is as 211 to +252; whilst in two Europeans measured at the same +<span class='pagenum'><a name="Page_120" id="Page_120">120</a></span>time, the femora to the tibiæ were as 244 to 230; and +in three Negroes as 258 to 241. The humerus is likewise +shorter relatively to the fore-arm. This shortening +of that part of the limb which is nearest to the body, +appears to be, as suggested to me by Mr. Forbes, a case +of compensation in relation with the greatly increased +length of the trunk. The Aymaras present some other +singular points of structure, for instance, the very small +projection of the heel.</p> + +<p>These men are so thoroughly acclimatised to their +cold and lofty abode, that when formerly carried down +by the Spaniards to the low Eastern plains, and when +now tempted down by high wages to the gold-washings, +they suffer a frightful rate of mortality. Nevertheless +Mr. Forbes found a few pure families which had survived +during two generations; and he observed that +they still inherited their characteristic peculiarities. +But it was manifest, even without measurement, that +these peculiarities had all decreased; and on measurement +their bodies were found not to be so much elongated +as those of the men on the high plateau; whilst +their femora had become somewhat lengthened, as had +their tibiæ but in a less degree. The actual measurements +may be seen by consulting Mr. Forbes‘ memoir. +From these valuable observations, there can, I think, +be no doubt that residence during many generations at +a great elevation tends, both directly and indirectly, to +induce inherited modifications in the proportions of the +body.<a name="FNanchor_172" id="FNanchor_172"></a><a href="#Footnote_172" class="fnanchor">172</a></p> + +<p class="tb">Although man may not have been much modified +during the latter stages of his existence through the +<span class='pagenum'><a name="Page_121" id="Page_121">121</a></span>increased or decreased use of parts, the facts now given +shew that his liability in this respect has not been lost; +and we positively know that the same law holds good +with the lower animals. Consequently we may infer, +that when at a remote epoch the progenitors of man +were in a transitional state, and were changing from +quadrupeds into bipeds, natural selection would probably +have been greatly aided by the inherited effects of the +increased or diminished use of the different parts of the +body.</p> + +<p class="tb"><i>Arrests of Development.</i>—Arrested development differs +from arrested growth, as parts in the former state continue +to grow whilst still retaining their early condition. +Various monstrosities come under this head, and some +are known to be occasionally inherited, as a cleft-palate. +It will suffice for our purpose to refer to the arrested +brain-development of microcephalous idiots, as described +in Vogt’s great memoir.<a name="FNanchor_173" id="FNanchor_173"></a><a href="#Footnote_173" class="fnanchor">173</a> Their skulls are smaller, and +the convolutions of the brain are less complex than in +normal men. The frontal sinus, or the projection over +the eyebrows, is largely developed, and the jaws are +prognathous to an “<i>effrayant</i>” degree; so that these idiots +somewhat resemble the lower types of mankind. Their +intelligence and most of their mental faculties are +extremely feeble. They cannot acquire the power of +speech, and are wholly incapable of prolonged attention, +but are much given to imitation. They are strong and +remarkably active, continually gamboling and jumping +about, and making grimaces. They often ascend stairs +on all-fours; and are curiously fond of climbing up +furniture or trees. We are thus reminded of the delight +<span class='pagenum'><a name="Page_122" id="Page_122">122</a></span>shewn by almost all boys in climbing trees; and this +again reminds us how lambs and kids, originally alpine +animals, delight to frisk on any hillock, however small.</p> + +<p><i>Reversion.</i>—Many of the cases to be here given +might have been introduced under the last heading. +Whenever a structure is arrested in its development, +but still continues growing until it closely resembles a +corresponding structure in some lower and adult member +of the same group, we may in one sense consider it as a case +of reversion. The lower members in a group give us +some idea how the common progenitor of the group was +probably constructed; and it is hardly credible that a +part arrested at an early phase of embryonic development +should be enabled to continue growing so as ultimately +to perform its proper function, unless it had +acquired this power of continued growth during some +earlier state of existence, when the present exceptional +or arrested structure was normal. The simple brain of +a microcephalous idiot, in as far as it resembles that +of an ape, may in this sense be said to offer a case of +reversion. There are other cases which come more +strictly under our present heading of reversion. Certain +structures, regularly occurring in the lower members of +the group to which man belongs, occasionally make +their appearance in him, though not found in the normal +human embryo; or, if present in the normal human +embryo, they become developed in an abnormal manner, +though this manner of development is proper to the +lower members of the same group. These remarks will +be rendered clearer by the following illustrations.</p> + +<p>In various mammals the uterus graduates from a +double organ with two distinct orifices and two passages, +as in the marsupials, into a single organ, showing no +signs of doubleness except a slight internal fold, as in<span class='pagenum'><a name="Page_123" id="Page_123">123</a></span> +the higher apes and man. The rodents exhibit a perfect +series of gradations between these two extreme +states. In all mammals the uterus is developed from +two simple primitive tubes, the inferior portions of +which form the cornua; and it is in the words of +Dr. Farre “by the coalescence of the two cornua at +their lower extremities that the body of the uterus is +formed in man; while in those animals in which no +middle portion or body exists, the cornua remain ununited. +As the development of the uterus proceeds, +the two cornua become gradually shorter, until at +length they are lost, or, as it were, absorbed into the +body of the uterus.” The angles of the uterus are +still produced into cornua, even so high in the scale as +in the lower apes, and their allies the lemurs.</p> + +<p>Now in women anomalous cases are not very infrequent, +in which the mature uterus is furnished with +cornua, or is partially divided into two organs; and +such cases, according to Owen, repeat “the grade of concentrative +development,” attained by certain rodents. +Here perhaps we have an instance of a simple arrest of +embryonic development, with subsequent growth and +perfect functional development, for either side of the +partially double uterus is capable of performing the +proper office of gestation. In other and rarer cases, +two distinct uterine cavities are formed, each having +its proper orifice and passage.<a name="FNanchor_174" id="FNanchor_174"></a><a href="#Footnote_174" class="fnanchor">174</a> No such stage is passed +through during the ordinary development of the embryo, +and it is difficult to believe, though perhaps not impossible, +that the two simple, minute, primitive tubes +could know how (if such an expression may be used) to +<span class='pagenum'><a name="Page_124" id="Page_124">124</a></span>grow into two distinct uteri, each with a well-constructed +orifice and passage, and each furnished with numerous +muscles, nerves, glands and vessels, if they had not +formerly passed through a similar course of development, +as in the case of existing marsupials. No one will +pretend that so perfect a structure as the abnormal +double uterus in woman could be the result of mere +chance. But the principle of reversion, by which long-lost +dormant structures are called back into existence, +might serve as the guide for the full development of +the organ, even after the lapse of an enormous interval +of time.</p> + +<p>Professor Canestrini,<a name="FNanchor_175" id="FNanchor_175"></a><a href="#Footnote_175" class="fnanchor">175</a> after discussing the foregoing +and various analogous cases, arrives at the same conclusion +as that just given. He adduces, as another +instance, the malar bone, which, in some of the Quadrumana +and other mammals, normally consists of two +portions. This is its condition in the two-months-old +human fœtus; and thus it sometimes remains, through +arrested development, in man when adult, more especially +in the lower prognathous races. Hence Canestrini concludes +that some ancient progenitor of man must have +possessed this bone normally divided into two portions, +which subsequently became fused together. In man +the frontal bone consists of a single piece, but in the +embryo and in children, and in almost all the lower +mammals, it consists of two pieces separated by a distinct +suture. This suture occasionally persists, more +or less distinctly, in man after maturity, and more fre<span class='pagenum'><a name="Page_125" id="Page_125">125</a></span>quently +in ancient than in recent crania, especially as +Canestrini has observed in those exhumed from the +Drift and belonging to the brachycephalic type. Here +again he comes to the same conclusion as in the analogous +case of the malar bones. In this and other +instances presently to be given, the cause of ancient +races approaching the lower animals in certain characters +more frequently than do the modern races, +appears to be that the latter stand at a somewhat greater +distance in the long line of descent from their early +semi-human progenitors.</p> + +<p>Various other anomalies in man, more or less analogous +with the foregoing, have been advanced by different +authors<a name="FNanchor_176" id="FNanchor_176"></a><a href="#Footnote_176" class="fnanchor">176</a> as cases of reversion; but these seem +not a little doubtful, for we have to descend extremely +low in the mammalian series before we find such structures +normally present.<a name="FNanchor_177" id="FNanchor_177"></a><a href="#Footnote_177" class="fnanchor">177</a></p> + +<p><span class='pagenum'><a name="Page_126" id="Page_126">126</a></span></p><p>In man the canine teeth are perfectly efficient instruments +for mastication. But their true canine character, +as Owen<a name="FNanchor_178" id="FNanchor_178"></a><a href="#Footnote_178" class="fnanchor">178</a> remarks, “is indicated by the conical form +of the crown, which terminates in an obtuse point, is +convex outward and flat or sub-concave within, at the +base of which surface there is a feeble prominence. +The conical form is best expressed in the Melanian +races, especially the Australian. The canine is more +deeply implanted, and by a stronger fang than the +incisors.” Nevertheless this tooth no longer serves +man as a special weapon for tearing his enemies or prey; +it may, therefore, as far as its proper function is concerned, +be considered as rudimentary. In every large +collection of human skulls some may be found, as +Häckel<a name="FNanchor_179" id="FNanchor_179"></a><a href="#Footnote_179" class="fnanchor">179</a> observes, with the canine teeth projecting considerably +beyond the others in the same manner, but +in a less degree, as in the anthropomorphous apes. In +these cases, open spaces between the teeth in the one +jaw are left for the reception of the canines belonging +to the opposite jaw. An interspace of this kind in a +Kaffir skull, figured by Wagner, is surprisingly wide.<a name="FNanchor_180" id="FNanchor_180"></a><a href="#Footnote_180" class="fnanchor">180</a> +Considering how few ancient skulls have been examined +in comparison with recent skulls, it is an interesting +fact that in at least three cases the canines project +largely; and in the Naulette jaw they are spoken of as +enormous.<a name="FNanchor_181" id="FNanchor_181"></a><a href="#Footnote_181" class="fnanchor">181</a></p> + +<p><span class='pagenum'><a name="Page_127" id="Page_127">127</a></span></p><p>The males alone of the anthropomorphous apes have +their canines fully developed; but in the female gorilla, +and in a less degree in the female orang, these teeth +project considerably beyond the others; therefore the +fact that women sometimes have, as I have been assured, +considerably projecting canines, is no serious objection +to the belief that their occasional great development in +man is a case of reversion to an ape-like progenitor. +He who rejects with scorn the belief that the shape of +his own canines, and their occasional great development +in other men, are due to our early progenitors having +been provided with these formidable weapons, will probably +reveal by sneering the line of his descent. For +though he no longer intends, nor has the power, to use +these teeth as weapons, he will unconsciously retract his +“snarling muscles” (thus named by Sir C. Bell)<a name="FNanchor_182" id="FNanchor_182"></a><a href="#Footnote_182" class="fnanchor">182</a> so as +to expose them ready for action, like a dog prepared to +fight.</p> + +<p>Many muscles are occasionally developed in man, +which are proper to the Quadrumana or other mammals. +Professor Vlacovich<a name="FNanchor_183" id="FNanchor_183"></a><a href="#Footnote_183" class="fnanchor">183</a> examined forty male subjects, +and found a muscle, called by him the ischiopubic, +in nineteen of them; in three others there was +a ligament which represented this muscle; and in the +remaining eighteen no trace of it. Out of thirty female +subjects this muscle was developed on both sides in only +two, but in three others the rudimentary ligament was +present. This muscle, therefore, appears to be much +more common in the male than in the female sex; and +on the principle of the descent of man from some lower +form, its presence can be understood; for it has been +detected in several of the lower animals, and in all of +<span class='pagenum'><a name="Page_128" id="Page_128">128</a></span>these it serves exclusively to aid the male in the act +of reproduction.</p> + +<p>Mr. J. Wood, in his valuable series of papers,<a name="FNanchor_184" id="FNanchor_184"></a><a href="#Footnote_184" class="fnanchor">184</a> has +minutely described a vast number of muscular variations +in man, which resemble normal structures in the +lower animals. Looking only to the muscles which +closely resemble those regularly present in our nearest +allies, the Quadrumana, they are too numerous to be +here even specified. In a single male subject, having +a strong bodily frame and well-formed skull, no less +than seven muscular variations were observed, all of +which plainly represented muscles proper to various +kinds of apes. This man, for instance, had on both +sides of his neck a true and powerful “<i>levator claviculæ</i>,” +such as is found in all kinds of apes, and which +is said to occur in about one out of sixty human subjects.<a name="FNanchor_185" id="FNanchor_185"></a><a href="#Footnote_185" class="fnanchor">185</a> +Again, this man had “a special abductor of +the metatarsal bone of the fifth digit, such as Professor +Huxley and Mr. Flower have shewn to exist +uniformly in the higher and lower apes.” The hands +and arms of man are eminently characteristic structures, +but their muscles are extremely liable to vary, so as to +resemble the corresponding muscles in the lower animals.<a name="FNanchor_186" id="FNanchor_186"></a><a href="#Footnote_186" class="fnanchor">186</a> +Such resemblances are either complete and per<span class='pagenum'><a name="Page_129" id="Page_129">129</a></span>fect +or imperfect, yet in this latter case manifestly of +a transitional nature. Certain variations are more common +in man, and others in woman, without our being +able to assign any reason. Mr. Wood, after describing +numerous cases, makes the following pregnant remark: +“Notable departures from the ordinary type of the +muscular structures run in grooves or directions, which +must be taken to indicate some unknown factor, of +much importance to a comprehensive knowledge of +general and scientific anatomy.”<a name="FNanchor_187" id="FNanchor_187"></a><a href="#Footnote_187" class="fnanchor">187</a></p> + +<p>That this unknown factor is reversion to a former +state of existence may be admitted as in the highest +degree probable. It is quite incredible that a man +should through mere accident abnormally resemble, in +no less than seven of his muscles, certain apes, if there +had been no genetic connection between them. On the +other hand, if man is descended from some ape-like +creature, no valid reason can be assigned why certain +muscles should not suddenly reappear after an interval of +many thousand generations, in the same manner as with +horses, asses, and mules, dark-coloured stripes suddenly +reappear on the legs and shoulders, after an interval of +hundreds, or more probably thousands, of generations.</p> + +<p class="tb">These various cases of reversion are so closely related +<span class='pagenum'><a name="Page_130" id="Page_130">130</a></span>to those of rudimentary organs given in the first chapter, +that many of them might have been indifferently introduced +in either chapter. Thus a human uterus furnished +with cornua may be said to represent in a rudimentary +condition the same organ in its normal state in certain +mammals. Some parts which are rudimental in man, +as the os coccyx in both sexes and the mammæ in the +male sex, are always present; whilst others, such as +the supra-condyloid foramen, only occasionally appear, +and therefore might have been introduced under the +head of reversion. These several reversionary, as well +as the strictly rudimentary, structures reveal the descent +of man from some lower form in an unmistakeable +manner.</p> + +<p><i>Correlated Variation.</i>—In man, as in the lower animals, +many structures are so intimately related, that +when one part varies so does another, without our being +able, in most cases, to assign any reason. We cannot say +whether the one part governs the other, or whether both +are governed by some earlier developed part. Various +monstrosities, as I. Geoffroy repeatedly insists, are thus +intimately connected. Homologous structures are particularly +liable to change together, as we see on the +opposite sides of the body, and in the upper and lower +extremities. Meckel long ago remarked that when the +muscles of the arm depart from their proper type, they +almost always imitate those of the leg; and so conversely +with the muscles of the legs. The organs of sight and +hearing, the teeth and hair, the colour of the skin and +hair, colour and constitution, are more or less correlated.<a name="FNanchor_188" id="FNanchor_188"></a><a href="#Footnote_188" class="fnanchor">188</a> +Professor Schaaffhausen first drew attention to the rela<span class='pagenum'><a name="Page_131" id="Page_131">131</a></span>tion +apparently existing between a muscular frame and +strongly-pronounced supra-orbital ridges, which are so +characteristic of the lower races of man.</p> + +<p class="tb">Besides the variations which can be grouped with +more or less probability under the foregoing heads, +there is a large class of variations which may be provisionally +called spontaneous, for they appear, owing to our +ignorance, to arise without any exciting cause. It can, +however, be shewn that such variations, whether consisting +of slight individual differences, or of strongly-marked +and abrupt deviations of structure, depend +much more on the constitution of the organism than +on the nature of the conditions to which it has been +subjected.<a name="FNanchor_189" id="FNanchor_189"></a><a href="#Footnote_189" class="fnanchor">189</a></p> + +<p><i>Rate of Increase.</i>—Civilised populations have been +known under favourable conditions, as in the United +States, to double their number in twenty-five years; +and according to a calculation by Euler, this might +occur in a little over twelve years.<a name="FNanchor_190" id="FNanchor_190"></a><a href="#Footnote_190" class="fnanchor">190</a> At the former rate +the present population of the United States, namely, +thirty millions, would in 657 years cover the whole +terraqueous globe so thickly, that four men would have +to stand on each square yard of surface. The primary +or fundamental check to the continued increase of man +is the difficulty of gaining subsistence and of living in +comfort. We may infer that this is the case from what +we see, for instance, in the United States, where subsistence +is easy and there is plenty of room. If such means +were suddenly doubled in Great Britain, our number +would be quickly doubled. With civilised nations the +<span class='pagenum'><a name="Page_132" id="Page_132">132</a></span>above primary check acts chiefly by restraining marriages. +The greater death-rate of infants in the poorest +classes is also very important; as well as the greater +mortality at all ages, and from various diseases, of the +inhabitants of crowded and miserable houses. The +effects of severe epidemics and wars are soon counterbalanced, +and more than counterbalanced, in nations +placed under favourable conditions. Emigration also +comes in aid as a temporary check, but not to any +great extent with the extremely poor classes.</p> + +<p>There is reason to suspect, as Malthus has remarked, +that the reproductive power is actually less in barbarous +than in civilised races. We know nothing positively on +this head, for with savages no census has been taken; +but from the concurrent testimony of missionaries, and +of others who have long resided with such people, it +appears that their families are usually small, and large +ones rare. This may be partly accounted for, as it is +believed, by the women suckling their infants for a prolonged +period; but it is highly probable that savages, +who often suffer much hardship, and who do not obtain so +much nutritious food as civilised men, would be actually +less prolific. I have shewn in a former work,<a name="FNanchor_191" id="FNanchor_191"></a><a href="#Footnote_191" class="fnanchor">191</a> that +all our domesticated quadrupeds and birds, and all +our cultivated plants, are more fertile than the corresponding +species in a state of nature. It is no valid +objection to this conclusion that animals suddenly +supplied with an excess of food, or when rendered very +fat, and that most plants when suddenly removed from +very poor to very rich soil, are rendered more or less +sterile. We might, therefore, expect that civilised +men, who in one sense are highly domesticated, would +<span class='pagenum'><a name="Page_133" id="Page_133">133</a></span>be more prolific than wild men. It is also probable +that the increased fertility of civilised nations would +become, as with our domestic animals, an inherited +character: it is at least known that with mankind a +tendency to produce twins runs in families.<a name="FNanchor_192" id="FNanchor_192"></a><a href="#Footnote_192" class="fnanchor">192</a></p> + +<p>Notwithstanding that savages appear to be less prolific +than civilised people, they would no doubt rapidly +increase if their numbers were not by some means +rigidly kept down. The Santali, or hill-tribes of India, +have recently afforded a good illustration of this fact; +for they have increased, as shewn by Mr. Hunter,<a name="FNanchor_193" id="FNanchor_193"></a><a href="#Footnote_193" class="fnanchor">193</a> +at an extraordinary rate since vaccination has been +introduced, other pestilences mitigated, and war sternly +repressed. This increase, however, would not have been +possible had not these rude people spread into the +adjoining districts and worked for hire. Savages almost +always marry; yet there is some prudential restraint, +for they do not commonly marry at the earliest possible +age. The young men are often required to show that +they can support a wife, and they generally have first +to earn the price with which to purchase her from her +parents. With savages the difficulty of obtaining subsistence +occasionally limits their number in a much +more direct manner than with civilised people, for all +tribes periodically suffer from severe famines. At such +times savages are forced to devour much bad food, +and their health can hardly fail to be injured. Many +accounts have been published of their protruding stomachs +and emaciated limbs after and during famines. +They are then, also, compelled to wander much about, +and their infants, as I was assured in Australia, perish +<span class='pagenum'><a name="Page_134" id="Page_134">134</a></span>in large numbers. As famines are periodical, depending +chiefly on extreme seasons, all tribes must fluctuate in +number. They cannot steadily and regularly increase, +as there is no artificial increase in the supply of food. +Savages when hardly pressed encroach on each other’s +territories, and war is the result; but they are indeed +almost always at war with their neighbours. They are +liable to many accidents on land and water in their search +for food; and in some countries they must suffer much +from the larger beasts of prey. Even in India, districts +have been depopulated by the ravages of tigers.</p> + +<p>Malthus has discussed these several checks, but he +does not lay stress enough on what is probably the most +important of all, namely infanticide, especially of female +infants, and the habit of procuring abortion. These +practices now prevail in many quarters of the world, +and infanticide seems formerly to have prevailed, as +Mr. M’Lennan<a name="FNanchor_194" id="FNanchor_194"></a><a href="#Footnote_194" class="fnanchor">194</a> has shewn, on a still more extensive +scale. These practices appear to have originated in +savages recognising the difficulty, or rather the impossibility +of supporting all the infants that are born. +Licentiousness may also be added to the foregoing +checks; but this does not follow from failing means of +subsistence; though there is reason to believe that in +some cases (as in Japan) it has been intentionally +encouraged as a means of keeping down the population.</p> + +<p class="tb">If we look back to an extremely remote epoch, before +man had arrived at the dignity of manhood, he would +have been guided more by instinct and less by reason +than are savages at the present time. Our early semi-human +progenitors would not have practised infanticide, +for the instincts of the lower animals are never so perverted +as to lead them regularly to destroy their own +<span class='pagenum'><a name="Page_135" id="Page_135">135</a></span>offspring. There would have been no prudential restraint +from marriage, and the sexes would have freely +united at an early age. Hence the progenitors of +man would have tended to increase rapidly, but checks +of some kind, either periodical or constant, must have +kept down their numbers, even more severely than with +existing savages. What the precise nature of these +checks may have been, we cannot say, any more than +with most other animals. We know that horses and +cattle, which are not highly prolific animals, when first +turned loose in South America, increased at an enormous +rate. The slowest breeder of all known animals, namely +the elephant, would in a few thousand years stock the +whole world. The increase of every species of monkey +must be checked by some means; but not, as Brehm +remarks, by the attacks of beasts of prey. No one +will assume that the actual power of reproduction in +the wild horses and cattle of America, was at first in +any sensible degree increased; or that, as each district +became fully stocked, this same power was diminished. +No doubt in this case and in all others, many checks +concur, and different checks under different circumstances; +periodical dearths, depending on unfavourable +seasons, being probably the most important of all. So +it will have been with the early progenitors of man.</p> + +<p><i>Natural Selection.</i>—We have now seen that man is +variable in body and mind; and that the variations +are induced, either directly or indirectly, by the same +general causes, and obey the same general laws, as with +the lower animals. Man has spread widely over the +face of the earth, and must have been exposed, during +his incessant migrations,<a name="FNanchor_195" id="FNanchor_195"></a><a href="#Footnote_195" class="fnanchor">195</a> to the most diversified con<span class='pagenum'><a name="Page_136" id="Page_136">136</a></span>ditions. +The inhabitants of Tierra del Fuego, the Cape +of Good Hope, and Tasmania in the one hemisphere, +and of the Arctic regions in the other, must have passed +through many climates and changed their habits many +times, before they reached their present homes.<a name="FNanchor_196" id="FNanchor_196"></a><a href="#Footnote_196" class="fnanchor">196</a> The +early progenitors of man must also have tended, like all +other animals, to have increased beyond their means of +subsistence; they must therefore occasionally have been +exposed to a struggle for existence, and consequently to +the rigid law of natural selection. Beneficial variations +of all kinds will thus, either occasionally or habitually, +have been preserved, and injurious ones eliminated. I +do not refer to strongly-marked deviations of structure, +which occur only at long intervals of time, but to mere +individual differences. We know, for instance, that the +muscles of our hands and feet, which determine our +powers of movement, are liable, like those of the lower +animals,<a name="FNanchor_197" id="FNanchor_197"></a><a href="#Footnote_197" class="fnanchor">197</a> to incessant variability. If then the ape-like +progenitors of man which inhabited any district, especially +one undergoing some change in its conditions, were +divided into two equal bodies, the one half which included +all the individuals best adapted by their powers +of movement for gaining subsistence or for defending +themselves, would on an average survive in greater +number and procreate more offspring than the other +and less well endowed half.</p> + +<p>Man in the rudest state in which he now exists is +the most dominant animal that has ever appeared +on the earth. He has spread more widely than any +<span class='pagenum'><a name="Page_137" id="Page_137">137</a></span>other highly organised form; and all others have +yielded before him. He manifestly owes this immense +superiority to his intellectual faculties, his social habits, +which lead him to aid and defend his fellows, and to +his corporeal structure. The supreme importance of +these characters has been proved by the final arbitrament +of the battle for life. Through his powers of intellect, +articulate language has been evolved; and on +this his wonderful advancement has mainly depended. +He has invented and is able to use various weapons, +tools, traps, &c., with which he defends himself, kills or +catches prey, and otherwise obtains food. He has made +rafts or canoes on which to fish or cross over to neighbouring +fertile islands. He has discovered the art of +making fire, by which hard and stringy roots can be +rendered digestible, and poisonous roots or herbs innocuous. +This last discovery, probably the greatest, +excepting language, ever made by man, dates from +before the dawn of history. These several inventions, +by which man in the rudest state has become so preeminent, +are the direct result of the development of +his powers of observation, memory, curiosity, imagination, +and reason. I cannot, therefore, understand how +it is that Mr. Wallace<a name="FNanchor_198" id="FNanchor_198"></a><a href="#Footnote_198" class="fnanchor">198</a> maintains, that “natural selection +could only have endowed the savage with a brain +a little superior to that of an ape.”</p> +<p><span class='pagenum'><a name="Page_138" id="Page_138">138</a></span></p> +<p>Although the intellectual powers and social habits of +man are of paramount importance to him, we must not +underrate the importance of his bodily structure, to which +subject the remainder of this chapter will be devoted. +The development of the intellectual and social or moral +faculties will be discussed in the following chapter.</p> + +<p>Even to hammer with precision is no easy matter, as +every one who has tried to learn carpentry will admit. +To throw a stone with as true an aim as can a Fuegian in +defending himself, or in killing birds, requires the most +consummate perfection in the correlated action of the +muscles of the hand, arm, and shoulder, not to mention +a fine sense of touch. In throwing a stone or spear, and +in many other actions, a man must stand firmly on his +feet; and this again demands the perfect coadaptation of +numerous muscles. To chip a flint into the rudest tool, +or to form a barbed spear or hook from a bone, demands +the use of a perfect hand; for, as a most capable judge, +Mr. Schoolcraft,<a name="FNanchor_199" id="FNanchor_199"></a><a href="#Footnote_199" class="fnanchor">199</a> remarks, the shaping fragments of +stone into knives, lances, or arrow-heads, shews “extraordinary +ability and long practice.” We have evidence +of this in primeval men having practised a division of +labour; each man did not manufacture his own flint +tools or rude pottery; but certain individuals appear to +have devoted themselves to such work, no doubt receiving +in exchange the produce of the chase. Archæologists +are convinced that an enormous interval of time +<span class='pagenum'><a name="Page_139" id="Page_139">139</a></span>elapsed before our ancestors thought of grinding chipped +flints into smooth tools. A man-like animal who possessed +a hand and arm sufficiently perfect to throw a +stone with precision or to form a flint into a rude tool, +could, it can hardly be doubted, with sufficient practice make +almost anything, as far as mechanical skill alone +is concerned, which a civilised man can make. The +structure of the hand in this respect may be compared +with that of the vocal organs, which in the apes are +used for uttering various signal-cries, or, as in one species, +musical cadences; but in man closely similar vocal +organs have become adapted through the inherited +effects of use for the utterance of articulate language.</p> + +<p>Turning now to the nearest allies of man, and therefore +to the best representatives of our early progenitors, +we find that the hands in the Quadrumana are constructed +on the same general pattern as in us, but are +far less perfectly adapted for diversified uses. Their +hands do not serve so well as the feet of a dog for locomotion; +as may be seen in those monkeys which walk +on the outer margins of the palms, or on the backs of +their bent fingers, as in the chimpanzee and orang.<a name="FNanchor_200" id="FNanchor_200"></a><a href="#Footnote_200" class="fnanchor">200</a> +Their hands, however, are admirably adapted for climbing +trees. Monkeys seize thin branches or ropes, with +the thumb on one side and the fingers and palm on +the other side, in the same manner as we do. They +can thus also carry rather large objects, such as the +neck of a bottle, to their mouths. Baboons turn over +stones and scratch up roots with their hands. They +seize nuts, insects, or other small objects with the +thumb in opposition to the fingers, and no doubt they +thus extract eggs and the young from the nests of +birds. American monkeys beat the wild oranges on the +<span class='pagenum'><a name="Page_140" id="Page_140">140</a></span>branches until the rind is cracked, and then tear it off +with the fingers of the two hands. Other monkeys open +mussel-shells with the two thumbs. With their fingers +they pull out thorns and burrs, and hunt for each other’s +parasites. In a state of nature they break open hard +fruits with the aid of stones. They roll down stones +or throw them at their enemies; nevertheless, they +perform these various actions clumsily, and they are +quite unable, as I have myself seen, to throw a stone +with precision.</p> + +<p>It seems to me far from true that because “objects +are grasped clumsily” by monkeys, “a much less +specialised organ of prehension” would have served +them<a name="FNanchor_201" id="FNanchor_201"></a><a href="#Footnote_201" class="fnanchor">201</a> as well as their present hands. On the contrary, +I see no reason to doubt that a more perfectly +constructed hand would have been an advantage to them, +provided, and it is important to note this, that their +hands had not thus been rendered less well adapted for +climbing trees. We may suspect that a perfect hand +would have been disadvantageous for climbing; as the +most arboreal monkeys in the world, namely Ateles in +America and Hylobates in Asia, either have their thumbs +much reduced in size and even rudimentary, or their +fingers partially coherent, so that their hands are converted +into mere grasping-hooks.<a name="FNanchor_202" id="FNanchor_202"></a><a href="#Footnote_202" class="fnanchor">202</a></p> + +<p>As soon as some ancient member in the great series +of the Primates came, owing to a change in its manner +of procuring subsistence, or to a change in the conditions +of its native country, to live somewhat less on +trees and more on the ground, its manner of progression +would have been modified; and in this case it +<span class='pagenum'><a name="Page_141" id="Page_141">141</a></span>would have had to become either more strictly quadrupedal +or bipedal. Baboons frequent hilly and rocky +districts, and only from necessity climb up high trees;<a name="FNanchor_203" id="FNanchor_203"></a><a href="#Footnote_203" class="fnanchor">203</a> +and they have acquired almost the gait of a dog. Man +alone has become a biped; and we can, I think, partly +see how he has come to assume his erect attitude, which +forms one of the most conspicuous differences between +him and his nearest allies. Man could not have +attained his present dominant position in the world +without the use of his hands which are so admirably +adapted to act in obedience to his will. As Sir C. Bell<a name="FNanchor_204" id="FNanchor_204"></a><a href="#Footnote_204" class="fnanchor">204</a> +insists “the hand supplies all instruments, and by its +correspondence with the intellect gives him universal +dominion.” But the hands and arms could hardly +have become perfect enough to have manufactured +weapons, or to have hurled stones and spears with a +true aim, as long as they were habitually used for +locomotion and for supporting the whole weight of the +body, or as long as they were especially well adapted, +as previously remarked, for climbing trees. Such rough +treatment would also have blunted the sense of touch, +on which their delicate use largely depends. From +these causes alone it would have been an advantage to +man to have become a biped; but for many actions it is +almost necessary that both arms and the whole upper +part of the body should be free; and he must for this +end stand firmly on his feet. To gain this great +advantage, the feet have been rendered flat, and the +great toe peculiarly modified, though this has entailed +the loss of the power of prehension. It accords with +the principle of the division of physiological labour, +which prevails throughout the animal kingdom, that +<span class='pagenum'><a name="Page_142" id="Page_142">142</a></span>as the hands became perfected for prehension, the +feet should have become perfected for support and +locomotion. With some savages, however, the foot has +not altogether lost its prehensile power, as shewn by +their manner of climbing trees and of using them in +other ways.<a name="FNanchor_205" id="FNanchor_205"></a><a href="#Footnote_205" class="fnanchor">205</a></p> + +<p>If it be an advantage to man to have his hands and +arms free and to stand firmly on his feet, of which there +can be no doubt from his preeminent success in the +battle of life, then I can see no reason why it should +not have been advantageous to the progenitors of man +to have become more and more erect or bipedal. They +would thus have been better able to have defended +themselves with stones or clubs, or to have attacked +their prey, or otherwise obtained food. The best constructed +individuals would in the long run have succeeded +best, and have survived in larger numbers. If the +gorilla and a few allied forms had become extinct, it +might have been argued with great force and apparent +truth, that an animal could not have been gradually +converted from a quadruped into a biped; as all the +individuals in an intermediate condition would have +been miserably ill-fitted for progression. But we know +(and this is well worthy of reflection) that several kinds +of apes are now actually in this intermediate condition; +and no one doubts that they are on the whole well +adapted for their conditions of life. Thus the gorilla +runs with a sidelong shambling gait, but more commonly +<span class='pagenum'><a name="Page_143" id="Page_143">143</a></span>progresses by resting on its bent hands. The long-armed +apes occasionally use their arms like crutches, +swinging their bodies forward between them, and some +kinds of Hylobates, without having been taught, can +walk or run upright with tolerable quickness; yet they +move awkwardly, and much less securely than man. +We see, in short, with existing monkeys various gradations +between a form of progression strictly like that of +a quadruped and that of a biped or man.</p> + +<p>As the progenitors of man became more and more +erect, with their hands and arms more and more modified +for prehension and other purposes, with their feet +and legs at the same time modified for firm support +and progression, endless other changes of structure +would have been necessary. The pelvis would have +had to be made broader, the spine peculiarly curved +and the head fixed in an altered position, and all these +changes have been attained by man. Prof. Schaaffhausen<a name="FNanchor_206" id="FNanchor_206"></a><a href="#Footnote_206" class="fnanchor">206</a> +maintains that “the powerful mastoid processes +of the human skull are the result of his erect position;” +and these processes are absent in the orang, chimpanzee, +&c., and are smaller in the gorilla than in man. +Various other structures might here have been specified, +which appear connected with man’s erect position. It +is very difficult to decide how far all these correlated +modifications are the result of natural selection, and +how far of the inherited effects of the increased use of +certain parts, or of the action of one part on another. +No doubt these means of change act and react on each +other: thus when certain muscles, and the crests of +bone to which they are attached, become enlarged by +<span class='pagenum'><a name="Page_144" id="Page_144">144</a></span>habitual use, this shews that certain fictions are +habitually performed and must be serviceable. Hence +the individuals which performed them best, would tend +to survive in greater numbers.</p> + +<p>The free use of the arms and hands, partly the cause +and partly the result of man’s erect position, appears to +have led in an indirect manner to other modifications of +structure. The early male progenitors of man were, as +previously stated, probably furnished with great canine +teeth; but as they gradually acquired the habit of using +stones, clubs, or other weapons, for fighting with their +enemies, they would have used their jaws and teeth less +and less. In this case, the jaws, together with the +teeth, would have become reduced in size, as we may +feel sure from innumerable analogous cases. In a future +chapter we shall meet with a closely-parallel case, in +the reduction or complete disappearance of the canine +teeth in male ruminants, apparently in relation with the +development of their horns; and in horses, in relation +with their habit of fighting with their incisor teeth and +hoofs.</p> + +<p>In the adult male anthropomorphous apes, as Rütimeyer,<a name="FNanchor_207" id="FNanchor_207"></a><a href="#Footnote_207" class="fnanchor">207</a> +and others have insisted, it is precisely the effect +which the jaw-muscles by their great development have +produced on the skull, that causes it to differ so greatly +in many respects from that of man, and has given to +it “a truly frightful physiognomy.” Therefore as the +jaws and teeth in the progenitors of man gradually +become reduced in size, the adult skull would have +presented nearly the same characters which it offers in +the young of the anthropomorphous apes, and would +thus have come to resemble more nearly that of existing +<span class='pagenum'><a name="Page_145" id="Page_145">145</a></span>man. A great reduction of the canine teeth in the +males would almost certainly, as we shall hereafter see, +have affected through inheritance the teeth of the +females.</p> + +<p>As the various mental faculties were gradually developed, +the brain would almost certainly have become +larger. No one, I presume, doubts that the large size +of the brain in man, relatively to his body, in comparison +with that of the gorilla or orang, is closely connected +with his higher mental powers. We meet with +closely analogous facts with insects, in which the cerebral +ganglia are of extraordinary dimensions in ants; these +ganglia in all the Hymenoptera being many times larger +than in the less intelligent orders, such as beetles.<a name="FNanchor_208" id="FNanchor_208"></a><a href="#Footnote_208" class="fnanchor">208</a> +On the other hand, no one supposes that the intellect of +any two animals or of any two men can be accurately +gauged by the cubic contents of their skulls. It is +certain that there may be extraordinary mental activity +with an extremely small absolute mass of nervous +matter: thus the wonderfully diversified instincts, +mental powers, and affections of ants are generally +known, yet their cerebral ganglia are not so large as the +quarter of a small pin’s head. Under this latter point +of view, the brain of an ant is one of the most marvellous +atoms of matter in the world, perhaps more marvellous +than the brain of man.</p> + +<p>The belief that there exists in man some close relation +between the size of the brain and the development of +the intellectual faculties is supported by the comparison +of the skulls of savage and civilised races, of ancient and +modern people, and by the analogy of the whole verte<span class='pagenum'><a name="Page_146" id="Page_146">146</a></span>brate +series. Dr. J. Barnard Davis has proved<a name="FNanchor_209" id="FNanchor_209"></a><a href="#Footnote_209" class="fnanchor">209</a> by +many careful measurements, that the mean internal +capacity of the skull in Europeans is 92·3 cubic inches; +in Americans 87·5; in Asiatics 87·1; and in Australians +only 81·9 inches. Professor Broca<a name="FNanchor_210" id="FNanchor_210"></a><a href="#Footnote_210" class="fnanchor">210</a> found that skulls +from graves in Paris of the nineteenth century, were +larger than those from vaults of the twelfth century, in +the proportion of 1484 to 1426; and Prichard is persuaded +that the present inhabitants of Britain have +“much more capacious brain-cases” than the ancient +inhabitants. Nevertheless it must be admitted that +some skulls of very high antiquity, such as the famous +one of Neanderthal, are well developed and capacious. +With respect to the lower animals, M. E. Lartet,<a name="FNanchor_211" id="FNanchor_211"></a><a href="#Footnote_211" class="fnanchor">211</a> by comparing +the crania of tertiary and recent mammals, belonging +to the same groups, has come to the remarkable +conclusion that the brain is generally larger and the +convolutions more complex in the more recent form. +On the other hand I have shewn<a name="FNanchor_212" id="FNanchor_212"></a><a href="#Footnote_212" class="fnanchor">212</a> that the brains of +domestic rabbits are considerably reduced in bulk, in +comparison with those of the wild rabbit or hare; and +this may be attributed to their having been closely confined +during many generations, so that they have exerted +but little their intellect, instincts, senses, and voluntary +movements.</p> + +<p>The gradually increasing weight of the brain and +skull in man must have influenced the development of +the supporting spinal column, more especially whilst +he was becoming erect. As this change of position was +<span class='pagenum'><a name="Page_147" id="Page_147">147</a></span>being brought about, the internal pressure of the brain, +will, also, have influenced the form of the skull; for +many facts shew how easily the skull is thus affected. +Ethnologists believe that it is modified by the kind of +cradle in which infants sleep. Habitual spasms of the +muscles and a cicatrix from a severe burn have permanently +modified the facial bones. In young persons +whose heads from disease have become fixed either +sideways or backwards, one of the eyes has changed +its position, and the bones of the skull have been +modified; and this apparently results from the brain +pressing in a new direction.<a name="FNanchor_213" id="FNanchor_213"></a><a href="#Footnote_213" class="fnanchor">213</a> I have shewn that with +long-eared rabbits, even so trifling a cause as the lopping +forward of one ear drags forward on that side almost +every bone of the skull; so that the bones on the opposite +sides no longer strictly correspond. Lastly, if any +animal were to increase or diminish much in general +size, without any change in its mental powers; or if +the mental powers were to be much increased or +diminished without any great change in the size of the +body; the shape of the skull would almost certainly be +altered. I infer this from my observations on domestic +rabbits, some kinds of which have become very much +larger than the wild animal, whilst others have retained +nearly the same size, but in both cases the brain has +been much reduced relatively to the size of the body. +Now I was at first much surprised by finding that in all +these rabbits the skull had become elongated or dolicho<span class='pagenum'><a name="Page_148" id="Page_148">148</a></span>cephalic; +for instance, of two skulls of nearly equal +breadth, the one from a wild rabbit and the other from +a large domestic kind, the former was only 3·15 and the +latter 4·3 inches in length.<a name="FNanchor_214" id="FNanchor_214"></a><a href="#Footnote_214" class="fnanchor">214</a> One of the most marked +distinctions in different races of man is that the skull +in some is elongated, and in others rounded; and here +the explanation suggested by the case of the rabbits +may partially hold good; for Welcker finds that short +“men incline more to brachycephaly, and tall men to +dolichocephaly;”<a name="FNanchor_215" id="FNanchor_215"></a><a href="#Footnote_215" class="fnanchor">215</a> and tall men may be compared with +the larger and longer-bodied rabbits, all of which have +elongated skulls, or are dolichocephalic.</p> + +<p>From these several facts we can to a certain extent +understand the means through which the great +size and more or less rounded form of the skull has +been acquired by man; and these are characters eminently +distinctive of him in comparison with the lower +animals.</p> + +<p>Another most conspicuous difference between man and +the lower animals is the nakedness of his skin. Whales +and dolphins (Cetacea), dugongs (Sirenia) and the hippopotamus +are naked; and this may be advantageous +to them for gliding through the water; nor would it +be injurious to them from the loss of warmth, as the +species which inhabit the colder regions are protected +by a thick layer of blubber, serving the same purpose +as the fur of seals and otters. Elephants and rhinoceroses +are almost hairless; and as certain extinct +species which formerly lived under an arctic climate +were covered with long wool or hair, it would almost +appear as if the existing species of both genera had lost +<span class='pagenum'><a name="Page_149" id="Page_149">149</a></span>their hairy covering from exposure to heat. This appears +the more probable, as the elephants in India which live +on elevated and cool districts are more hairy<a name="FNanchor_216" id="FNanchor_216"></a><a href="#Footnote_216" class="fnanchor">216</a> than those +on the lowlands. May we then infer that man became +divested of hair from having aboriginally inhabited some +tropical land? The fact of the hair being chiefly +retained in the male sex on the chest and face, and in +both sexes at the junction of all four limbs with the +trunk, favours this inference, assuming that the hair was +lost before man became erect; for the parts which now +retain most hair would then have been most protected +from the heat of the sun. The crown of the head, +however, offers a curious exception, for at all times it +must have been one of the most exposed parts, yet +it is thickly clothed with hair. In this respect man +agrees with the great majority of quadrupeds, which +generally have their upper and exposed surfaces more +thickly clothed than the lower surface. Nevertheless, +the fact that the other members of the order of Primates, +to which man belongs, although inhabiting various +hot regions, are well clothed with hair, generally +thickest on the upper surface,<a name="FNanchor_217" id="FNanchor_217"></a><a href="#Footnote_217" class="fnanchor">217</a> is strongly opposed +to the supposition that man became naked through the +action of the sun. I am inclined to believe, as we +shall see under sexual selection, that man, or rather +primarily woman, became divested of hair for ornamental +purposes; and according to this belief it is not +<span class='pagenum'><a name="Page_150" id="Page_150">150</a></span>surprising that man should differ so greatly in hairiness +from all his lower brethren, for characters gained +through sexual selection often differ in closely-related +forms to an extraordinary degree.</p> + +<p class="tb">According to a popular impression, the absence of a +tail is eminently distinctive of man; but as those apes +which come nearest to man are destitute of this organ, +its disappearance does not especially concern us. Nevertheless +it may be well to own that no explanation, as +far as I am aware, has ever been given of the loss of the +tail by certain apes and man. Its loss, however, is not +surprising, for it sometimes differs remarkably in length +in species of the same genera: thus in some species +of Macacus the tail is longer than the whole body, consisting +of twenty-four vertebræ; in others it consists of a +scarcely visible stump, containing only three or four +vertebræ. In some kinds of baboons there are twenty-five, +whilst in the mandrill there are ten very small +stunted caudal vertebræ, or, according to Cuvier,<a name="FNanchor_218" id="FNanchor_218"></a><a href="#Footnote_218" class="fnanchor">218</a> sometimes +only five. This great diversity in the structure and +length of the tail in animals belonging to the same genera, +and following nearly the same habits of life, renders it +probable that the tail is not of much importance to +them; and if so, we might have expected that it would +sometimes have become more or less rudimentary, in +accordance with what we incessantly see with other structures. +The tail almost always tapers towards the end +whether it be long or short; and this, I presume, results +from the atrophy, through disuse, of the terminal +muscles together with their arteries and nerves, leading +to the atrophy of the terminal bones. With respect +<span class='pagenum'><a name="Page_151" id="Page_151">151</a></span>to the os coccyx, which in man and the higher apes +manifestly consists of the few basal and tapering segments +of an ordinary tail, I have heard it asked how +could these have become completely embedded within +the body; but there is no difficulty in this respect, +for in many monkeys the basal segments of the true +tail are thus embedded. For instance, Mr. Murie informs +me that in the skeleton of a not full-grown +<i>Macacus inornatus</i>, he counted nine or ten caudal vertebræ, +which altogether were only 1·8 inch in length. +Of these the three basal ones appeared to have been +embedded; the remainder forming the free part of the +tail, which was only one inch in length, and half an +inch in diameter. Here, then, the three embedded +caudal vertebræ plainly correspond with the four coalesced +vertebræ of the human os coccyx.</p> + +<p>I have now endeavoured to shew that some of the +most distinctive characters of man have in all probability +been acquired, either directly, or more commonly +indirectly, through natural selection. We should bear +in mind that modifications in structure or constitution, +which are of no service to an organism in adapting +it to its habits of life, to the food which it consumes, +or passively to the surrounding conditions, cannot +have been thus acquired. We must not, however, +be too confident in deciding what modifications are of +service to each being: we should remember how little +we know about the use of many parts, or what changes +in the blood or tissues may serve to fit an organism for +a new climate or some new kind of food. Nor must +we forget the principle of correlation, by which, as +Isidore Geoffroy has shewn in the case of man, many +fit-range deviations of structure are tied together. Independently +of correlation, a change in one part often leads<span class='pagenum'><a name="Page_152" id="Page_152">152</a></span> +through the increased or decreased use of other parts, +to other changes of a quite unexpected nature. It is +also well to reflect on such facts, as the wonderful +growth of galls on plants caused by the poison of an +insect, and on the remarkable changes of colour in the +plumage of parrots when fed on certain fishes, or inoculated +with the poison of toads;<a name="FNanchor_219" id="FNanchor_219"></a><a href="#Footnote_219" class="fnanchor">219</a> for we can thus +see that the fluids of the system, if altered for some +special purpose, might induce other strange changes. +We should especially bear in mind that modifications +acquired and continually used during past ages for +some useful purpose would probably become firmly +fixed and might be long inherited.</p> + +<p>Thus a very large yet undefined extension may safely +be given to the direct and indirect results of natural +selection; but I now admit, after reading the essay by +Nägeli on plants, and the remarks by various authors +with respect to animals, more especially those recently +made by Professor Broca, that in the earlier editions of +my ‘Origin of Species’ I probably attributed too much +to the action of natural selection or the survival of the +fittest. I have altered the fifth edition of the Origin +so as to confine my remarks to adaptive changes of +structure. I had not formerly sufficiently considered +the existence of many structures which appear to be, +as far as we can judge, neither beneficial nor injurious; +and this I believe to be one of the greatest oversights as +yet detected in my work. I may be permitted to say +as some excuse, that I had two distinct objects in view, +firstly, to shew that species had not been separately +created, and secondly, that natural selection had been +the chief agent of change, though largely aided by the +<span class='pagenum'><a name="Page_153" id="Page_153">153</a></span>inherited effects of habit, and slightly by the direct +action of the surrounding conditions. Nevertheless I +was not able to annul the influence of my former +belief, then widely prevalent, that each species had +been purposely created; and this led to my tacitly +assuming that every detail of structure, excepting rudiments, +was of some special, though unrecognised, service. +Any one with this assumption in his mind would +naturally extend the action of natural selection, either +during past or present times, too far. Some of those +who admit the principle of evolution, but reject natural +selection, seem to forget, when criticising my book, that +I had the above two objects in view; hence if I have +erred in giving to natural selection great power, which +I am far from admitting, or in having exaggerated its +power, which is in itself probable, I have at least, as I +hope, done good service in aiding to overthrow the +dogma of separate creations.</p> + +<p class="tb">That all organic beings, including man, present many +modifications of structure which are of no service to +them at present, nor have been formerly, is, as I can +now see, probable. We know not what produces the +numberless slight differences between the individuals of +each species, for reversion only carries the problem a few +steps backwards; but each peculiarity must have had +its own efficient cause. If these causes, whatever they +may be, were to act more uniformly and energetically +during a lengthened period (and no reason can be +assigned why this should not sometimes occur), the +result would probably be not mere slight individual +differences, but well-marked, constant modifications. +Modifications which are in no way beneficial cannot +have been kept uniform through natural selection, +though any which were injurious would have been thus +eliminated. Uniformity of character would, however,<span class='pagenum'><a name="Page_154" id="Page_154">154</a></span> +naturally follow from, the assumed uniformity of the +exciting causes, and likewise from the free intercrossing +of many individuals. The same organism might +acquire in this manner during successive periods successive +modifications, and these would be transmitted +in a nearly uniform state as long as the exciting causes +remained the same and there was free intercrossing. +With respect to the exciting causes we can only say, as +when speaking of so-called spontaneous variations, that +they relate much more closely to the constitution of the +varying organism, than to the nature of the conditions +to which it has been subjected.</p> + +<p><i>Conclusion.</i>—In this chapter we have seen that as man +at the present day is liable, like every other animal, to +multiform individual differences or slight variations, so +no doubt were the early progenitors of man; the variations +being then as now induced by the same general +causes, and governed by the same general and complex +laws. As all animals tend to multiply beyond their +means of subsistence, so it must have been with the +progenitors of man; and this will inevitably have led +to a struggle for existence and to natural selection. +This latter process will have been greatly aided by +the inherited effects of the increased use of parts; +these two processes incessantly reacting on each other. +It appears, also, as we shall hereafter see, that various +unimportant characters have been acquired by man +through sexual selection. An unexplained residuum +of change, perhaps a large one, must be left to the +assumed uniform action of those unknown agencies, +which occasionally induce strongly-marked and abrupt +deviations of structure in our domestic productions.</p> + +<p>Judging from the habits of savages and of the greater +number of the Quadrumana, primeval men, and even<span class='pagenum'><a name="Page_155" id="Page_155">155</a></span> +the ape-like progenitors of man, probably lived in +society. With strictly social animals, natural selection +sometimes acts indirectly on the individual, through +the preservation of variations which are beneficial only +to the community. A community including a large +number of well-endowed individuals increases in number +and is victorious over other and less well-endowed communities; +although each separate member may gain no +advantage over the other members of the same community. +With associated insects many remarkable +structures, which are of little or no service to the individual +or its own offspring, such as the pollen-collecting +apparatus, or the sting of the worker-bee, or the +great jaws of soldier-ants, have been thus acquired. +With the higher social animals, I am not aware that +any structure has been modified solely for the good of +the community, though some are of secondary service +to it. For instance, the horns of ruminants and the +great canine teeth of baboons appear to have been +acquired by the males as weapons for sexual strife, but +they are used in defence of the herd or troop. In +regard to certain mental faculties the case, as we shall +see in the following chapter, is wholly different; for +these faculties have been chiefly, or even exclusively, +gained for the benefit of the community; the individuals +composing the community being at the same +time indirectly benefited.</p> + +<p>It has often been objected to such views as the foregoing, +that man is one of the most helpless and defenceless +creatures in the world; and that during his early +and less well-developed condition he would have been +still more helpless. The Duke of Argyll, for instance, +insists<a name="FNanchor_220" id="FNanchor_220"></a><a href="#Footnote_220" class="fnanchor">220</a> that “the human frame has diverged from +<span class='pagenum'><a name="Page_156" id="Page_156">156</a></span>the structure of brutes, in the direction of greater +physical helplessness and weakness. That is to say, +it is a divergence which of all others it is most +impossible to ascribe to mere natural selection.” He +adduces the naked and unprotected state of the body, +the absence of great teeth or claws for defence, the +little strength of man, his small speed in running, and +his slight power of smell, by which to discover food or +to avoid danger. To these deficiencies there might +have been added the still more serious loss of the power +of quickly climbing trees, so as to escape from enemies. +Seeing that the unclothed Fuegians can exist under +their wretched climate, the loss of hair would not +have been a great injury to primeval man, if he inhabited +a warm country. When we compare defenceless +man with the apes, many of which are provided with +formidable canine teeth, we must remember that these +in their fully-developed condition are possessed by the +males alone, being chiefly used by them for fighting +with their rivals; yet the females which are not thus +provided, are able to survive.</p> + +<p>In regard to bodily size or strength, we do not know +whether man is descended from some comparatively +small species, like the chimpanzee, or from one as +powerful as the gorilla; and, therefore, we cannot say +whether man has become larger and stronger, or smaller +and weaker, in comparison with his progenitors. We +should, however, bear in mind that an animal possessing +great size, strength, and ferocity, and which, like the +gorilla, could defend itself from all enemies, would +probably, though not necessarily, have failed to become +social; and this would most effectually have checked +the acquirement by man of his higher mental qualities, +such as sympathy and the love of his fellow-creatures. +Hence it might have been an immense<span class='pagenum'><a name="Page_157" id="Page_157">157</a></span> +advantage to man to have sprung from some comparatively +weak creature.</p> + +<p>The slight corporeal strength of man, his little speed, +his want of natural weapons, &c., are more than counterbalanced, +firstly by his intellectual powers, through +which he has, whilst still remaining in a barbarous state, +formed for himself weapons, tools, &c., and secondly by +his social qualities which lead him to give aid to his +fellow-men and to receive it in return. No country +in the world abounds in a greater degree with dangerous +beasts than Southern Africa; no country presents +more fearful physical hardships than the Arctic +regions; yet one of the puniest races, namely, the +Bushmen, maintain themselves in Southern Africa, as +do the dwarfed Esquimaux in the Arctic regions. The +early progenitors of man were, no doubt, inferior in +intellect, and probably in social disposition, to the +lowest existing savages; but it is quite conceivable that +they might have existed, or even flourished, if, whilst +they gradually lost their brute-like powers, such as +climbing trees, &c., they at the same time advanced +in intellect. But granting that the progenitors of man +were far more helpless and defenceless than any existing +savages, if they had inhabited some warm continent +or large island, such as Australia or New Guinea, or +Borneo (the latter island being now tenanted by the +orang), they would not have been exposed to any special +danger. In an area as large as one of these islands, +the competition between tribe and tribe would have +been sufficient, under favourable conditions, to have +raised man, through the survival of the fittest, combined +with the inherited effects of habit, to his present high +position in the organic scale.</p> + +<hr /> +<p><span class='pagenum'><a name="Page_158" id="Page_158">158</a></span></p> + +<h3>CHAPTER V.</h3> + +<h4><span class="smcap">On the Development of the Intellectual and Moral +Faculties during Primeval and Civilised Times.</span></h4> + +<div class="blockquot"><p>The advancement of the intellectual powers through natural selection—Importance +of imitation—Social and moral faculties—Their +development within the limits of the same tribe—Natural +selection as affecting civilised nations—Evidence that civilised +nations were once barbarous.</p></div> + +<p>The subjects to be discussed in this chapter are of +the highest interest, but are treated by me in a most +imperfect and fragmentary manner. Mr. Wallace, +in an admirable paper before referred to,<a name="FNanchor_221" id="FNanchor_221"></a><a href="#Footnote_221" class="fnanchor">221</a> argues that +man after he had partially acquired those intellectual +and moral faculties which distinguish him from the +lower animals, would have been but little liable to +have had his bodily structure modified through natural +selection or any other means. For man is enabled +through his mental faculties “to keep with an unchanged +body in harmony with the changing universe.” +He has great power of adapting his habits to new +conditions of life. He invents weapons, tools and +various stratagems, by which he procures food and +defends himself. When he migrates into a colder +climate he uses clothes, builds sheds, and makes fires; +and, by the aid of fire, cooks food otherwise indigestible. +He aids his fellow-men in many ways, and anticipates +future events. Even at a remote period he practised +some subdivision of labour.</p> +<p><span class='pagenum'><a name="Page_159" id="Page_159">159</a></span></p> +<p>The lower animals, on the other hand, must have +their bodily structure modified in order to survive under +greatly changed conditions. They must be rendered +stronger, or acquire more effective teeth or claws, in +order to defend themselves from new enemies; or they +must be reduced in size so as to escape detection and +danger. When they migrate into a colder climate they +must become clothed with thicker fur, or have their +constitutions altered. If they fail to be thus modified, +they will cease to exist.</p> + +<p>The case, however, is widely different, as Mr. Wallace +has with justice insisted, in relation to the intellectual +and moral faculties of man. These faculties are +variable; and we have every reason to believe that the +variations tend to be inherited. Therefore, if they were +formerly of high importance to primeval man and to +his ape-like progenitors, they would have been perfected +or advanced through natural selection. Of the +high importance of the intellectual faculties there can +be no doubt, for man mainly owes to them his preeminent +position in the world. We can see that, in +the rudest state of society, the individuals who were the +most sagacious, who invented and used the best weapons +or traps, and who were best able to defend themselves, +would rear the greatest number of offspring. The tribes +which included the largest number of men thus endowed +would increase in number and supplant other tribes. +Numbers depend primarily on the means of subsistence, +and this, partly on the physical nature of the country, +but in a much higher degree on the arts which are there +practised. As a tribe increases and is victorious, it is +often still further increased by the absorption of other +tribes.<a name="FNanchor_222" id="FNanchor_222"></a><a href="#Footnote_222" class="fnanchor">222</a> The stature and strength of the men of a tribe +<span class='pagenum'><a name="Page_160" id="Page_160">160</a></span>are likewise of some importance for its success, and +these depend in part on the nature and amount of the +food which can be obtained. In Europe the men of the +Bronze period were supplanted by a more powerful and, +judging from their sword-handles, larger-handed race;<a name="FNanchor_223" id="FNanchor_223"></a><a href="#Footnote_223" class="fnanchor">223</a> +but their success was probably due in a much higher +degree to their superiority in the arts.</p> + +<p>All that we know about savages, or may infer from +their traditions and from old monuments, the history +of which is quite forgotten by the present inhabitants, +shew that from the remotest times successful tribes have +supplanted other tribes. Relics of extinct or forgotten +tribes have been discovered throughout the civilised +regions of the earth, on the wild plains of America, and +on the isolated islands in the Pacific Ocean. At the +present day civilised nations are everywhere supplanting +barbarous nations, excepting where the climate opposes +a deadly barrier; and they succeed mainly, though not +exclusively, through their arts, which are the products +of the intellect. It is, therefore, highly probable that +with mankind the intellectual faculties have been +gradually perfected through natural selection; and this +conclusion is sufficient for our purpose. Undoubtedly +it would have been very interesting to have traced the +development of each separate faculty from the state in +which it exists in the lower animals to that in which it +exists in man; but neither my ability nor knowledge +permit the attempt.</p> + +<p>It deserves notice that as soon as the progenitors of +man became social (and this probably occurred at a +very early period), the advancement of the intellectual +faculties will have been aided and modified in an +important manner, of which we see only traces in +<span class='pagenum'><a name="Page_161" id="Page_161">161</a></span>the lower animals, namely, through the principle +of imitation, together with reason and experience. +Apes are much given to imitation, as are the lowest +savages; and the simple fact previously referred to, +that after a time no animal can be caught in the same +place by the same sort of trap, shews that animals learn +by experience, and imitate each others’ caution. Now, +if some one man in a tribe, more sagacious than the +others, invented a new snare or weapon, or other means +of attack or defence, the plainest self-interest, without +the assistance of much reasoning power, would prompt +the other members to imitate him; and all would thus +profit. The habitual practice of each new art must +likewise in some slight degree strengthen the intellect. +If the new invention were an important one, the tribe +would increase in number, spread, and supplant other +tribes. In a tribe thus rendered more numerous there +would always be a rather better chance of the birth of +other superior and inventive members. If such men +left children to inherit their mental superiority, the +chance of the birth of still more ingenious members +would be somewhat better, and in a very small tribe +decidedly better. Even if they left no children, the +tribe would still include their blood-relations; and it has +been ascertained by agriculturists<a name="FNanchor_224" id="FNanchor_224"></a><a href="#Footnote_224" class="fnanchor">224</a> that by preserving +and breeding from the family of an animal, which +when slaughtered was found to be valuable, the desired +character has been obtained.</p> + +<p>Turning now to the social and moral faculties. In +order that primeval men, or the ape-like progenitors +of man, should have become social, they must have +<span class='pagenum'><a name="Page_162" id="Page_162">162</a></span>acquired the same instinctive feelings which impel other +animals to live in a body; and they no doubt exhibited +the same general disposition. They would have +felt uneasy when separated from their comrades, for +whom they would have felt some degree of love; they +would have warned each other of danger, and have +given mutual aid in attack or defence. All this implies +some degree of sympathy, fidelity, and courage. Such +social qualities, the paramount importance of which +to the lower animals is disputed by no one, were no +doubt acquired by the progenitors of man in a similar +manner, namely, through natural selection, aided by +inherited habit. When two tribes of primeval man, +living in the same country, came into competition, +if the one tribe included (other circumstances being +equal) a greater number of courageous, sympathetic, +and faithful members, who were always ready to warn +each other of danger, to aid and defend each other, this +tribe would without doubt succeed best and conquer the +other. Let it be borne in mind how all-important, in +the never-ceasing wars of savages, fidelity and courage +must be. The advantage which disciplined soldiers +have over undisciplined hordes follows chiefly from the +confidence which each man feels in his comrades. +Obedience, as Mr. Bagehot has well shewn,<a name="FNanchor_225" id="FNanchor_225"></a><a href="#Footnote_225" class="fnanchor">225</a> is of the +highest value, for any form of government is better +than none. Selfish and contentious people will not +cohere, and without coherence nothing can be effected. +A tribe possessing the above qualities in a high degree +would spread and be victorious over other tribes; +but in the course of time it would, judging from all +past history, be in its turn overcome by some other +<span class='pagenum'><a name="Page_163" id="Page_163">163</a></span>and still more highly endowed tribe. Thus the social +and moral qualities would tend slowly to advance and +be diffused throughout the world.</p> + +<p>But it may be asked, how within the limits of the +same tribe did a large number of members first become +endowed with these social and moral qualities, and +how was the standard of excellence raised? It is +extremely doubtful whether the offspring of the more +sympathetic and benevolent parents, or of those which +were the most faithful to their comrades, would be +reared in greater number than the children of selfish +and treacherous parents of the same tribe. He who +was ready to sacrifice his life, as many a savage has +been, rather than betray his comrades, would often leave +no offspring to inherit his noble nature. The bravest +men, who were always willing to come to the front in +war, and who freely risked their lives for others, would +on an average perish in larger number than other men. +Therefore it seems scarcely possible (bearing in mind +that we are not here speaking of one tribe being victorious +over another) that the number of men gifted +with such virtues, or that the standard of their excellence, +could be increased through natural selection, that +is, by the survival of the fittest.</p> + +<p>Although the circumstances which lead to an increase +in the number of men thus endowed within the same +tribe are too complex to be clearly followed out, we can +trace some of the probable steps. In the first place, as +the reasoning powers and foresight of the members +became improved, each man would soon learn from +experience that if he aided his fellow-men, he would +commonly receive aid in return. From this low motive +he might acquire the habit of aiding his fellows; and +the habit of performing benevolent actions certainly +strengthens the feeling of sympathy, which gives the<span class='pagenum'><a name="Page_164" id="Page_164">164</a></span> +first impulse to benevolent actions. Habits, moreover, +followed during many generations probably tend to be +inherited.</p> + +<p>But there is another and much more powerful stimulus +to the development of the social virtues, namely, +the praise and the blame of our fellow-men. The love +of approbation and the dread of infamy, as well as the +bestowal of praise or blame, are primarily due, as we +have seen in the third chapter, to the instinct of sympathy; +and this instinct no doubt was originally acquired, +like all the other social instincts, through natural selection. +At how early a period the progenitors of man, in +the course of their development, became capable of feeling +and being impelled by the praise or blame of their +fellow-creatures, we cannot, of course, say. But it appears +that even dogs appreciate encouragement, praise, and +blame. The rudest savages feel the sentiment of glory, +as they clearly show by preserving the trophies of their +prowess, by their habit of excessive boasting, and even +by the extreme care which they take of their personal +appearance and decorations; for unless they regarded +the opinion of their comrades, such habits would be +senseless.</p> + +<p>They certainly feel shame at the breach of some of +their lesser rules; but how far they experience remorse +is doubtful. I was at first surprised that I could not recollect +any recorded instances of this feeling in savages; +and Sir J. Lubbock<a name="FNanchor_226" id="FNanchor_226"></a><a href="#Footnote_226" class="fnanchor">226</a> states that he knows of none. +But if we banish from our minds all cases given in +novels and plays and in death-bed confessions made +to priests, I doubt whether many of us have actually +witnessed remorse; though we may have often seen +shame and contrition for smaller offences. Remorse is +<span class='pagenum'><a name="Page_165" id="Page_165">165</a></span>a deeply hidden feeling. It is incredible that a savage, +who will sacrifice his life rather than betray his tribe, +or one who will deliver himself up as a prisoner rather +than break his parole,<a name="FNanchor_227" id="FNanchor_227"></a><a href="#Footnote_227" class="fnanchor">227</a> would not feel remorse in his +inmost soul, though he might conceal it, if he had failed +in a duty which he held sacred.</p> + +<p>We may therefore conclude that primeval man, at a +very remote period, would have been influenced by the +praise and blame of his fellows. It is obvious, that the +members of the same tribe would approve of conduct +which appeared to them to be for the general good, and +would reprobate that which appeared evil. To do good +unto others—to do unto others as ye would they should +do unto you,—is the foundation-stone of morality. It +is, therefore, hardly possible to exaggerate the importance +during rude times of the love of praise and the +dread of blame. A man who was not impelled by any +deep, instinctive feeling, to sacrifice his life for the good +of others, yet was roused to such actions by a sense +of glory, would by his example excite the same wish +for glory in other men, and would strengthen by exercise +the noble feeling of admiration. He might thus +do far more good to his tribe than by begetting offspring +with a tendency to inherit his own high character.</p> + +<p>With increased experience and reason, man perceives +the more remote consequences of his actions, and the +self-regarding virtues, such as temperance, chastity, &c., +which during early times are, as we have before seen, +utterly disregarded, come to be highly esteemed or even +held sacred. I need not, however, repeat what I have +said on this head in the third chapter. Ultimately a +highly complex sentiment, having its first origin in the +<span class='pagenum'><a name="Page_166" id="Page_166">166</a></span>social instincts, largely guided by the approbation of +our fellow-men, ruled by reason, self-interest, and in +later times by deep religious feelings, confirmed by +instruction and habit, all combined, constitute our moral +sense or conscience.</p> + +<p>It must not be forgotten that although a high standard +of morality gives but a slight or no advantage to +each individual man and his children over the other +men of the same tribe, yet that an advancement in the +standard of morality and an increase in the number +of well-endowed men will certainly give an immense +advantage to one tribe over another. There can be no +doubt that a tribe including many members who, from +possessing in a high degree the spirit of patriotism, +fidelity, obedience, courage, and sympathy, were always +ready to give aid to each other and to sacrifice themselves +for the common good, would be victorious over +most other tribes; and this would be natural selection. +At all times throughout the world tribes have supplanted +other tribes; and as morality is one element +in their success, the standard of morality and the number +of well-endowed men will thus everywhere tend to +rise and increase.</p> + +<p class="tb">It is, however, very difficult to form any judgment why +one particular tribe and not another has been successful +and has risen in the scale of civilisation. Many savages +are in the same condition as when first discovered several +centuries ago. As Mr. Bagehot has remarked, we are +apt to look at progress as the normal rule in human +society; but history refutes this. The ancients did not +even entertain the idea; nor do the oriental nations at +the present day. According to another high authority, +Mr. Maine,<a name="FNanchor_228" id="FNanchor_228"></a><a href="#Footnote_228" class="fnanchor">228</a> “the greatest part of mankind has never +<span class='pagenum'><a name="Page_167" id="Page_167">167</a></span>shewn a particle of desire that its civil institutions +should be improved.” Progress seems to depend on +many concurrent favourable conditions, far too complex +to be followed out. But it has often been remarked, that +a cool climate from leading to industry and the various +arts has been highly favourable, or even indispensable +for this end. The Esquimaux, pressed by hard necessity, +have succeeded in many ingenious inventions, but their +climate has been too severe for continued progress. +Nomadic habits, whether over wide plains, or through +the dense forests of the tropics, or along the shores of +the sea, have in every case been highly detrimental. +Whilst observing the barbarous inhabitants of Tierra +del Fuego, it struck me that the possession of some +property, a fixed abode, and the union of many families +under a chief, were the indispensable requisites for +civilisation. Such habits almost necessitate the cultivation +of the ground; and the first steps in cultivation +would probably result, as I have elsewhere shewn,<a name="FNanchor_229" id="FNanchor_229"></a><a href="#Footnote_229" class="fnanchor">229</a> from +some such accident as the seeds of a fruit-tree falling +on a heap of refuse and producing an unusually fine +variety. The problem, however, of the first advance of +savages towards civilisation is at present much too difficult +to be solved.</p> + +<p><i>Natural Selection as affecting Civilised Nations.</i>—In +the last and present chapters I have considered the +advancement of man from a former semi-human condition +to his present state as a barbarian. But some +remarks on the agency of natural selection on civilised +nations may be here worth adding. This subject has +been ably discussed by Mr. W. R. Greg,<a name="FNanchor_230" id="FNanchor_230"></a><a href="#Footnote_230" class="fnanchor">230</a> and previously +<span class='pagenum'><a name="Page_168" id="Page_168">168</a></span>by Mr. Wallace and Mr. Galton.<a name="FNanchor_231" id="FNanchor_231"></a><a href="#Footnote_231" class="fnanchor">231</a> Most of my remarks +are taken from these three authors. With savages, the +weak in body or mind are soon eliminated; and those +that survive commonly exhibit a vigorous state of +health. We civilised men, on the other hand, do our +utmost to check the process of elimination; we build +asylums for the imbecile, the maimed, and the sick; we +institute poor-laws; and our medical men exert their +utmost skill to save the life of every one to the last +moment. There is reason to believe that vaccination +has preserved thousands, who from a weak constitution +would formerly have succumbed to small-pox. Thus +the weak members of civilised societies propagate their +kind. No one who has attended to the breeding of +domestic animals will doubt that this must be highly +injurious to the race of man. It is surprising how soon +a want of care, or care wrongly directed, leads to the +degeneration of a domestic race; but excepting in the +case of man himself, hardly any one is so ignorant as +to allow his worst animals to breed.</p> + +<p>The aid which we feel impelled to give to the helpless +is mainly an incidental result of the instinct of +sympathy, which was originally acquired as part of +the social instincts, but subsequently rendered, in the +manner previously indicated, more tender and more +widely diffused. Nor could we check our sympathy, if +so urged by hard reason, without deterioration in the +<span class='pagenum'><a name="Page_169" id="Page_169">169</a></span>noblest part of our nature. The surgeon may harden +himself whilst performing an operation, for he knows +that he is acting for the good of his patient; but if we +were intentionally to neglect the weak and helpless, it +could only be for a contingent benefit, with a certain +and great present evil. Hence we must bear without +complaining the undoubtedly bad effects of the weak +surviving and propagating their kind; but there appears +to be at least one check in steady action, namely the +weaker and inferior members of society not marrying +so freely as the sound; and this check might be indefinitely +increased, though this is more to be hoped for +than expected, by the weak in body or mind refraining +from marriage.</p> + +<p>In all civilised countries man accumulates property +and bequeaths it to his children. So that the children +in the same country do not by any means start fair in +the race for success. But this is far from an unmixed +evil; for without the accumulation of capital the arts +could not progress; and it is chiefly through their power +that the civilised races have extended, and are now +everywhere extending, their range, so as to take the place +of the lower races. Nor does the moderate accumulation +of wealth interfere with the process of selection. When +a poor man becomes rich, his children enter trades or +professions in which there is struggle enough, so that +the able in body and mind succeed best. The presence +of a body of well-instructed men, who have not to +labour for their daily bread, is important to a degree +which cannot be over-estimated; as all high intellectual +work is carried on by them, and on such work material +progress of all kinds mainly depends, not to mention +other and higher advantages. No doubt wealth when +very great tends to convert men into useless drones, but +their number is never large; and some degree of elimi<span class='pagenum'><a name="Page_170" id="Page_170">170</a></span>nation +here occurs, as we daily see rich men, who happen +to be fools or profligate, squandering away all their +wealth.</p> + +<p>Primogeniture with entailed estates is a more direct +evil, though it may formerly have been a great advantage +by the creation of a dominant class, and any +government is better than anarchy. The eldest sons, +though they may be weak in body or mind, generally +marry, whilst the younger sons, however superior in these +respects, do not so generally marry. Nor can worthless +eldest sons with entailed estates squander their wealth. +But here, as elsewhere, the relations of civilised life are +so complex that some compensatory checks intervene. +The men who are rich through primogeniture are able +to select generation after generation the more beautiful +and charming women; and these must generally be +healthy in body and active in mind. The evil consequences, +such as they may be, of the continued preservation +of the same line of descent, without any +selection, are checked by men of rank always wishing to +increase their wealth and power; and this they effect +by marrying heiresses. But the daughters of parents +who have produced single children, are themselves, as +Mr. Galton has shewn,<a name="FNanchor_232" id="FNanchor_232"></a><a href="#Footnote_232" class="fnanchor">232</a> apt to be sterile; and thus +noble families are continually cut off in the direct line, +and their wealth flows into some side channel; but +unfortunately this channel is not determined by superiority +of any kind.</p> + +<p>Although civilisation thus checks in many ways the +action of natural selection, it apparently favours, by +means of improved food and the freedom from occasional +hardships, the better development of the body. +This may be inferred from civilised men having been +<span class='pagenum'><a name="Page_171" id="Page_171">171</a></span>found, wherever compared, to be physically stronger than +savages. They appear also to have equal powers of +endurance, as has been proved in many adventurous +expeditions. Even the great luxury of the rich can be +but little detrimental; for the expectation of life of our +aristocracy, at all ages and of both sexes, is very little +inferior to that of healthy English lives in the lower +classes.<a name="FNanchor_233" id="FNanchor_233"></a><a href="#Footnote_233" class="fnanchor">233</a></p> + +<p>We will now look to the intellectual faculties alone. +If in each grade of society the members were divided +into two equal bodies, the one including the intellectually +superior and the other the inferior, there can +be little doubt that the former would succeed best in +all occupations and rear a greater number of children. +Even in the lowest walks of life, skill and ability must +be of some advantage, though in many occupations, +owing to the great division of labour, a very small +one. Hence in civilised nations there will be some +tendency to an increase both in the number and in +the standard of the intellectually able. But I do not +wish to assert that this tendency may not be more than +counterbalanced in other ways, as by the multiplication +of the reckless and improvident; but even to such as +these, ability must be some advantage.</p> + +<p>It has often been objected to views like the foregoing, +that the most eminent men who have ever lived +have left no offspring to inherit their great intellect. +Mr. Galton says,<a name="FNanchor_234" id="FNanchor_234"></a><a href="#Footnote_234" class="fnanchor">234</a> “I regret I am unable to solve the +simple question whether, and how far, men and women +who are prodigies of genius are infertile. I have, however, +shewn that men of eminence are by no means so.”</p> + +<p><span class='pagenum'><a name="Page_172" id="Page_172">172</a></span></p><p>Great lawgivers, the founders of beneficent religions, +great philosophers and discoverers in science, aid the +progress of mankind in a far higher degree by their +works than by leaving a numerous progeny. In the +case of corporeal structures, it is the selection of +the slightly better-endowed and the elimination of the +slightly less well-endowed individuals, and not the preservation +of strongly-marked and rare anomalies, that +leads to the advancement of a species.<a name="FNanchor_235" id="FNanchor_235"></a><a href="#Footnote_235" class="fnanchor">235</a> So it will be +with the intellectual faculties, namely from the somewhat +more able men in each grade of society succeeding +rather better than the less able, and consequently increasing +in number, if not otherwise prevented. When +in any nation the standard of intellect and the number +of intellectual men have increased, we may expect from +the law of the deviation from an average, as shewn by +Mr. Galton, that prodigies of genius will appear somewhat +more frequently than before.</p> + +<p>In regard to the moral qualities, some elimination of +the worst dispositions is always in progress even in the +most civilised nations. Malefactors are executed, or +imprisoned for long periods, so that they cannot freely +transmit their bad qualities. Melancholic and insane +persons are confined, or commit suicide. Violent and +quarrelsome men often come to a bloody end. Restless +men who will not follow any steady occupation—and +this relic of barbarism is a great check to civilisation<a name="FNanchor_236" id="FNanchor_236"></a><a href="#Footnote_236" class="fnanchor">236</a>—emigrate +to newly-settled countries, where they prove +useful pioneers. Intemperance is so highly destructive, +that the expectation of life of the intemperate, at the +age, for instance, of thirty, is only 13.8 years; whilst for +the rural labourers of England at the same age it is +<span class='pagenum'><a name="Page_173" id="Page_173">173</a></span>40·59 years.<a name="FNanchor_237" id="FNanchor_237"></a><a href="#Footnote_237" class="fnanchor">237</a> Profligate women bear few children, and +profligate men rarely marry; both suffer from disease. +In the breeding of domestic animals, the elimination of +those individuals, though few in number, which are in +any marked manner inferior, is by no means an unimportant +element towards success. This especially holds +good with injurious characters which tend to reappear +through reversion, such as blackness in sheep; and +with mankind some of the worst dispositions, which +occasionally without any assignable cause make their +appearance in families, may perhaps be reversions to +a savage state, from which we are not removed by very +many generations. This view seems indeed recognised +in the common expression that such men are the black +sheep of the family.</p> + +<p>With civilised nations, as far as an advanced standard +of morality, and an increased number of fairly +well-endowed men are concerned, natural selection apparently +effects but little; though the fundamental +social instincts were originally thus gained. But I +have already said enough, whilst treating of the lower +races, on the causes which lead to the advance of +morality, namely, the approbation of our fellow-men—the +strengthening of our sympathies by habit—example +and imitation—reason—experience and even self-interest—instruction +during youth, and religious feelings.</p> + +<p>A most important obstacle in civilised countries to +an increase in the number of men of a superior class has +been strongly urged by Mr. Greg and Mr. Galton,<a name="FNanchor_238" id="FNanchor_238"></a><a href="#Footnote_238" class="fnanchor">238</a> +<span class='pagenum'><a name="Page_174" id="Page_174">174</a></span>namely, the fact that the very poor and reckless, who +are often degraded by vice, almost invariably marry +early, whilst the careful and frugal, who are generally +otherwise virtuous, marry late in life, so that they may +be able to support themselves and their children in +comfort. Those who marry early produce within a +given period not only a greater number of generations, +but, as shewn by Dr. Duncan,<a name="FNanchor_239" id="FNanchor_239"></a><a href="#Footnote_239" class="fnanchor">239</a> they produce many more +children. The children, moreover, that are born by +mothers during the prime of life are heavier and larger, +and therefore probably more vigorous, than those born +at other periods. Thus the reckless, degraded, and +often vicious members of society, tend to increase at a +quicker rate than the provident and generally virtuous +members. Or as Mr. Greg puts the case: “The careless, +squalid, unaspiring Irishman multiplies like +rabbits: the frugal, foreseeing, self-respecting, ambitious +Scot, stern in his morality, spiritual in his +faith, sagacious and disciplined in his intelligence, +passes his best years in struggle and in celibacy, +marries late, and leaves few behind him. Given a +land originally peopled by a thousand Saxons and a +thousand Celts—and in a dozen generations five-sixths +of the population would be Celts, but five-sixths of +the property, of the power, of the intellect, would +belong to the one-sixth of Saxons that remained. +In the eternal ‘struggle for existence,’ it would be +the inferior and <i>less</i> favoured race that had prevailed—and +prevailed by virtue not of its good qualities +but of its faults.”</p> + +<p>There are, however, some checks to this downward +tendency. We have seen that the intemperate suffer +<span class='pagenum'><a name="Page_175" id="Page_175">175</a></span>from a high rate of mortality, and the extremely profligate +leave few offspring. The poorest classes crowd +into towns, and it has been proved by Dr. Stark from +the statistics of ten years in Scotland,<a name="FNanchor_240" id="FNanchor_240"></a><a href="#Footnote_240" class="fnanchor">240</a> that at all ages +the death-rate is higher in towns than in rural districts, +“and during the first five years of life the town death-rate +is almost exactly double that of the rural districts.” +As these returns include both the rich and the poor, no +doubt more than double the number of births would be +requisite to keep up the number of the very poor inhabitants +in the towns, relatively to those in the country. +With women, marriage at too early an age is highly +injurious; for it has been found in France that, “twice +as many wives under twenty die in the year, as died out +of the same number of the unmarried.” The mortality, +also, of husbands under twenty is “excessively high,”<a name="FNanchor_241" id="FNanchor_241"></a><a href="#Footnote_241" class="fnanchor">241</a> +but what the cause of this may be seems doubtful. +Lastly, if the men who prudently delay marrying until +they can bring up their families in comfort, were to +select, as they often do, women in the prime of life, the +rate of increase in the better class would be only slightly +lessened.</p> + +<p>It was established from an enormous body of statistics, +taken during 1853, that the unmarried men throughout +France, between the ages of twenty and eighty, die in a +much larger proportion than the married: for instance, +out of every 1000 unmarried men, between the ages of +twenty and thirty, 11·3 annually died, whilst of the +married only 6·5 died.<a name="FNanchor_242" id="FNanchor_242"></a><a href="#Footnote_242" class="fnanchor">242</a> A similar law was proved to +<span class='pagenum'><a name="Page_176" id="Page_176">176</a></span>hold good, during the years 1863 and 1864, with the +entire population above the age of twenty in Scotland: +for instance, out of every 1000 unmarried men, between +the ages of twenty and thirty, 14·97 annually died, +whilst of the married only 7·24 died, that is less than +half.<a name="FNanchor_243" id="FNanchor_243"></a><a href="#Footnote_243" class="fnanchor">243</a> Dr. Stark remarks on this, “Bachelorhood is +more destructive to life than the most unwholesome +trades, or than residence in an unwholesome house or +district where there has never been the most distant +attempt at sanitary improvement.” He considers that +the lessened mortality is the direct result of “marriage, +and the more regular domestic habits which attend that +state.” He admits, however, that the intemperate, +profligate, and criminal classes, whose duration of life +is low, do not commonly marry; and it must likewise +be admitted that men with a weak constitution, +ill health, or any great infirmity in body or mind, will +often not wish to marry, or will be rejected. Dr. Stark +seems to have come to the conclusion that marriage in +itself is a main cause of prolonged life, from finding +that aged married men still have a considerable advantage +in this respect over the unmarried of the same +advanced age; but every one must have known instances +of men, who with weak health during youth did not +marry, and yet have survived to old age, though +remaining weak and therefore always with a lessened +chance of life. There is another remarkable circumstance +which seems to support Dr. Stark’s conclusion, +namely, that widows and widowers in France suffer in +comparison with the married a very heavy rate of mortality; +but Dr. Farr attributes this to the poverty and +<span class='pagenum'><a name="Page_177" id="Page_177">177</a></span>evil habits consequent on the disruption of the family, +and to grief. On the whole we may conclude with Dr. +Farr that the lesser mortality of married than of unmarried +men, which seems to be a general law, “is mainly +due to the constant elimination of imperfect types, and +to the skilful selection of the finest individuals out of +each successive generation;” the selection relating only +to the marriage state, and acting on all corporeal, intellectual, +and moral qualities. We may, therefore, +infer that sound and good men who out of prudence +remain for a time unmarried do not suffer a high rate +of mortality.</p> + +<p>If the various checks specified in the two last paragraphs, +and perhaps others as yet unknown, do not +prevent the reckless, the vicious and otherwise inferior +members of society from increasing at a quicker rate +than the better class of men, the nation will retrograde, +as has occurred too often in the history of the +world. We must remember that progress is no invariable +rule. It is most difficult to say why one civilised +nation rises, becomes more powerful, and spreads more +widely, than another; or why the same nation progresses +more at one time than at another. We can only say +that it depends on an increase in the actual number of +the population, on the number of the men endowed +with high intellectual and moral faculties, as well as +on their standard of excellence. Corporeal structure, +except so far as vigour of body leads to vigour of mind, +appears to have little influence.</p> + +<p>It has been urged by several writers that as high +intellectual powers are advantageous to a nation, the +old Greeks, who stood some grades higher in intellect +than any race that has ever existed,<a name="FNanchor_244" id="FNanchor_244"></a><a href="#Footnote_244" class="fnanchor">244</a> ought to have +<span class='pagenum'><a name="Page_178" id="Page_178">178</a></span>risen, if the power of natural selection were real, still +higher in the scale, increased in number, and stocked +the whole of Europe. Here we have the tacit assumption, +so often made with respect to corporeal structures, +that there is some innate tendency towards continued +development in mind and body. But development of +all kinds depends on many concurrent favourable circumstances. +Natural selection acts only in a tentative +manner. Individuals and races may have acquired +certain indisputable advantages, and yet have perished +from failing in other characters. The Greeks may have +retrograded from a want of coherence between the many +small states, from the small size of their whole country, +from the practice of slavery, or from extreme sensuality; +for they did not succumb until “they were enervated +and corrupt to the very core.”<a name="FNanchor_245" id="FNanchor_245"></a><a href="#Footnote_245" class="fnanchor">245</a> The western nations +of Europe, who now so immeasurably surpass their +former savage progenitors and stand at the summit of +civilisation, owe little or none of their superiority to +direct inheritance from the old Greeks; though they +owe much to the written works of this wonderful people.</p> + +<p>Who can positively say why the Spanish nation, +so dominant at one time, has been distanced in the +race. The awakening of the nations of Europe from +the dark ages is a still more perplexing problem. At +this early period, as Mr. Galton<a name="FNanchor_246" id="FNanchor_246"></a><a href="#Footnote_246" class="fnanchor">246</a> has remarked, almost +all the men of a gentle nature, those given to meditation +or culture of the mind, had no refuge except in +the bosom of the Church which demanded celibacy; +<span class='pagenum'><a name="Page_179" id="Page_179">179</a></span>and this could hardly fail to have had a deteriorating +influence on each successive generation. During this +same period the Holy Inquisition selected with extreme +care the freest and boldest men in order to burn or +imprison them. In Spain alone some of the best men—those +who doubted and questioned, and without doubting +there can be no progress—were eliminated during three +centuries at the rate of a thousand a year. The evil +which the Catholic Church has thus effected, though +no doubt counterbalanced to a certain, perhaps large +extent in other ways, is incalculable; nevertheless, +Europe has progressed at an unparalleled rate.</p> + +<p>The remarkable success of the English as colonists +over other European nations, which is well illustrated by +comparing the progress of the Canadians of English and +French extraction, has been ascribed to their “daring +and persistent energy;” but who can say how the +English gained their energy. There is apparently +much truth in the belief that the wonderful progress +of the United States, as well as the character of the +people, are the results of natural selection; the more +energetic, restless, and courageous men from all parts +of Europe having emigrated during the last ten or +twelve generations to that great country, and having there +succeeded best.<a name="FNanchor_247" id="FNanchor_247"></a><a href="#Footnote_247" class="fnanchor">247</a> Looking to the distant future, I do +not think that the Rev. Mr. Zincke takes an exaggerated +view when he says:<a name="FNanchor_248" id="FNanchor_248"></a><a href="#Footnote_248" class="fnanchor">248</a> “All other series of events—as +that which resulted in the culture of mind in Greece, +and that which resulted in the empire of Rome—only +appear to have purpose and value when viewed in +connection with, or rather as subsidiary to ... the +great stream of Anglo-Saxon emigration to the west.”</p> + +<p><span class='pagenum'><a name="Page_180" id="Page_180">180</a></span></p><p>Obscure as is the problem of the advance of civilisation, +we can at least see that a nation which produced during +a lengthened period the greatest number of highly intellectual, +energetic, brave, patriotic, and benevolent men, +would generally prevail over less favoured nations.</p> + +<p class="tb">Natural selection follows from the struggle for existence; +and this from a rapid rate of increase. It is +impossible not bitterly to regret, but whether wisely +is another question, the rate at which man tends to +increase; for this leads in barbarous tribes to infanticide +and many other evils, and in civilised nations to +abject poverty, celibacy, and to the late marriages of +the prudent. But as man suffers from the same physical +evils with the lower animals, he has no right to expect +an immunity from the evils consequent on the struggle +for existence. Had he not been subjected to natural +selection, assuredly he would never have attained to +the rank of manhood. When we see in many parts +of the world enormous areas of the most fertile land +peopled by a few wandering savages, but which are +capable of supporting numerous happy homes, it might +be argued that the struggle for existence had not been +sufficiently severe to force man upwards to his highest +standard. Judging from all that we know of man and +the lower animals, there has always been sufficient +variability in the intellectual and moral faculties, for +their steady advancement through natural selection. +No doubt such advancement demands many favourable +concurrent circumstances; but it may well be doubted +whether the most favourable would have sufficed, had +not the rate of increase been rapid, and the consequent +struggle for existence severe to an extreme degree.</p> + +<p><i>On the evidence that all civilised nations were once barbarous.</i>—As +we have had to consider the steps by which<span class='pagenum'><a name="Page_181" id="Page_181">181</a></span> +some semi-human creature has been gradually raised to +the rank of man in his most perfect state, the present +subject cannot be quite passed over. But it has been +treated in so full and admirable a manner by Sir J. +Lubbock,<a name="FNanchor_249" id="FNanchor_249"></a><a href="#Footnote_249" class="fnanchor">249</a> Mr. Tylor, Mr. M’Lennan, and others, that +I need here give only the briefest summary of their +results. The arguments recently advanced by the +Duke of Argyll<a name="FNanchor_250" id="FNanchor_250"></a><a href="#Footnote_250" class="fnanchor">250</a> and formerly by Archbishop Whately, +in favour of the belief that man came into the world +as a civilised being and that all savages have since +undergone degradation, seem to me weak in comparison +with those advanced on the other side. Many nations, +no doubt, have fallen away in civilisation, and some +may have lapsed into utter barbarism, though on this +latter head I have not met with any evidence. The +Fuegians were probably compelled by other conquering +hordes to settle in their inhospitable country, and they +may have become in consequence somewhat more +degraded; but it would be difficult to prove that they +have fallen much below the Botocudos who inhabit the +finest parts of Brazil.</p> + +<p>The evidence that all civilised nations are the descendants +of barbarians, consists, on the one side, of +clear traces of their former low condition in still-existing +customs, beliefs, language, &c.; and on the other side, +of proofs that savages are independently able to raise +themselves a few steps in the scale of civilisation, and +have actually thus risen. The evidence on the first +head is extremely curious, but cannot be here given: +I refer to such cases as that, for instance, of the art of +enumeration, which, as Mr. Tylor clearly shews by the +words still used in some places, originated in counting +<span class='pagenum'><a name="Page_182" id="Page_182">182</a></span>the fingers, first of one hand and then of the other, +and lastly of the toes. We have traces of this in our +own decimal system, and in the Roman numerals, which +after reaching to the number V., change into VI., &c., +when the other hand no doubt was used. So again, +“when we speak of three-score and ten, we are counting +by the vigesimal system, each score thus ideally +made, standing for 20—for ‘one man’ as a Mexican +or Carib would put it.”<a name="FNanchor_251" id="FNanchor_251"></a><a href="#Footnote_251" class="fnanchor">251</a> According to a large and +increasing school of philologists, every language bears +the marks of its slow and gradual evolution. So it +is with the art of writing, as letters are rudiments of +pictorial representations. It is hardly possible to read +Mr. M’Lennan’s work<a name="FNanchor_252" id="FNanchor_252"></a><a href="#Footnote_252" class="fnanchor">252</a> and not admit that almost all +civilised nations still retain some traces of such rude +habits as the forcible capture of wives. What ancient +nation, as the same author asks, can be named that was +originally monogamous? The primitive idea of justice, +as shewn by the law of battle and other customs of +which traces still remain, was likewise most rude. Many +existing superstitions are the remnants of former false +religious beliefs. The highest form of religion—the +grand idea of God hating sin and loving righteousness—was +unknown during primeval times.</p> + +<p>Turning to the other kind of evidence: Sir J. Lubbock +has shewn that some savages have recently improved +a little in some of their simpler arts. From the +<span class='pagenum'><a name="Page_183" id="Page_183">183</a></span>extremely curious account which he gives of the weapons, +tools, and arts, used or practised by savages in various +parts of the world, it cannot be doubted that these have +nearly all been independent discoveries, excepting perhaps +the art of making fire.<a name="FNanchor_253" id="FNanchor_253"></a><a href="#Footnote_253" class="fnanchor">253</a> The Australian boomerang +is a good instance of one such independent discovery. +The Tahitians when first visited had advanced in many +respects beyond the inhabitants of most of the other +Polynesian islands. There are no just grounds for the +belief that the high culture of the native Peruvians +and Mexicans was derived from any foreign source;<a name="FNanchor_254" id="FNanchor_254"></a><a href="#Footnote_254" class="fnanchor">254</a> +many native plants were there cultivated, and a few +native animals domesticated. We should bear in mind +that a wandering crew from some semi-civilised land, +if washed to the shores of America, would not, judging +from the small influence of most missionaries, have produced +any marked effect on the natives, unless they had +already become somewhat advanced. Looking to a very +remote period in the history of the world, we find, to +use Sir J. Lubbock’s well-known terms, a paleolithic +and neolithic period; and no one will pretend that the +art of grinding rough flint tools was a borrowed one. +In all parts of Europe, as far east as Greece, in Palestine, +India, Japan, New Zealand, and Africa, including Egypt, +flint tools have been discovered in abundance; and of +their use the existing inhabitants retain no tradition. +There is also indirect evidence of their former use by the +Chinese and ancient Jews. Hence there can hardly be a +doubt that the inhabitants of these many countries, which +include nearly the whole civilised world, were once in a +barbarous condition. To believe that man was aboriginally +<span class='pagenum'><a name="Page_184" id="Page_184">184</a></span> +civilised and then suffered utter degradation in +so many regions, is to take a pitiably low view of +human nature. It is apparently a truer and more +cheerful view that progress has been much more general +than retrogression; that man has risen, though by slow +and interrupted steps, from a lowly condition to the +highest standard as yet attained by him in knowledge, +morals, and religion.</p> +<hr /> +<p><span class='pagenum'><a name="Page_185" id="Page_185">185</a></span></p> + +<h3>CHAPTER VI.</h3> + +<h4><span class="smcap">On the Affinities and Genealogy of Man.</span></h4> + +<div class="blockquot"><p>Position of man in the animal series—The natural system genealogical—Adaptive +characters of slight value—Various small +points of resemblance between man and the Quadrumana—Rank +of man in the natural system—Birthplace and antiquity +of man—Absence of fossil connecting-links—Lower stages in +the genealogy of man, as inferred, firstly from his affinities and +secondly from his structure—Early androgynous condition of +the Vertebrata—Conclusion.</p></div> + +<p>Even if it be granted that the difference between man +and his nearest allies is as great in corporeal structure as +some naturalists maintain, and although we must grant +that the difference between them is immense in mental +power, yet the facts given in the previous chapters +declare, as it appears to me, in the plainest manner, +that man is descended from some lower form, notwithstanding +that connecting-links have not hitherto been +discovered.</p> + +<p>Man is liable to numerous, slight, and diversified +variations, which are induced by the same general +causes, are governed and transmitted in accordance +with the same general laws, as in the lower animals. +Man tends to multiply at so rapid a rate that his offspring +are necessarily exposed to a struggle for existence, +and consequently to natural selection. He has given +rise to many races, some of which are so different that +they have often been ranked by naturalists as distinct +species. His body is constructed on the same homological +plan as that of other mammals, independently +of the uses to which the several parts may be put. He<span class='pagenum'><a name="Page_186" id="Page_186">186</a></span> +passes through the same phases of embryological development. +He retains many rudimentary and useless +structures, which no doubt were once serviceable. Characters +occasionally make their reappearance in him, +which we have every reason to believe were possessed by +his early progenitors. If the origin of man had been +wholly different from that of all other animals, these +various appearances would be mere empty deceptions; +but such an admission is incredible. These appearances, +on the other hand, are intelligible, at least to a large +extent, if man is the co-descendant with other mammals +of some unknown and lower form.</p> + +<p>Some naturalists, from being deeply impressed with +the mental and spiritual powers of man, have divided the +whole organic world into three kingdoms, the Human, +the Animal, and the Vegetable, thus giving to man a +separate kingdom.<a name="FNanchor_255" id="FNanchor_255"></a><a href="#Footnote_255" class="fnanchor">255</a> Spiritual powers cannot be compared +or classed by the naturalist; but he may endeavour +to shew, as I have done, that the mental faculties +of man and the lower animals do not differ in kind, +although immensely in degree. A difference in degree, +however great, does not justify us in placing man in a +distinct kingdom, as will perhaps be best illustrated +by comparing the mental powers of two insects, namely, +a coccus or scale-insect and an ant, which undoubtedly +belong to the same class. The difference is here +greater, though of a somewhat different kind, than +that between man and the highest mammal. The +female coccus, whilst young, attaches itself by its proboscis +to a plant; sucks the sap but never moves again; +is fertilised and lays eggs; and this is its whole history. +On the other hand, to describe the habits and mental +<span class='pagenum'><a name="Page_187" id="Page_187">187</a></span>powers of a female ant, would require, as Pierre Huber +has shewn, a large volume; I may, however, briefly +specify a few points. Ants communicate information +to each other, and several unite for the same work, +or games of play. They recognise their fellow-ants +after months of absence. They build great edifices, +keep them clean, close the doors in the evening, and +post sentries. They make roads, and even tunnels +under rivers. They collect food for the community, +and when an object, too large for entrance, is brought +to the nest, they enlarge the door, and afterwards +build it up again.<a name="FNanchor_256" id="FNanchor_256"></a><a href="#Footnote_256" class="fnanchor">256</a> They go out to battle in regular +bands, and freely sacrifice their lives for the common +weal. They emigrate in accordance with a preconcerted +plan. They capture slaves. They keep Aphides +as milch-cows. They move the eggs of their aphides, +as well as their own eggs and cocoons, into warm parts +of the nest, in order that they may be quickly hatched; +and endless similar facts could be given. On the +whole, the difference in mental power between an ant +and a coccus is immense; yet no one has ever dreamed +of placing them in distinct classes, much less in distinct +kingdoms. No doubt this interval is bridged over by +the intermediate mental powers of many other insects; +and this is not the case with man and the higher apes. +But we have every reason to believe that breaks in the +series are simply the result of many forms having become +extinct.</p> + +<p>Professor Owen, relying chiefly on the structure of +the brain, has divided the mammalian series into four +sub-classes. One of these he devotes to man; in another +he places both the marsupials and the monotremata; +so that he makes man as distinct from all other mam<span class='pagenum'><a name="Page_188" id="Page_188">188</a></span>mals +as are these two latter groups conjoined. This +view has not been accepted, as far as I am aware, by +any naturalist capable of forming an independent judgment, +and therefore need not here be further considered.</p> + +<p>We can understand why a classification founded on +any single character or organ—even an organ so wonderfully +complex and important as the brain—or on the +high development of the mental faculties, is almost sure +to prove unsatisfactory. This principle has indeed been +tried with hymenopterous insects; but when thus classed +by their habits or instincts, the arrangement proved +thoroughly artificial.<a name="FNanchor_257" id="FNanchor_257"></a><a href="#Footnote_257" class="fnanchor">257</a> Classifications may, of course, be +based on any character whatever, as on size, colour, or +the element inhabited; but naturalists have long felt a +profound conviction that there is a natural system. This +system, it is now generally admitted, must be, as far +as possible, genealogical in arrangement,—that is, the +co-descendants of the same form must be kept together +in one group, separate from the co-descendants of any +other form; but if the parent-forms are related, so will +be their descendants, and the two groups together will +form a larger group. The amount of difference between +the several groups—that is the amount of modification +which each has undergone—will be expressed by such +terms as genera, families, orders, and classes. As we +have no record of the lines of descent, these lines can +be discovered only by observing the degrees of resemblance +between the beings which are to be classed. +For this object numerous points of resemblance are of +much more importance than the amount of similarity +or dissimilarity in a few points. If two languages +were found to resemble each other in a multitude of +<span class='pagenum'><a name="Page_189" id="Page_189">189</a></span>words and points of construction, they would be universally +recognised as having sprung from a common +source, notwithstanding that they differed greatly in +some few words or points of construction. But with +organic beings the points of resemblance must not consist +of adaptations to similar habits of life: two animals +may, for instance, have had their whole frames modified +for living in the water, and yet they will not be brought +any nearer to each other in the natural system. Hence +we can see how it is that resemblances in unimportant +structures, in useless and rudimentary organs, and in +parts not as yet fully developed or functionally active, +are by far the most serviceable for classification; for +they can hardly be due to adaptations within a late +period; and thus they reveal the old lines of descent +or of true affinity.</p> + +<p>We can further see why a great amount of modification +in some one character ought not to lead us to +separate widely any two organisms. A part which +already differs much from the same part in other allied +forms has already, according to the theory of evolution, +varied much; consequently it would (as long as the +organism remained exposed to the same exciting conditions) +be liable to further variations of the same kind; +and these, if beneficial, would be preserved, and thus +continually augmented. In many cases the continued +development of a part, for instance, of the beak of a +bird, or of the teeth of a mammal, would not be advantageous +to the species for gaining its food, or for any +other object; but with man we can see no definite limit, +as far as advantage is concerned, to the continued development +of the brain and mental faculties. Therefore +in determining the position of man in the natural or +genealogical system, the extreme development of his +brain ought not to outweigh a multitude of resem<span class='pagenum'><a name="Page_190" id="Page_190">190</a></span>blances +in other less important or quite unimportant +points.</p> + +<p>The greater number of naturalists who have taken +into consideration the whole structure of man, including +his mental faculties, have followed Blumenbach and +Cuvier, and have placed man in a separate Order, under +the title of the Bimana, and therefore on an equality +with the Orders of the Quadrumana, Carnivora, &c. +Recently many of our best naturalists have recurred to +the view first propounded by Linnæus, so remarkable for +his sagacity, and have placed man in the same Order +with the Quadrumana, under the title of the Primates. +The justice of this conclusion will be admitted if, in +the first place, we bear in mind the remarks just +made on the comparatively small importance for classification +of the great development of the brain in man; +bearing, also, in mind that the strongly-marked differences +between the skulls of man and the Quadrumana +(lately insisted upon by Bischoff, Aeby, and others) +apparently follow from their differently developed brains. +In the second place, we must remember that nearly all +the other and more important differences between man +and the Quadrumana are manifestly adaptive in their +nature, and relate chiefly to the erect position of man; +such as the structure of his hand, foot, and pelvis, the +curvature of his spine, and the position of his head. +The family of seals offers a good illustration of the +small importance of adaptive characters for classification. +These animals differ from all other Carnivora in the +form of their bodies and in the structure of their limbs, +far more than does man from the higher apes; yet in +every system, from that of Cuvier to the most recent +one by Mr. Flower,<a name="FNanchor_258" id="FNanchor_258"></a><a href="#Footnote_258" class="fnanchor">258</a> seals are ranked as a mere family +<span class='pagenum'><a name="Page_191" id="Page_191">191</a></span>in the Order of the Carnivora. If man had not been +his own classifier, he would never have thought of +founding a separate order for his own reception.</p> + +<p>It would be beyond my limits, and quite beyond my +knowledge, even to name the innumerable points of +structure in which man agrees with the other Primates. +Our great anatomist and philosopher, Prof. Huxley, has +fully discussed this subject,<a name="FNanchor_259" id="FNanchor_259"></a><a href="#Footnote_259" class="fnanchor">259</a> and has come to the conclusion +that man in all parts of his organisation differs +less from the higher apes, than these do from the lower +members of the same group. Consequently there “is +no justification for placing man in a distinct order.”</p> + +<p>In an early part of this volume I brought forward +various facts, shewing how closely man agrees in constitution +with the higher mammals; and this agreement, +no doubt, depends on our close similarity in minute +structure and chemical composition. I gave, as +instances, our liability to the same diseases, and to the +attacks of allied parasites; our tastes in common for the +same stimulants, and the similar effects thus produced, +as well as by various drugs; and other such facts.</p> + +<p>As small unimportant points of resemblance between +man and the higher apes are not commonly noticed in +systematic works, and as, when numerous, they clearly +reveal our relationship, I will specify a few such points. +The relative position of the features are manifestly the +same in man and the Quadrumana; and the various +emotions are displayed by nearly similar movements of +the muscles and skin, chiefly above the eyebrows and +round the mouth. Some few expressions are, indeed, +almost the same, as in the weeping of certain kinds of +monkeys, and in the laughing noise made by others, +during which the corners of the mouth are drawn back<span class='pagenum'><a name="Page_192" id="Page_192">192</a></span>wards, +and the lower eyelids wrinkled. The external +ears are curiously alike. In man the nose is much +more prominent than in most monkeys; but we may +trace the commencement of an aquiline curvature in +the nose of the Hoolock Gibbon; and this in the <i>Semnopithecus +nasica</i> is carried to a ridiculous extreme.</p> + +<p>The faces of many monkeys are ornamented with +beards, whiskers, or moustaches. The hair on the head +grows to a great length in some species of Semnopithecus;<a name="FNanchor_260" id="FNanchor_260"></a><a href="#Footnote_260" class="fnanchor">260</a> +and in the Bonnet monkey (<i>Macacus +radiatus</i>) it radiates from a point on the crown, with a +parting down the middle, as in man. It is commonly +said that the forehead gives to man his noble and intellectual +appearance; but the thick hair on the head of +the Bonnet monkey terminates abruptly downwards, +and is succeeded by such short and fine hair, or down, +that at a little distance the forehead, with the exception +of the eyebrows, appears quite naked. It has been +erroneously asserted that eyebrows are not present in +any monkey. In the species just named the degree of +nakedness of the forehead differs in different individuals; +and Eschricht states<a name="FNanchor_261" id="FNanchor_261"></a><a href="#Footnote_261" class="fnanchor">261</a> that in our children the limit +between the hairy scalp and the naked forehead is +sometimes not well defined; so that here we seem to +have a trifling case of reversion to a progenitor, in whom +the forehead had not as yet become quite naked.</p> + +<p>It is well known that the hair on our arms tends to +converge from above and below to a point at the elbow. +This curious arrangement, so unlike that in most of the +lower mammals, is common to the gorilla, chimpanzee, +orang, some species of Hylobates, and even to some few +American monkeys. But in <i>Hylobates agilis</i> the hair +<span class='pagenum'><a name="Page_193" id="Page_193">193</a></span>on the fore-arm is directed downwards or towards the +wrist in the ordinary manner; and in <i>H. lar</i> it is nearly +erect, with only a very slight forward inclination; so +that in this latter species it is in a transitional state. +It can hardly be doubted that with most mammals the +thickness of the hair and its direction on the back is +adapted to throw off the rain; even the transverse hairs +on the fore-legs of a dog may serve for this end when he +is coiled up asleep. Mr. Wallace remarks that the convergence +of the hair towards the elbow on the arms +of the orang (whose habits he has so carefully studied) +serves to throw off the rain, when, as is the custom +of this animal, the arms are bent, with the hands +clasped round a branch or over its own head. We +should, however, bear in mind that the attitude of an +animal may perhaps be in part determined by the +direction of the hair; and not the direction of the hair +by the attitude. If the above explanation is correct in +the case of the orang, the hair on our fore-arms offers a +curious record of our former state; for no one supposes +that it is now of any use in throwing off the rain, nor in +our present erect condition is it properly directed for this +purpose.</p> + +<p>It would, however, be rash to trust too much to the +principle of adaptation in regard to the direction of the +hair in man or his early progenitors; for it is impossible +to study the figures given by Eschricht of the arrangement +of the hair on the human fœtus (this being the +same as in the adult) and not agree with this excellent +observer that other and more complex causes have +intervened. The points of convergence seem to stand +in some relation to those points in the embryo which +are last closed in during development. There appears, +also, to exist some relation between the arrangement<span class='pagenum'><a name="Page_194" id="Page_194">194</a></span> +of the hair on the limbs, and the course of the medullary +arteries.<a name="FNanchor_262" id="FNanchor_262"></a><a href="#Footnote_262" class="fnanchor">262</a></p> + +<p>It must not be supposed that the resemblances between +man and certain apes in the above and many +other points—such as in having a naked forehead, +long tresses on the head, &c.—are all necessarily the +result of unbroken inheritance from a common progenitor +thus characterised, or of subsequent reversion. +Many of these resemblances are more probably due +to analogous variation, which follows, as I have elsewhere +attempted to shew,<a name="FNanchor_263" id="FNanchor_263"></a><a href="#Footnote_263" class="fnanchor">263</a> from co-descended organisms +having a similar constitution and having been acted +on by similar causes inducing variability. With respect +to the similar direction of the hair on the fore-arms +of man and certain monkeys, as this character is +common to almost all the anthropomorphous apes, it +may probably be attributed to inheritance; but not +certainly so, as some very distinct American monkeys +are thus characterised. The same remark is applicable +to the tailless condition of man; for the tail is absent +in all the anthropomorphous apes. Nevertheless this +character cannot with certainty be attributed to inheritance, +as the tail, though not absent, is rudimentary +in several other Old World and in some New World +species, and is quite absent in several species belonging +to the allied group of Lemurs.</p> + +<p>Although, as we have now seen, man has no just right +to form a separate Order for his own reception, he may +<span class='pagenum'><a name="Page_195" id="Page_195">195</a></span>perhaps claim a distinct Sub-order or Family. Prof. +Huxley, in his last work,<a name="FNanchor_264" id="FNanchor_264"></a><a href="#Footnote_264" class="fnanchor">264</a> divides the Primates into +three Sub-orders; namely, the Anthropidæ with man +alone, the Simiadæ including monkeys of all kinds, and +the Lemuridæ with the diversified genera of lemurs. As +far as differences in certain important points of structure +are concerned, man may no doubt rightly claim the +rank of a Sub-order; and this rank is too low, if we look +chiefly to his mental faculties. Nevertheless, under a +genealogical point of view it appears that this rank is +too high, and that man ought to form merely a Family, +or possibly even only a Sub-family. If we imagine +three lines of descent proceeding from a common source, +it is quite conceivable that two of them might after +the lapse of ages be so slightly changed as still to +remain as species of the same genus; whilst the third +line might become so greatly modified as to deserve +to rank as a distinct Sub-family, Family, or even +Order. But in this case it is almost certain that +the third line would still retain through inheritance +numerous small points of resemblance with the other +two lines. Here then would occur the difficulty, at +present insoluble, how much weight we ought to assign +in our classifications to strongly-marked differences in +some few points,—that is to the amount of modification +undergone; and how much to close resemblance in +numerous unimportant points, as indicating the lines of +descent or genealogy. The former alternative is the +most obvious, and perhaps the safest, though the latter +appears the most correct as giving a truly natural +classification.</p> + +<p>To form a judgment on this head, with reference +to man we must glance at the classification of the +<span class='pagenum'><a name="Page_196" id="Page_196">196</a></span>Simiadæ. This family is divided by almost all naturalists +into the Catarhine group, or Old World monkeys, +all of which are characterised (as their name expresses) +by the peculiar structure of their nostrils and by having +four premolars in each jaw; and into the Platyrhine +group or New World monkeys (including two very +distinct sub-groups), all of which are characterised by +differently-constructed nostrils and by having six premolars +in each jaw. Some other small differences might +be mentioned. Now man unquestionably belongs in +his dentition, in the structure of his nostrils, and some +other respects, to the Catarhine or Old World division; +nor does he resemble the Platyrhines more closely than +the Catarhines in any characters, excepting in a few +of not much importance and apparently of an adaptive +nature. Therefore it would be against all probability +to suppose that some ancient New World species had +varied, and had thus produced a man-like creature with +all the distinctive characters proper to the Old World +division; losing at the same time all its own distinctive +characters. There can consequently hardly be a doubt +that man is an offshoot from the Old World Simian stem; +and that under a genealogical point of view, he must +be classed with the Catarhine division.<a name="FNanchor_265" id="FNanchor_265"></a><a href="#Footnote_265" class="fnanchor">265</a></p> + +<p>The anthropomorphous apes, namely the gorilla, +chimpanzee, orang, and hylobates, are separated as a +distinct sub-group from the other Old World monkeys +by most naturalists. I am aware that Gratiolet, relying +on the structure of the brain, does not admit the exist<span class='pagenum'><a name="Page_197" id="Page_197">197</a></span>ence +of this sub-group, and no doubt it is a broken +one; thus the orang, as Mr. St. G. Mivart remarks,<a name="FNanchor_266" id="FNanchor_266"></a><a href="#Footnote_266" class="fnanchor">266</a> +“is one of the most peculiar and aberrant forms to be +found in the Order.” The remaining, non-anthropomorphous, +Old World monkeys, are again divided by +some naturalists into two or three smaller sub-groups; +the genus Semnopithecus, with its peculiar sacculated +stomach, being the type of one such sub-group. But +it appears from M. Gaudry’s wonderful discoveries in +Attica, that during the Miocene period a form existed +there, which connected Semnopithecus and Macacus; +and this probably illustrates the manner in which the +other and higher groups were once blended together.</p> + +<p>If the anthropomorphous apes be admitted to form +a natural sub-group, then as man agrees with them, +not only in all those characters which he possesses in +common with the whole Catarhine group, but in other +peculiar characters, such as the absence of a tail and +of callosities and in general appearance, we may infer +that some ancient member of the anthropomorphous +sub-group gave birth to man. It is not probable +that a member of one of the other lower sub-groups +should, through the law of analogous variation, have +given rise to a man-like creature, resembling the higher +anthropomorphous apes in so many respects. No +doubt man, in comparison with most of his allies, has +undergone an extraordinary amount of modification, +chiefly in consequence of his greatly developed brain +and erect position; nevertheless we should bear in +mind that he “is but one of several exceptional forms +of Primates.”<a name="FNanchor_267" id="FNanchor_267"></a><a href="#Footnote_267" class="fnanchor">267</a></p> + +<p>Every naturalist, who believes in the principle of +<span class='pagenum'><a name="Page_198" id="Page_198">198</a></span>evolution, will grant that the two main divisions of the +Simiadæ, namely the Catarhine and Platyrhine monkeys, +with their sub-groups, have all proceeded from +some one extremely ancient progenitor. The early +descendants of this progenitor, before they had diverged +to any considerable extent from each other, would still +have formed a single natural group; but some of the +species or incipient genera would have already begun +to indicate by their diverging characters the future +distinctive marks of the Catarhine and Platyrhine divisions. +Hence the members of this supposed ancient +group would not have been so uniform in their dentition +or in the structure of their nostrils, as are the existing +Catarhine monkeys in one way and the Platyrhines in +another way, but would have resembled in this respect +the allied Lemuridæ which differ greatly from each +other in the form of their muzzles,<a name="FNanchor_268" id="FNanchor_268"></a><a href="#Footnote_268" class="fnanchor">268</a> and to an extraordinary +degree in their dentition.</p> + +<p class="tb">The Catarhine and Platyrhine monkeys agree in +a multitude of characters, as is shewn by their unquestionably +belonging to one and the same Order. The +many characters which they possess in common can +hardly have been independently acquired by so many +distinct species; so that these characters must have +been inherited. But an ancient form which possessed +many characters common to the Catarhine and Platyrhine +monkeys, and others in an intermediate condition, +and some few perhaps distinct from those now present +in either group, would undoubtedly have been ranked, +if seen by a naturalist, as an ape or monkey. And as +man under a genealogical point of view belongs to the +Catarhine or Old World stock, we must conclude, how<span class='pagenum'><a name="Page_199" id="Page_199">199</a></span>ever +much the conclusion may revolt our pride, that +our early progenitors would have been properly thus +designated.<a name="FNanchor_269" id="FNanchor_269"></a><a href="#Footnote_269" class="fnanchor">269</a> But we must not fall into the error of +supposing that the early progenitor of the whole Simian +stock, including man, was identical with, or even closely +resembled, any existing ape or monkey.</p> + +<p><i>On the Birthplace and Antiquity of Man.</i>—We are +naturally led to enquire where was the birthplace of +man at that stage of descent when our progenitors +diverged from the Catarhine stock. The fact that +they belonged to this stock clearly shews that they +inhabited the Old World; but not Australia nor any +oceanic island, as we may infer from the laws of geographical +distribution. In each great region of the world +the living mammals are closely related to the extinct +species of the same region. It is therefore probable +that Africa was formerly inhabited by extinct apes +closely allied to the gorilla and chimpanzee; and as +these two species are now man’s nearest allies, it is +somewhat more probable that our early progenitors +lived on the African continent than elsewhere. But +it is useless to speculate on this subject, for an ape +nearly as large as a man, namely the Dryopithecus +of Lartet, which was closely allied to the anthropomorphous +Hylobates, existed in Europe during the +Upper Miocene period; and since so remote a period +the earth has certainly undergone many great revolutions, +and there has been ample time for migration +on the largest scale.</p> +<p><span class='pagenum'><a name="Page_200" id="Page_200">200</a></span></p> +<p>At the period and place, whenever and wherever it +may have been, when man first lost his hairy covering, +he probably inhabited a hot country; and this would +have been favourable for a frugiferous diet, on which, +judging from analogy, he subsisted. We are far from +knowing how long ago it was when man first diverged +from the Catarhine stock; but this may have occurred +at an epoch as remote as the Eocene period; for the +higher apes had diverged from the lower apes as +early as the Upper Miocene period, as shewn by the +existence of the Dryopithecus. We are also quite +ignorant at how rapid a rate organisms, whether high +or low in the scale, may under favourable circumstances +be modified: we know, however, that some have retained +the same form during an enormous lapse of time. From +what we see going on under domestication, we learn that +within the same period some of the co-descendants of +the same species may be not at all changed, some a +little, and some greatly changed. Thus it may have +been with man, who has undergone a great amount of +modification in certain characters in comparison with +the higher apes.</p> + +<p>The great break in the organic chain between man +and his nearest allies, which cannot be bridged over +by any extinct or living species, has often been advanced +as a grave objection to the belief that man +is descended from some lower form; but this objection +will not appear of much weight to those who, convinced +by general reasons, believe in the general principle +of evolution. Breaks incessantly occur in all parts +of the series, some being wide, sharp and defined, others +less so in various degrees; as between the orang and +its nearest allies—between the Tarsius and the other +Lemuridæ—between the elephant and in a more +striking manner between the Ornithorhynchus or<span class='pagenum'><a name="Page_201" id="Page_201">201</a></span> +Echidna, and other mammals. But all these breaks +depend merely on the number of related forms which +have become extinct. At some future period, not +very distant as measured by centuries, the civilised races +of man will almost certainly exterminate and replace +throughout the world the savage races. At the +same time the anthropomorphous apes, as Professor +Schaaffhausen has remarked,<a name="FNanchor_270" id="FNanchor_270"></a><a href="#Footnote_270" class="fnanchor">270</a> will no doubt be exterminated. +The break will then be rendered wider, for +it will intervene between man in a more civilised state, +as we may hope, than the Caucasian, and some ape as +low as a baboon, instead of as at present between the +negro or Australian and the gorilla.</p> + +<p class="tb">With respect to the absence of fossil remains, serving to +connect man with his ape-like progenitors, no one will +lay much stress on this fact, who will read Sir C. Lyell’s +discussion,<a name="FNanchor_271" id="FNanchor_271"></a><a href="#Footnote_271" class="fnanchor">271</a> in which he shews that in all the vertebrate +classes the discovery of fossil remains has been an +extremely slow and fortuitous process. Nor should it +be forgotten that those regions which are the most +likely to afford remains connecting man with some +extinct ape-like creature, have not as yet been searched +by geologists.</p> + +<p class="tb"><i>Lower Stages in the Genealogy of Man.</i>—We have seen +that man appears to have diverged from the Catarhine +or Old World division of the Simiadæ, after these had +diverged from the New World division. We will now +endeavour to follow the more remote traces of his +genealogy, trusting in the first place to the mutual +affinities between the various classes and orders, with +some slight aid from the periods, as far as ascertained, +<span class='pagenum'><a name="Page_202" id="Page_202">202</a></span>of their successive appearance on the earth. The +Lemuridæ stand below and close to the Simiadæ, constituting +a very distinct family of the Primates, or, +according to Häckel, a distinct Order. This group is +diversified and broken to an extraordinary degree, and +includes many aberrant forms. It has, therefore, probably +suffered much extinction. Most of the remnants +survive on islands, namely in Madagascar and in the +islands of the Malayan archipelago, where they have +not been exposed to such severe competition as they +would have been on well-stocked continents. This +group likewise presents many gradations, leading, as +Huxley remarks,<a name="FNanchor_272" id="FNanchor_272"></a><a href="#Footnote_272" class="fnanchor">272</a> “insensibly from the crown and +summit of the animal creation down to creatures +from which there is but a step, as it seems, to the +lowest, smallest, and least intelligent of the placental +mammalia.” From these various considerations it is +probable that the Simiadæ were originally developed +from the progenitors of the existing Lemuridæ; and +these in their turn from forms standing very low in the +mammalian series.</p> + +<p>The Marsupials stand in many important characters +below the placental mammals. They appeared at an +earlier geological period, and their range was formerly +much more extensive than what it now is. Hence the +Placentata are generally supposed to have been derived +from the Implacentata or Marsupials; not, however, from +forms closely like the existing Marsupials, but from +their early progenitors. The Monotremata are plainly +allied to the Marsupials; forming a third and still +lower division in the great mammalian series. They +are represented at the present day solely by the Ornithorhynchus +and Echidna; and these two forms may +<span class='pagenum'><a name="Page_203" id="Page_203">203</a></span>be safely considered as relics of a much larger group +which have been preserved in Australia through some +favourable concurrence of circumstances. The Monotremata +are eminently interesting, as in several +important points of structure they lead towards the +class of reptiles.</p> + +<p>In attempting to trace the genealogy of the Mammalia, +and therefore of man, lower down in the series, +we become involved in greater and greater obscurity. +He who wishes to see what ingenuity and knowledge +can effect, may consult Prof. Häckel’s works.<a name="FNanchor_273" id="FNanchor_273"></a><a href="#Footnote_273" class="fnanchor">273</a> I will +content myself with a few general remarks. Every +evolutionist will admit that the five great vertebrate +classes, namely, mammals, birds, reptiles, amphibians, +and fishes, are all descended from some one prototype; +for they have much in common, especially during their +embryonic state. As the class of fishes is the most +lowly organised and appeared before the others, we may +conclude that all the members of the vertebrate kingdom +are derived from some fish-like animal, less highly +organised than any as yet found in the lowest known +formations. The belief that animals so distinct as a +monkey or elephant and a humming-bird, a snake, frog, +and fish, &c., could all have sprung from the same +parents, will appear monstrous to those who have not +attended to the recent progress of natural history. For +this belief implies the former existence of links closely +binding together all these forms, now so utterly unlike.</p> +<p><span class='pagenum'><a name="Page_204" id="Page_204">204</a></span></p> +<p>Nevertheless it is certain that groups of animals have +existed, or do now exist, which serve to connect more or +less closely the several great vertebrate classes. We +have seen that the Ornithorhynchus graduates towards +reptiles; and Prof. Huxley has made the remarkable +discovery, confirmed by Mr. Cope and others, that the +old Dinosaurians are intermediate in many important +respects between certain reptiles and certain birds—the +latter consisting of the ostrich-tribe (itself evidently a +widely-diffused remnant of a larger group) and of the +Archeopteryx, that strange Secondary bird having a +long tail like that of the lizard. Again, according to +Prof. Owen,<a name="FNanchor_274" id="FNanchor_274"></a><a href="#Footnote_274" class="fnanchor">274</a> the Ichthyosaurians—great sea-lizards furnished +with paddles—present many affinities with fishes, +or rather, according to Huxley, with amphibians. This +latter class (including in its highest division frogs and +toads) is plainly allied to the Ganoid fishes. These +latter fishes swarmed during the earlier geological +periods, and were constructed on what is called a highly +generalised type, that is they presented diversified affinities +with other groups of organisms. The amphibians +and fishes are also so closely united by the Lepidosiren, +that naturalists long disputed in which of these two +classes it ought to be placed. The Lepidosiren and +some few Ganoid fishes have been preserved from utter +extinction by inhabiting our rivers, which are harbours +of refuge, bearing the same relation to the great waters +of the ocean that islands bear to continents.</p> + +<p class="tb">Lastly, one single member of the immense and diversified +class of fishes, namely the lancelet or amphioxus, +is so different from all other fishes, that Häckel maintains +that it ought to form a distinct class in the +vertebrate kingdom. This fish is remarkable for its +<span class='pagenum'><a name="Page_205" id="Page_205">205</a></span>negative characters; it can hardly be said to possess a +brain, vertebral column, or heart, &c.; so that it was +classed by the older naturalists amongst the worms. +Many years ago Prof. Goodsir perceived that the +lancelet presented some affinities with the Ascidians, +which are invertebrate, hermaphrodite, marine creatures +permanently attached to a support. They hardly +appear like animals, and consist of a simple, tough, +leathery sack, with two small projecting orifices. They +belong to the Molluscoida of Huxley—a lower division +of the great kingdom of the Mollusca; but they have +recently been placed by some naturalists amongst the +Vermes or worms. Their larvæ somewhat resemble +tadpoles in shape,<a name="FNanchor_275" id="FNanchor_275"></a><a href="#Footnote_275" class="fnanchor">275</a> and have the power of swimming +freely about. Some observations lately made by M. +Kowalevsky,<a name="FNanchor_276" id="FNanchor_276"></a><a href="#Footnote_276" class="fnanchor">276</a> since confirmed by Prof. Kuppfer, will +form a discovery of extraordinary interest, if still further +extended, as I hear from M. Kowalevsky in Naples he +has now effected. The discovery is that the larvæ of +Ascidians are related to the Vertebrata, in their manner +of development, in the relative position of the nervous +system, and in possessing a structure closely like the +<i>chorda dorsalis</i> of vertebrate animals. It thus appears, +if we may rely on embryology, which has always proved +the safest guide in classification, that we have at last +gained a clue to the source whence the Vertebrata have +<span class='pagenum'><a name="Page_206" id="Page_206">206</a></span>been derived. We should thus be justified in believing +that at an extremely remote period a group of animals +existed, resembling in many respects the larvæ of our +present Ascidians, which diverged into two great +branches—the one retrograding in development and +producing the present class of Ascidians, the other rising +to the crown and summit of the animal kingdom by +giving birth to the Vertebrata.</p> + +<p>We have thus far endeavoured rudely to trace the +genealogy of the Vertebrata by the aid of their mutual +affinities. We will now look to man as he exists; and +we shall, I think, be able partially to restore during +successive periods, but not in due order of time, the +structure of our early progenitors. This can be effected +by means of the rudiments which man still retains, by +the characters which occasionally make their appearance +in him through reversion, and by the aid of the +principles of morphology and embryology. The various +facts, to which I shall here allude, have been given in +the previous chapters. The early progenitors of man +were no doubt once covered with hair, both sexes +having beards; their ears were pointed and capable of +movement; and their bodies were provided with a tail, +having the proper muscles. Their limbs and bodies +were also acted on by many muscles which now +only occasionally reappear, but are normally present +in the Quadrumana. The great artery and nerve of +the humerus ran through a supra-condyloid foramen. +At this or some earlier period, the intestine gave forth +a much larger diverticulum or cæcum than that now +existing. The foot, judging from the condition of the +great toe in the fœtus, was then prehensile; and our +progenitors, no doubt, were arboreal in their habits, +frequenting some warm, forest-clad land. The males<span class='pagenum'><a name="Page_207" id="Page_207">207</a></span> +were provided with great canine teeth, which served +them as formidable weapons.</p> + +<p>At a much earlier period the uterus was double; the +excreta were voided through a cloaca; and the eye +was protected by a third eyelid or nictitating membrane. +At a still earlier period the progenitors of +man must have been aquatic in their habits; for +morphology plainly tells us that our lungs consist of a +modified swim-bladder, which once served as a float. +The clefts on the neck in the embryo of man show +where the branchiæ once existed. At about this period +the true kidneys were replaced by the corpora Wolffiana. +The heart existed as a simple pulsating vessel; and +the chorda dorsalis took the place of a vertebral column. +These early predecessors of man, thus seen in the dim +recesses of time, must have been as lowly organised +as the lancelet or amphioxus, or even still more lowly +organised.</p> + +<p>There is one other point deserving a fuller notice. +It has long been known that in the vertebrate kingdom +one sex bears rudiments of various accessory +parts, appertaining to the reproductive system, which +properly belong to the opposite sex; and it has now +been ascertained that at a very early embryonic period +both sexes possess true male and female glands. Hence +some extremely remote progenitor of the whole vertebrate +kingdom appears to have been hermaphrodite or +androgynous.<a name="FNanchor_277" id="FNanchor_277"></a><a href="#Footnote_277" class="fnanchor">277</a> But here we encounter a singular +<span class='pagenum'><a name="Page_208" id="Page_208">208</a></span>difficulty. In the mammalian class the males possess +in their vesiculæ prostraticæ rudiments of a uterus with +the adjacent passage; they bear also rudiments of +mammæ, and some male marsupials have rudiments +of a marsupial sack.<a name="FNanchor_278" id="FNanchor_278"></a><a href="#Footnote_278" class="fnanchor">278</a> Other analogous facts could be +added. Are we, then, to suppose that some extremely +ancient mammal possessed organs proper to both sexes, +that is, continued androgynous after it had acquired +the chief distinctions of its proper class, and therefore +after it had diverged from the lower classes of the +vertebrate kingdom? This seems improbable in the +highest degree; for had this been the case, we might +have expected that some few members of the two +lower classes, namely fishes<a name="FNanchor_279" id="FNanchor_279"></a><a href="#Footnote_279" class="fnanchor">279</a> and amphibians, would +still have remained androgynous. We must, on the +contrary, believe that when the five vertebrate classes +diverged from their common progenitor the sexes +had already become separated. To account, however, +for male mammals possessing rudiments of the accessory +female organs, and for female mammals possessing +rudiments of the masculine organs, we need not suppose +that their early progenitors were still androgynous after +they had assumed their chief mammalian characters. +It is quite possible that as the one sex gradually +acquired the accessory organs proper to it, some of the +successive steps or modifications were transmitted to +the opposite sex. When we treat of sexual selection, +we shall meet with innumerable instances of this form +of transmission,—as in the case of the spurs, plumes, +<span class='pagenum'><a name="Page_209" id="Page_209">209</a></span>and brilliant colours, acquired by male birds for battle +or ornament, and transferred to the females in an imperfect +or rudimentary condition.</p> + +<p>The possession by male mammals of functionally +imperfect mammary organs is, in some respects, especially +curious. The Monotremata have the proper milk-secreting +glands with orifices, but no nipples; and as +these animals stand at the very base of the mammalian +series, it is probable that the progenitors of the +class possessed, in like manner, the milk-secreting +glands, but no nipples. This conclusion is supported +by what is known of their manner of development; +for Professor Turner informs me, on the authority of +Kölliker and Lauger, that in the embryo the mammary +glands can be distinctly traced before the nipples are +in the least visible; and it should be borne in mind that +the development of successive parts in the individual +generally seems to represent and accord with the development +of successive beings in the same line of descent. +The Marsupials differ from the Monotremata by possessing +nipples; so that these organs were probably first +acquired by the Marsupials after they had diverged +from, and risen above, the Monotremata, and were +then transmitted to the placental mammals. No one +will suppose that after the Marsupials had approximately +acquired their present structure, and therefore +at a rather late period in the development of the +mammalian series, any of its members still remained +androgynous. We seem, therefore, compelled to recur +to the foregoing view, and to conclude that the nipples +were first developed in the females of some very early +marsupial form, and were then, in accordance with a +common law of inheritance, transferred in a functionally +imperfect condition to the males.</p> + +<p class="tb">Nevertheless a suspicion has sometimes crossed my<span class='pagenum'><a name="Page_210" id="Page_210">210</a></span> +mind that long after the progenitors of the whole +mammalian class had ceased to be androgynous, both +sexes might have yielded milk and thus nourished +their young; and in the case of the Marsupials, that +both sexes might have carried their young in marsupial +sacks. This will not appear utterly incredible, if +we reflect that the males of syngnathous fishes receive +the eggs of the females in their abdominal pouches, +hatch them, and afterwards, as some believe, nourish +the young;<a name="FNanchor_280" id="FNanchor_280"></a><a href="#Footnote_280" class="fnanchor">280</a>—that certain other male fishes hatch the +eggs within their mouths or branchial cavities;—that +certain male toads take the chaplets of eggs from the +females and wind them round their own thighs, keeping +them there until the tadpoles are born;—that certain +male birds undertake the whole duty of incubation, +and that male pigeons, as well as the females, feed their +nestlings with a secretion from their crops. But the +above suspicion first occurred to me from the mammary +glands in male mammals being developed so much more +perfectly than the rudiments of those other accessory +reproductive parts, which are found in the one sex +though proper to the other. The mammary glands +and nipples, as they exist in male mammals, can indeed +hardly be called rudimentary; they are simply not +fully developed and not functionally active. They are +sympathetically affected under the influence of certain +diseases, like the same organs in the female. At birth +they often secrete a few drops of milk; and they have +<span class='pagenum'><a name="Page_211" id="Page_211">211</a></span>been known occasionally in man and other mammals to +become well developed, and to yield a fair supply of +milk. Now if we suppose that during a former prolonged +period male mammals aided the females in +nursing their offspring, and that afterwards from some +cause, as from a smaller number of young being produced, +the males ceased giving this aid, disuse of the +organs during maturity would lead to their becoming +inactive; and from two well-known principles of inheritance +this state of inactivity would probably be +transmitted to the males at the corresponding age of +maturity. But at all earlier ages these organs would +be left unaffected, so that they would be equally well +developed in the young of both sexes.</p> + +<p class="tb"><i>Conclusion.</i>—The best definition of advancement or +progress in the organic scale ever given, is that by +Von Baer; and this rests on the amount of differentiation +and specialisation of the several parts of +the same being, when arrived, as I should be inclined +to add, at maturity. Now as organisms have become +slowly adapted by means of natural selection for +diversified lines of life, their parts will have become, +from the advantage gained by the division of physiological +labour, more and more differentiated and specialised +for various functions. The same part appears +often to have been modified first for one purpose, and +then long afterwards for some other and quite distinct +purpose; and thus all the parts are rendered more and +more complex. But each organism will still retain the +general type of structure of the progenitor from which +it was aboriginally derived. In accordance with this +view it seems, if we turn to geological evidence, that +organisation on the whole has advanced throughout the +world by slow and interrupted steps. In the great<span class='pagenum'><a name="Page_212" id="Page_212">212</a></span> +kingdom of the Vertebrata it has culminated in man. +It must not, however, be supposed that groups of organic +beings are always supplanted and disappear as soon as +they have given birth to other and more perfect groups. +The latter, though victorious over their predecessors, +may not have become better adapted for all places in +the economy of nature. Some old forms appear to have +survived from inhabiting protected sites, where they +have not been exposed to very severe competition; and +these often aid us in constructing our genealogies, +by giving us a fair idea of former and lost populations. +But we must not fall into the error of looking at the +existing members of any lowly-organised group as perfect +representatives of their ancient predecessors.</p> + +<p>The most ancient progenitors in the kingdom of the +Vertebrata, at which we are able to obtain an obscure +glance, apparently consisted of a group of marine animals,<a name="FNanchor_281" id="FNanchor_281"></a><a href="#Footnote_281" class="fnanchor">281</a> +resembling the larvæ of existing Ascidians. These +animals probably gave rise to a group of fishes, as lowly +organised as the lancelet; and from these the Ganoids, +and other fishes like the Lepidosiren, must have been +developed. From such fish a very small advance would +<span class='pagenum'><a name="Page_213" id="Page_213">213</a></span>carry us on to the amphibians. We have seen that +birds and reptiles were once intimately connected +together; and the Monotremata now, in a slight degree, +connect mammals with reptiles. But no one can at +present say by what line of descent the three higher +and related classes, namely, mammals, birds, and reptiles, +were derived from either of the two lower vertebrate +classes, namely amphibians and fishes. In the +class of mammals the steps are not difficult to conceive +which led from the ancient Monotremata to +the ancient Marsupials; and from these to the early +progenitors of the placental mammals. We may thus +ascend to the Lemuridæ; and the interval is not wide +from these to the Simiadæ. The Simiadæ then branched +off into two great stems, the New World and Old World +monkeys; and from the latter, at a remote period, Man, +the wonder and glory of the Universe, proceeded.</p> + +<p>Thus we have given to man a pedigree of prodigious +length, but not, it may be said, of noble quality. The +world, it has often been remarked, appears as if it had +long been preparing for the advent of man; and this, in +one sense is strictly true, for he owes his birth to a long +line of progenitors. If any single link in this chain +had never existed, man would not have been exactly +what he now is. Unless we wilfully close our eyes, we +may, with our present knowledge, approximately recognise +our parentage; nor need we feel ashamed of it. +The most humble organism is something much higher +than the inorganic dust under our feet; and no one +with an unbiassed mind can study any living creature, +however humble, without being struck with enthusiasm +at its marvellous structure and properties.</p> + +<hr /> +<p><span class='pagenum'><a name="Page_214" id="Page_214">214</a></span></p> + +<h3>CHAPTER VII.</h3> + +<h4><span class="smcap">On the Races of Man.</span></h4> + +<div class="blockquot"><p>The nature and value of specific characters—Application to the races +of man—Arguments in favour of, and opposed to, ranking the +so-called races of man as distinct species—Sub-species—Monogenists +and polygenists—Convergence of character—Numerous +points of resemblance in body and mind between the most distinct +races of man—The state of man when he first spread over the +earth—Each race not descended from a single pair—The extinction +of races—The formation of races—The effects of crossing—Slight +influence of the direct action of the conditions of life—Slight +or no influence of natural selection—Sexual selection.</p></div> + +<p>It is not my intention here to describe the several +so-called races of men; but to inquire what is the +value of the differences between them under a classificatory +point of view, and how they have originated. +In determining whether two or more allied forms +ought to be ranked as species or varieties, naturalists +are practically guided by the following considerations; +namely, the amount of difference between +them, and whether such differences relate to few or +many points of structure, and whether they are of +physiological importance; but more especially whether +they are constant. Constancy of character is what is +chiefly valued and sought for by naturalists. Whenever +it can be shewn, or rendered probable, that the forms +in question have remained distinct for a long period, +this becomes an argument of much weight in favour +of treating them as species. Even a slight degree of +sterility between any two forms when first crossed, or +in their offspring, is generally considered as a decisive<span class='pagenum'><a name="Page_215" id="Page_215">215</a></span> +test of their specific distinctness; and their continued +persistence without blending within the same area, is +usually accepted as sufficient evidence, either of some +degree of mutual sterility, or in the case of animals of +some repugnance to mutual pairing.</p> + +<p>Independently of blending from intercrossing, the +complete absence, in a well-investigated region, of +varieties linking together any two closely-allied forms, +is probably the most important of all the criterions +of their specific distinctness; and this is a somewhat +different consideration from mere constancy of character, +for two forms may be highly variable and yet not +yield intermediate varieties. Geographical distribution +is often unconsciously and sometimes consciously brought +into play; so that forms living in two widely separated +areas, in which most of the other inhabitants are specifically +distinct, are themselves usually looked at as distinct; +but in truth this affords no aid in distinguishing +geographical races from so-called good or true species.</p> + +<p>Now let us apply these generally-admitted principles +to the races of man, viewing him in the same spirit as +a naturalist would any other animal. In regard to the +amount of difference between the races, we must make +some allowance for our nice powers of discrimination +gained by the long habit of observing ourselves. In +India, as Elphinstone remarks,<a name="FNanchor_282" id="FNanchor_282"></a><a href="#Footnote_282" class="fnanchor">282</a> although a newly-arrived +European cannot at first distinguish the various native +races, yet they soon appear to him extremely dissimilar; +and the Hindoo cannot at first perceive any difference +between the several European nations. Even the most +distinct races of man, with the exception of certain +negro tribes, are much more like each other in form +<span class='pagenum'><a name="Page_216" id="Page_216">216</a></span>than would at first be supposed. This is well shewn by +the French photographs in the Collection Anthropologique +du Muséum of the men belonging to various +races, the greater number of which, as many persons +to whom I have shown them have remarked, might +pass for Europeans. Nevertheless, these men if seen +alive would undoubtedly appear very distinct, so that +we are clearly much influenced in our judgment by +the mere colour of the skin and hair, by slight differences +in the features, and by expression.</p> + +<p>There is, however, no doubt that the various races, +when carefully compared and measured, differ much +from each other,—as in the texture of the hair, the +relative proportions of all parts of the body,<a name="FNanchor_283" id="FNanchor_283"></a><a href="#Footnote_283" class="fnanchor">283</a> the capacity +of the lungs, the form and capacity of the skull, +and even in the convolutions of the brain.<a name="FNanchor_284" id="FNanchor_284"></a><a href="#Footnote_284" class="fnanchor">284</a> But it +would be an endless task to specify the numerous points +of structural difference. The races differ also in constitution, +in acclimatisation, and in liability to certain +diseases. Their mental characteristics are likewise very +distinct; chiefly as it would appear in their emotional, +but partly in their intellectual, faculties. Every one +who has had the opportunity of comparison, must have +been struck with the contrast between the taciturn, +even morose, aborigines of S. America and the light-hearted, +talkative negroes. There is a nearly similar +contrast between the Malays and the Papuans,<a name="FNanchor_285" id="FNanchor_285"></a><a href="#Footnote_285" class="fnanchor">285</a> who live +<span class='pagenum'><a name="Page_217" id="Page_217">217</a></span>under the same physical conditions, and are separated +from each other only by a narrow space of sea.</p> + +<p>We will first consider the arguments which may be +advanced in favour of classing the races of man as +distinct species, and then those on the other side. If a +naturalist, who had never before seen such beings, were +to compare a Negro, Hottentot, Australian, or Mongolian, +he would at once perceive that they differed in a multitude +of characters, some of slight and some of considerable +importance. On inquiry he would find that they +were adapted to live under widely different climates, and +that they differed somewhat in bodily constitution and +mental disposition. If he were then told that hundreds +of similar specimens could be brought from the same +countries, he would assuredly declare that they were as +good species as many to which he had been in the +habit of affixing specific names. This conclusion would +be greatly strengthened as soon as he had ascertained +that these forms had all retained the same character for +many centuries; and that negroes, apparently identical +with existing negroes, had lived at least 4000 years +ago.<a name="FNanchor_286" id="FNanchor_286"></a><a href="#Footnote_286" class="fnanchor">286</a> He would also hear from an excellent observer, +<span class='pagenum'><a name="Page_218" id="Page_218">218</a></span> +Dr. Lund,<a name="FNanchor_287" id="FNanchor_287"></a><a href="#Footnote_287" class="fnanchor">287</a> that the human skulls found in the caves of +Brazil, entombed with many extinct mammals, belonged +to the same type as that now prevailing throughout the +American Continent.</p> + +<p>Our naturalist would then perhaps turn to geographical +distribution, and he would probably declare that +forms differing not only in appearance, but fitted for +the hottest and dampest or driest countries, as well as +for the arctic regions, must be distinct species. He +might appeal to the fact that no one species in the +group next to man, namely the Quadrumana, can resist +a low temperature or any considerable change of climate; +and that those species which come nearest to man have +never been reared to maturity, even under the temperate +climate of Europe. He would be deeply impressed with +the fact, first noticed by Agassiz,<a name="FNanchor_288" id="FNanchor_288"></a><a href="#Footnote_288" class="fnanchor">288</a> that the different +races of man are distributed over the world in the same +zoological provinces, as those inhabited by undoubtedly +distinct species and genera of mammals. This is manifestly +the case with the Australian, Mongolian, and +Negro races of man; in a less well-marked manner with +the Hottentots; but plainly with the Papuans and +Malays, who are separated, as Mr. Wallace has shewn, +by nearly the same line which divides the great Malayan +and Australian zoological provinces. The aborigines +of America range throughout the Continent; and this +at first appears opposed to the above rule, for most of +the productions of the Southern and Northern halves +differ widely; yet some few living forms, as the +<span class='pagenum'><a name="Page_219" id="Page_219">219</a></span>opossum, range from the one into the other, as did +formerly some of the gigantic Edentata. The Esquimaux, +like other Arctic animals, extend round the +whole polar regions. It should be observed that the +mammalian forms which inhabit the several zoological +provinces, do not differ from each other in the same +degree; so that it can hardly be considered as an +anomaly that the Negro differs more, and the American +much less, from the other races of man than do the +mammals of the same continents from those of the other +provinces. Man, it may be added, does not appear to +have aboriginally inhabited any oceanic island; and in +this respect he resembles the other members of his class.</p> + +<p>In determining whether the varieties of the same +kind of domestic animal should be ranked as specifically +distinct, that is, whether any of them are descended from +distinct wild species, every naturalist would lay much +stress on the fact, if established, of their external parasites +being specifically distinct. All the more stress would be +laid on this fact, as it would be an exceptional one, for +I am informed by Mr. Denny that the most different +kinds of dogs, fowls, and pigeons, in England, are infested +by the same species of Pediculi or lice. Now Mr. A. +Murray has carefully examined the Pediculi collected in +different countries from the different races of man;<a name="FNanchor_289" id="FNanchor_289"></a><a href="#Footnote_289" class="fnanchor">289</a> +and he finds that they differ, not only in colour, but +in the structure of their claws and limbs. In every +case in which numerous specimens were obtained the +differences were constant. The surgeon of a whaling +ship in the Pacific assured me that when the Pediculi, +with which some Sandwich Islanders on board swarmed, +strayed on to the bodies of the English sailors, they +died in the course of three or four days. These Pediculi +<span class='pagenum'><a name="Page_220" id="Page_220">220</a></span>were darker coloured and appeared different from those +proper to the natives of Chiloe in South America, of +which he gave me specimens. These, again, appeared +larger and much softer than European lice. Mr. Murray +procured four kinds from Africa, namely from the +Negroes of the Eastern and Western coasts, from the +Hottentots and Caffres; two kinds from the natives +of Australia; two from North, and two from South +America. In these latter cases it may be presumed +that the Pediculi came from natives inhabiting different +districts. With insects slight structural differences, if +constant, are generally esteemed of specific value: and +the fact of the races of man being infested by parasites, +which appear to be specifically distinct, might fairly +be urged as an argument that the races themselves +ought to be classed as distinct species.</p> + +<p>Our supposed naturalist having proceeded thus far +in his investigation, would next inquire whether the +races of men, when crossed, were in any degree sterile. +He might consult the work<a name="FNanchor_290" id="FNanchor_290"></a><a href="#Footnote_290" class="fnanchor">290</a> of a cautious and philosophical +observer, Professor Broca; and in this he would +find good, evidence that some races were quite fertile +together; but evidence of an opposite nature in regard to +other races. Thus it has been asserted that the native +women of Australia and Tasmania rarely produce +children to European men; the evidence, however, on +this head has now been shewn to be almost valueless. +The half-castes are killed by the pure blacks; and an +account has lately been published of eleven half-caste +youths murdered and burnt at the same time, whose +remains were found by the police.<a name="FNanchor_291" id="FNanchor_291"></a><a href="#Footnote_291" class="fnanchor">291</a> Again, it has often +<span class='pagenum'><a name="Page_221" id="Page_221">221</a></span>been said that when mulattoes intermarry they produce +few children; on the other hand, Dr. Bachman of +Charlestown<a name="FNanchor_292" id="FNanchor_292"></a><a href="#Footnote_292" class="fnanchor">292</a> positively asserts that he has known +mulatto families which have intermarried for several +generations, and have continued on an average as fertile +as either pure whites or pure blacks. Inquiries formerly +made by Sir C. Lyell on this subject led him, as he +informs me, to the same conclusion. In the United +States the census for the year 1854 included, according +to Dr. Bachman, 405,751 mulattoes; and this number, +considering all the circumstances of the case, seems +small; but it may partly be accounted for by the degraded +and anomalous position of the class, and by the +profligacy of the women. A certain amount of absorption +of mulattoes into negroes must always be in progress; +and this would lead to an apparent diminution +of the former. The inferior vitality of mulattoes is +spoken of in a trustworthy work<a name="FNanchor_293" id="FNanchor_293"></a><a href="#Footnote_293" class="fnanchor">293</a> as a well-known +phenomenon; but this is a different consideration from +their lessened fertility; and can hardly be advanced as +a proof of the specific distinctness of the parent races. +No doubt both animal and vegetable hybrids, when +produced from extremely distinct species, are liable to +premature death; but the parents of mulattoes cannot +be put under the category of extremely distinct species. +The common Mule, so notorious for long life and vigour, +and yet so sterile, shews how little necessary connection +<span class='pagenum'><a name="Page_222" id="Page_222">222</a></span>there is in hybrids between lessened fertility and vitality: +other analogous cases could be added.</p> + +<p>Even if it should hereafter be proved that all the +races of men were perfectly fertile together, he who was +inclined from other reasons to rank them as distinct +species, might with justice argue that fertility and +sterility are not safe criterions of specific distinctness. +We know that these qualities are easily affected by +changed conditions of life or by close inter-breeding, +and that they are governed by highly complex laws, for +instance that of the unequal fertility of reciprocal crosses +between the same two species. With forms which must +be ranked as undoubted species, a perfect series exists +from those which are absolutely sterile when crossed, +to those which are almost or quite fertile. The degrees +of sterility do not coincide strictly with the degrees of +difference in external structure or habits of life. Man +in many respects may be compared with those animals +which have long been domesticated, and a large body +of evidence can be advanced in favour of the Pallasian +doctrine<a name="FNanchor_294" id="FNanchor_294"></a><a href="#Footnote_294" class="fnanchor">294</a> that domestication tends to eliminate the +<span class='pagenum'><a name="Page_223" id="Page_223">223</a></span>sterility which is so general a result of the crossing of +species in a state of nature. From these several considerations, +it may be justly urged that the perfect fertility +of the intercrossed races of man, if established, +would not absolutely preclude us from ranking them as +distinct species.</p> + +<p>Independently of fertility, the character of the offspring +from a cross has sometimes been thought to +afford evidence whether the parent-forms ought to be +ranked as species or varieties; but after carefully studying +the evidence, I have come to the conclusion that no +general rules of this kind can be trusted. Thus with +mankind the offspring of distinct races resemble in all +respects the offspring of true species and of varieties. +This is shewn, for instance, by the manner in which +the characters of both parents are blended, and by +one form absorbing another through repeated crosses. +In this latter case the progeny both of crossed species +and varieties retain for a long period a tendency to +revert to their ancestors, especially to that one which +is prepotent in transmission. When any character has +suddenly appeared in a race or species as the result of a +<span class='pagenum'><a name="Page_224" id="Page_224">224</a></span>single act of variation, as is general with monstrosities,<a name="FNanchor_295" id="FNanchor_295"></a><a href="#Footnote_295" class="fnanchor">295</a> +and this race is crossed with another not thus characterised, +the characters in question do not commonly +appear in a blended condition in the young, but are +transmitted to them either perfectly developed or not at +all. As with the crossed races of man cases of this kind +rarely or never occur, this may be used as an argument +against the view suggested by some ethnologists, namely +that certain characters, for instance the blackness of the +negro, first appeared as a sudden variation or sport. +Had this occurred, it is probable that mulattoes would +often have been born, either completely black or completely +white.</p> + +<p class="tb">We have now seen that a naturalist might feel himself +fully justified in ranking the races of man as distinct +species; for he has found that they are distinguished by +many differences in structure and constitution, some +being of importance. These differences have, also, remained +nearly constant for very long periods of time. +He will have been in some degree influenced by the +enormous range of man, which is a great anomaly in +the class of mammals, if mankind be viewed as a single +species. He will have been struck with the distribution +of the several so-called races, in accordance with that +of other undoubtedly distinct species of mammals. +Finally he might urge that the mutual fertility of all +the races has not as yet been fully proved; and even if +proved would not be an absolute proof of their specific +identity.</p> + +<p>On the other side of the question, if our supposed +naturalist were to enquire whether the forms of man +kept distinct like ordinary species, when mingled to<span class='pagenum'><a name="Page_225" id="Page_225">225</a></span>gether +in large numbers in the same country, he would +immediately discover that this was by no means the +case. In Brazil he would behold an immense mongrel +population of Negroes and Portuguese; in Chiloe and +other parts of South America, he would behold the +whole population consisting of Indians and Spaniards +blended in various degrees.<a name="FNanchor_296" id="FNanchor_296"></a><a href="#Footnote_296" class="fnanchor">296</a> In many parts of the +same continent he would meet with the most complex +crosses between Negroes, Indians, and Europeans; and +such triple crosses afford the severest test, judging from +the vegetable kingdom, of the mutual fertility of the +parent-forms. In one island of the Pacific he would +find a small population of mingled Polynesian and +English blood; and in the Viti Archipelago a population +of Polynesians and Negritos crossed in all degrees. +Many analogous cases could be added, for instance, in +South Africa. Hence the races of man are not sufficiently +distinct to co-exist without fusion; and this it +is, which in all ordinary cases affords the usual test of +specific distinctness.</p> + +<p>Our naturalist would likewise be much disturbed as +soon as he perceived that the distinctive characters of +every race of man were highly variable. This strikes +every one when he first beholds the negro-slaves in +Brazil, who have been imported from all parts of Africa. +The same remark holds good with the Polynesians, and +with many other races. It may be doubted whether +any character can be named which is distinctive of a +race and is constant. Savages, even within the limits of +the same tribe, are not nearly so uniform in character, +as has often been said. Hottentot women offer certain +<span class='pagenum'><a name="Page_226" id="Page_226">226</a></span>peculiarities, more strongly marked than those occurring +in any other race, but these are known not to +be of constant occurrence. In the several American +tribes, colour and hairyness differ considerably; as does +colour to a certain degree, and the shape of the features +greatly, in the Negroes of Africa. The shape of the +skull varies much in some races;<a name="FNanchor_297" id="FNanchor_297"></a><a href="#Footnote_297" class="fnanchor">297</a> and so it is with +every other character. Now all naturalists have learnt +by dearly-bought experience, how rash it is to attempt +to define species by the aid of inconstant characters.</p> + +<p>But the most weighty of all the arguments against +treating the races of man as distinct species, is that they +graduate into each other, independently in many cases, +as far as we can judge, of their having intercrossed. +Man has been studied more carefully than any other +organic being, and yet there is the greatest possible +diversity amongst capable judges whether he should be +classed as a single species or race, or as two (Virey), as +three (Jacquinot), as four (Kant), five (Blumenbach), +six (Buffon), seven (Hunter), eight (Agassiz), eleven +(Pickering), fifteen (Bory St. Vincent), sixteen (Desmoulins), +twenty-two (Morton), sixty (Crawfurd), or as +sixty-three, according to Burke.<a name="FNanchor_298" id="FNanchor_298"></a><a href="#Footnote_298" class="fnanchor">298</a> This diversity of +judgment does not prove that the races ought not to be +ranked as species, but it shews that they graduate into +each other, and that it is hardly possible to discover +clear distinctive characters between them.</p> + +<p>Every naturalist who has had the misfortune to under<span class='pagenum'><a name="Page_227" id="Page_227">227</a></span>take +the description of a group of highly varying +organisms, has encountered cases (I speak after experience) +precisely like that of man; and if of a cautious +disposition, he will end by uniting all the forms which +graduate into each other as a single species; for he will +say to himself that he has no right to give names to +objects which he cannot define. Cases of this kind occur +in the Order which includes man, namely in certain +genera of monkeys; whilst in other genera, as in Cercopithecus, +most of the species can be determined with +certainty. In the American genus Cebus, the various +forms are ranked by some naturalists as species, by +others as mere geographical races. Now if numerous +specimens of Cebus were collected from all parts of +South America, and those forms which at present appear +to be specifically distinct, were found to graduate +into each other by close steps, they would be ranked by +most naturalists as mere varieties or races; and thus the +greater number of naturalists have acted with respect +to the races of man. Nevertheless it must be confessed +that there are forms, at least in the vegetable kingdom,<a name="FNanchor_299" id="FNanchor_299"></a><a href="#Footnote_299" class="fnanchor">299</a> +which we cannot avoid naming as species, but +which are connected together, independently of intercrossing, +by numberless gradations.</p> + +<p class="tb">Some naturalists have lately employed the term +“sub-species” to designate forms which possess many of +the characteristics of true species, but which hardly deserve +so high a rank. Now if we reflect on the weighty +arguments, above given, for raising the races of man to +the dignity of species, and the insuperable difficulties +on the other side in defining them, the term “sub-species”<span class='pagenum'><a name="Page_228" id="Page_228">228</a></span> +might here be used with much propriety. But +from long habit the term “race” will perhaps always +be employed. The choice of terms is only so far important +as it is highly desirable to use, as far as that +may be possible, the same terms for the same degrees of +difference. Unfortunately this is rarely possible; for +within the same family the larger genera generally +include closely-allied forms, which can be distinguished +only with much difficulty, whilst the smaller genera +include forms that are perfectly distinct; yet all must +equally be ranked as species. So again the species +within the same large genus by no means resemble +each other to the same degree: on the contrary, in +most cases some of them can be arranged in little groups +round other species, like satellites round planets.<a name="FNanchor_300" id="FNanchor_300"></a><a href="#Footnote_300" class="fnanchor">300</a></p> + +<p>The question whether mankind consists of one or +several species has of late years been much agitated by +anthropologists, who are divided into two schools of +monogenists and polygenists. Those who do not admit +the principle of evolution, must look at species either +as separate creations or as in some manner distinct +entities; and they must decide what forms to rank as +species by the analogy of other organic beings which +are commonly thus received. But it is a hopeless endeavour +to decide this point on sound grounds, until +some definition of the term “species” is generally accepted; +and the definition must not include an element +which cannot possibly be ascertained, such as an act of +creation. We might as well attempt without any definition +to decide whether a certain number of houses +should be called a village, or town, or city. We have a +practical illustration of the difficulty in the never-<span class='pagenum'><a name="Page_229" id="Page_229">229</a></span>ending +doubts whether many closely-allied mammals, +birds, insects, and plants, which represent each other in +North America and Europe, should be ranked species +or geographical races; and so it is with the productions +of many islands situated at some little distance from the +nearest continent.</p> + +<p>Those naturalists, on the other hand, who admit the +principle of evolution, and this is now admitted by the +greater number of rising men, will feel no doubt that +all the races of man are descended from a single primitive +stock; whether or not they think fit to designate +them as distinct species, for the sake of expressing their +amount of difference.<a name="FNanchor_301" id="FNanchor_301"></a><a href="#Footnote_301" class="fnanchor">301</a> With our domestic animals the +question whether the various races have arisen from +one or more species is different. Although all such +races, as well as all the natural species within the same +genus, have undoubtedly sprung from the same primitive +stock, yet it is a fit subject for discussion, whether, +for instance, all the domestic races of the dog have +acquired their present differences since some one species +was first domesticated and bred by man; or whether they +owe some of their characters to inheritance from distinct +species, which had already been modified in a state of +nature. With mankind no such question can arise, for +he cannot be said to have been domesticated at any +particular period.</p> + +<p>When the races of man diverged at an extremely +remote epoch from their common progenitor, they will +have differed but little from each other, and been few +in number; consequently they will then, as far as their +distinguishing characters are concerned, have had less +claim to rank as distinct species, than the existing so<span class='pagenum'><a name="Page_230" id="Page_230">230</a></span>-called +races. Nevertheless such early races would perhaps +have been ranked by some naturalists as distinct +species, so arbitrary is the term, if their differences, +although extremely slight, had been more constant than +at present, and had not graduated into each other.</p> + +<p>It is, however, possible, though far from probable, +that the early progenitors of man might at first have +diverged much in character, until they became more +unlike each other than are any existing races; but that +subsequently, as suggested by Vogt,<a name="FNanchor_302" id="FNanchor_302"></a><a href="#Footnote_302" class="fnanchor">302</a> they converged +in character. When man selects for the same object +the offspring of two distinct species, he sometimes +induces, as far as general appearance is concerned, +a considerable amount of convergence. This is the +case, as shewn by Von Nathusius,<a name="FNanchor_303" id="FNanchor_303"></a><a href="#Footnote_303" class="fnanchor">303</a> with the improved +breeds of pigs, which are descended from two distinct +species; and in a less well-marked manner with the +improved breeds of cattle. A great anatomist, Gratiolet, +maintains that the anthropomorphous apes do not form +a natural sub-group; but that the orang is a highly +developed gibbon or semnopithecus; the chimpanzee +a highly developed macacus; and the gorilla a highly +developed mandrill. If this conclusion, which rests +almost exclusively on brain-characters, be admitted, +we should have a case of convergence at least in +external characters, for the anthropomorphous apes are +certainly more like each other in many points than +they are to other apes. All analogical resemblances, +as of a whale to a fish, may indeed be said to be +cases of convergence; but this term has never been +applied to superficial and adaptive resemblances. It +<span class='pagenum'><a name="Page_231" id="Page_231">231</a></span>would be extremely rash in most cases to attribute to +convergence close similarity in many points of structure +in beings which had once been widely different. +The form of a crystal is determined solely by the +molecular forces, and it is not surprising that dissimilar +substances should sometimes assume the same form; +but with organic beings we should bear in mind that +the form of each depends on an infinitude of complex +relations, namely on the variations which have arisen, +these being due to causes far too intricate to be followed +out,—on the nature of the variations which have been +preserved, and this depends on the surrounding physical +conditions, and in a still higher degree on the surrounding +organisms with which each has come into +competition,—and lastly, on inheritance (in itself a +fluctuating element) from innumerable progenitors, all +of which have had their forms determined through +equally complex relations. It appears utterly incredible +that two organisms, if differing in a marked manner, +should ever afterwards converge so closely as to lead +to a near approach to identity throughout their whole +organisation. In the case of the convergent pigs above +referred to, evidence of their descent from two primitive +stocks is still plainly retained, according to Von Nathusius, +in certain bones of their skulls. If the races of man +were descended, as supposed by some naturalists, from +two or more distinct species, which had differed as much, +or nearly as much, from each other, as the orang differs +from the gorilla, it can hardly be doubted that marked +differences in the structure of certain bones would still +have been discoverable in man as he now exists.</p> + +<p>Although the existing races of man differ in many +respects, as in colour, hair, shape of skull, proportions +of the body, &c., yet if their whole organisation be taken<span class='pagenum'><a name="Page_232" id="Page_232">232</a></span> +into consideration they are found to resemble each +other closely in a multitude of points. Many of these +points are of so unimportant or of so singular a nature, +that it is extremely improbable that they should have +been independently acquired by aboriginally distinct +species or races. The same remark holds good with +equal or greater force with respect to the numerous +points of mental similarity between the most distinct +races of man. The American aborigines, Negroes and +Europeans differ as much from each other in mind as any +three races that can be named; yet I was incessantly +struck, whilst living with the Fuegians on board the +“Beagle,” with the many little traits of character, +shewing how similar their minds were to ours; and so +it was with a full-blooded negro with whom I happened +once to be intimate.</p> + +<p>He who will carefully read Mr. Tylor’s and Sir +J. Lubbock’s interesting works<a name="FNanchor_304" id="FNanchor_304"></a><a href="#Footnote_304" class="fnanchor">304</a> can hardly fail to be +deeply impressed with the close similarity between +the men of all races in tastes, dispositions and habits. +This is shewn by the pleasure which they all take in +dancing, rude music, acting, painting, tattooing, and +otherwise decorating themselves,—in their mutual +comprehension of gesture-language—and, as I shall be +able to shew in a future essay, by the same expression +in their features, and by the same inarticulate cries, +when they are excited by various emotions. This +similarity, or rather identity, is striking, when contrasted +with the different expressions which may be observed +in distinct species of monkeys. There is good evidence +that the art of shooting with bows and arrows has +not been handed down from any common progenitor of +<span class='pagenum'><a name="Page_233" id="Page_233">233</a></span>mankind, yet the stone arrow-heads, brought from the +most distant parts of the world and manufactured at the +most remote periods, are, as Nilsson has shewn,<a name="FNanchor_305" id="FNanchor_305"></a><a href="#Footnote_305" class="fnanchor">305</a> almost +identical; and this fact can only be accounted for by +the various races having similar inventive or mental +powers. The same observation has been made by +archæologists<a name="FNanchor_306" id="FNanchor_306"></a><a href="#Footnote_306" class="fnanchor">306</a> with respect to certain widely-prevalent +ornaments, such as zigzags, &c.; and with respect to +various simple beliefs and customs, such as the burying +of the dead under megalithic structures. I remember +observing in South America,<a name="FNanchor_307" id="FNanchor_307"></a><a href="#Footnote_307" class="fnanchor">307</a> that there, as in so many +other parts of the world, man has generally chosen the +summits of lofty hills, on which to throw up piles of +stones, either for the sake of recording some remarkable +event, or for burying his dead.</p> + +<p>Now when naturalists observe a close agreement in +numerous small details of habits, tastes and dispositions +between two or more domestic races, or between nearly-allied +natural forms, they use this fact as an argument +that all are descended from a common progenitor who +was thus endowed; and consequently that all should be +classed under the same species. The same argument +may be applied with much force to the races of man.</p> + +<p>As it is improbable that the numerous and unimportant +points of resemblance between the several races +of man in bodily structure and mental faculties (I do +not here refer to similar customs) should all have been +independently acquired, they must have been inherited +from progenitors who were thus characterised. We +thus gain some insight into the early state of man, +<span class='pagenum'><a name="Page_234" id="Page_234">234</a></span>before he had spread step by step over the face of the +earth. The spreading of man to regions widely separated +by the sea, no doubt, preceded any considerable +amount of divergence of character in the several races; +for otherwise we should sometimes meet with the same +race in distinct continents; and this is never the case. +Sir J. Lubbock, after comparing the arts now practised +by savages in all parts of the world, specifies those +which man could not have known, when he first wandered +from his original birthplace; for if once learnt +they would never have been forgotten.<a name="FNanchor_308" id="FNanchor_308"></a><a href="#Footnote_308" class="fnanchor">308</a> He thus shews +that “the spear, which is but a development of the +knife-point, and the club, which is but a long hammer, +are the only things left.” He admits, however, that +the art of making fire probably had already been discovered, +for it is common to all the races now existing, +and was known to the ancient cave-inhabitants of +Europe. Perhaps the art of making rude canoes or +rafts was likewise known; but as man existed at a remote +epoch, when the land in many places stood at a +very different level, he would have been able, without +the aid of canoes, to have spread widely. Sir J. Lubbock +further remarks how improbable it is that our earliest +ancestors could have “counted as high as ten, considering +that so many races now in existence cannot get +beyond four.” Nevertheless, at this early period, the +intellectual and social faculties of man could hardly have +been inferior in any extreme degree to those now possessed +by the lowest savages; otherwise primeval man +could not have been so eminently successful in the +struggle for life, as proved by his early and wide +diffusion.</p> + +<p>From the fundamental differences between certain +<span class='pagenum'><a name="Page_235" id="Page_235">235</a></span>languages, some philologists have inferred that when +man first became widely diffused he was not a speaking +animal; but it may be suspected that languages, far +less perfect than any now spoken, aided by gestures, +might have been used, and yet have left no traces +on subsequent and more highly-developed tongues. +Without the use of some language, however imperfect, +it appears doubtful whether man’s intellect could have +risen to the standard implied by his dominant position +at an early period.</p> + +<p class="tb">Whether primeval man, when he possessed very few +arts of the rudest kind, and when his power of language +was extremely imperfect, would have deserved to be +called man, must depend on the definition which we +employ. In a series of forms graduating insensibly +from some ape-like creature to man as he now exists, +it would be impossible to fix on any definite point when +the term “man” ought to be used. But this is a matter +of very little importance. So again it is almost a +matter of indifference whether the so-called races of +man are thus designated, or are ranked as species +or sub-species; but the latter term appears the most +appropriate. Finally, we may conclude that when +the principles of evolution are generally accepted, as +they surely will be before long, the dispute between the +monogenists and the polygenists will die a silent and +unobserved death.</p> + +<p class="tb">One other question ought not to be passed over +without notice, namely, whether, as is sometimes +assumed, each sub-species or race of man has sprung +from a single pair of progenitors. With our domestic +animals a new race can readily be formed from a single +pair possessing some new character, or even from a +single individual thus characterised, by carefully match<span class='pagenum'><a name="Page_236" id="Page_236">236</a></span>ing +the varying offspring; but most of our races have +been formed, not intentionally from a selected pair, +but unconsciously by the preservation of many individuals +which have varied, however slightly, in some +useful or desired manner. If in one country stronger +and heavier horses, and in another country lighter and +fleeter horses, were habitually preferred, we may feel +sure that two distinct sub-breeds would, in the course +of time, be produced, without any particular pairs +or individuals having been separated and bred from +in either country. Many races have been thus formed, +and their manner of formation is closely analogous with +that of natural species. We know, also, that the +horses which have been brought to the Falkland +Islands have become, during successive generations, +smaller and weaker, whilst those which have run wild +on the Pampas have acquired larger and coarser +heads; and such changes are manifestly due, not to +any one pair, but to all the individuals having been +subjected to the same conditions, aided, perhaps, by +the principle of reversion. The new sub-breeds in +none of these cases are descended from any single +pair, but from many individuals which have varied in +different degrees, but in the same general manner; +and we may conclude that the races of man have been +similarly produced, the modifications being either the +direct result of exposure to different conditions, or the +indirect result of some form of selection. But to this +latter subject we shall presently return.</p> + +<p><i>On the Extinction of the Races of Man.</i>—The partial +and complete extinction of many races and sub-races +of man are historically known events. Humboldt saw +in South America a parrot which was the sole living +creature that could speak the language of a lost tribe.<span class='pagenum'><a name="Page_237" id="Page_237">237</a></span> +Ancient monuments and stone implements found in +all parts of the world, of which no tradition is preserved +by the present inhabitants, indicate much +extinction. Some small and broken tribes, remnants +of former races, still survive in isolated and generally +mountainous districts. In Europe the ancient +races were all, according to Schaaffhausen,<a name="FNanchor_309" id="FNanchor_309"></a><a href="#Footnote_309" class="fnanchor">309</a> “lower in +the scale than the rudest living savages;” they must +therefore have differed, to a certain extent, from any +existing race. The remains described by Professor +Broca<a name="FNanchor_310" id="FNanchor_310"></a><a href="#Footnote_310" class="fnanchor">310</a> from Les Eyzies, though they unfortunately +appear to have belonged to a single family, indicate a +race with a most singular combination of low or simious +and high characteristics, and is “entirely different +from any other race, ancient or modern, that we have +ever heard of.” It differed, therefore, from the quaternary +race of the caverns of Belgium.</p> + +<p>Unfavourable physical conditions appear to have had +but little effect in the extinction of races.<a name="FNanchor_311" id="FNanchor_311"></a><a href="#Footnote_311" class="fnanchor">311</a> Man has +long lived in the extreme regions of the North, with +no wood wherewith to make his canoes or other implements, +and with blubber alone for burning and giving +him warmth, but more especially for melting the snow. +In the Southern extremity of America the Fuegians +survive without the protection of clothes, or of any +building worthy to be called a hovel. In South Africa +the aborigines wander over the most arid plains, where +dangerous beasts abound. Man can withstand the +deadly influence of the Terai at the foot of the Himalaya, +and the pestilential shores of tropical Africa.</p> + +<p><span class='pagenum'><a name="Page_238" id="Page_238">238</a></span></p><p>Extinction follows chiefly from the competition of +tribe with tribe, and race with race. Various checks +are always in action, as specified in a former chapter, +which serve to keep down the numbers of each savage +tribe,—such as periodical famines, the wandering of +the parents and the consequent deaths of infants, prolonged +suckling, the stealing of women, wars, accidents, +sickness, licentiousness, especially infanticide, and, +perhaps, lessened fertility from less nutritious food, and +many hardships. If from any cause any one of these +checks is lessened, even in a slight degree, the tribe +thus favoured will tend to increase; and when one +of two adjoining tribes becomes more numerous and +powerful than the other, the contest is soon settled by +war, slaughter, cannibalism, slavery, and absorption. +Even when a weaker tribe is not thus abruptly swept +away, if it once begins to decrease, it generally goes on +decreasing until it is extinct.<a name="FNanchor_312" id="FNanchor_312"></a><a href="#Footnote_312" class="fnanchor">312</a></p> + +<p>When civilised nations come into contact with barbarians +the struggle is short, except where a deadly climate +gives its aid to the native race. Of the causes which +lead to the victory of civilised nations, some are plain +and some very obscure. We can see that the cultivation +of the land will be fatal in many ways to savages, for +they cannot, or will not, change their habits. New +diseases and vices are highly destructive; and it appears +that in every nation a new disease causes much death, +until those who are most susceptible to its destructive +influence are gradually weeded out;<a name="FNanchor_313" id="FNanchor_313"></a><a href="#Footnote_313" class="fnanchor">313</a> and so it may be +with the evil effects from spirituous liquors, as well as +with the unconquerably strong taste for them shewn by +so many savages. It further appears, mysterious as is +<span class='pagenum'><a name="Page_239" id="Page_239">239</a></span>the fact, that the first meeting of distinct and separated +people generates disease.<a name="FNanchor_314" id="FNanchor_314"></a><a href="#Footnote_314" class="fnanchor">314</a> Mr. Sproat, who in Vancouver +Island closely attended to the subject of extinction, +believes that changed habits of life, which always +follow from the advent of Europeans, induces much ill-health. +He lays, also, great stress on so trifling a cause +as that the natives become “bewildered and dull by the +new life around them; they lose the motives for exertion, +and get no new ones in their place.”<a name="FNanchor_315" id="FNanchor_315"></a><a href="#Footnote_315" class="fnanchor">315</a></p> + +<p>The grade of civilisation seems a most important +element in the success of nations which come in competition. +A few centuries ago Europe feared the inroads +of Eastern barbarians; now, any such fear would be ridiculous. +It is a more curious fact, that savages did not +formerly waste away, as Mr. Bagehot has remarked, +before the classical nations, as they now do before +modern civilised nations; had they done so, the old +moralists would have mused over the event; but there +is no lament in any writer of that period over the perishing +barbarians.<a name="FNanchor_316" id="FNanchor_316"></a><a href="#Footnote_316" class="fnanchor">316</a></p> + +<p class="tb">Although the gradual decrease and final extinction +of the races of man is an obscure problem, we can see +that it depends on many causes, differing in different +places and at different times. It is the same difficult +problem as that presented by the extinction of one of +the higher animals—of the fossil horse, for instance, +which disappeared from South America, soon afterwards +to be replaced, within the same districts, by countless +troops of the Spanish horse. The New Zealander seems +<span class='pagenum'><a name="Page_240" id="Page_240">240</a></span>conscious of this parallelism, for he compares his future +fate with that of the native rat almost exterminated by +the European rat. The difficulty, though great to our +imagination, and really great if we wish to ascertain +the precise causes, ought not to be so to our reason, +as long as we keep steadily in mind that the increase of +each species and each race is constantly hindered by +various checks; so that if any new check, or cause of +destruction, even a slight one, be superadded, the race +will surely decrease in number; and as it has everywhere +been observed that savages are much opposed to +any change of habits, by which means injurious checks +could be counterbalanced, decreasing numbers will +sooner or later lead to extinction; the end, in most +cases, being promptly determined by the inroads of +increasing and conquering tribes.</p> + +<p class="tb"><i>On the Formation of the Races of Man.</i>—It may be +premised that when we find the same race, though +broken up into distinct tribes, ranging over a great +area, as over America, we may attribute their general +resemblance to descent from a common stock. In some +cases the crossing of races already, distinct has led to +the formation of new races. The singular fact that +Europeans and Hindoos, who belong to the same Aryan +stock and speak a language fundamentally the same, +differ widely in appearance, whilst Europeans differ +but little from Jews, who belong to the Semitic stock +and speak quite another language, has been accounted +for by Broca<a name="FNanchor_317" id="FNanchor_317"></a><a href="#Footnote_317" class="fnanchor">317</a> through the Aryan branches having been +largely crossed during their wide diffusion by various +indigenous tribes. When two races in close contact +<span class='pagenum'><a name="Page_241" id="Page_241">241</a></span>cross, the first result is a heterogeneous mixture: +thus Mr. Hunter, in describing the Santali or hill-tribes +of India, says that hundreds of imperceptible +gradations may be traced “from the black, squat tribes +of the mountains to the tall olive-coloured Brahman, +with his intellectual brow, calm eyes, and high but +narrow head;” so that it is necessary in courts of +justice to ask the witnesses whether they are Santalis +or Hindoos.<a name="FNanchor_318" id="FNanchor_318"></a><a href="#Footnote_318" class="fnanchor">318</a> Whether a heterogeneous people, such +as the inhabitants of some of the Polynesian islands, +formed by the crossing of two distinct races, with few +or no pure members left, would ever become homogeneous, +is not known from direct evidence. But as +with our domesticated animals, a crossed breed can +certainly, in the course of a few generations, be fixed +and made uniform by careful selection,<a name="FNanchor_319" id="FNanchor_319"></a><a href="#Footnote_319" class="fnanchor">319</a> we may infer +that the free and prolonged intercrossing during many +generations of a heterogeneous mixture would supply +the place of selection, and overcome any tendency to +reversion, so that a crossed race would ultimately become +homogeneous, though it might not partake in an +equal degree of the characters of the two parent-races.</p> + +<p>Of all the differences between the races of man, the +colour of the skin is the most conspicuous and one of +the best marked. Differences of this kind, it was formerly +thought, could be accounted for by long exposure +under different climates; but Pallas first shewed +that this view is not tenable, and he has been followed +by almost all anthropologists.<a name="FNanchor_320" id="FNanchor_320"></a><a href="#Footnote_320" class="fnanchor">320</a> The view has been +<span class='pagenum'><a name="Page_242" id="Page_242">242</a></span>rejected chiefly because the distribution of the variously +coloured races, most of whom must have long inhabited +their present homes, does not coincide with corresponding +differences of climate. Weight must also be given +to such cases as that of the Dutch families, who, as +we hear on excellent authority,<a name="FNanchor_321" id="FNanchor_321"></a><a href="#Footnote_321" class="fnanchor">321</a> have not undergone +the least change of colour, after residing for three centuries +in South Africa. The uniform appearance in +various parts of the world of gypsies and Jews, though +the uniformity of the latter has been somewhat exaggerated,<a name="FNanchor_322" id="FNanchor_322"></a><a href="#Footnote_322" class="fnanchor">322</a> +is likewise an argument on the same side. A +very damp or a very dry atmosphere has been supposed +to be more influential in modifying the colour of the +skin than mere heat; but as D’Orbigny in South +America, and Livingstone in Africa, arrived at diametrically +opposite conclusions with respect to dampness +and dryness, any conclusion on this head must be considered +as very doubtful.<a name="FNanchor_323" id="FNanchor_323"></a><a href="#Footnote_323" class="fnanchor">323</a></p> + +<p>Various facts, which I have elsewhere given, prove +that the colour of the skin and hair is sometimes correlated +in a surprising manner with a complete immunity +from the action of certain vegetable poisons and from +the attacks of certain parasites. Hence it occurred to +me, that negroes and other dark races might have +acquired their dark tints by the darker individuals +escaping during a long series of generations from +the deadly influence of the miasmas of their native +countries.</p> + +<p>I afterwards found that the same idea had long ago +<span class='pagenum'><a name="Page_243" id="Page_243">243</a></span>occurred to Dr. Wells.<a name="FNanchor_324" id="FNanchor_324"></a><a href="#Footnote_324" class="fnanchor">324</a> That negroes, and even mulattoes, +are almost completely exempt from the yellow-fever, +which is so destructive in tropical America, has +long been known.<a name="FNanchor_325" id="FNanchor_325"></a><a href="#Footnote_325" class="fnanchor">325</a> They likewise escape to a large +extent the fatal intermittent fevers that prevail along, +at least, 2600 miles of the shores of Africa, and which +annually cause one-fifth of the white settlers to die, and +another fifth to return home invalided.<a name="FNanchor_326" id="FNanchor_326"></a><a href="#Footnote_326" class="fnanchor">326</a> This immunity +in the negro seems to be partly inherent, depending +on some unknown peculiarity of constitution, +and partly the result of acclimatisation. Pouchet<a name="FNanchor_327" id="FNanchor_327"></a><a href="#Footnote_327" class="fnanchor">327</a> +states that the negro regiments, borrowed from the +Viceroy of Egypt for the Mexican war, which had been +recruited near the Soudan, escaped the yellow-fever +almost equally well with the negroes originally brought +from various parts of Africa, and accustomed to the +climate of the West Indies. That acclimatisation plays +a part is shewn by the many cases in which negroes, +after having resided for some time in a colder climate, +have become to a certain extent liable to tropical +fevers.<a name="FNanchor_328" id="FNanchor_328"></a><a href="#Footnote_328" class="fnanchor">328</a> The nature of the climate under which the +white races have long resided, likewise has some influence +on them; for during the fearful epidemic of +yellow-fever in Demerara during 1837, Dr. Blair found +that the death-rate of the immigrants was proportional +<span class='pagenum'><a name="Page_244" id="Page_244">244</a></span>to the latitude of the country whence they had come. +With the negro the immunity, as far as it is the result +of acclimatisation, implies exposure during a prodigious +length of time; for the aborigines of tropical America, +who have resided there from time immemorial, are not +exempt from yellow-fever; and the Rev. B. Tristram +states, that there are districts in Northern Africa which +the native inhabitants are compelled annually to leave, +though the negroes can remain with safety.</p> + +<p>That the immunity of the negro is in any degree +correlated with the colour of his skin is a mere conjecture: +it may be correlated with some difference in his +blood, nervous system, or other tissues. Nevertheless, +from the facts above alluded to, and from some connection +apparently existing between complexion and a +tendency to consumption, the conjecture seemed to me +not improbable. Consequently I endeavoured, with but +little success,<a name="FNanchor_329" id="FNanchor_329"></a><a href="#Footnote_329" class="fnanchor">329</a> to ascertain how far it held good. The +<span class='pagenum'><a name="Page_245" id="Page_245">245</a></span>late Dr. Daniell, who had long lived on the West Coast +of Africa, told me that he did not believe in any such +relation. He was himself unusually fair, and had withstood +the climate in a wonderful manner. When he +first arrived as a boy on the coast, an old and experienced +negro chief predicted from his appearance that +this would prove the case. Dr. Nicholson, of Antigua, +after having attended to this subject, wrote to me that +he did not think that dark-coloured Europeans escaped +the yellow-fever better than those that were light-coloured. +Mr. J. M. Harris altogether denies<a name="FNanchor_330" id="FNanchor_330"></a><a href="#Footnote_330" class="fnanchor">330</a> that +Europeans with dark hair withstand a hot climate +better than other men; on the contrary, experience has +taught him in making a selection of men for service +on the coast of Africa, to choose those with red hair. +As far, therefore, as these slight indications serve, there +seems no foundation for the hypothesis, which has been +accepted by several writers, that the colour of the black +races may have resulted from darker and darker individuals +having survived in greater numbers, during +their exposure to the fever-generating miasmas of their +native countries.</p> + +<p>Although with our present knowledge we cannot +account for the strongly-marked differences in colour +between the races of man, either through correlation +with constitutional peculiarities, or through the direct +action of climate; yet we must not quite ignore the +<span class='pagenum'><a name="Page_246" id="Page_246">246</a></span>latter agency, for there is good reason to believe that +some inherited effect is thus produced.<a name="FNanchor_331" id="FNanchor_331"></a><a href="#Footnote_331" class="fnanchor">331</a></p> + +<p>We have seen in our third chapter that the conditions +of life, such as abundant food and general comfort, +affect in a direct manner the development of the bodily +frame, the effects being transmitted. Through the +combined influences of climate and changed habits of +life, European settlers, in the United States undergo, as +is generally admitted, a slight but extraordinarily rapid +change of appearance. There is, also, a considerable +body of evidence shewing that in the Southern States +the house-slaves of the third generation present a +markedly different appearance from the field-slaves.<a name="FNanchor_332" id="FNanchor_332"></a><a href="#Footnote_332" class="fnanchor">332</a></p> + +<p>If, however, we look to the races of man, as distributed +over the world, we must infer that their characteristic +differences cannot be accounted for by the +direct action of different conditions of life, even after +exposure to them for an enormous period of time. The +Esquimaux live exclusively on animal food; they are +clothed in thick fur, and are exposed to intense cold +and to prolonged darkness; yet they do not differ in +any extreme degree from the inhabitants of Southern +China, who live entirely on vegetable food and are exposed +almost naked to a hot, glaring climate. The unclothed +Fuegians live on the marine productions of their +inhospitable shores; the Botocudos of Brazil wander +<span class='pagenum'><a name="Page_247" id="Page_247">247</a></span>about the hot forests of the interior and live chiefly on +vegetable productions; yet these tribes resemble each +other so closely that the Fuegians on board the “Beagle” +were mistaken by some Brazilians for Botocudos. The +Botocudos again, as well as the other inhabitants of +tropical America, are wholly different from the Negroes +who inhabit the opposite shores of the Atlantic, are +exposed to a nearly similar climate, and follow nearly +the same habits of life.</p> + +<p>Nor can the differences between the races of man be +accounted for, except to a quite insignificant degree, by +the inherited effects of the increased or decreased use of +parts. Men who habitually live in canoes, may have +their legs somewhat stunted; those who inhabit lofty +regions have their chests enlarged; and those who constantly +use certain sense-organs have the cavities in +which they are lodged somewhat increased in size, and +their features consequently a little modified. With +civilised nations, the reduced size of the jaws from +lessened use, the habitual play of different muscles +serving to express different emotions, and the increased +size of the brain from greater intellectual activity, have +together produced a considerable effect on their general +appearance in comparison with savages.<a name="FNanchor_333" id="FNanchor_333"></a><a href="#Footnote_333" class="fnanchor">333</a> It is also +possible that increased bodily stature, with no corresponding +increase in the size of the brain, may have +given to some races (judging from the previously adduced +cases of the rabbits) an elongated skull of the +dolichocephalic type.</p> + +<p class="tb">Lastly, the little-understood principle of correlation +will almost certainly have come into action, as in the +case of great muscular development and strongly pro<span class='pagenum'><a name="Page_248" id="Page_248">248</a></span>jecting +supra-orbital ridges. It is not improbable that +the texture of the hair, which differs much in the different +races, may stand in some kind of correlation with +the structure of the skin; for the colour of the hair +and skin are certainly correlated, as is its colour and +texture with the Mandans.<a name="FNanchor_334" id="FNanchor_334"></a><a href="#Footnote_334" class="fnanchor">334</a> The colour of the skin +and the odour emitted by it are likewise in some manner +connected. With the breeds of sheep the number +of hairs within a given space and the number of the +excretory pores stand in some relation to each other.<a name="FNanchor_335" id="FNanchor_335"></a><a href="#Footnote_335" class="fnanchor">335</a> +If we may judge from the analogy of our domesticated +animals, many modifications of structure in man probably +come under this principle of correlated growth.</p> + +<p>We have now seen that the characteristic differences +between the races of man cannot be accounted for in a +satisfactory manner by the direct action of the conditions +of life, nor by the effects of the continued use of +parts, nor through the principle of correlation. We +are therefore led to inquire whether slight individual +differences, to which man is eminently liable, may +not have been preserved and augmented during a long +series of generations through natural selection. But +here we are at once met by the objection that beneficial +variations alone can be thus preserved; and as far as +we are enabled to judge (although always liable to +error on this head) not one of the external differences +between the races of man are of any direct or +<span class='pagenum'><a name="Page_249" id="Page_249">249</a></span>special service to him. The intellectual and moral or +social faculties must of course be excepted from this remark; +but differences in these faculties can have had +little or no influence on external characters. The variability +of all the characteristic differences between the +races, before referred to, likewise indicates that these +differences cannot be of much importance; for, had +they been important, they would long ago have been +either fixed and preserved, or eliminated. In this +respect man resembles those forms, called by naturalists +protean or polymorphic, which have remained extremely +variable, owing, as it seems, to their variations being of +an indifferent nature, and consequently to their having +escaped the action of natural selection.</p> + +<p>We have thus far been baffled in all our attempts +to account for the differences between the races of man; +but there remains one important agency, namely Sexual +Selection, which appears to have acted as powerfully +on man, as on many other animals. I do not intend +to assert that sexual selection will account for all the +differences between the races. An unexplained residuum +is left, about which we can in our ignorance +only say, that as individuals are continually born with, +for instance, heads a little rounder or narrower, and +with noses a little longer or shorter, such slight differences +might become fixed and uniform, if the unknown +agencies which induced them were to act in a +more constant manner, aided by long-continued intercrossing. +Such modifications come under the provisional +class, alluded to in our fourth chapter, which for +the want of a better term have been called spontaneous +variations. Nor do I pretend that the effects of sexual +selection can be indicated with scientific precision; but +it can be shewn that it would be an inexplicable fact if +man had not been modified by this agency, which has<span class='pagenum'><a name="Page_250" id="Page_250">250</a></span> +acted so powerfully on innumerable animals, both high +and low in the scale. It can further be shewn that the +differences between the races of man, as in colour, hairyness, +form of features, &c., are of the nature which it +might have been expected would have been acted on by +sexual selection. But in order to treat this subject in a +fitting manner, I have found it necessary to pass the +whole animal kingdom in review; I have therefore devoted +to it the Second Part of this work. At the close +I shall return to man, and, after attempting to shew +how far he has been modified through sexual selection, +will give a brief summary of the chapters in this First +Part.</p> + +<hr /> +<p><span class='pagenum'><a name="Page_253" id="Page_253">253</a></span></p> + +<h2><span class="smcap">Part II.—SEXUAL SELECTION.</span></h2> + +<hr /> +<h3>CHAPTER VIII.</h3> + +<h4><span class="smcap">Principles of Sexual Selection.</span></h4> + +<div class="blockquot"><p>Secondary sexual characters—Sexual selection—Manner of action—Excess +of males—Polygamy—The male alone generally +modified through sexual selection—Eagerness of the male—Variability +of the male—Choice exerted by the female—Sexual +compared with natural selection—Inheritance, at corresponding +periods of life, at corresponding seasons of the year, and as limited +by sex—Relations between the several forms of inheritance—Causes +why one sex and the young are not modified through +sexual selection—Supplement on the proportional numbers of +the two sexes throughout the animal kingdom—On the limitation +of the numbers of the two sexes through natural selection.</p></div> + +<p>With animals which have their sexes separated, the +males necessarily differ from the females in their organs +of reproduction; and these afford the primary sexual +characters. But the sexes often differ in what Hunter +has called secondary sexual characters, which are not +directly connected with the act of reproduction; for +instance, in the male possessing certain organs of sense +or locomotion, of which the female is quite destitute, or +in having them more highly-developed, in order that +he may readily find or reach her; or again, in the male +having special organs of prehension so as to hold her +securely. These latter organs of infinitely diversified +kinds graduate into, and in some cases can hardly be +distinguished from, those which are commonly ranked +as primary, such as the complex appendages at the +apex of the abdomen in male insects. Unless indeed<span class='pagenum'><a name="Page_254" id="Page_254">254</a></span> +we confine the term “primary” to the reproductive +glands, it is scarcely possible to decide, as far as the +organs of prehension are concerned, which ought to +be called primary and which secondary.</p> + +<p>The female often differs from the male in having +organs for the nourishment or protection of her young, +as the mammary glands of mammals, and the abdominal +sacks of the marsupials. The male, also, in +some few cases differs from the female in possessing +analogous organs, as the receptacles for the ova possessed +by the males of certain fishes, and those temporarily +developed in certain male frogs. Female bees +have a special apparatus for collecting and carrying +pollen, and their ovipositor is modified into a sting for +the defence of their larvæ and the community. In the +females of many insects the ovipositor is modified in +the most complex manner for the safe placing of the +eggs. Numerous similar cases could be given, but they +do not here concern us. There are, however, other +sexual differences quite disconnected with the primary +organs with which we are more especially concerned—such +as the greater size, strength, and pugnacity of the +male, his weapons of offence or means of defence +against rivals, his gaudy colouring and various ornaments, +his power of song, and other such characters.</p> + +<p>Besides the foregoing primary and secondary sexual +differences, the male and female sometimes differ in +structures connected with different habits of life, and +not at all, or only indirectly, related to the reproductive +functions. Thus the females of certain flies (Culicidæ +and Tabanidæ) are blood-suckers, whilst the males live +on flowers and have their mouths destitute of mandibles.<a name="FNanchor_336" id="FNanchor_336"></a><a href="#Footnote_336" class="fnanchor">336</a> +The males alone of certain moths and of some +<span class='pagenum'><a name="Page_255" id="Page_255">255</a></span>crustaceans (<i>e.g.</i> Tanais) have imperfect, closed mouths, +and cannot feed. The Complemental males of certain +cirripedes live like epiphytic plants either on the female +or hermaphrodite form, and are destitute of a mouth +and prehensile limbs. In these cases it is the male +which has been modified and has lost certain important +organs, which the other members of the same +group possess. In other cases it is the female which +has lost such parts; for instance, the female glow-worm +is destitute of wings, as are many female moths, some +of which never leave their cocoons. Many female parasitic +crustaceans have lost their natatory legs. In +some weevil-beetles (Curculionidæ) there is a great +difference between the male and female in the length +of the rostrum or snout;<a name="FNanchor_337" id="FNanchor_337"></a><a href="#Footnote_337" class="fnanchor">337</a> but the meaning of this and +of many analogous differences, is not at all understood. +Differences of structure between the two sexes in relation +to different habits of life are generally confined to +the lower animals; but with some few birds the beak +of the male differs from that of the female. No doubt +in most, but apparently not in all these cases, the differences +are indirectly connected with the propagation +of the species: thus a female which has to nourish a +multitude of ova will require more food than the male, +and consequently will require special means for procuring +it. A male animal which lived for a very short +time might without detriment lose through disuse its +organs for procuring food; but he would retain his +locomotive organs in a perfect state, so that he might +reach the female. The female, on the other hand, +might safely lose her organs for flying, swimming, +<span class='pagenum'><a name="Page_256" id="Page_256">256</a></span>or walking, if she gradually acquired habits which rendered +such powers useless.</p> + +<p>We are, however, here concerned only with that kind +of selection, which I have called sexual selection. This +depends on the advantage which certain individuals have +over other individuals of the same sex and species, in +exclusive relation to reproduction. When the two sexes +differ in structure in relation to different habits of life, +as in the cases above mentioned, they have no doubt +been modified through natural selection, accompanied +by inheritance limited to one and the same sex. So +again the primary sexual organs, and those for nourishing +or protecting the young, come under this same head; +for those individuals which generated or nourished their +offspring best, would leave, <i>cæteris paribus</i>, the greatest +number to inherit their superiority; whilst those which +generated or nourished their offspring badly, would leave +but few to inherit their weaker powers. As the male +has to search for the female, he requires for this purpose +organs of sense and locomotion, but if these organs are +necessary for the other purposes of life, as is generally +the case, they will have been developed through natural +selection. When the male has found the female he +sometimes absolutely requires prehensile organs to hold +her; thus Dr. Wallace informs me that the males of certain +moths cannot unite with the females if their tarsi +or feet are broken. The males of many oceanic crustaceans +have their legs and antennæ modified in an extraordinary +manner for the prehension of the female; +hence we may suspect that owing to these animals +being washed about by the waves of the open sea, they +absolutely require these organs in order to propagate +their kind, and if so their development will have been +the result of ordinary or natural selection.</p> + +<p>When the two sexes follow exactly the same habits<span class='pagenum'><a name="Page_257" id="Page_257">257</a></span> +of life, and the male has more highly developed sense +or locomotive organs than the female, it may be that +these in their perfected state are indispensable to the +male for finding the female; but in the vast majority +of cases, they serve only to give one male an advantage +over another, for the less well-endowed males, +if time were allowed them, would succeed in pairing +with the females; and they would in all other +respects, judging from the structure of the female, be +equally well adapted for their ordinary habits of life. +In such cases sexual selection must have come into +action, for the males have acquired their present structure, +not from being better fitted to survive in the +struggle for existence, but from having gained an advantage +over other males, and from having transmitted +this advantage to their male offspring alone. It was the +importance of this distinction which led me to designate +this form of selection as sexual selection. So again, +if the chief service rendered to the male by his prehensile +organs is to prevent the escape of the female +before the arrival of other males, or when assaulted by +them, these organs will have been perfected through +sexual selection, that is by the advantage acquired by +certain males over their rivals. But in most cases it +is scarcely possible to distinguish between the effects +of natural and sexual selection. Whole chapters could +easily be filled with details on the differences between +the sexes in their sensory, locomotive, and prehensile +organs. As, however, these structures are not more +interesting than others adapted for the ordinary purposes +of life, I shall almost pass them over, giving only +a few instances under each class.</p> + +<p>There are many other structures and instincts which +must have been developed through sexual selection—such +as the weapons of offence and the means of defence<span class='pagenum'><a name="Page_258" id="Page_258">258</a></span> +possessed by the males for fighting with and driving +away their rivals—their courage and pugnacity—their +ornaments of many kinds—their organs for producing +vocal or instrumental music—and their glands for +emitting odours; most of these latter structures serving +only to allure or excite the female. That these +characters are the result of sexual and not of ordinary +selection is clear, as unarmed, unornamented, or unattractive +males would succeed equally well in the +battle for life and in leaving a numerous progeny, if +better endowed males were not present. We may infer +that this would be the case, for the females, which are +unarmed and unornamented, are able to survive and +procreate their kind. Secondary sexual characters of +the kind just referred to, will be fully discussed in the +following chapters, as they are in many respects interesting, +but more especially as they depend on the +will, choice, and rivalry of the individuals of either sex. +When we behold two males fighting for the possession +of the female, or several male birds displaying their +gorgeous plumage, and performing the strangest antics +before an assembled body of females, we cannot doubt +that, though led by instinct, they know what they are +about, and consciously exert their mental and bodily +powers.</p> + +<p>In the same manner as man can improve the breed +of his game-cocks by the selection of those birds which +are victorious in the cockpit, so it appears that the +strongest and most vigorous males, or those provided +with the best weapons, have prevailed under nature, +and have led to the improvement of the natural breed +or species. Through repeated deadly contests, a slight +degree of variability, if it led to some advantage, however +slight, would suffice for the work of sexual selection; +and it is certain that secondary sexual characters<span class='pagenum'><a name="Page_259" id="Page_259">259</a></span> +are eminently variable. In the same manner as man +can give beauty, according to his standard of taste, to his +male poultry—can give to the Sebright bantam a new +and elegant plumage, an erect and peculiar carriage—so +it appears that in a state of nature female birds, by +having long selected the more attractive males, have +added to their beauty. No doubt this implies powers +of discrimination and taste on the part of the female +which will at first appear extremely improbable; but +I hope hereafter to shew that this is not the case.</p> + +<p>From our ignorance on several points, the precise +manner in which sexual selection acts is to a certain +extent uncertain. Nevertheless if those naturalists who +already believe in the mutability of species, will read +the following chapters, they will, I think, agree with +me that sexual selection has played an important part +in the history of the organic world. It is certain that +with almost all animals there is a struggle between the +males for the possession of the female. This fact is so +notorious that it would be superfluous to give instances. +Hence the females, supposing that their mental capacity +sufficed for the exertion of a choice, could select one out +of several males. But in numerous cases it appears as +if it had been specially arranged that there should be +a struggle between many males. Thus with migratory +birds, the males generally arrive before the females at +their place of breeding, so that many males are ready +to contend for each female. The bird-catchers assert +that this is invariably the case with the nightingale +and blackcap, as I am informed by Mr. Jenner Weir, +who confirms the statement with respect to the latter +species.</p> + +<p>Mr. Swaysland of Brighton, who has been in the habit, +during the last forty years, of catching our migratory +birds on their first arrival, writes to me that he has<span class='pagenum'><a name="Page_260" id="Page_260">260</a></span> +never known the females of any species to arrive before +their males. During one spring he shot thirty-nine +males of Ray’s wagtail (<i>Budytes Raii</i>) before he saw a +single female. Mr. Gould has ascertained by dissection, +as he informs me, that male snipes arrive in this +country before the females. In the case of fish, at the +period when the salmon ascend our rivers, the males in +large numbers are ready to breed before the females. +So it apparently is with frogs and toads. Throughout +the great class of insects the males almost always +emerge from the pupal state before the other sex, so +that they generally swarm for a time before any females +can be seen.<a name="FNanchor_338" id="FNanchor_338"></a><a href="#Footnote_338" class="fnanchor">338</a> The cause of this difference between +the males and females in their periods of arrival and +maturity is sufficiently obvious. Those males which +annually first migrated into any country, or which in +the spring were first ready to breed, or were the most +eager, would leave the largest number of offspring; +and these would tend to inherit similar instincts and +constitutions. On the whole there can be no doubt that +with almost all animals, in which the sexes are separate, +there is a constantly recurrent struggle between the +males for the possession of the females.</p> + +<p>Our difficulty in regard to sexual selection lies in +understanding how it is that the males which conquer +other males, or those which prove the most attractive +to the females, leave a greater number of offspring +to inherit their superiority than the beaten and less +<span class='pagenum'><a name="Page_261" id="Page_261">261</a></span>attractive males. Unless this result followed, the characters +which gave to certain males an advantage over +others, could not be perfected and augmented through +sexual selection. When the sexes exist in exactly +equal numbers, the worst-endowed males will ultimately +find females (excepting where polygamy prevails), and +leave as many offspring, equally well fitted for their +general habits of life, as the best-endowed males. From +various facts and considerations, I formerly inferred +that with most animals, in which secondary sexual +characters were well developed, the males considerably +exceeded the females in number; and this does hold +good in some few cases. If the males were to the +females as two to one, or as three to two, or even in +a somewhat lower ratio, the whole affair would be +simple; for the better-armed or more attractive males +would leave the largest number of offspring. But after +investigating, as far as possible, the numerical proportions +of the sexes, I do not believe that any great +inequality in number commonly exists. In most cases +sexual selection appears to have been effective in the +following manner.</p> + +<p>Let us take any species, a bird for instance, and +divide the females inhabiting a district into two equal +bodies: the one consisting of the more vigorous and +better-nourished individuals, and the other of the less +vigorous and healthy. The former, there can be little +doubt, would be ready to breed in the spring before the +others; and this is the opinion of Mr. Jenner Weir, who +has during many years carefully attended to the habits +of birds. There can also be no doubt that the most +vigorous, healthy, and best-nourished females would on +an average succeed in rearing the largest number of +offspring. The males, as we have seen, are generally +ready to breed before the females; of the males the<span class='pagenum'><a name="Page_262" id="Page_262">262</a></span> +strongest, and with some species the best armed, drive +away the weaker males; and the former would then +unite with the more vigorous and best-nourished females, +as these are the first to breed. Such vigorous +pairs would surely rear a larger number of offspring +than the retarded females, which would be compelled, +supposing the sexes to be numerically equal, to unite +with the conquered and less powerful males; and this +is all that is wanted to add, in the course of successive +generations, to the size, strength and courage of the +males, or to improve their weapons.</p> + +<p>But in a multitude of cases the males which conquer +other males, do not obtain possession of the females, +independently of choice on the part of the latter. The +courtship of animals is by no means so simple and short +an affair as might be thought. The females are most +excited by, or prefer pairing with, the more ornamented +males, or those which are the best songsters, or play the +best antics; but it is obviously probable, as has been +actually observed in some cases, that they would at the +same time prefer the more vigorous and lively males.<a name="FNanchor_339" id="FNanchor_339"></a><a href="#Footnote_339" class="fnanchor">339</a> +Thus the more vigorous females, which are the first to +breed, will have the choice of many males; and though +they may not always select the strongest or best armed, +they will select those which are vigorous and well armed, +and in other respects the most attractive. Such early +pairs would have the same advantage in rearing offspring +on the female side as above explained, and nearly +the same advantage on the male side. And this apparently +has sufficed during a long course of generations +to add not only to the strength and fighting-powers of +<span class='pagenum'><a name="Page_263" id="Page_263">263</a></span>the males, but likewise to their various ornaments or +other attractions.</p> + +<p>In the converse and much rarer case of the males +selecting particular females, it is plain that those which +were the most vigorous and had conquered others, would +have the freest choice; and it is almost certain that they +would select vigorous as well as attractive females. Such +pairs would have an advantage in rearing offspring, more +especially if the male had the power to defend the +female during the pairing-season, as occurs with some +of the higher animals, or aided in providing for the +young. The same principles would apply if both sexes +mutually preferred and selected certain individuals of +the opposite sex; supposing that they selected not only +the more attractive, but likewise the more vigorous +individuals.</p> + +<p class="tb"><i>Numerical Proportion of the Two Sexes.</i>—I have +remarked that sexual selection would be a simple +affair if the males considerably exceeded in number +the females. Hence I was led to investigate, as far +as I could, the proportions between the two sexes of +as many animals as possible; but the materials are +scanty. I will here give only a brief abstract of the +results, retaining the details for a supplementary discussion, +so as not to interfere with the course of my +argument. Domesticated animals alone afford the +opportunity of ascertaining the proportional numbers +at birth; but no records have been specially kept for +this purpose. By indirect means, however, I have +collected a considerable body of statistical data, from +which it appears that with most of our domestic +animals the sexes are nearly equal at birth. Thus with +race-horses, 25,560 births have been recorded during +twenty-one years, and the male births have been to the +female births as 99·7 to 100. With greyhounds the<span class='pagenum'><a name="Page_264" id="Page_264">264</a></span> +inequality is greater than with any other animal, for +during twelve years, out of 6878 births, the male births +have been as 110·1 to 100 female births. It is, however, +in some degree doubtful whether it is safe to infer that +the same proportional numbers would hold good under +natural conditions as under domestication; for slight +and unknown differences in the conditions affect to a +certain extent the proportion of the sexes. Thus with +mankind, the male births in England are as 104·5, +in Russia as 108·9, and with the Jews of Livornia as +120 <i>to</i> 100 females. The proportion is also mysteriously +affected by the circumstance of the births being legitimate +or illegitimate.</p> + +<p>For our present purpose we are concerned with the +proportion of the sexes, not at birth, but at maturity, +and this adds another element of doubt; for it is a well +ascertained fact that with man a considerably larger +proportion of males than of females die before or during +birth, and during the first few years of infancy. So it +almost certainly is with male lambs, and so it may be +with the males of other animals. The males of some +animals kill each other by fighting; or they drive +each other about until they become greatly emaciated. +They must, also, whilst wandering about in eager search +for the females, be often exposed to various dangers. +With many kinds of fish the males are much smaller +than the females, and they are believed often to be +devoured by the latter or by other fishes. With some +birds the females appear to die in larger proportion +than the males: they are also liable to be destroyed on +their nests, or whilst in charge of their young. With +insects the female larvæ are often larger than those of +the males, and would consequently be more likely to be +devoured: in some cases the mature females are less +active and less rapid in their movements than the males,<span class='pagenum'><a name="Page_265" id="Page_265">265</a></span> +and would not be so well able to escape from danger. +Hence, with animals in a state of nature, in order to +judge of the proportions of the sexes at maturity, we +must rely on mere estimation; and this, except perhaps +when the inequality is strongly marked, is but little +trustworthy. Nevertheless, as far as a judgment can +be formed, we may conclude from the facts given in the +supplement, that the males of some few mammals, of +many birds, of some fish and insects, considerably +exceed in number the females.</p> + +<p>The proportion between the sexes fluctuates slightly +during successive years: thus with race-horses, for every +100 females born, the males varied from 1O7.1 in one +year to 92.6 in another year, and with greyhounds from +116.3 to 95.3. But had larger numbers been tabulated +throughout a more extensive area than England, these +fluctuations would probably have disappeared; and such +as they are, they would hardly suffice to lead under +a state of nature to the effective action of sexual selection. +Nevertheless with some few wild animals, the +proportions seem, as shewn in the supplement, to fluctuate +either during different seasons or in different +localities in a sufficient degree to lead to such action. +For it should be observed that any advantage gained +during certain years or in certain localities by those +males which were able to conquer other males, or were +the most attractive to the females, would probably be +transmitted to the offspring and would not subsequently +be eliminated. During the succeeding seasons, when +from the equality of the sexes every male was everywhere +able to procure a female, the stronger or more +attractive males previously produced would still have +at least as good a chance of leaving offspring as the +less strong or less attractive.</p> + +<p class="tb"><i>Polygamy.</i>—The practice of polygamy leads to the<span class='pagenum'><a name="Page_266" id="Page_266">266</a></span> +same results as would follow from an actual inequality +in the number of the sexes; for if each male secures +two or more females, many males will not be able to +pair; and the latter assuredly will be the weaker or +less attractive individuals. Many mammals and some +few birds are polygamous, but with animals belonging to +the lower classes I have found no evidence of this habit. +The intellectual powers of such animals are, perhaps, +not sufficient to lead them to collect and guard a harem +of females. That some relation exists between polygamy +and the development of secondary sexual characters, +appears nearly certain; and this supports the +view that a numerical preponderance of males would +be eminently favourable to the action of sexual selection. +Nevertheless many animals, especially birds, which are +strictly monogamous, display strongly-marked secondary +sexual characters; whilst some few animals, which are +polygamous, are not thus characterised.</p> + +<p>We will first briefly run through the class of mammals, +and then turn to birds. The gorilla seems to be +a polygamist, and the male differs considerably from +the female; so it is with some baboons which live in +herds containing twice as many adult females as males. +In South America the <i>Mycetes caraya</i> presents well-marked +sexual differences in colour, beard, and vocal +organs, and the male generally lives with two or three +wives: the male of the <i>Cebus capucinus</i> differs somewhat +from the female, and appears to be polygamous.<a name="FNanchor_340" id="FNanchor_340"></a><a href="#Footnote_340" class="fnanchor">340</a> +Little is known on this head with respect to most other +monkeys, but some species are strictly monogamous. +The ruminants are eminently polygamous, and they +<span class='pagenum'><a name="Page_267" id="Page_267">267</a></span>more frequently present sexual differences than almost +any other group of mammals, especially in their weapons, +but likewise in other characters. Most deer, cattle, and +sheep are polygamous; as are most antelopes, though +some of the latter are monogamous. Sir Andrew +Smith, in speaking of the antelopes of South Africa, +says that in herds of about a dozen there was rarely +more than one mature male. The Asiatic <i>Antilope +saiga</i> appears to be the most inordinate polygamist +in the world; for Pallas<a name="FNanchor_341" id="FNanchor_341"></a><a href="#Footnote_341" class="fnanchor">341</a> states that the male drives +away all rivals, and collects a herd of about a hundred, +consisting of females and kids: the female is hornless +and has softer hair, but does not otherwise differ much +from the male. The horse is polygamous, but, except +in his greater size and in the proportions of his body, +differs but little from the mare. The wild boar, in his +great tusks and some other characters, presents well-marked +sexual characters; in Europe and in India he +leads a solitary life, except during the breeding-season; +but at this season he consorts in India with several +females, as Sir W. Elliot, who has had large experience +in observing this animal, believes: whether this holds +good in Europe is doubtful, but is supported by some +statements. The adult male Indian elephant, like the +boar, passes much of his time in solitude; but when +associating with others, “it is rare to find,” as Dr. +Campbell states, “more than one male with a whole +herd of females.” The larger males expel or kill the +smaller and weaker ones. The male differs from the +female by his immense tusks and greater size, strength, +and endurance; so great is the difference in these latter +<span class='pagenum'><a name="Page_268" id="Page_268">268</a></span>respects, that the males when caught are valued at +twenty per cent. above the females.<a name="FNanchor_342" id="FNanchor_342"></a><a href="#Footnote_342" class="fnanchor">342</a> With other pachydermatous +animals the sexes differ very little or not at +all, and they are not, as far as known, polygamists. +Hardly a single species amongst the Cheiroptera and +Edentata, or in the great Orders of the Rodents and +Insectivora, presents well-developed secondary sexual +differences; and I can find no account of any species +being polygamous, excepting, perhaps, the common rat, +the males of which, as some rat-catchers affirm, live +with several females.</p> + +<p>The lion in South Africa, as I hear from Sir Andrew +Smith, sometimes lives with a single female, but generally +with more than one, and, in one case, was found +with as many as five females, so that he is polygamous. +He is, as far as I can discover, the sole polygamist in +the whole group of the terrestrial Carnivora, and he +alone presents well-marked sexual characters. If, however, +we turn to the marine Carnivora, the case is widely +different; for many species of seals offer, as we shall +hereafter see, extraordinary sexual differences, and they +are eminently polygamous. Thus the male sea-elephant +of the Southern Ocean, always possesses, according +to Péron, several females, and the sea-lion of Forster +is said to be surrounded by from twenty to thirty females. +In the North, the male sea-bear of Steller is accompanied +by even a greater number of females.</p> + +<p>With respect to birds, many species, the sexes of +which differ greatly from each other, are certainly +monogamous. In Great Britain we see well-marked +sexual differences in, for instance, the wild-duck which +pairs with a single female, with the common blackbird, +<span class='pagenum'><a name="Page_269" id="Page_269">269</a></span>and with the bullfinch which is said to pair for life. So +it is, as I am informed by Mr. Wallace, with the Chatterers +or Cotingidæ of South America, and numerous +other birds. In several groups I have not been able to +discover whether the species are polygamous or monogamous. +Lesson says that birds of paradise, so remarkable +for their sexual differences, are polygamous, +but Mr. Wallace doubts whether he had sufficient evidence. +Mr. Salvin informs me that he has been led +to believe that humming-birds are polygamous. The +male widow-bird; remarkable for his caudal plumes, +certainly seems to be a polygamist.<a name="FNanchor_343" id="FNanchor_343"></a><a href="#Footnote_343" class="fnanchor">343</a> I have been +assured by Mr. Jenner Weir and by others, that three +starlings not rarely frequent the same nest; but whether +this is a case of polygamy or polyandry has not been +ascertained.</p> + +<p>The Gallinaceæ present almost as strongly marked +sexual differences as birds of paradise or humming-birds, +and many of the species are, as is well known, +polygamous; others being strictly monogamous. What +a contrast is presented between the sexes by the polygamous +peacock or pheasant, and the monogamous +guinea-fowl or partridge! Many similar cases could +be given, as in the grouse tribe, in which the males +of the polygamous capercailzie and black-cock differ +greatly from the females; whilst the sexes of the monogamous +red grouse and ptarmigan differ very little. +Amongst the Cursores, no great number of species +offer strongly-marked sexual differences, except the +bustards, and the great bustard (<i>Otis tarda</i>), is said to +<span class='pagenum'><a name="Page_270" id="Page_270">270</a></span>be polygamous. With the Grallatores, extremely few +species differ sexually, but the ruff (<i>Machetes pugnax</i>) +affords a strong exception, and this species is believed +by Montagu to be a polygamist. Hence it appears +that with birds there often exists a close relation +between polygamy and the development of strongly-marked +sexual differences. On asking Mr. Bartlett, at +the Zoological Gardens, who has had such large experience +with birds, whether the male tragopan (one of +the Gallinaceæ) was polygamous, I was struck by his +answering, “I do not know, but should think so from +his splendid colours.”</p> + +<p>It deserves notice that the instinct of pairing with a +single female is easily lost under domestication. The +wild-duck is strictly monogamous, the domestic-duck +highly polygamous. The Rev. W. D. Fox informs me +that with some half-tamed wild-ducks, kept on a large +pond in his neighbourhood, so many mallards were shot +by the gamekeeper that only one was left for every +seven or eight females; yet unusually large broods +were reared. The guinea-fowl is strictly monogamous; +but Mr. Fox finds that his birds succeed best when he +keeps one cock to two or three hens.<a name="FNanchor_344" id="FNanchor_344"></a><a href="#Footnote_344" class="fnanchor">344</a> Canary-birds +pair in a state of nature, but the breeders in England +successfully put one male to four or five females; nevertheless +the first female, as Mr. Fox has been assured, +is alone treated as the wife, she and her young ones +being fed by him; the others are treated as concubines. +I have noticed these cases, as it renders it in some +degree probable that monogamous species, in a state of +nature, might readily become either temporarily or permanently +polygamous.</p> +<p><span class='pagenum'><a name="Page_271" id="Page_271">271</a></span></p> +<p>With respect to reptiles and fishes, too little is known +of their habits to enable us to speak of their marriage +arrangements. The stickle-back Gasterosteus, however, +is said to be a polygamist;<a name="FNanchor_345" id="FNanchor_345"></a><a href="#Footnote_345" class="fnanchor">345</a> and the male during the +breeding-season differs conspicuously from the female.</p> + +<p class="tb">To sum up on the means through which, as far as +we can judge, sexual selection has led to the development +of secondary sexual characters. It has been shewn +that the largest number of vigorous offspring will be +reared from the pairing of the strongest and best-armed +males, which have conquered other males, with the +most vigorous and best-nourished females, which are +the first to breed in the spring. Such females, if +they select the more attractive, and at the same time +vigorous, males, will rear a larger number of offspring +than the retarded females, which must pair with the +less vigorous and less attractive males. So it will be +if the more vigorous males select the more attractive +and at the same time healthy and vigorous females; +and this will especially hold good if the male defends +the female, and aids in providing food for the young. +The advantage thus gained by the more vigorous pairs +in rearing a larger number of offspring has apparently +sufficed to render sexual selection efficient. But a large +preponderance in number of the males over the females +would be still more efficient; whether the preponderance +was only occasional and local, or permanent; +whether it occurred at birth, or subsequently from the +greater destruction of the females; or whether it indirectly +followed from the practice of polygamy.</p> + +<p class="tb"><i>The Male generally more modified than the Female.</i>—Throughout +the animal kingdom, when the sexes differ +<span class='pagenum'><a name="Page_272" id="Page_272">272</a></span>from each other in external appearance, it is the male +which, with rare exceptions, has been chiefly modified; +for the female still remains more like the young of her +own species, and more like the other members of the +same group. The cause of this seems to lie in the +males of almost all animals having stronger passions +than the females. Hence it is the males that fight +together and sedulously display their charms before +the females; and those which are victorious transmit +their superiority to their male offspring. Why the +males do not transmit their characters to both sexes +will hereafter be considered. That the males of all +mammals eagerly pursue the females is notorious to +every one. So it is with birds; but many male birds +do not so much pursue the female, as display their +plumage, perform strange antics, and pour forth their +song, in her presence. With the few fish which have +been observed, the male seems much more eager than +the female; and so it is with alligators, and apparently +with Batrachians. Throughout the enormous class of +insects, as Kirby remarks,<a name="FNanchor_346" id="FNanchor_346"></a><a href="#Footnote_346" class="fnanchor">346</a> “the law is, that the male +shall seek the female.” With spiders and crustaceans, +as I hear from two great authorities, Mr. Blackwall and +Mr. C. Spence Bate, the males are more active and more +erratic in their habits than the females. With insects and +crustaceans, when the organs of sense or locomotion are +present in the one sex and absent in the other, or when, +as is frequently the case, they are more highly developed +in the one than the other, it is almost invariably the male, +as far as I can discover, which retains such organs, or has +them most developed; and this shews that the male is +the more active member in the courtship of the sexes.<a name="FNanchor_347" id="FNanchor_347"></a><a href="#Footnote_347" class="fnanchor">347</a></p> + +<p><span class='pagenum'><a name="Page_273" id="Page_273">273</a></span></p><p>The female, on the other hand, with the rarest exception, +is less eager than the male. As the illustrious +Hunter<a name="FNanchor_348" id="FNanchor_348"></a><a href="#Footnote_348" class="fnanchor">348</a> long ago observed, she generally “requires to +be courted;” she is coy, and may often be seen endeavouring +for a long time to escape from the male. +Every one who has attended to the habits of animals +will be able to call to mind instances of this kind. +Judging from various facts, hereafter to be given, and +from the results which may fairly be attributed to +sexual selection, the female, though comparatively +passive, generally exerts some choice and accepts one +male in preference to others. Or she may accept, as +appearances would sometimes lead us to believe, not +the male which is the most attractive to her, but the +one which is the least distasteful. The exertion of +some choice on the part of the female seems almost as +general a law as the eagerness of the male.</p> + +<p>We are naturally led to enquire why the male in so +many and such widely distinct classes has been rendered +more eager than the female, so that he searches +for her and plays the more active part in courtship. +It would be no advantage and some loss of power if +both sexes were mutually to search for each other; but +why should the male almost always be the seeker? +With plants, the ovules after fertilisation have to be +nourished for a time; hence the pollen is necessarily +brought to the female organs—being placed on the +stigma, through the agency of insects or of the wind, +<span class='pagenum'><a name="Page_274" id="Page_274">274</a></span>or by the spontaneous movements of the stamens; and +with the Algæ, &c., by the locomotive power of the +antherozooids. With lowly-organised animals permanently +affixed to the same spot and having their sexes +separate, the male element is invariably brought to +the female; and we can see the reason; for the ova, +even if detached before being fertilised and not requiring +subsequent nourishment or protection, would +be, from their larger relative size, less easily transported +than the male element. Hence plants<a name="FNanchor_349" id="FNanchor_349"></a><a href="#Footnote_349" class="fnanchor">349</a> and many of +the lower animals are, in this respect, analogous. In +the case of animals not affixed to the same spot, but +enclosed within a shell with no power of protruding any +part of their bodies, and in the case of animals having +little power of locomotion, the males must trust the +fertilising element to the risk of at least a short transit +through the waters of the sea. It would, therefore, be +a great advantage to such animals, as their organisation +became perfected, if the males when ready to emit the +fertilising element, were to acquire the habit of approaching +the female as closely as possible. The males +of various lowly-organised animals having thus aboriginally +acquired the habit of approaching and seeking +the females, the same habit would naturally be transmitted +to their more highly developed male descendants; +and in order that they should become efficient +seekers, they would have to be endowed with strong +passions. The acquirement of such passions would +naturally follow from the more eager males leaving a +larger number of offspring than the less eager.</p> + +<p>The great eagerness of the male has thus indirectly +<span class='pagenum'><a name="Page_275" id="Page_275">275</a></span>led to the much more frequent development of secondary +sexual characters in the male than in the female. +But the development of such characters will have been +much aided, if the conclusion at which I arrived after +studying domesticated animals, can be trusted, namely, +that the male is more liable to vary than the female. +I am aware how difficult it is to verify a conclusion of +this kind. Some slight evidence, however, can be gained +by comparing the two sexes in mankind, as man has +been more carefully observed than any other animal. +During the Novara Expedition<a name="FNanchor_350" id="FNanchor_350"></a><a href="#Footnote_350" class="fnanchor">350</a> a vast number of measurements +of various parts of the body in different races +were made, and the men were found in almost every +case to present a greater range of variation than the +women; but I shall have to recur to this subject in +a future chapter. Mr. J. Wood,<a name="FNanchor_351" id="FNanchor_351"></a><a href="#Footnote_351" class="fnanchor">351</a> who has carefully +attended to the variation of the muscles in man, puts +in italics the conclusion that “the greatest number of +abnormalities in each subject is found in the males.” +He had previously remarked that “altogether in 102 +subjects the varieties of redundancy were found to +be half as many again as in females, contrasting +widely with the greater frequency of deficiency in +females before described.” Professor Macalister like +wise remarks<a name="FNanchor_352" id="FNanchor_352"></a><a href="#Footnote_352" class="fnanchor">352</a> that variations in the muscles “are +probably more common in males than females.” +Certain muscles which are not normally present in mankind +are also more frequently developed in the male +than in the female sex, although exceptions to this rule +<span class='pagenum'><a name="Page_276" id="Page_276">276</a></span>are said to occur. Dr. Burt Wilder<a name="FNanchor_353" id="FNanchor_353"></a><a href="#Footnote_353" class="fnanchor">353</a> has tabulated +the cases of 152 individuals with supernumerary digits, +of which 86 were males, and 39, or less than half, +females; the remaining 27 being of unknown sex. It +should not, however, be overlooked that women would +more frequently endeavour to conceal a deformity of +this kind than men. Whether the large proportional +number of deaths of the male offspring of man and +apparently of sheep, compared with the female offspring, +before, during, and shortly after birth (see supplement), +has any relation to a stronger tendency in the organs +of the male to vary and thus to become abnormal in +structure or function, I will not pretend to conjecture.</p> + +<p>In various classes of animals a few exceptional cases +occur, in which the female instead of the male has +acquired well pronounced secondary sexual characters, +such as brighter colours, greater size, strength, or pugnacity. +With birds, as we shall hereafter see, there +has sometimes been a complete transposition of the +ordinary characters proper to each sex; the females +having become the more eager in courtship, the males +remaining comparatively passive, but apparently selecting, +as we may infer from the results, the more attractive +females. Certain female birds have thus been rendered +more highly coloured or otherwise ornamented, as well +as more powerful and pugnacious than the males, these +characters being transmitted to the female offspring +alone.</p> + +<p>It may be suggested that in some cases a double +process of selection has been carried on; the males +having selected the more attractive females, and the +latter the more attractive males. This process however, +though it might lead to the modification of both sexes, +<span class='pagenum'><a name="Page_277" id="Page_277">277</a></span>would not make the one sex different from the other, +unless indeed their taste for the beautiful differed; but +this is a supposition too improbable in the case of any +animal, excepting man, to be worth considering. There +are, however, many animals, in which the sexes resemble +each other, both being furnished with the same ornaments, +which analogy would lead us to attribute to the +agency of sexual selection. In such cases it may be +suggested with more plausibility, that there has been a +double or mutual process of sexual selection; the more +vigorous and precocious females having selected the +more attractive and vigorous males, the latter having +rejected all except the more attractive females. But +from what we know of the habits of animals, this view is +hardly probable, the male being generally eager to +pair with any female. It is more probable that the +ornaments common to both sexes were acquired by one +sex, generally the male, and then transmitted to the offspring +of both sexes. If, indeed, during a lengthened +period the males of any species were greatly to exceed +the females in number, and then during another +lengthened period under different conditions the reverse +were to occur, a double, but not simultaneous, process +of sexual selection might easily be carried on, by which +the two sexes might be rendered widely different.</p> + +<p>We shall hereafter see that many animals exist, of +which neither sex is brilliantly coloured or provided +with special ornaments, and yet the members of both +sexes or of one alone have probably been modified +through sexual selection. The absence of bright tints +or other ornaments may be the result of variations of +the right kind never having occurred, or of the animals +themselves preferring simple colours, such as plain black +or white. Obscure colours have often been acquired +through natural selection for the sake of protection, and<span class='pagenum'><a name="Page_278" id="Page_278">278</a></span> +the acquirement through sexual selection of conspicuous +colours, may have been checked from the danger thus +incurred. But in other cases the males have probably +struggled together during long ages, through brute +force, or by the display of their charms, or by both +means combined, and yet no effect will have been produced +unless a larger number of offspring were left by +the more successful males to inherit their superiority, +than by the less successful males; and this, as previously +shewn, depends on various complex contingencies.</p> + +<p>Sexual selection acts in a less rigorous manner than +natural selection. The latter produces its effects by the +life or death at all ages of the more or less successful +individuals. Death, indeed, not rarely ensues from the +conflicts of rival males. But generally the less successful +male merely fails to obtain a female, or obtains later +in the season a retarded and less vigorous female, or, +if polygamous, obtains fewer females; so that they +leave fewer, or less vigorous, or no offspring. In +regard to structures acquired through ordinary or +natural selection, there is in most cases, as long as the +conditions of life remain the same, a limit to the amount +of advantageous modification in relation to certain special +ends; but in regard to structures adapted to make one +male victorious over another, either in fighting or in +charming the female, there is no definite limit to the +amount of advantageous modification; so that as long as +the proper variations arise the work of sexual selection +will go on. This circumstance may partly account for +the frequent and extraordinary amount of variability +presented by secondary sexual characters. Nevertheless, +natural selection will determine that characters of this +kind shall not be acquired by the victorious males, +which would be injurious to them in any high degree, +either by expending too much of their vital powers, or<span class='pagenum'><a name="Page_279" id="Page_279">279</a></span> +by exposing them to any great danger. The development, +however, of certain structures—of the horns, for +instance, in certain stags—has been carried to a +wonderful extreme; and in some instances to an +extreme which, as far as the general conditions of life +are concerned, must be slightly injurious to the male. +From this fact we learn that the advantages which +favoured males have derived from conquering other +males in battle or courtship, and thus leaving a +numerous progeny, have been in the long run greater +than those derived from rather more perfect adaptation +to the external conditions of life. We shall further see, +and this could never have been anticipated, that the +power to charm the female has been in some few instances +more important than the power to conquer other +males in battle.</p> + +<h4>LAWS OF INHERITANCE.</h4> + +<p class="tb">In order to understand how sexual selection has +acted, and in the course of ages has produced conspicuous +results with many animals of many classes, it is necessary +to bear in mind the laws of inheritance, as far as +they are known. Two distinct elements are included +under the term “inheritance,” namely the transmission +and the development of characters; but as these +generally go together, the distinction is often overlooked. +We see this distinction in those characters +which are transmitted through the early years of life, +but are developed only at maturity or during old +age. We see the same distinction more clearly with +secondary sexual characters, for these are transmitted +through both sexes, though developed in one alone. +That they are present in both sexes, is manifest when +two species, having strongly-marked sexual characters, +are crossed, for each transmits the characters proper to<span class='pagenum'><a name="Page_280" id="Page_280">280</a></span> +its own male and female sex to the hybrid offspring of +both sexes. The same fact is likewise manifest, when +characters proper to the male are occasionally developed +in the female when she grows old or becomes +diseased; and so conversely with the male. Again, +characters occasionally appear, as if transferred from +the male to the female, as when, in certain breeds of the +fowl, spurs regularly appear in the young and healthy +females; but in truth they are simply developed in the +female; for in every breed each detail in the structure +of the spur is transmitted through the female to her +male offspring. In all cases of reversion, characters +are transmitted through two, three, or many generations, +and are then under certain unknown favourable conditions +developed. This important distinction between +transmission and development will be easiest kept in +mind by the aid of the hypothesis of pangenesis, whether +or not it be accepted as true. According to this hypothesis, +every unit or cell of the body throws off gemmules +or undeveloped atoms, which are transmitted to the +offspring of both sexes, and are multiplied by self-division. +They may remain undeveloped during the +early years of life or during successive generations; +their development into units or cells, like those from +which they were derived, depending on their affinity +for, and union with, other units or cells previously +developed in the due order of growth.</p> + +<p class="tb"><i>Inheritance at Corresponding Periods of Life.</i>—This +tendency is well established. If a new character appears +in an animal whilst young, whether it endures throughout +life or lasts only for a time, it will reappear, as a +general rule, at the same age and in the same manner +in the offspring. If, on the other hand, a new character +appears at maturity, or even during old age, it tends<span class='pagenum'><a name="Page_281" id="Page_281">281</a></span> +to reappear in the offspring at the same advanced age. +When deviations from this rule occur, the transmitted +characters much oftener appear before than after the +corresponding age. As I have discussed this subject +at sufficient length in another work,<a name="FNanchor_354" id="FNanchor_354"></a><a href="#Footnote_354" class="fnanchor">354</a> I will here merely +give two or three instances, for the sake of recalling the +subject to the reader’s mind. In several breeds of the +Fowl, the chickens whilst covered with down, in their +first true plumage, and in their adult plumage, differ +greatly from each other, as well as from their common +parent-form, the <i>Gallus bankiva</i>; and these characters +are faithfully transmitted by each breed to their +offspring at the corresponding period of life. For +instance, the chickens of spangled Hamburghs, whilst +covered with down, have a few dark spots on the head +and rump, but are not longitudinally striped, as in +many other breeds; in their first true plumage, “they +are beautifully pencilled,” that is each feather is +transversely marked by numerous dark bars; but in +their second plumage the feathers all become spangled +or tipped with a dark round spot.<a name="FNanchor_355" id="FNanchor_355"></a><a href="#Footnote_355" class="fnanchor">355</a> Hence in this +breed variations have occurred and have been transmitted +at three distinct periods of life. The Pigeon offers +a more remarkable case, because the aboriginal parent-species +does not undergo with advancing age any change +of plumage, excepting that at maturity the breast +becomes more iridescent; yet there are breeds which +do not acquire their characteristic colours until they +<span class='pagenum'><a name="Page_282" id="Page_282">282</a></span>have moulted two, three, or four times; and these +modifications of plumage are regularly transmitted.</p> + +<p class="tb"><i>Inheritance at Corresponding Seasons of the Year.</i>—With +animals in a state of nature innumerable +instances occur of characters periodically appearing at +different seasons. We see this with the horns of the +stag, and with the fur of arctic animals which becomes +thick and white during the winter. Numerous birds +acquire bright colours and other decorations during the +breeding-season alone. I can throw but little light on +this form of inheritance from facts observed under +domestication. Pallas states,<a name="FNanchor_356" id="FNanchor_356"></a><a href="#Footnote_356" class="fnanchor">356</a> that in Siberia domestic +cattle and horses periodically become lighter-coloured +during the winter; and I have observed a similar +marked change of colour in certain ponies in England. +Although I do not know that this tendency to assume a +differently coloured coat during different seasons of the +year is transmitted, yet it probably is so, as all shades of +colour are strongly inherited by the horse. Nor is this +form of inheritance, as limited by season, more remarkable +than inheritance as limited by age or sex.</p> + +<p><i>Inheritance as Limited by Sex.</i>—The equal transmission +of characters to both, sexes is the commonest +form of inheritance, at least with those animals which +do not present strongly-marked sexual differences, and +indeed with many of these. But characters are not +rarely transferred exclusively to that sex, in which they +first appeared. Ample evidence on this head has been +advanced in my work on Variation under Domestica<span class='pagenum'><a name="Page_283" id="Page_283">283</a></span>tion; +but a few instances may here be given. There +are breeds of the sheep and goat, in which the horns +of the male differ greatly in shape from those of the +female; and these differences, acquired under domestication, +are regularly transmitted to the same sex. +With tortoise-shell cats the females alone, as a general +rule, are thus coloured, the males being rusty-red. +With most breeds of the fowl, the characters proper +to each sex are transmitted to the same sex alone. So +general is this form of transmission that it is an anomaly +when we see in certain breeds variations transmitted +equally to both sexes. There are also certain +sub-breeds of the fowl in which the males can hardly +be distinguished from each other, whilst the females +differ considerably in colour. With the pigeon the +sexes of the parent-species do not differ in any external +character; nevertheless in certain domesticated breeds +the male is differently coloured from the female.<a name="FNanchor_357" id="FNanchor_357"></a><a href="#Footnote_357" class="fnanchor">357</a> +The wattle in the English Carrier pigeon and the crop +in the Pouter are more highly developed in the male +than in the female; and although these characters have +been gained through long-continued selection by man, +the difference between the two sexes is wholly due to +the form of inheritance which has prevailed; for it +has arisen, not from, but rather in opposition to, the +wishes of the breeder.</p> + +<p>Most of our domestic races have been formed by the +accumulation of many slight variations; and as some +of the successive steps have been transmitted to one +sex alone, and some to both sexes, we find in the different +breeds of the same species all gradations between +great sexual dissimilarity and complete similarity. In<span class='pagenum'><a name="Page_284" id="Page_284">284</a></span>stances +have already been given with the breeds of the +fowl and pigeon; and under nature analogous cases are +of frequent occurrence. With animals under domestication, +but whether under nature I will not venture to +say, one sex may lose characters proper to it, and may +thus come to resemble to a certain extent the opposite +sex; for instance, the males of some breeds of the fowl +have lost their masculine plumes and hackles. On the +other hand the differences between the sexes may be +increased under domestication, as with merino sheep, in +which the ewes have lost their horns. Again, characters +proper to one sex may suddenly appear in the other +sex; as with those sub-breeds of the fowl in which the +hens whilst young acquire spurs; or, as in certain +Polish sub-breeds, in which the females, as there is +reason to believe, originally acquired a crest, and subsequently +transferred it to the males. All these cases +are intelligible on the hypothesis of pangenesis; for +they depend on the gemmules of certain units of the +body, although present in both sexes, becoming through +the influence of domestication dormant in the one sex; +or if naturally dormant, becoming developed.</p> + +<p class="tb">There is one difficult question which it will be convenient +to defer to a future chapter; namely, whether +a character at first developed in both sexes, can be rendered +through selection limited in its development to +one sex alone. If, for instance, a breeder observed that +some of his pigeons (in which species characters are +usually transferred in an equal degree to both sexes) +varied into pale blue; could he by long-continued +selection make a breed, in which the males alone should +be of this tint, whilst the females remained unchanged? +I will here only say, that this, though perhaps not +impossible, would be extremely difficult; for the natural +result of breeding from the pale-blue males would be<span class='pagenum'><a name="Page_285" id="Page_285">285</a></span> +to change his whole stock, including both sexes, into +this tint. If, however, variations of the desired tint +appeared, which were from the first limited in their +development to the male sex, there would not be the +least difficulty in making a breed characterised by the +two sexes being of a different colour, as indeed has been +effected with a Belgian breed, in which the males alone +are streaked with black. In a similar manner, if any +variation appeared in a female pigeon, which was from +the first sexually limited in its development, it would +be easy to make a breed with the females alone thus +characterised; but if the variation was not thus originally +limited, the process would be extremely difficult, perhaps +impossible.</p> + +<p><i>On the Relation between the period of Development of a +Character and its transmission to one sex or to both sexes.</i>—Why +certain characters should be inherited by both +sexes, and other characters by one sex alone, namely by +that sex in which the character first appeared, is in most +cases quite unknown. We cannot even conjecture why +with certain sub-breeds of the pigeon, black striæ, though +transmitted through the female, should be developed in +the male alone, whilst every other character is equally +transferred to both sexes. Why, again, with cats, the +tortoise-shell colour should, with rare exceptions, be +developed in the female alone. The very same characters, +such as deficient or supernumerary digits, colour-blindness, +&c., may with mankind be inherited by the +males alone of one family, and in another family by the +females alone, though in both cases transmitted through +the opposite as well as the same sex.<a name="FNanchor_358" id="FNanchor_358"></a><a href="#Footnote_358" class="fnanchor">358</a> Although we +are thus ignorant, two rules often hold good, namely +<span class='pagenum'><a name="Page_286" id="Page_286">286</a></span>that variations which, first appear in either sex at a late +period of life, tend to be developed in the same sex +alone; whilst variations which first appear early in life +in either sex tend to be developed in both sexes. I am, +however, far from supposing that this is the sole determining +cause. As I have not elsewhere discussed +this subject, and as it has an important bearing on +sexual selection, I must here enter into lengthy and +somewhat intricate details.</p> + +<p>It is in itself probable that any character appearing +at an early age would tend to be inherited equally by +both sexes, for the sexes do not differ much in constitution, +before the power of reproduction is gained. On +the other hand, after this power has been gained and the +sexes have come to differ in constitution, the gemmules +(if I may again use the language of pangenesis) which +are cast off from each varying part in the one sex would +be much more likely to possess the proper affinities +for uniting with the tissues of the same sex, and thus +becoming developed, than with those of the opposite +sex.</p> + +<p>I was first led to infer that a relation of this kind +exists, from the fact that whenever and in whatever +manner the adult male has come to differ from the +adult female, he differs in the same manner from the +young of both sexes. The generality of this fact is quite +remarkable: it holds good with almost all mammals, +birds, amphibians, and fishes; also with many crustaceans, +spiders and some few insects, namely certain +orthoptera and libellulæ. In all these cases the variations, +through the accumulation of which the male acquired +his proper masculine characters, must have occurred +at a somewhat late period of life; otherwise the +young males would have been similarly characterised; +and conformably with our rule, they are transmitted to<span class='pagenum'><a name="Page_287" id="Page_287">287</a></span> +and developed in the adult males alone. When, on the +other hand, the adult male closely resembles the young +of both sexes (these, with rare exceptions, being alike), +he generally resembles the adult female; and in most of +these cases the variations through which the young and +old acquired their present characters, probably occurred +in conformity with our rule during youth. But there is +here room for doubt, as characters are sometimes transferred +to the offspring at an earlier age than that at +which they first appeared in the parents, so that the +parents may have varied when adult, and have transferred +their characters to their offspring whilst young. +There are, moreover, many animals, in which the two +sexes closely resemble each other, and yet both differ +from their young; and here the characters of the adults +must have been acquired late in life; nevertheless, +these characters in apparent contradiction to our rule, +are transferred to both sexes. We must not, however, +overlook the possibility or even probability of successive +variations of the same nature sometimes occurring, +under exposure to similar conditions, simultaneously in +both sexes at a rather late period of life; and in this case +the variations would be transferred to the offspring of +both sexes at a corresponding late age; and there would +be no real contradiction to our rule of the variations +which occur late in life being transferred exclusively to +the sex in which they first appeared. This latter rule +seems to hold true more generally than the second rule, +namely, that variations which occur in either sex early +in life tend to be transferred to both sexes. As it was +obviously impossible even to estimate in how large a +number of cases throughout the animal kingdom these +two propositions hold good, it occurred to me to investigate +some striking or crucial instances, and to rely +on the result.</p> + +<p><span class='pagenum'><a name="Page_288" id="Page_288">288</a></span>An excellent case for investigation is afforded by the +Deer Family. In all the species, excepting one, the +horns are developed in the male alone, though certainly +transmitted through the female, and capable of occasional +abnormal development in her. In the reindeer, on the +other hand, the female is provided with horns; so that +in this species, the horns ought, according to our rule, +to appear early in life, long before the two sexes had +arrived at maturity and had come to differ much in +constitution. In all the other species of deer the horns +ought to appear later in life, leading to their development +in that sex alone, in which they first appeared +in the progenitor of the whole Family. Now in seven +species, belonging to distinct sections of the family and +inhabiting different regions, in which the stags alone +bear horns, I find that the horns first appear at periods +varying from nine months after birth in the roebuck to +ten or twelve or even more months in the stags of the +six other larger species.<a name="FNanchor_359" id="FNanchor_359"></a><a href="#Footnote_359" class="fnanchor">359</a> But with the reindeer the +case is widely different, for as I hear from Prof. Nilsson, +who kindly made special enquiries for me in Lapland, +the horns appear in the young animals within four or +five weeks after birth, and at the same time in both +sexes. So that here we have a structure, developed at +a most unusually early age in one species of the family, +and common to both sexes in this one species.</p> + +<p>In several kinds of antelopes the males alone are +<span class='pagenum'><a name="Page_289" id="Page_289">289</a></span>provided with horns, whilst in the greater number both +sexes have horns. With respect to the period of development, +Mr. Blyth informs me that there lived +at one time in the Zoological Gardens a young koodoo +(<i>Ant. strepsiceros</i>), in which species the males alone +are horned, and the young of a closely-allied species, +viz. the eland (<i>Ant. oreas</i>), in which both sexes are +horned. Now in strict conformity with our rule, in the +young male koodoo, although arrived at the age of ten +months, the horns were remarkably small considering +the size ultimately attained by them: whilst in the +young male eland, although only three months old, the +horns were already very much larger than in the koodoo. +It is also worth notice that in the prong-horned antelope,<a name="FNanchor_360" id="FNanchor_360"></a><a href="#Footnote_360" class="fnanchor">360</a> +in which species the horns, though present in both +sexes, are almost rudimentary in the female, they do not +appear until about five or six months after birth. With +sheep, goats, and cattle, in which the horns are well +developed in both sexes, though not quite equal in size, +they can be felt, or even seen, at birth or soon afterwards.<a name="FNanchor_361" id="FNanchor_361"></a><a href="#Footnote_361" class="fnanchor">361</a> +Our rule, however, fails in regard to some +breeds of sheep, for instance merinos, in which the rams +alone are horned; for I cannot find on enquiry,<a name="FNanchor_362" id="FNanchor_362"></a><a href="#Footnote_362" class="fnanchor">362</a> that +<span class='pagenum'><a name="Page_290" id="Page_290">290</a></span>the horns are developed later in life in this breed than +in ordinary sheep in which both sexes are horned. But +with domesticated sheep the presence or absence of +horns is not a firmly fixed character; a certain proportion +of the merino ewes bearing small horns, and some of the +rams being hornless; whilst with ordinary sheep hornless +ewes are occasionally produced.</p> + +<p>In most of the species of the splendid family of the +Pheasants, the males differ conspicuously from the +females, and they acquire their ornaments at a rather +late period of life. The eared pheasant (<i>Crossoptilon +auritum</i>), however, offers a remarkable exception, for +both sexes possess the fine caudal plumes, the large ear-tufts +and the crimson velvet about the head; and I find +on enquiry in the Zoological Gardens that all these +characters, in accordance with our rule, appear very +early in life. The adult male can, however, be distinguished +from the adult female by one character, namely +by the presence of spurs; and conformably with our +rule, these do not begin to be developed, as I am assured +by Mr. Bartlett, before the age of six months, and even +at this age, can hardly be distinguished in the two +sexes.<a name="FNanchor_363" id="FNanchor_363"></a><a href="#Footnote_363" class="fnanchor">363</a> The male and female Peacock differ con<span class='pagenum'><a name="Page_291" id="Page_291">291</a></span>spicuously +from each other in almost every part of their +plumage, except in the elegant head-crest, which is +common to both sexes; and this is developed very early +in life, long before the other ornaments which are confined +to the male. The wild-duck offers an analogous +case, for the beautiful green speculum on the wings +is common to both sexes, though duller and somewhat +smaller in the female, and it is developed early in life, +whilst the curled tail-feathers and other ornaments +peculiar to the male are developed later.<a name="FNanchor_364" id="FNanchor_364"></a><a href="#Footnote_364" class="fnanchor">364</a> Between +such extreme cases of close sexual resemblance and +wide dissimilarity, as those of the Crossoptilon and +peacock, many intermediate ones could be given, in +which the characters follow in their order of development +our two rules.</p> + +<p>As most insects emerge from their pupal state in a +mature condition, it is doubtful whether the period of +development determines the transference of their characters +to one or both sexes. But we do not know that +the coloured scales, for instance, in two species of butterflies, +in one of which the sexes differ in colour, whilst +in the other they are alike, are developed at the same +relative age in the cocoon. Nor do we know whether +all the scales are simultaneously developed on the wings +<span class='pagenum'><a name="Page_292" id="Page_292">292</a></span>of the same species of butterfly, in which certain coloured +marks are confined to one sex, whilst other marks are +common to both sexes. A difference of this kind in the +period of development is not so improbable as it may +at first appear; for with the Orthoptera, which assume +their adult state, not by a single metamorphosis, but by +a succession of moults, the young males of some species +at first resemble the females, and acquire their distinctive +masculine characters only during a later moult. +Strictly analogous cases occur during the successive +moults of certain male crustaceans.</p> + +<p>We have as yet only considered the transference of +characters, relatively to their period of development, with +species in a natural state; we will now turn to domesticated +animals; first touching on monstrosities and +diseases. The presence of supernumerary digits, and +the absence of certain phalanges, must be determined +at an early embryonic period—the tendency to profuse +bleeding is at least congenital, as is probably colour-blindness—yet +these peculiarities, and other similar +ones, are often limited in their transmission to one sex; +so that the rule that characters which are developed +at an early period tend to be transmitted to both sexes, +here wholly fails. But this rule, as before remarked, +does not appear to be nearly so generally true as the +converse proposition, namely, that characters which +appear late in life in one sex are transmitted exclusively +to the same sex. From the fact of the above +abnormal peculiarities becoming attached to one sex. +long before the sexual functions are active, we may +infer that there must be a difference of some kind +between the sexes at an extremely early age. With +respect to sexually-limited diseases, we know too little +of the period at which they originate, to draw any +fair conclusion. Gout, however, seems to fall under<span class='pagenum'><a name="Page_293" id="Page_293">293</a></span> +our rule; for it is generally caused by intemperance +after early youth, and is transmitted from the father +to his sons in a much more marked manner than to his +daughters.</p> + +<p>In the various domestic breeds of sheep, goats, and +cattle, the males differ from their respective females +in the shape or development of their horns, forehead, +mane, dewlap, tail, and hump on the shoulders; and +these peculiarities, in accordance with our rule, are not +fully developed until rather late in life. With dogs, +the sexes do not differ, except that in certain breeds, +especially in the Scotch deer-hound, the male is much +larger and heavier than the female; and as we shall see +in a future chapter, the male goes on increasing in size +to an unusually late period of life, which will account, +according to our rule, for his increased size being transmitted +to his male offspring alone. On the other hand, +the tortoise-shell colour of the hair, which is confined +to female cats, is quite distinct at birth, and this case +violates our rule. There is a breed of pigeons in which +the males alone are streaked with black, and the streaks +can be detected even in the nestlings; but they become +more conspicuous at each successive moult, so that this +case partly opposes and partly supports the rule. With +the English Carrier and Pouter pigeon the full development +of the wattle and the crop occurs rather late in +life, and these characters, conformably with our rule, +are transmitted in full perfection to the males alone. +The following cases perhaps come within the class previously +alluded to, in which the two sexes have varied +in the same manner at a rather late period of life, and +have consequently transferred their new characters to +both sexes at a corresponding late period; and if so, +such cases are not opposed to our rule. Thus there +are sub-breeds of the pigeon, described by Neumeis<span class='pagenum'><a name="Page_294" id="Page_294">294</a></span>ter,<a name="FNanchor_365" id="FNanchor_365"></a><a href="#Footnote_365" class="fnanchor">365</a> +both sexes of which change colour after moulting +twice or thrice, as does likewise the Almond Tumbler; +nevertheless these changes, though occurring rather +late in life, are common to both sexes. One variety +of the Canary-bird, namely the London Prize, offers a +nearly analogous case.</p> + +<p>With the breeds of the Fowl the inheritance of various +characters by one sex or by both sexes, seems generally +determined by the period at which such characters are +developed. Thus in all the many breeds in which the +adult male differs greatly in colour from the female and +from the adult male parent-species, he differs from the +young male, so that the newly acquired characters must +have appeared at a rather late period of life. On the +other hand with most of the breeds in which the two sexes +resemble each other, the young are coloured in nearly +the same manner as their parents, and this renders it +probable that their colours first appeared early in life. +We have instances of this fact in all black and white +breeds, in which the young and old of both sexes are +alike; nor can it be maintained that there is something +peculiar in a black or white plumage, leading to its +transference to both sexes; for the males alone of many +natural species are either black or white, the females +being very differently coloured. With the so-called +Cuckoo sub-breeds of the fowl, in which the feathers are +transversely pencilled with dark stripes, both sexes and +the chickens are coloured in nearly the same manner. +The laced plumage of the Sebright bantam is the same +in both sexes, and in the chickens the feathers are tipped +with black, which makes a near approach to lacing. +Spangled Hamburghs, however, offer a partial exception, +<span class='pagenum'><a name="Page_295" id="Page_295">295</a></span>for the two sexes, though not quite alike, resemble each +other more closely than do the sexes of the aboriginal +parent-species, yet they acquire their characteristic +plumage late in life, for the chickens are distinctly +pencilled. Turning to other characters besides colour: +the males alone of the wild parent-species and of most +domestic breeds possess a fairly well developed comb, but +in the young of the Spanish fowl it is largely developed +at a very early age, and apparently in consequence of +this it is of unusual size in the adult females. In the +Game breeds pugnacity is developed at a wonderfully +early age, of which curious proofs could be given; and +this character is transmitted to both sexes, so that the +hens, from their extreme pugnacity, are now generally +exhibited in separate pens. With the Polish breeds the +bony protuberance of the skull which supports the crest +is partially developed even before the chickens are +hatched, and the crest itself soon begins to grow, though +at first feebly;<a name="FNanchor_366" id="FNanchor_366"></a><a href="#Footnote_366" class="fnanchor">366</a> and in this breed a great bony protuberance +and an immense crest characterise the adults of +both sexes.</p> + +<p class="tb">Finally, from what we have now seen of the relation +which exists in many natural species and domesticated +races, between the period of the development of their +characters and the manner of their transmission—for +example the striking fact of the early growth of the +horns in the reindeer, in which both sexes have horns, +in comparison with their much later growth in the +other species in which the male alone bears horns—we +may conclude that one cause, though not the sole +<span class='pagenum'><a name="Page_296" id="Page_296">296</a></span>cause, of characters being exclusively inherited by one +sex, is their development at a late age. And secondly, +that one, though apparently a less efficient, cause of +characters being inherited by both sexes is their development +at an early age, whilst the sexes differ but +little in constitution. It appears, however, that some +difference must exist between the sexes even during an +early embryonic period, for characters developed at this +age not rarely become attached to one sex.</p> + +<p><i>Summary and concluding remarks.</i>—From the foregoing +discussion on the various laws of inheritance, we +learn that characters often or even generally tend to +become developed in the same sex, at the same age, +and periodically at the same season of the year, in +which they first appeared in the parents. But these +laws, from unknown causes, are very liable to change. +Hence the successive steps in the modification of a +species might readily be transmitted in different ways; +some of the steps being transmitted to one sex, +and some to both; some to the offspring at one age, +and some at all ages. Not only are the laws of inheritance +extremely complex, but so are the causes which +induce and govern variability. The variations thus +caused are preserved and accumulated by sexual selection, +which is in itself an extremely complex affair, +depending, as it does, on ardour in love, courage, and +the rivalry of the males, and on the powers of perception, +taste, and will of the female. Sexual selection will +also be dominated by natural selection for the general +welfare of the species. Hence the manner in which the +individuals of either sex or of both sexes are affected +through sexual selection cannot fail to be complex in +the highest degree.</p> + +<p>When variations occur late in life in one sex, and are<span class='pagenum'><a name="Page_297" id="Page_297">297</a></span> +transmitted to the same sex at the same age, the other +sex and the young are necessarily left unmodified. +When they occur late in life, but are transmitted to +both sexes at the same age, the young alone are left unmodified. +Variations, however, may occur at any period +of life in one sex or in both, and be transmitted to both +sexes at all ages, and then all the individuals of the +species will be similarly modified. In the following chapters +it will be seen that all these cases frequently +occur under nature.</p> + +<p>Sexual selection can never act on any animal whilst +young, before the age for reproduction has arrived. +From the great eagerness of the male it has generally +acted on this sex and not on the females. The +males have thus become provided with weapons for +fighting with their rivals, or with organs for discovering +and securely holding the female, or for exciting and +charming her. When the sexes differ in these respects, +it is also, as we have seen, an extremely general law +that the adult male differs more or less from the young +male; and we may conclude from this fact that the +successive variations, by which the adult male became +modified, cannot have occurred much before the age +for reproduction. How then are we to account for this +general and remarkable coincidence between the period +of variability and that of sexual selection,—principles +which are quite independent of each other? I think +we can see the cause: it is not that the males have +never varied at an early age, but that such variations +have commonly been lost, whilst those occurring at a +later age have been preserved.</p> + +<p>All animals produce more offspring than can survive +to maturity; and we have every reason to believe that +death falls heavily on the weak and inexperienced +young. If then a certain proportion of the offspring<span class='pagenum'><a name="Page_298" id="Page_298">298</a></span> +were to vary at birth or soon afterwards, in some +manner which at this age was of no service to them, +the chance of the preservation of such variations would +be small. We have good evidence under domestication +how soon variations of all kinds are lost, if not selected. +But variations which occurred at or near maturity, and +which were of immediate service to either sex, would +probably be preserved; as would similar variations +occurring at an earlier period in any individuals which +happened to survive. As this principle has an important +bearing on sexual selection, it may be advisable to +give an imaginary illustration. We will take a pair of +animals, neither very fertile nor the reverse, and assume +that after arriving at maturity they live on an average +for five years, producing each year five young. They +would thus produce 25 offspring; and it would not, I +think, be an unfair estimate to assume that 18 or 20 +out of the 25 would perish before maturity, whilst still +young and inexperienced; the remaining seven or five +sufficing to keep up the stock of mature individuals. +If so, we can see that variations which occurred during +youth, for instance in brightness, and which were not +of the least service to the young, would run a good +chance of being utterly lost. Whilst similar variations, +which occurring at or near maturity in the comparatively +few individuals surviving to this age, and which +immediately gave an advantage to certain males, by +rendering them more attractive to the females, would be +likely to be preserved. No doubt some of the variations +in brightness which occurred at an earlier age +would by chance be preserved, and eventually give to +the male the same advantage as those which appeared +later; and this will account for the young males commonly +partaking to a certain extent (as may be observed +with many birds) of the bright colours of their<span class='pagenum'><a name="Page_299" id="Page_299">299</a></span> +adult male parents. If only a few of the successive +variations in brightness were to occur at a late age, +the adult male would be only a little brighter than the +young male; and such cases are common.</p> + +<p>In this illustration I have assumed that the young +varied in a manner which was of no service to them; +but many characters proper to the adult male would be +actually injurious to the young,—as bright colours from +making them conspicuous, or horns of large size from +expending much vital force. Such variations in the +young would promptly be eliminated through natural +selection. With the adult and experienced males, on +the other hand, the advantage thus derived in their +rivalry with other males would often more than counterbalance +exposure to some degree of danger. Thus we +can understand how it is that variations which must +originally have appeared rather late in life have alone +or in chief part been preserved for the development of +secondary sexual characters; and the remarkable coincidence +between the periods of variability and of sexual +selection is intelligible.</p> + +<p>As variations which give to the male an advantage +in lighting with other males, or in finding, securing, +or charming the female, would be of no use to the +female, they will not have been preserved in this sex +either during youth or maturity. Consequently such +variations would be extremely liable to be lost; and the +female, as far as these characters are concerned, would be +left unmodified, excepting in so far as she may have received +them by transference from the male. No doubt if +the female varied and transferred serviceable characters +to her male offspring, these would be favoured through +sexual selection; and then both sexes would thus far be +modified in the same manner. But I shall hereafter +have to recur to these more intricate contingencies.</p> + +<p class="tb"><span class='pagenum'><a name="Page_300" id="Page_300">300</a></span>In the following chapters, I shall treat of the +secondary sexual characters in animals of all classes, +and shall endeavour in each case to apply the principles +explained in the present chapter. The lowest +classes will detain us for a very short time, but the +higher animals, especially birds, must be treated at +considerable length. It should be borne in mind that +for reasons already assigned, I intend to give only a few +illustrative instances of the innumerable structures by +the aid of which the male finds the female, or, when +found, holds her. On the other hand, all structures +and instincts by which the male conquers other males, +and by which he allures or excites the female, will be +fully discussed, as these are in many ways the most +interesting.</p> + +<h4><i>Supplement on the proportional numbers of the two sexes +in animals belonging to various classes.</i></h4> + +<p class="tb">As no one, as far as I can discover, has paid attention +to the relative numbers of the two sexes throughout +the animal kingdom, I will here give such materials +as I have been able to collect, although they are extremely +imperfect. They consist in only a few instances +of actual enumeration, and the numbers are not very +large. As the proportions are known with certainty on +a large scale in the case of man alone, I will first give +them, as a standard of comparison.</p> + +<p><i>Man.</i>—In England during ten years (from 1857 to +1866) 707,120 children on an annual average have +been born alive, in the proportion of 104.5 males to +100 females. But in 1857 the male births throughout +England were as 105.2, and in 1865 as 104.0 to +100. Looking to separate districts, in Buckinghamshire +(where on an average 5000 children are annually born)<span class='pagenum'><a name="Page_301" id="Page_301">301</a></span> +the <i>mean</i> proportion of male to female births, during +the whole period of the above ten years, was as 102.8 +to 100; whilst in N. Wales (where the average annual +births are 12,873) it was as high as 106.2 to 100. +Taking a still smaller district, viz., Rutlandshire (where +the annual births average only 739), in 1864 the male +births were as 114.6, and in 1862 as 97.0 to 100; but +even in this small district the average of the 7385 +births during the whole ten years was as 104.5 to 100; +that is in the same ratio as throughout England.<a name="FNanchor_367" id="FNanchor_367"></a><a href="#Footnote_367" class="fnanchor">367</a> +The proportions are sometimes slightly disturbed by +unknown causes; thus Prof. Faye states “that in +some districts of Norway there has been during a +decennial period a steady deficiency of boys, whilst +in others the opposite condition has existed.” In +France during forty-four years the male to the female +births have been as 106.2 to 100; but during this +period it has occurred five times in one department, +and six times in another, that the female births have +exceeded the males. In Russia the average proportion +is as high as 108.9 to 100.<a name="FNanchor_368" id="FNanchor_368"></a><a href="#Footnote_368" class="fnanchor">368</a> It is a singular fact that +with Jews the proportion of male births is decidedly +larger than with Christians: thus in Prussia the proportion +is as 113, in Breslau as 114, and in Livonia as 120 +to 100; the Christian births in these countries being +the same as usual, for instance, in Livonia as 104 to +100.<a name="FNanchor_369" id="FNanchor_369"></a><a href="#Footnote_369" class="fnanchor">369</a> It is a still more singular fact that in different +nations, under different conditions and climates, in +Naples, Prussia, Westphalia, France and England, the +<span class='pagenum'><a name="Page_302" id="Page_302">302</a></span>excess of male over female births is less when they are +illegitimate than when legitimate.<a name="FNanchor_370" id="FNanchor_370"></a><a href="#Footnote_370" class="fnanchor">370</a></p> + +<p>In various parts of Europe, according to Prof. Faye +and other authors, “a still greater preponderance of +males would be met with, if death struck both sexes +in equal proportion in the womb and during birth. +But the fact is, that for every 100 still-born females, +we have in several countries from 134.6 to 144.9 +still-born males.” Moreover during the first four or +five years of life more male children die than females; +“for example in England, during the first year, 126 +boys die for every 100 girls,—a proportion which in +France is still more unfavourable.”<a name="FNanchor_371" id="FNanchor_371"></a><a href="#Footnote_371" class="fnanchor">371</a> As a consequence +of this excess in the death-rate of male children, and of +the exposure of men when adult to various dangers, and of +their tendency to emigrate, the females in all old-settled +countries, where statistical records have been kept,<a name="FNanchor_372" id="FNanchor_372"></a><a href="#Footnote_372" class="fnanchor">372</a> are +found to preponderate considerably over the males.</p> + +<p class="tb">It has often been supposed that the relative ages +of the parents determine the sex of the offspring; +and Prof. Leuckart<a name="FNanchor_373" id="FNanchor_373"></a><a href="#Footnote_373" class="fnanchor">373</a> has advanced what he considers +<span class='pagenum'><a name="Page_303" id="Page_303">303</a></span>sufficient evidence, with respect to man and certain +domesticated animals, to shew that this is one important +factor in the result. So again the period of +impregnation has been thought to be the efficient cause; +but recent observations discountenance this belief. +Again, with mankind polygamy has been supposed to +lead to the birth of a greater proportion of female +infants; but Dr. J. Campbell<a name="FNanchor_374" id="FNanchor_374"></a><a href="#Footnote_374" class="fnanchor">374</a> carefully attended to +this subject in the harems of Siam, and he concludes +that the proportion of male to female births is the +same as from monogamous unions. Hardly any animal +has been rendered so highly polygamous as our English +race-horses, and we shall immediately see that their +male and female offspring are almost exactly equal in +number.</p> + +<div class="blockquot2"><p class="tb"><i>Horses.</i>—Mr. Tegetmeier has been so kind as to tabulate for me +from the ‘Racing Calendar’ the births of race-horses during a period +of twenty-one years, viz. from 1846 to 1867; 1849 being omitted, +as no returns were that year published. The total births have +been 25,560,<a name="FNanchor_375" id="FNanchor_375"></a><a href="#Footnote_375" class="fnanchor">375</a> consisting of 12,763 males and 12,797 females, or in +the proportion of 99.7 males to 100 females. As these numbers are +tolerably large, and as they are drawn from all parts of England, +during several years, we may with much confidence conclude that +with the domestic horse, or at least with the race-horse, the two +sexes are produced in almost equal numbers. The fluctuations in +the proportions during successive years are closely like those which +occur with mankind, when a small and thinly-populated area is considered: +thus in 1856 the male horses were as 107.1, and in 1867 +as only 92.6 to 100 females. In the tabulated returns the proportions +vary in cycles, for the males exceeded the females during six +successive years; and the females exceeded the males during two +<span class='pagenum'><a name="Page_304" id="Page_304">304</a></span>periods each of four years: this, however, may be accidental; at +least I can detect nothing of the kind with man in the decennial +table in the Registrar’s Report for 1866. I may add that certain, +mares, and this holds good with certain cows and with women, +tend to produce more of one sex than of the other; Mr. Wright of +Yeldersley House, informs me that one of his Arab mares, though +put seven times to different horses, produced seven fillies.</p> + +<p class="tb"><i>Dogs.</i>—During a period of twelve years, from 1857 to 1868, the +births of a large number of greyhounds, throughout England, have +been sent to the ‘Field’ newspaper; and I am again indebted to +Mr. Tegetmeier for carefully tabulating the results. The recorded, +births have been 6878, consisting of 3605 males and 3273 females, +that is, in the proportion of 110.1 males to 100 females. The +greatest fluctuations occurred in 1864, when the proportion was as +95.3 males, and in 1867, as 116.3 males to 100 females. The above +average proportion of 110.1 to 100 is probably nearly correct in the +case of the greyhound, but whether it would hold with other domesticated +breeds is in some degree doubtful. Mr. Cupples has enquired +from several great breeders of dogs, and finds that all without +exception believe that females are produced in excess; he suggests +that this belief may have arisen from females being less valued and +the consequent disappointment producing a stronger impression on +the mind.</p> + +<p class="tb"><i>Sheep.</i>—The sexes of sheep are not ascertained by agriculturists +until several months after birth, at the period when the males are +castrated; so that the following returns do not give the proportions +at birth. Moreover, I find that several great breeders in Scotland, +who annually raise some thousand sheep, are firmly convinced that +a larger proportion of males than of females die during the first one +or two years; therefore the proportion of males would be somewhat +greater at birth than at the age of castration. This is a remarkable +coincidence with what occurs, as we have seen, with mankind, and +both cases probably depend on some common cause. I have received +returns from four gentlemen in England who have bred lowland +sheep, chiefly Leicesters, during the last ten or sixteen years; +they amount altogether to 8965 births, consisting of 4407 males +and 4558 females; that is in the proportion of 96.7 males to 100 +females. With respect to Cheviot and black-faced sheep bred in +Scotland, I have received returns from six breeders, two of them on +a large scale, chiefly for the years 1867-1869, but some of the +returns extending back to 1862. The total number recorded +amounts to 50,685, consisting of 25,071 males and 25,614 females, +or in the proportion of 97.9 males to 100 females. If we take the +English and Scotch returns together, the total number amounts +<span class='pagenum'><a name="Page_305" id="Page_305">305</a></span>to 59,650, consisting of 29,478 males and 30,172 females, or as +97·7 to 100. So that with sheep at the age of castration the females +are certainly in excess of the males; but whether this would hold +good at birth is doubtful, owing to the greater liability in the males +to early death.<a name="FNanchor_376" id="FNanchor_376"></a><a href="#Footnote_376" class="fnanchor">376</a></p> + +<p class="tb">Of <i>Cattle</i> I have received returns from nine gentlemen of 982 +births, too few to be trusted; these consisted of 477 bull-calves and +505 cow-calves; <i>i.e.</i> in the proportion of 94·4 males to 100 females. +The Rev. W. D. Fox informs me that in 1867 out of 34 calves born +on a farm in Derbyshire only one was a bull. Mr. Harrison Weir +writes to me that he has enquired from several breeders of <i>Pigs</i>, and +most of them estimate the male to the female births as about +7 to 6. This same gentleman has bred <i>Rabbits</i> for many years, +and has noticed that a far greater number of bucks are produced +than does.</p> + +<p>Of mammalia in a state of nature I have been able to learn very +little. In regard to the common rat, I have received conflicting +statements. Mr. R. Elliot of Laighwood, informs me that a rat-catcher +assured him that he had always found the males in +great excess, even with the young in the nest. In consequence +of this, Mr. Elliot himself subsequently examined some hundred +old ones, and found the statement true. Mr. F. Buckland has +bred a large number of white rats, and he also believes that the +males greatly exceed the females. In regard to Moles, it is said +that “the males are much more numerous than the females;”<a name="FNanchor_377" id="FNanchor_377"></a><a href="#Footnote_377" class="fnanchor">377</a> +and as the catching of these animals is a special occupation, the +statement may perhaps be trusted. Sir A. Smith, in describing +an antelope of S. Africa<a name="FNanchor_378" id="FNanchor_378"></a><a href="#Footnote_378" class="fnanchor">378</a> (<i>Kobus ellipsiprymnus</i>), remarks, that +in the herds of this and other species, the males are few in number +compared with the females: the natives believe that they are born +in this proportion; others believe that the younger males are expelled +from the herds, and Sir A. Smith says, that though he has +himself never seen herds consisting of young males alone, others +affirm that this does occur. It appears probable that the young +males when expelled from the herd, would be likely to fell a prey +to the many beasts of prey of the country.</p></div> + +<p><span class='pagenum'><a name="Page_306" id="Page_306">306</a></span></p> +<h4>BIRDS.</h4> + +<div class="blockquot2"><p>With respect to the <i>Fowl</i>, I have received only one account, +namely, that out of 1001 chickens of a highly-bred stock of Cochins, +reared during eight years by Mr. Stretch, 487 proved males and 514 +females: <i>i.e.</i> as 94.7 to 100. In regard to domestic pigeons there is +good evidence that the males are produced in excess, or that their +lives are longer; for these birds invariably pair, and single males, +as Mr. Tegetmeier informs me, can always be purchased cheaper +than females. Usually the two birds reared from the two eggs +laid in the same nest consist of a male and female; but Mr. Harrison +Weir, who has been so large a breeder, says that he has often bred +two cocks from the same nest, and seldom two hens; moreover the +hen is generally the weaker of the two, and more liable to perish.</p> + +<p>With respect to birds in a state of nature, Mr. Gould and others<a name="FNanchor_379" id="FNanchor_379"></a><a href="#Footnote_379" class="fnanchor">379</a> +are convinced that the males are generally the more numerous; and +as the young males of many species resemble the females, the latter +would naturally appear to be the most numerous. Large numbers of +pheasants are reared by Mr. Baker of Leadenhall from eggs laid +by wild birds, and he informs Mr. Jenner Weir that four or five +males to one female are generally produced. An experienced observer +remarks<a name="FNanchor_380" id="FNanchor_380"></a><a href="#Footnote_380" class="fnanchor">380</a> that in Scandinavia the broods of the capercailzie +and black-cock contain more males than females; and that with the +Dal-ripa (a kind of ptarmigan) more males than females attend the +<i>leks</i> or places of courtship; but this latter circumstance is accounted +for by some observers by a greater number of hen birds being killed +by vermin. From various facts given by White of Selbourne,<a name="FNanchor_381" id="FNanchor_381"></a><a href="#Footnote_381" class="fnanchor">381</a> it +seems clear that the males of the partridge must be in considerable excess +in the south of England; and I have been assured that this +is the case in Scotland. Mr. Weir on enquiring from the dealers +who receive at certain seasons large numbers of ruffs (<i>Machetes +pugnax</i>) was told that the males are much the most numerous. +This same naturalist has also enquired for me from the bird-catchers, +who annually catch an astonishing number of various small +species alive for the London market, and he was unhesitatingly +answered by an old and trustworthy man, that with the chaffinch +the males are in large excess; he thought as high as 2 males to +<span class='pagenum'><a name="Page_307" id="Page_307">307</a></span>1 female, or at least as high as 5 to 3.<a name="FNanchor_382" id="FNanchor_382"></a><a href="#Footnote_382" class="fnanchor">382</a> The males of the blackbird, +he likewise maintained, were by far the most numerous, whether +caught by traps or by netting at night. These statements +may apparently be trusted, because the same man said that the +sexes are about equal with the lark, the twite (<i>Linaria montana</i>), +and goldfinch. On the other hand he is certain that with the +common linnet, the females preponderate greatly, but unequally +during different years; during some years he has found the females +to the males as four to one. It should, however, be borne in mind, +that the chief season for catching birds does not begin till September, +so that with some species partial migrations may have begun, +and the flocks at this period often consist of hens alone. Mr. Salvin +paid particular attention to the sexes of the humming-birds in +Central America, and he is convinced that with most of the species +the males are in excess; thus one year he procured 204 specimens +belonging to ten species, and these consisted of 166 males and of +38 females. With two other species the females were in excess: +but the proportions apparently vary either during different seasons +or in different localities; for on one occasion the males of <i>Campylopterus +hemileucurus</i> were to the females as five to two, and +on another occasion<a name="FNanchor_383" id="FNanchor_383"></a><a href="#Footnote_383" class="fnanchor">383</a> in exactly the reversed ratio. As bearing on +this latter point, I may add, that Mr. Powys found in Corfu and +Epirus the sexes of the chaffinch keeping apart, and “the females +by far the most numerous;” whilst in Palestine Mr. Tristram +found “the male flocks appearing greatly to exceed the female in +number.”<a name="FNanchor_384" id="FNanchor_384"></a><a href="#Footnote_384" class="fnanchor">384</a> So again with the <i>Quiscalus major</i>, Mr. G. Taylor<a name="FNanchor_385" id="FNanchor_385"></a><a href="#Footnote_385" class="fnanchor">385</a> +says, that in Florida there were “very few females in proportion to +the males,” whilst in Honduras the proportion was the other way, +the species there having the character of a polygamist.</p></div> + +<h4>FISH.</h4> + +<div class="blockquot2"><p>With Fish the proportional numbers of the sexes can be ascertained +only by catching them in the adult or nearly adult state; and there +<span class='pagenum'><a name="Page_308" id="Page_308">308</a></span>are many difficulties in arriving at any just conclusion.<a name="FNanchor_386" id="FNanchor_386"></a><a href="#Footnote_386" class="fnanchor">386</a> Infertile +females might readily be mistaken for males, as Dr. Günther has +remarked to me in regard to trout. With some species the males +are believed to die soon after fertilising the ova. With many species +the males are of much smaller size than the females, so that a large +number of males would escape from the same net by which the females +were caught. M. Carbonnier,<a name="FNanchor_387" id="FNanchor_387"></a><a href="#Footnote_387" class="fnanchor">387</a> who has especially attended +to the natural history of the pike (<i>Esox lucius</i>) states that many +males, owing to their small size, are devoured by the larger females; +and he believes that the males of almost all fish are exposed from +the same cause to greater danger than the females. Nevertheless +in the few cases in which the proportional numbers have been +actually observed, the males appear to be largely in excess. Thus +Mr. R. Buist, the superintendent of the Stormontfield experiments, +says that in 1865, out of 70 salmon first landed for the +purpose of obtaining the ova, upwards of 60 were males. In +1867 he again “calls attention to the vast disproportion of the +males to the females. We had at the outset at least ten males +to one female.” Afterwards sufficient females for obtaining ova +were procured. He adds, “from the great proportion of the +males, they are constantly fighting and tearing each other on the +spawning-beds.”<a name="FNanchor_388" id="FNanchor_388"></a><a href="#Footnote_388" class="fnanchor">388</a> This disproportion, no doubt, can be accounted +for in part, but whether wholly is very doubtful, by the males +ascending the rivers before the females. Mr. F. Buckland remarks +in regard to trout, that “it is a curious fact that the males preponderate +very largely in number over the females. It <i>invariably</i> +happens that when the first rush of fish is made to the net, there +will be at least seven or eight males to one female found captive. +I cannot quite account for this; either the males are more numerous +than the females, or the latter seek safety by concealment +rather than flight.” He then adds, that by carefully searching the +banks, sufficient females for obtaining ova can be found.<a name="FNanchor_389" id="FNanchor_389"></a><a href="#Footnote_389" class="fnanchor">389</a> Mr. H. +Lee informs me that out of 212 trout, taken for this purpose in Lord +Portsmouth’s park, 150 were males and 62 females.</p> + +<p>With the Cyprinidæ the males likewise seem to be in excess; +but several members of this Family, viz., the carp, tench, bream +and minnow, appear regularly to follow the practice, rare in the +<span class='pagenum'><a name="Page_309" id="Page_309">309</a></span>animal kingdom, of polyandry; for the female whilst spawning is +always attended by two males, one on each side, and in the case of +the bream by three or four males. This fact is so well known, that +it is always recommended to stock a pond with two male tenches +to one female, or at least with three males to two females. With +the minnow, an excellent observer states, that on the spawning-beds +the males are ten times as numerous as the females; when a +female comes amongst the males, “she is immediately pressed closely +by a male on each side; and when they have been in that situation +for a time, are superseded by other two males.”<a name="FNanchor_390" id="FNanchor_390"></a><a href="#Footnote_390" class="fnanchor">390</a></p></div> + +<h4>INSECTS.</h4> + +<div class="blockquot2"><p>In this class, the Lepidoptera alone afford the means of judging +of the proportional numbers of the sexes; for they have been collected +with special care by many good observers, and have been +largely bred from the egg or caterpillar state. I had hoped that +some breeders of silk-moths might have kept an exact record, but +after writing to France and Italy, and consulting various treatises, +I cannot find that this has ever been done. The general opinion +appears to be that the sexes are nearly equal, but in Italy as I hear +from Professor Canestrini, many breeders are convinced that the +females are produced in excess. The same naturalist, however, +informs me, that in the two yearly broods of the Ailanthus silk-moth +(<i>Bombyx cynthia</i>), the males greatly preponderate in the +first, whilst in the second the two sexes are nearly equal, or the +females rather in excess.</p> + +<p>In regard to Butterflies in a state of nature, several observers +have been much struck by the apparently enormous preponderance +of the males.<a name="FNanchor_391" id="FNanchor_391"></a><a href="#Footnote_391" class="fnanchor">391</a> Thus Mr. Bates,<a name="FNanchor_392" id="FNanchor_392"></a><a href="#Footnote_392" class="fnanchor">392</a> in speaking of the species, no +less than about a hundred in number, which inhabit the Upper +Amazons, says that the males are much more numerous than the +females, even in the proportion of a hundred to one. In North +America, Edwards, who had great experience, estimates in the +genus Papilio the males to the females as four to one; and Mr. +<span class='pagenum'><a name="Page_310" id="Page_310">310</a></span>Walsh, who informed me of this statement, says that with <i>P. +turnus</i> this is certainly the case. In South Africa, Mr. R. Trimen +found the males in excess in 19 species;<a name="FNanchor_393" id="FNanchor_393"></a><a href="#Footnote_393" class="fnanchor">393</a> and in one of these, +which swarms in open places, he estimated the number of males as +fifty to one female. With another species, in which the males are +numerous in certain localities, he collected during seven years only +five females. In the island of Bourbon, M. Maillard states that +the males of one species of Papilio are twenty times as numerous +as the females.<a name="FNanchor_394" id="FNanchor_394"></a><a href="#Footnote_394" class="fnanchor">394</a> Mr. Trimen informs me that as far as he has +himself seen, or heard from others, it is rare for the females of any +butterfly to exceed in number the males; but this is perhaps the +case with three South African species. Mr. Wallace<a name="FNanchor_395" id="FNanchor_395"></a><a href="#Footnote_395" class="fnanchor">395</a> states that +the females of <i>Ornithoptera crœsus</i>, in the Malay archipelago, are +more common and more easily caught than the males; but this is +a rare butterfly. I may here add, that in Hyperythra, a genus of +moths, Guenée says, that from four to five females are sent in +collections from India for one male.</p> + +<p>When this subject of the proportional numbers of the sexes of +insects was brought before the Entomological Society,<a name="FNanchor_396" id="FNanchor_396"></a><a href="#Footnote_396" class="fnanchor">396</a> it was +generally admitted that the males of most Lepidoptera, in the +adult or imago state, are caught in greater numbers than the +females; but this fact was attributed by various observers to the +more retiring habits of the females, and to the males emerging +earlier from the cocoon. This latter circumstance is well known to +occur with most Lepidoptera, as well as with other insects. So +that, as M. Personnat remarks, the males of the domesticated +<i>Bombyx yamamai</i>, are lost at the beginning of the season, and +the females at the end, from the want of mates.<a name="FNanchor_397" id="FNanchor_397"></a><a href="#Footnote_397" class="fnanchor">397</a> I cannot however +persuade myself that these causes suffice to explain the great +excess of males in the cases, above given, of butterflies which are +extremely common in their native countries. Mr. Stainton, who +has paid such close attention during many years to the smaller +moths, informs me that when he collected them in the imago state, +he thought that the males were ten times as numerous as the +females, but that since he has reared them on a large scale from the +caterpillar state, he is convinced that the females are the most +<span class='pagenum'><a name="Page_311" id="Page_311">311</a></span>numerous. Several entomologists concur in this view. Mr. Doubleday, +however, and some others, take an opposite view, and are convinced +that they have reared from the egg and caterpillar states a +larger proportion of males than of females.</p> + +<p>Besides the more active habits of the males, their earlier emergence +from the cocoon, and their frequenting in some cases more +open stations, other causes may be assigned for an apparent or real +difference in the proportional numbers of the sexes of Lepidoptera, +when captured in the imago state, and when reared from the +egg or caterpillar state. It is believed by many breeders in Italy, +as I hear from Professor Canestrini, that the female caterpillar of +the silk-moth suffers more from the recent disease than the male; +and Dr. Staudinger informs me that in rearing Lepidoptera more +females die in the cocoon than males. With many species the +female caterpillar is larger than the male, and a collector would +naturally choose the finest specimens, and thus unintentionally +collect a larger number of females. Three collectors have told me +that this was their practice; but Dr. Wallace is sure that most +collectors take all the specimens which they can find of the rarer +kinds, which alone are worth the trouble of rearing. Birds when +surrounded by caterpillars would probably devour the largest; and +Professor Canestrini informs me that in Italy some breeders believe, +though on insufficient evidence, that in the first brood of the +Ailanthus silk-moth, the wasps destroy a larger number of the +female than of the male caterpillars. Dr. Wallace further remarks +that female caterpillars, from being larger than the males, require +more time for their development and consume more food and moisture; +and thus they would be exposed during a longer time to +danger from ichneumons, birds, &c., and in times of scarcity would +perish in greater numbers. Hence it appears quite possible that, +in a state of nature, fewer female Lepidoptera may reach maturity +than males; and for our special object we are concerned with the +numbers at maturity, when the sexes are ready to propagate their +kind.</p> + +<p>The manner in which the males of certain moths congregate in +extraordinary numbers round a single female, apparently indicates +a great excess of males, though this fact may perhaps be accounted +for by the earlier emergence of the males from their cocoons. Mr. +Stainton informs me that from twelve to twenty males may often be +seen congregated round a female <i>Elachista rufocinerea</i>. It is well +known that if a virgin <i>Lasiocampa quercus</i> or <i>Saturnia carpini</i> +be exposed in a cage, vast numbers of males collect round her, +and if confined in a room will even come down the chimney to her. +<span class='pagenum'><a name="Page_312" id="Page_312">312</a></span>Mr. Doubleday believes that he has seen from fifty to a hundred +males of both these species attracted in the course of a single day +by a female under confinement. Mr. Trimen exposed in the Isle +of Wight a box in which a female of the Lasiocampa had been +confined on the previous day, and five males soon endeavoured +to gain admittance. M. Verreaux, in Australia, having placed +the female of a small Bombyx in a box in his pocket, was followed +by a crowd of males, so that about 200 entered the house +with him.<a name="FNanchor_398" id="FNanchor_398"></a><a href="#Footnote_398" class="fnanchor">398</a></p> + +<p>Mr. Doubleday has called my attention to Dr. Staudinger’s<a name="FNanchor_399" id="FNanchor_399"></a><a href="#Footnote_399" class="fnanchor">399</a> list +of Lepidoptera, which gives the prices of the males and females of +300 species or well-marked varieties of (Rhopalocera) butterflies. +The prices for both sexes of the very common species are of +course the same; but with 114 of the rarer species they differ; the +males being in all cases, excepting one, the cheapest. On an average +of the prices of the 113 species, the price of the male to that +of the female is as 100 to 149; and this apparently indicates that +inversely the males exceed the females in number in the same +proportion. About 2000 species or varieties of moths (Heterocera) +are catalogued, those with wingless females being here excluded on +account of the difference in habits of the two sexes: of these 2000 +species, 141 differ in price according to sex, the males of 130 being +cheaper, and the males of only 11 being dearer than the females. +The average price of the males of the 130 species, to that of the +females, is as 100 to 143. With respect to the butterflies in this +priced list, Mr. Doubleday thinks (and no man in England has had +more experience), that there is nothing in the habits of the species +which can account for the difference in the prices of the two sexes, +and that it can be accounted for only by an excess in the numbers of +the males. But I am bound to add that Dr. Staudinger himself, as +he informs me, is of a different opinion. He thinks that the less +active habits of the females and the earlier emergence of the males will +account for his collectors securing a larger number of males than +of females, and consequently for the lower prices of the former +With respect to specimens reared from the caterpillar-state, Dr. +Staudinger believes, as previously stated, that a greater number of +females than of males die under confinement in the cocoons. He +adds that with certain species one sex seems to preponderate over +the other during certain years.</p> + +<p>Of direct observations on the sexes of Lepidoptera, reared either +<span class='pagenum'><a name="Page_313" id="Page_313">313</a></span>from eggs or caterpillars, I have received only the few following +cases:—</p> + +<table width="100%" summary="Lepidoptera"> +<tr> +<td class="left80"> </td> +<td class="right10">Males.</td> +<td class="right10">Females.</td> +</tr> +<tr> +<td class="left80">The Rev. J. Hellins<a name="FNanchor_400" id="FNanchor_400"></a><a href="#Footnote_400" class="fnanchor">400</a> of Exeter reared, during +1868, imagos of 73 species, which consisted of</td> +<td class="right10">153</td> +<td class="right10">137</td> +</tr> +<tr> +<td class="left80">Mr. Albert Jones of Eltham reared, during 1868, +imagos of 9 species, which, consisted of</td> +<td class="right10">159</td> +<td class="right10">126</td> +</tr> +<tr> +<td class="left80">During 1869 he reared imagos from 4 species, +consisting of</td> +<td class="right10">114</td> +<td class="right10">112</td> +</tr> +<tr> +<td class="left80">Mr. Buckler of Emsworth, Hants, during 1869, +reared imagos from 74 species, consisting of</td> +<td class="right10">180</td> +<td class="right10">169</td> +</tr> +<tr> +<td class="left80">Dr. Wallace of Colchester reared from one brood +of Bombyx cynthia</td> +<td class="right10">52</td> +<td class="right10">48</td> +</tr> +<tr> +<td class="left80">Dr. Wallace raised, from cocoons of Bombyx +Pernyi sent from China, during 1869</td> +<td class="right10">224</td> +<td class="right10">123</td> +</tr> +<tr> +<td class="left80">Dr. Wallace raised, during 1868 and 1869, +from two lots of cocoons of Bombyx Yamamai</td> +<td class="right10">52</td> +<td class="right10">46</td> +</tr> +<tr> +<td class="left80"> </td> +<td class="right10">——</td> +<td class="right10">——</td> +</tr> +<tr> +<td class="left80"> Total</td> +<td class="right10">934</td> +<td class="right10">761</td> +</tr> +</table> + +<p>So that in these eight lots of cocoons and eggs, males were produced +in excess. Taken together the proportion of males is as +122.7 to 100 females. But the numbers are hardly large enough +to be trustworthy.</p> + +<p>On the whole, from the above various sources of evidence, all +pointing to the same direction, I infer that with most species of +Lepidoptera, the males in the imago state generally exceed the +females in number, whatever the proportions may be at their first +emergence from the egg.</p> + +<p>With reference to the other Orders of insects, I have been able +to collect very little reliable information. With the stag-beetle +(<i>Lucanus cervus</i>) “the males appear to be much more numerous +than the females;” but when, as Cornelius remarked during 1867, +an unusual number of these beetles appeared in one part of Germany, +the females appeared to exceed the males as six so one. +With one of the Elateridæ, the males are said to be much more +numerous than the females, and “two or three are often found +united with one female;”<a name="FNanchor_401" id="FNanchor_401"></a><a href="#Footnote_401" class="fnanchor">401</a> so that here polyandry seems to prevail. + +<span class='pagenum'><a name="Page_314" id="Page_314">314</a></span> + +With Siagonium (Staphylinidæ), in which the males are furnished +with horns, “the females are far more numerous than the opposite +sex.” Mr. Janson stated at the Entomological Society that the +females of the bark-feeding <i>Tomicus villosus</i> are so common as to +be a plague, whilst the males are so rare as to be hardly known. +In other Orders, from unknown causes, but apparently in some instances +owing to parthenogenesis, the males of certain species have +never been discovered or are excessively rare, as with several of the +Cynipidæ.<a name="FNanchor_402" id="FNanchor_402"></a><a href="#Footnote_402" class="fnanchor">402</a> In all the gall-making Cynipidæ known to Mr. Walsh, +the females are four or five times as numerous as the males; and so +it is, as he informs me, with the gall-making Cecidomyiiæ (Diptera). +With some common species of Saw-flies (Tenthredinæ) Mr. F. +Smith has reared hundreds of specimens from larvæ of all sizes, +but has never reared a single male: on the other hand Curtis says,<a name="FNanchor_403" id="FNanchor_403"></a><a href="#Footnote_403" class="fnanchor">403</a> +that with certain species (Athalia), bred by him, the males to the +females were as six to one; whilst exactly the reverse occurred with +the mature insects of the same species caught in the fields. With +the Neuroptera, Mr. Walsh states that in many, but by no means +in all, the species of the Odonatous groups (Ephemerina), there is a +great overplus of males: in the genus Hetærina, also, the males are +generally at least four times as numerous as the females. In certain +species in the genus Gomphus the males are equally numerous, +whilst in two other species, the females are twice or thrice as +numerous as the males. In some European species of Psocus thousands +of females may be collected without a single male, whilst +with other species of the same genus both sexes are common.<a name="FNanchor_404" id="FNanchor_404"></a><a href="#Footnote_404" class="fnanchor">404</a> In +England, Mr. MacLachlan has captured hundreds of the female +<i>Apatania muliebris</i>, but has never seen the male; and of <i>Boreus +hyemalis</i> only four or five males have been here seen.<a name="FNanchor_405" id="FNanchor_405"></a><a href="#Footnote_405" class="fnanchor">405</a> With most +of these species (excepting, as I have heard, with the Tenthredinæ) +there is no reason to suppose that the females are subject to parthenogenesis; +and thus we see how ignorant we are on the causes of the +apparent discrepancy in the proportional numbers of the two sexes.</p> + +<p>In the other Classes of the Articulata I have been able to collect +still less information. With Spiders, Mr. Blackwall, who has carefully +attended to this class during many years, writes to me that +the males from their more erratic habits are more commonly seen, +<span class='pagenum'><a name="Page_315" id="Page_315">315</a></span>and therefore appear to be the more numerous. This is actually the +case with a few species; but he mentions several species in six +genera, in which the females appear to be much more numerous +than the males.<a name="FNanchor_406" id="FNanchor_406"></a><a href="#Footnote_406" class="fnanchor">406</a> The small size of the males in comparison with +the females, which is sometimes carried to an extreme degree, and +their widely different appearance, may account in some instances +for their rarity in collections.<a name="FNanchor_407" id="FNanchor_407"></a><a href="#Footnote_407" class="fnanchor">407</a></p> + +<p class="tb">Some of the lower Crustaceans are able to propagate their kind +asexually, and this will account for the extreme rarity of the males. +With some other forms (as with Tanais and Cypris) there is reason +to believe, as Fritz Müller informs me, that the male is much shorter-lived +than the female, which, supposing the two sexes to be at first +equal in number, would explain the scarcity of the males. On the +other hand this same naturalist has invariably taken, on the shores +of Brazil, far more males than females of the Diastylidæ and of +Cypridina; thus with a species in the latter genus, 63 specimens +caught the same day, included 57 males; but he suggests that this +preponderance may be due to some unknown difference in the habits +of the two sexes. With one of the higher Brazilian crabs, namely +a Gelasimus, Fritz Müller found the males to be more numerous +than the females. The reverse seems to be the case, according to +the large experience of Mr. C. Spence Bate, with six common British +crabs, the names of which he has given me.</p></div> + +<p><i>On the Power of Natural Selection to regulate the proportional +Numbers of the Sexes, and General Fertility.</i>—In +some peculiar cases, an excess in the number of one +sex over the other might be a great advantage to a +species, as with the sterile females of social insects, or +with those animals in which more than one male is +requisite to fertilise the female, as with certain cirripedes +and perhaps certain fishes. An inequality between +the sexes in these cases might have been acquired +through natural selection, but from their rarity they +need not here be further considered. In all ordinary +<span class='pagenum'><a name="Page_316" id="Page_316">316</a></span>cases an inequality would be no advantage or disadvantage +to certain individuals more than to others; and +therefore it could hardly have resulted from natural +selection. We must attribute the inequality to the +direct action of those unknown conditions, which with +mankind lead to the males being born in a somewhat +larger excess in certain countries than in others, or +which cause the proportion between the sexes to differ +slightly in legitimate and illegitimate births.</p> + +<p>Let us now take the case of a species producing from +the unknown causes just alluded to, an excess of one +sex—we will say of males—these being superfluous and +useless, or nearly useless. Could the sexes be equalised +through natural selection? We may feel sure, from all +characters being variable, that certain pairs would produce +a somewhat less excess of males over females than +other pairs. The former, supposing the actual number +of the offspring to remain constant, would necessarily +produce more females, and would therefore be more productive. +On the doctrine of chances a greater number +of the offspring of the more productive pairs would survive; +and these would inherit a tendency to procreate +fewer males and more females. Thus a tendency towards +the equalisation of the sexes would be brought +about. But our supposed species would by this process +be rendered, as just remarked, more productive; and +this would in many cases be far from an advantage; +for whenever the limit to the numbers which exist, depends, +not on destruction by enemies, but on the amount +of food, increased fertility will lead to severer competition +and to most of the survivors being badly fed. In +this case, if the sexes were equalised by an increase in +the number of the females, a simultaneous decrease in +the total number of the offspring would be beneficial, +or even necessary, for the existence of the species; and<span class='pagenum'><a name="Page_317" id="Page_317">317</a></span> +this, I believe, could be effected through natural selection +in the manner hereafter to be described. The +same train of reasoning is applicable in the above, +as well as in the following case, if we assume that +females instead of males are produced in excess, for +such females from not uniting with males would be +superfluous and useless. So it would be with polygamous +species, if we assume the excess of females +to be inordinately great.</p> + +<p>An excess of either sex, we will again say of the +males, could, however, apparently be eliminated through +natural selection in another and indirect manner, namely +by an actual diminution of the males, without any increase +of the females, and consequently without any +increase in the productiveness of the species. From +the variability of all characters, we may feel assured +that some pairs, inhabiting any locality, would produce +a rather smaller excess of superfluous males, but an +equal number of productive females. When the offspring +from the more and the less male-productive +parents were all mingled together, none would have any +direct advantage over the others; but those that produced +few superfluous males would have one great +indirect advantage, namely that their ova or embryos +would probably be larger and finer, or their young +better nurtured in the womb and afterwards. We see +this principle illustrated with plants; as those which +bear a vast number of seed produce small ones; whilst +those which bear comparatively few seeds, often produce +large ones well-stocked with nutriment for the use of the +seedlings.<a name="FNanchor_408" id="FNanchor_408"></a><a href="#Footnote_408" class="fnanchor">408</a> Hence the offspring of the parents which +<span class='pagenum'><a name="Page_318" id="Page_318">318</a></span>had wasted least force in producing superfluous males +would be the most likely to survive, and would inherit +the same tendency not to produce superfluous males, +whilst retaining their full fertility in the production of +females. So it would be with the converse case of the +female sex. Any slight excess, however, of either sex +could hardly be checked in so indirect a manner. Nor +indeed has a considerable inequality between the sexes +been always prevented, as we have seen in some of the +cases given in the previous discussion. In these cases +the unknown causes which determine the sex of the +embryo, and which under certain conditions lead to +the production of one sex in excess over the other, have +not been mastered by the survival of those varieties +which were subjected to the least waste of organised +matter and force by the production of superfluous individuals +of either sex. Nevertheless we may conclude +that natural selection will always tend, though sometimes +inefficiently, to equalise the relative numbers of +the two sexes.</p> + +<p>Having said this much on the equalisation of the +sexes, it may be well to add a few remarks on the regulation +through natural selection of the ordinary fertility +of species. Mr. Herbert Spencer has shewn in an able +discussion<a name="FNanchor_409" id="FNanchor_409"></a><a href="#Footnote_409" class="fnanchor">409</a> that with all organisms a ratio exists between +what he calls individuation and genesis; whence +it follows that beings which consume much matter or +force in their growth, complicated structure or activity, +or which produce ova and embryos of large size, or +which expend much energy in nurturing their young, +cannot be so productive as beings of an opposite nature. +Mr. Spencer further shews that minor differences in fertility +will be regulated through natural selection. Thus +<span class='pagenum'><a name="Page_319" id="Page_319">319</a></span>the fertility of each species will tend to increase, from +the more fertile pairs producing a larger number of offspring, +and these from their mere number will have the +best chance of surviving, and will transmit their tendency +to greater fertility. The only check to a continued +augmentation of fertility in each organism seems +to be either the expenditure of more power and the +greater risks run by the parents that produce a more +numerous progeny, or the contingency of very numerous +eggs and young being produced of smaller size, or less +vigorous, or subsequently not so well nurtured. To +strike a balance in any case between the disadvantages +which follow from the production of a numerous progeny, +and the advantages (such as the escape of at least +some individuals from various dangers) is quite beyond +our power of judgment.</p> + +<p>When an organism has once been rendered extremely +fertile, how its fertility can be reduced through natural +selection is not so clear as how this capacity was first +acquired. Yet it is obvious that if individuals of a +species, from a decrease of their natural enemies, were +habitually reared in larger numbers than could be supported, +all the members would suffer. Nevertheless the +offspring from the less fertile parents would have no +direct advantage over the offspring from the more fertile +parents, when all were mingled together in the +same district. All the individuals would mutually tend +to starve each other. The offspring indeed of the less +fertile parents would lie under one great disadvantage, +for from the simple fact of being produced in smaller +numbers, they would be the most liable to extermination. +Indirectly, however, they would partake of one +great advantage; for under the supposed condition of +severe competition, when all were pressed for food, it is +extremely probable that those individuals which from<span class='pagenum'><a name="Page_320" id="Page_320">320</a></span> +some variation in their constitution produced fewer eggs +or young, would produce them of greater size or vigour; +and the adults reared from such eggs or young would +manifestly have the best chance of surviving, and +would inherit a tendency towards lessened fertility. +The parents, moreover, which had to nourish or provide +for fewer offspring would themselves be exposed to a +less severe strain in the struggle for existence, and +would have a better chance of surviving. By these +steps, and by no others as far as I can see, natural +selection under the above conditions of severe competition +for food, would lead to the formation of a new +race less fertile, but better adapted for survival, than +the parent-race.</p> + +<hr /> +<p><span class='pagenum'><a name="Page_321" id="Page_321">321</a></span></p> + +<h3>CHAPTER IX.</h3> + +<h4><span class="smcap">Secondary Sexual Characters in the Lower Classes of +the Animal Kingdom.</span></h4> + +<div class="blockquot"><p>These characters absent in the lowest classes—Brilliant colours—Mollusca—Annelids—Crustacea, +secondary sexual characters +strongly developed; dimorphism; colour; characters not acquired +before maturity—Spiders, sexual colours of; stridulation by the +males—Myriapoda.</p></div> + +<p>In the lowest classes the two sexes are not rarely united +in the same individual, and therefore secondary sexual +characters cannot be developed. In many cases in which +the two sexes are separate, both are permanently attached +to some support, and the one cannot search or +struggle for the other. Moreover it is almost certain +that these animals have too imperfect senses and +much too low mental powers to feel mutual rivalry, +or to appreciate each other’s beauty or other attractions.</p> + +<p>Hence in these classes, such as the Protozoa, Cœlenterata, +Echinodermata, Scolecida, true secondary sexual +characters do not occur; and this fact agrees with the +belief that such characters in the higher classes have +been acquired through sexual selection, which depends +on the will, desires, and choice of either sex. Nevertheless +some few apparent exceptions occur; thus, as I +hear from Dr. Baird, the males of certain Entozoa, or +internal parasitic worms, differ slightly in colour from +the females; but we have no reason to suppose that +such differences have been augmented through sexual +selection.</p> + +<p><span class='pagenum'><a name="Page_322" id="Page_322">322</a></span>Many of the lower animals, whether hermaphrodites +or with the sexes separate, are ornamented with the +most brilliant tints, or are shaded and striped in an +elegant manner. This is the case with many corals +and sea-anemonies (Actineæ), with some jelly-fish (Medusæ, +Porpita, &c.), with some Planariæ, Ascidians, +numerous Star-fishes, Echini, &c.; but we may conclude +from the reasons already indicated, namely the union +of the two sexes in some of these animals, the permanently +affixed condition of others, and the low +mental powers of all, that such colours do not serve +as a sexual attraction, and have not been acquired +through sexual selection. With the higher animals +the case is very different; for with them when one sex +is much more brilliantly or conspicuously coloured +than the other, and there is no difference in the +habits of the two sexes which will account for this +difference, we have reason to believe in the influence +of sexual selection; and this belief is strongly confirmed +when the more ornamented individuals, which +are almost always the males, display their attractions +before the other sex. We may also extend this conclusion +to both sexes, when coloured alike, if their +colours are plainly analogous to those of one sex alone +in certain other species of the same group.</p> + +<p>How, then, are we to account for the beautiful or +even gorgeous colours of many animals in the lowest +classes? It appears very doubtful whether such colours +usually serve as a protection; but we are extremely +liable to err in regard to characters of all kinds in +relation to protection, as will be admitted by every one +who has read Mr. Wallace’s excellent essay on this +subject. It would not, for instance, at first occur to +any one that the perfect transparency of the Medusæ, +or jelly-fishes, was of the highest service to them as a<span class='pagenum'><a name="Page_323" id="Page_323">323</a></span> +protection; but when we are reminded by Häckel that +not only the medusæ but many floating mollusca, crustaceans, +and even small oceanic fishes partake of this +same glass-like structure, we can hardly doubt that +they thus escape the notice of pelagic birds and other +enemies.</p> + +<p>Notwithstanding our ignorance how far colour in +many cases serves as a protection, the most probable +view in regard to the splendid tints of many of the +lowest animals seems to be that their colours are the +direct result either of the chemical nature or the minute +structure of their tissues, independently of any benefit +thus derived. Hardly any colour is finer than that of +arterial blood; but there is no reason to suppose that +the colour of the blood is in itself any advantage; and +though it adds to the beauty of the maiden’s cheek, no +one will pretend that it has been acquired for this purpose. +So again with many animals, especially the lower +ones, the bile is richly coloured; thus the extreme +beauty of the Eolidæ (naked sea-slugs) is chiefly due, as +I am informed by Mr. Hancock, to the biliary glands +seen through the translucent integuments; this beauty +being probably of no service to these animals. The +tints of the decaying leaves in an American forest are +described by every one as gorgeous; yet no one supposes +that these tints are of the least advantage to +the trees. Bearing in mind how many substances +closely analogous to natural organic compounds have +been recently formed by chemists, and which exhibit +the most splendid colours, it would have been a strange +fact if substances similarly coloured had not often +originated, independently of any useful end being +thus gained, in the complex laboratory of living +organisms.</p> + +<p><span class='pagenum'><a name="Page_324" id="Page_324">324</a></span><i>The sub-kingdom of the Mollusca.</i>—Throughout this +great division (taken in its largest acceptation) of the +animal kingdom, secondary sexual characters, such as +we are here considering, never, as far as I can discover, +occur. Nor could they be expected in the three lowest +classes, namely in the Ascidians, Polyzoa, and Brachiopods +(constituting the Molluscoida of Huxley), for most +of these animals are permanently affixed to a support +or have their sexes united in the same individual. In +the Lamellibranchiata, or bivalve shells, hermaphroditism +is not rare. In the next higher class of the +Gasteropoda, or marine univalve shells, the sexes are +either united or separate. But in this latter case the +males never possess special organs for finding, securing, +or charming the females, or for fighting with other +males. The sole external difference between the sexes +consists, as I am informed by Mr. Gwyn Jeffreys, in +the shell sometimes differing a little in form; for +instance, the shell of the male periwinkle (<i>Littorina +littorea</i>) is narrower and has a more elongated spire than +that of the female. But differences of this nature, it +may be presumed, are directly connected with the act +of reproduction or with the development of the ova.</p> + +<p>The Gasteropoda, though capable of locomotion and +furnished with imperfect eyes, do not appear to be endowed +with sufficient mental powers for the members +of the same sex to struggle together in rivalry, and +thus to acquire secondary sexual characters. Nevertheless +with the pulmoniferous gasteropods, or land-shells, +the pairing is preceded by courtship; for these +animals, though hermaphrodites, are compelled by their +structure to pair together. Agassiz remarks,<a name="FNanchor_410" id="FNanchor_410"></a><a href="#Footnote_410" class="fnanchor">410</a> “Quiconque +a eu l’occasion d’observer les amours des lima<span class='pagenum'><a name="Page_325" id="Page_325">325</a></span>çons, +ne saurait mettre en doute la séduction déployée +dans les mouvements et les allures qui préparent et +accomplissent le double embrassement de ces hermaphrodites.” +These animals appear also susceptible +of some degree of permanent attachment: an accurate +observer, Mr. Lonsdale, informs me that he placed a +pair of land-shells (<i>Helix pomatia</i>), one of which was +weakly, into a small and ill-provided garden. After a +short time the strong and healthy individual disappeared, +and was traced by its track of slime over a wall into an +adjoining well-stocked garden. Mr. Lonsdale concluded +that it had deserted its sickly mate; but after an +absence of twenty-four hours it returned, and apparently +communicated the result of its successful exploration, +for both then started along the same track and disappeared +over the wall.</p> + +<p>Even in the highest class of the Mollusca, namely the +Cephalopoda or cuttle-fishes, in which the sexes are +separate, secondary sexual characters of the kind which +we are here considering, do not, as far as I can discover, +occur. This is a surprising circumstance, as these +animals possess highly-developed sense-organs and have +considerable mental powers, as will be admitted by +every one who has watched their artful endeavours to +escape from an enemy.<a name="FNanchor_411" id="FNanchor_411"></a><a href="#Footnote_411" class="fnanchor">411</a> Certain Cephalopoda, however, +are characterised by one extraordinary sexual character, +namely, that the male element collects within one of +the arms or tentacles, which is then cast off, and, +clinging by its sucking-discs to the female, lives for +a time an independent life. So completely does the +cast-off arm resemble a separate animal, that it was +described by Cuvier as a parasitic worm under the name +<span class='pagenum'><a name="Page_326" id="Page_326">326</a></span>of Hectocotyle. But this marvellous structure may be +classed as a primary rather than as a secondary sexual +character.</p> + +<p>Although with the Mollusca sexual selection does not +seem to have come into play; yet many univalve and +bivalve shells, such as volutes, cones, scallops, &c., are +beautifully coloured and shaped. The colours do not +appear in most cases to be of any use as a protection; +they are probably the direct result, as in the lowest +classes, of the nature of the tissues; the patterns and +the sculpture of the shell depending on its manner of +growth. The amount of light seems to a certain extent +to be influential; for although, as repeatedly stated by +Mr. Gwyn Jeffreys, the shells of some species living at a +profound depth are brightly coloured, yet we generally +see the lower surfaces and the parts covered by the +mantle less highly coloured than the upper and exposed +surfaces.<a name="FNanchor_412" id="FNanchor_412"></a><a href="#Footnote_412" class="fnanchor">412</a> In some cases, as with shells living amongst +corals or brightly-tinted sea-weeds, the bright colours +may serve as a protection. But many of the nudibranch +mollusca, or sea-slugs, are as beautifully coloured as +any shells, as may be seen in Messrs. Alder and Hancock’s +magnificent work; and from information kindly +given me by Mr. Hancock, it is extremely doubtful +whether these colours usually serve as a protection. +With some species this may be the case, as with one +which lives on the green leaves of algæ, and is itself +bright-green. But many brightly-coloured, white or +otherwise conspicuous species, do not seek concealment; +whilst again some equally conspicuous species, as well +as other dull-coloured kinds, live under stones and in +<span class='pagenum'><a name="Page_327" id="Page_327">327</a></span>dark recesses. So that with these nudibranch molluscs, +colour apparently does not stand in any close relation +to the nature of the places which they inhabit.</p> + +<p class="tb">These naked sea-slugs are hermaphrodites, yet they +pair together, as do land-snails, many of which have +extremely pretty shells. It is conceivable that two +hermaphrodites, attracted by each others’ greater beauty, +might unite and leave offspring which would inherit +their parents’ greater beauty. But with such lowly-organised +creatures this is extremely improbable. Nor +is it at all obvious how the offspring from the more +beautiful pairs of hermaphrodites would have any advantage, +so as to increase in numbers, over the offspring +of the less beautiful, unless indeed vigour and beauty +generally coincided. We have not here a number of +males becoming mature before the females, and the +more beautiful ones selected by the more vigorous +females. If, indeed, brilliant colours were beneficial +to an hermaphrodite animal in relation to its general +habits of life, the more brightly-tinted individuals would +succeed best and would increase in number; but this +would be a case of natural and not of sexual selection.</p> + +<p><i>Sub-kingdom of the Vermes or Annulosa</i>: Class, +<i>Annelida (or Sea-worms)</i>.—In this class, although +the sexes (when separate) sometimes differ from each +other in characters of such importance that they have +been placed under distinct genera or even families, +yet the differences do not seem of the kind which can +be safely attributed to sexual selection. These animals, +like those in the preceding classes, apparently stand too +low in the scale, for the individuals of either sex to +exert any choice in selecting a partner, or for the individuals +of the same sex to struggle together in rivalry.</p> + +<p class="tb"><span class='pagenum'><a name="Page_328" id="Page_328">328</a></span><i>Sub-kingdom of the Arthropoda</i>: Class, <i>Crustacea</i>.—In +this great class we first meet with undoubted secondary +sexual characters, often developed in a remarkable +manner. Unfortunately the habits of crustaceans +are very imperfectly known, and we cannot explain the +uses of many structures peculiar to one sex. With +the lower parasitic species the males are of small size, +and they alone are furnished with perfect swimming-legs, +antennæ and sense-organs; the females being +destitute of these organs, with their bodies often consisting +of a mere distorted mass. But these extraordinary + +<span class="figright" style="width: 200px;"><a name="f3" id="f3"></a><img src="images/fig3.png" width="200" height="408" +alt="Fig. 3. Labidocera Darwinii." title="" /> + +<span class="indent2">Fig. 3. Labidocera Darwinii,<br /> (from Lubbock).<br /></span> + +<span class="indent2"><i>a.</i> Part of right-hand anterior antenna +of male, forming a prehensile organ.<br /></span> + +<span class="indent2"><i>b.</i> Posterior pair of thoracic legs of male.<br /></span> +<span class="indent2"><i>c.</i> Ditto of female.</span></span> + +differences between the two sexes are no doubt related +to their widely different habits of life, and consequently +do not concern us. In various crustaceans, belonging +to distinct families, the anterior antennæ are furnished +with peculiar thread-like bodies, which are believed to +act as smelling-organs, and these are much more numerous +in the males than in the females. As the males, +without any unusual development of their olfactory +organs, would almost certainly be able sooner or later +to find the females, the increased number of the smelling-threads +has probably been acquired through sexual +selection, by the better provided males having been the +most successful in finding partners and in leaving offspring. +Fritz Müller has described a remarkable dimorphic +species of Tanais, in which the male is represented +by two distinct forms, never graduating into each other. +In the one form the male is furnished with more +numerous smelling-threads, and in the other form with +more powerful and more elongated chelæ or pincers +which serve to hold the female. Fritz Müller suggests +that these differences between the two male forms of the +same species must have originated in certain individuals +having varied in the number of the smelling-threads, +whilst other individuals varied in the shape and size of<span class='pagenum'><a name="Page_329" id="Page_329">329</a></span> +their chelæ; so that of the former, those which were best +able to find the female, and of the latter, those which were +best able to hold her when found, +have left the greater number of +progeny to inherit their respective +advantages.<a name="FNanchor_413" id="FNanchor_413"></a><a href="#Footnote_413" class="fnanchor">413</a></p> + +<p>In some of the lower crustaceans, +the right-hand anterior +antenna of the male differs +greatly in structure from the +left-hand one, the latter resembling +in its simple tapering +joints the antennæ of the female. +In the male the modified +antenna is either swollen +in the middle or angularly bent, +or converted (fig. 3) into an +elegant, and sometimes wonderfully +complex, prehensile organ.<a name="FNanchor_414" id="FNanchor_414"></a><a href="#Footnote_414" class="fnanchor">414</a> +It serves, as I hear from Sir J. +Lubbock, to hold the female, +and for this same purpose one +of the two posterior legs (<i>b</i>) on +the same side of the body is +converted into a forceps. In +another family the inferior or +posterior antennæ are “curiously zigzagged” in the +males alone.</p> + +<p><span class='pagenum'><a name="Page_330" id="Page_330">330</a></span></p> + +<div class="figcenter" style="width: 550px;"><a name="f4" id="f4"></a><img src="images/fig4.png" width="550" height="332" +alt="Fig. 4. Anterior part of body of Callianassa." title="" /> + +<p class="center2">Fig. 4. Anterior part of body of Callianassa (from Milne-Edwards), showing the unequal +and differently-constructed right and left-hand chelæ of the male.</p> + +<p class="small">N.B.—The artist by mistake has reversed the drawing, and made the left-hand chela +the largest.</p></div> + +<table width="100%" summary="fig 5 and 6" border="0"> +<tr> +<td class="center50"><div class="figcenter" style="width: 250px;"><a name="f5" id="f5"></a><img src="images/fig5.png" width="250" height="279" +alt="Fig. 5. Second leg of male Orchestia Tucuratinga." title="" /></div></td> +<td class="center50"><div class="figcenter" style="width: 211px;"><a name="f6" id="f6"></a><img src="images/fig6.png" width="211" height="279" +alt="Fig. 6. Ditto of female." title="" /></div></td> +</tr> +<tr> +<td class="center50">Fig. 5.</td> +<td class="center50">Fig. 6.</td> +</tr> +</table> + +<p class="small">Fig. 5. Second leg of male Orchestia Tucuratinga (from Fritz Müller).</p> +<p class="small">Fig. 6. Ditto of female.</p> + +<p>In the higher crustaceans the anterior legs form a +pair of chelæ or pincers, and these are generally +larger in the male than in the female. In many species +the chelæ on the opposite sides of the body are of +unequal size, the right-hand one being, as I am informed +by Mr. C. Spence Bate, generally, though not +invariably, the largest. This inequality is often much +greater in the male than in the female. The two chelæ +also often differ in structure (figs. 4 and 5), the smaller +one resembling those of the female. What advantage<span class='pagenum'><a name="Page_331" id="Page_331">331</a></span> +is gained by their inequality in size on the opposite +sides of the body, and by the inequality being much +greater in the male than in the female; and why, when +they are of equal size, both are often much larger in +the male than in the female, is not known. The chelæ +are sometimes of such length and size that they cannot +possibly be used, as I hear from Mr. Spence Bate, for +carrying food to the mouth. In the males of certain freshwater +prawns (Palæmon) the right leg is actually longer +than the whole body.<a name="FNanchor_415" id="FNanchor_415"></a><a href="#Footnote_415" class="fnanchor">415</a> It is probable that the great size +of one leg with its chelæ may aid the male in fighting +with his rivals; but this use will not account for their +inequality in the female on the opposite sides of the body. +In Gelasimus, according to a statement quoted by Milne-Edwards,<a name="FNanchor_416" id="FNanchor_416"></a><a href="#Footnote_416" class="fnanchor">416</a> +the male and female live in the same burrow, +which is worth notice, as shewing that they pair, and +the male closes the mouth of the burrow with one of its +chelæ, which is enormously developed; so that here it +indirectly serves as a means of defence. Their main +use, however, probably is to seize and to secure the +female, and this in some instances, as with Gammarus, +is known to be the case. The sexes, however, of the +common shore-crab (<i>Carcinus mænas</i>), as Mr. Spence +Bate informs me, unite directly after the female has +moulted her hard shell, and when she is so soft that she +would be injured if seized by the strong pincers of the +male; but as she is caught and carried about by the +male previously to the act of moulting, she could then +be seized with impunity.</p> + +<p>Fritz Müller states that certain species of Melita are +<span class='pagenum'><a name="Page_332" id="Page_332">332</a></span>distinguished from all other amphipods by the females +having “the coxal lamellæ of the penultimate pair of +feet produced into hook-like processes, of which the +males lay hold with the hands of the first pair.” The +development of these hook-like processes probably +resulted from those females which were the most +securely held during the act of reproduction, having +left the largest number of offspring. Another Brazilian +amphipod (<i>Orchestia Darwinii</i>, fig. 7) is described +by Fritz Müller, as presenting a case of dimorphism, +like that of Tanais; for there are two male +forms, which differ in the structure of their chelæ.<a name="FNanchor_417" id="FNanchor_417"></a><a href="#Footnote_417" class="fnanchor">417</a> As +chelæ of either shape would certainly have sufficed to +hold the female, for both are now used for this purpose, +the two male forms probably originated, by some having +varied in one manner and some in another; both forms +having derived certain special, but nearly equal advantages, +from their differently shaped organs.</p> + +<p>It is not known that male crustaceans fight together +for the possession of the females, but this is probable; +for with most animals when the male is larger than the +female, he seems to have acquired his greater size by +having conquered during many generations other males. +Now Mr. Spence Bate informs me that in most of the +crustacean orders, especially in the highest or the +Brachyura, the male is larger than the female; the +parasitic genera, however, in which the sexes follow +different habits of life, and most of the Entomostraca +must be excepted. The chelæ of many crustaceans are +weapons well adapted for fighting. Thus a Devil-crab +(<i>Portunus puber</i>) was seen by a son of Mr. Bate fighting +with a <i>Carcinus mænas</i>, and the latter was soon thrown +on its back, and had every limb torn from its body. + +<span class='pagenum'><a name="Page_333" id="Page_333">333</a></span> + + +When several males of a Brazilian Gelasimus, a species +furnished with immense pincers, were placed together +by Fritz Müller in a glass vessel, they mutilated and +killed each other. Mr. Bate put a large male <i>Carcinus +mænas</i> into a pan of water, inhabited by a female paired +with a smaller male; the latter was soon dispossessed, +but, as Mr. Bate adds, “if they fought, the victory<span class='pagenum'><a name="Page_334" id="Page_334">334</a></span> +was a bloodless one, for I saw no wounds.” This +same naturalist separated a male sand-skipper (so common +on our sea-shores), <i>Gammarus marinus</i>, from its +female, both of which were imprisoned in the same +vessel with many individuals of the same species. The +female being thus divorced joined her comrades. After +an interval the male was again put into the same +vessel and he then, after swimming about for a time, +dashed into the crowd, and without any fighting at once +took away his wife. This fact shews that in the Amphipoda, +an order low in the scale, the males and females +recognise each other, and are mutually attached.</p> + +<div class="figcenter" style="width: 550px;"><a name="f7" id="f7"></a><img src="images/fig7.png" width="550" height="689" +alt="Fig. 7. Orchestia Darwinii." title="" /> + +<span class="center2">Fig. 7. Orchestia Darwinii (from Fritz Müller), showing the differently-constructed +chelæ of the two male forms.</span></div> + +<p>The mental powers of the Crustacea are probably +higher than might have been expected. Any one who +has tried to catch one of the shore-crabs, so numerous +on many tropical coasts, will have perceived how wary +and alert they are. There is a large crab (<i>Birgos +latro</i>), found on coral islands, which makes at the +bottom of a deep burrow a thick bed of the picked +fibres of the cocoa-nut. It feeds on the fallen fruit of +this tree by tearing off the husk, fibre by fibre; and +it always begins at that end where the three eye-like +depressions are situated. It then breaks through +one of these eyes by hammering with its heavy front +pincers, and turning round, extracts the albuminous +core with its narrow posterior pincers. But these actions +are probably instinctive, so that they would be performed +as well by a young as by an old animal. +The following case, however, can hardly be so considered: +a trustworthy naturalist, Mr. Gardner,<a name="FNanchor_418" id="FNanchor_418"></a><a href="#Footnote_418" class="fnanchor">418</a> whilst +watching a shore-crab (Gelasimus) making its burrow, +<span class='pagenum'><a name="Page_335" id="Page_335">335</a></span>threw some shells towards the hole. One rolled in, +and three other shells remained within a few inches of +the mouth. In about five minutes the crab brought +out the shell which had fallen in, and carried it away +to the distance of a foot; it then saw the three other +shells lying near, and evidently thinking that they +might likewise roll in, carried them to the spot where +it had laid the first. It would, I think, be difficult to +distinguish this act from one performed by man by the +aid of reason.</p> + +<p>With respect to colour which so often differs in the +two sexes of animals belonging to the higher classes, +Mr. Spence Bate does not know of any well-marked +instances with our British crustaceans. In some cases, +however, the male and female differ slightly in tint, +but Mr. Bate thinks not more than may be accounted +for by their different habits of life, such as by the +male wandering more about and being thus more exposed +to the light. In a curious Bornean crab, which +inhabits sponges, Mr. Bate could always distinguish the +sexes by the male not having the epidermis so much +rubbed off. Dr. Power tried to distinguish by colour +the sexes of the species which inhabit the Mauritius, but +always failed, except with one species of Squilla, probably +the <i>S. stylifera</i>, the male of which is described as +being “of a beautiful blueish-green,” with some of the +appendages cherry-red, whilst the female is clouded +with brown and grey, “with the red about her much +less vivid than in the male.”<a name="FNanchor_419" id="FNanchor_419"></a><a href="#Footnote_419" class="fnanchor">419</a> In this case, we may +suspect the agency of sexual selection. With Saphirina +(an oceanic genus of Entomostraca, and therefore +low in the scale) the males are furnished with +<span class='pagenum'><a name="Page_336" id="Page_336">336</a></span>minute shields or cell-like bodies, which exhibit beautiful +changing colours; these being absent in the +females, and in the case of one species in both sexes.<a name="FNanchor_420" id="FNanchor_420"></a><a href="#Footnote_420" class="fnanchor">420</a> +It would, however, be extremely rash to conclude that +these curious organs serve merely to attract the females. +In the female of a Brazilian species of Gelasimus, the +whole body, as I am informed by Fritz Müller, is of a +nearly uniform greyish-brown. In the male the posterior +part of the cephalo-thorax is pure white, with the +anterior part of a rich green, shading into dark brown; +and it is remarkable that these colours are liable to +change in the course of a few minutes—the white +becoming dirty grey or even black, the green “losing +much of its brilliancy.” The males apparently are +much more numerous than the females. It deserves +especial notice that they do not acquire their bright +colours until they become mature. They differ also +from the females in the larger size of their chelæ. +In some species of the genus, probably in all, the +sexes pair and inhabit the same burrow. They are +also, as we have seen, highly intelligent animals. +From these various considerations it seems highly +probable that the male in this species has become +gaily ornamented in order to attract or excite the +female.</p> + +<p class="tb">It has just been stated that the male Gelasimus does +not acquire his conspicuous colours until mature and +nearly ready to breed. This seems the general rule in +the whole class with the many remarkable differences +in structure between the two sexes. We shall hereafter +find the same law prevailing throughout the great +sub-kingdom of the Vertebrata, and in all cases it is +eminently distinctive of characters which have been +<span class='pagenum'><a name="Page_337" id="Page_337">337</a></span>acquired through sexual selection. Fritz Müller<a name="FNanchor_421" id="FNanchor_421"></a><a href="#Footnote_421" class="fnanchor">421</a> gives +some striking instances of this law; thus the male +sand-hopper (Orchestia) does not acquire his large +claspers, which are very differently constructed from +those of the female, until nearly full-grown; whilst +young his claspers resemble those of the female. Thus, +again, the male Brachyscelus possesses, like all other +amphipods, a pair of posterior antennæ; the female, +and this is a most extraordinary circumstance, is destitute +of them, and so is the male as long as he remains +immature.</p> + +<p>Class, <i>Arachnida</i> (Spiders).—The males are often +darker, but sometimes lighter than the females, as may +be seen in Mr. Blackwall’s magnificent work.<a name="FNanchor_422" id="FNanchor_422"></a><a href="#Footnote_422" class="fnanchor">422</a> In +some species the sexes differ conspicuously from each +other in colour; thus the female of <i>Sparassus smaragdulus</i> +is dullish-green; whilst the adult male has +the abdomen of a fine yellow, with three longitudinal +stripes of rich red. In some species of Thomisus the +two sexes closely resemble each other; in others they +differ much; thus in <i>T. citreus</i> the legs and body of +the female are pale-yellow or green, whilst the front +legs of the male are reddish-brown: in <i>T. floricolens</i>, +the legs of the female are pale-green, those of the +male being ringed in a conspicuous manner with various +tints. Numerous analogous cases could be given in the +genera Epeira, Nephila, Philodromus, Theridion, Linyphia, +&c. It is often difficult to say which of the two +sexes departs most from the ordinary coloration of the +genus to which the species belong; but Mr. Blackwall +<span class='pagenum'><a name="Page_338" id="Page_338">338</a></span>thinks that, as a general rule, it is the male. Both +sexes whilst young, as I am informed by the same +author, usually resemble each other; and both often +undergo great changes in colour during their successive +moults before arriving at maturity. In other cases +the male alone appears to change colour. Thus the +male of the above-mentioned brightly-coloured Sparassus +at first resembles the female and acquires his +peculiar tints only when nearly adult. Spiders are +possessed of acute senses, and exhibit much intelligence. +The females often shew, as is well known, the +strongest affection for their eggs, which they carry +about enveloped in a silken web. On the whole it +appears probable that well-marked differences in colour +between the sexes have generally resulted from sexual +selection, either on the male or female side. But doubts +may be entertained on this head from the extreme +variability in colour of some species, for instance of +<i>Theridion lineatum</i>, the sexes of which differ when +adult; this great variability indicates that their colours +have not been subjected to any form of selection.</p> + +<p>Mr. Blackwall does not remember to have seen the +males of any species fighting together for the possession +of the female. Nor, judging from analogy, is this +probable; for the males are generally much smaller +than the females, sometimes to an extraordinary degree.<a name="FNanchor_423" id="FNanchor_423"></a><a href="#Footnote_423" class="fnanchor">423</a> +Had the males been in the habit of fighting +together, they would, it is probable, have gradually +<span class='pagenum'><a name="Page_339" id="Page_339">339</a></span>acquired greater size and strength. Mr. Blackwall has +sometimes seen two or more males on the same web +with a single female; but their courtship is too tedious +and prolonged an affair to be easily observed. The male +is extremely cautious in making his advances, as the +female carries her coyness to a dangerous pitch. De +Geer saw a male that “in the midst of his preparatory +caresses was seized by the object of his attractions, +enveloped by her in a web and then devoured, a +sight which, as he adds, filled him with horror and +indignation.”<a name="FNanchor_424" id="FNanchor_424"></a><a href="#Footnote_424" class="fnanchor">424</a></p> + +<p class="tb">Westring has made the interesting discovery that +the males of several species of Theridion<a name="FNanchor_425" id="FNanchor_425"></a><a href="#Footnote_425" class="fnanchor">425</a> have the +power of making a stridulating sound (like that made +by many beetles and other insects, but feebler), whilst +the females are quite mute. The apparatus consists of +a serrated ridge at the base of the abdomen, against +which the hard hinder part of the thorax is rubbed; +and of this structure not a trace could be detected in +the females. From the analogy of the Orthoptera and +Homoptera, to be described in the next chapter, we +may feel almost sure that the stridulation serves, as +Westring remarks, either to call or to excite the +female; and this is the first case in the ascending scale +of the animal kingdom, known to me, of sounds emitted +for this purpose.</p> + +<p>Class, <i>Myriapoda</i>.—In neither of the two orders in +this class, including the millipedes and centipedes, +<span class='pagenum'><a name="Page_340" id="Page_340">340</a></span>can I find any well-marked instances of sexual differences +such as more particularly concern us. In +<i>Glomeris limbata</i>, however, and perhaps in some few +other species, the males differ slightly in colour from +the females; but this Glomeris is a highly variable +species. In the males of the Diplopoda, the legs belonging +to one of the anterior segments of the body, or +to the posterior segment, are modified into prehensile +hooks which serve to secure the female. In some +species of Iulus the tarsi of the male are furnished +with membranous suckers for the same purpose. It is +a much more unusual circumstance, as we shall see +when we treat of Insects, that it is the female in +Lithobius which is furnished with prehensile appendages +at the extremity of the body for holding the +male.<a name="FNanchor_426" id="FNanchor_426"></a><a href="#Footnote_426" class="fnanchor">426</a></p> + +<hr /> + +<p><span class='pagenum'><a name="Page_341" id="Page_341">341</a></span></p> + +<h3>CHAPTER X.</h3> + +<h4><span class="smcap">Secondary Sexual Characters of Insects</span>.</h4> + +<div class="blockquot"><p>Diversified structures possessed by the males for seizing the females—Differences +between the sexes, of which the meaning is not +understood—Difference in size between the sexes—Thysanura—Diptera—Hemiptera—Homoptera, +musical powers possessed +by the males alone—Orthoptera, musical instruments of the +males, much diversified in structure; pugnacity; colours—Neuroptera, +sexual differences in colour—Hymenoptera, pugnacity +and colours—Coleoptera, colours; furnished with great horns, +apparently as an ornament; battles; stridulating organs generally +common to both sexes.</p></div> + +<p>In the immense class of insects the sexes sometimes +differ in their organs for locomotion, and often in +their sense-organs, as in the pectinated and beautifully +plumose antennæ of the males of many species. +In one of the Ephemeræ, namely Chloëon, the male +has great pillared eyes, of which the female is entirely +destitute.<a name="FNanchor_427" id="FNanchor_427"></a><a href="#Footnote_427" class="fnanchor">427</a> The ocelli are absent in the females of +certain other insects, as in the Mutillidæ, which are +likewise destitute of wings. But we are chiefly concerned +with structures by which one male is enabled to +conquer another, either in battle or courtship, through +his strength, pugnacity, ornaments, or music. The +innumerable contrivances, therefore, by which the male +is able to seize the female, may be briefly passed over. +Besides the complex structures at the apex of the abdomen, +which ought perhaps to be ranked as primary +<span class='pagenum'><a name="Page_342" id="Page_342">342</a></span>organs,<a name="FNanchor_428" id="FNanchor_428"></a><a href="#Footnote_428" class="fnanchor">428</a> “it is astonishing,” as Mr. B. D. Walsh<a name="FNanchor_429" id="FNanchor_429"></a><a href="#Footnote_429" class="fnanchor">429</a> has +remarked, “how many different organs are worked in +by nature, for the seemingly insignificant object of +enabling the male to grasp the female firmly.” The +mandibles or jaws are sometimes used for this purpose; +thus the male <i>Corydalis cornutus</i> (a neuropterous insect +in some degree allied to the Dragon-flies, &c.) has immense +curved jaws, many times longer than those of the +female; and they are smooth instead of being toothed, +by which means he is enabled to seize her without +injury.<a name="FNanchor_430" id="FNanchor_430"></a><a href="#Footnote_430" class="fnanchor">430</a> One of the stag-beetles of North America +(<i>Lucanus elaphus</i>) uses his jaws, which are much larger +than those of the female, for the same purpose, but +probably likewise for fighting. In one of the sand-wasps +(<i>Ammophila</i>) the jaws in the two sexes are closely +alike, but are used for widely different purposes; the +males, as Professor Westwood observes, “are exceedingly +ardent, seizing their partners round the neck +with their sickle-shaped jaws;”<a name="FNanchor_431" id="FNanchor_431"></a><a href="#Footnote_431" class="fnanchor">431</a> whilst the females use +<span class='pagenum'><a name="Page_343" id="Page_343">343</a></span>these organs for burrowing in sand-banks and making +their nests.</p> + +<p>The tarsi of the front-legs are dilated in many male +beetles, or are furnished with broad cushions of hairs; +and in many genera of water-beetles they are armed +with a round flat sucker, so that the male may adhere +to the slippery body of the female. It is a much more +unusual circumstance that the females of some water-beetles + +<span class="figright2" style="width: 300px;"><a name="f8" id="f8"></a><img src="images/fig8.png" width="300" height="496" +alt="Fig. 8. Crabro cribrarius. Upper figure, +male: lower figure, female." title="" /> + +<span class="indent2">Fig. 8. Crabro cribrarius. Upper figure, +male: lower figure, female.</span></span> + +(Dytiscus) have their +elytra deeply grooved, and +in <i>Acilius sulcatus</i> thickly set +with hairs, as an aid to the +male. The females of some +other water-beetles (Hydroporus) +have their elytra +punctured for the same object.<a name="FNanchor_432" id="FNanchor_432"></a><a href="#Footnote_432" class="fnanchor">432</a> +In the male of <i>Crabro +cribrarius</i> (fig. 8.), it is the +tibia which is dilated into a +broad horny plate, with minute +membraneous dots, giving +to it a singular appearance +like that of a riddle.<a name="FNanchor_433" id="FNanchor_433"></a><a href="#Footnote_433" class="fnanchor">433</a> +In the male of Penthe (a +genus of beetles) a few of +the middle joints of the antennæ +are dilated and furnished on the inferior surface +<span class='pagenum'><a name="Page_344" id="Page_344">344</a></span>with cushions of hair, exactly like those on the tarsi of +the Carabidæ, “and obviously for the same end.” In +male dragon-flys, “the appendages at the tip of the tail +are modified in an almost infinite variety of curious +patterns to enable them to embrace +the neck of the female.” Lastly in +the males of many insects, the legs +are furnished with peculiar spines, +knobs or spurs; or the whole leg is +bowed or thickened, but this is by +no means invariably a sexual character; + +<span class="figleft2" style="width: 200px;"><a name="f9" id="f9"></a><img src="images/fig9.png" width="200" height="798" +alt="Fig. 9. Taphroderes distortus +(much enlarged). Upper +figure, male; lower figure, +female." title="" /> + +<span class="indent2">Fig. 9. Taphroderes distortus +(much enlarged). Upper +figure, male; lower figure, +female.</span></span> + +or one pair, or all three +pairs are elongated, sometimes to +an extravagant length.<a name="FNanchor_434" id="FNanchor_434"></a><a href="#Footnote_434" class="fnanchor">434</a></p> + +<p>In all the orders, the sexes of +many species present differences, of +which the meaning is not understood. +One curious case is that of +a beetle (fig. 9), the male of which +has the left mandible much enlarged; +so that the mouth is greatly +distorted. In another Carabidous +beetle, the Eurygnathus,<a name="FNanchor_435" id="FNanchor_435"></a><a href="#Footnote_435" class="fnanchor">435</a> we have +the unique case, as far as known to +Mr. Wollaston, of the head of the +female being much broader and +larger, though in a variable degree, +than that of the male. Any number +of such cases could be given. They +abound in the Lepidoptera: one +of the most extraordinary is that +certain male butterflies have their fore-legs more or +<span class='pagenum'><a name="Page_345" id="Page_345">345</a></span>less atrophied, with the tibiæ and tarsi reduced to mere +rudimentary knobs. The wings, also, in the two sexes +often differ in neuration,<a name="FNanchor_436" id="FNanchor_436"></a><a href="#Footnote_436" class="fnanchor">436</a> and sometimes considerably +in outline, as in the <i>Aricoris epitus</i>, which was shown +to me in the British Museum by Mr. A. Butler. The +males of certain South American butterflies have tufts +of hair on the margins of the wings, and horny excrescences +on the discs of the posterior pair.<a name="FNanchor_437" id="FNanchor_437"></a><a href="#Footnote_437" class="fnanchor">437</a> In several +British butterflies, the males alone, as shewn by Mr. +Wonfor, are in parts clothed with peculiar scales.</p> + +<p class="tb">The purpose of the luminosity in the female glow-worm +is likewise not understood; for it is very doubtful +whether the primary use of the light is to guide the +male to the female. It is no serious objection to this +latter belief that the males emit a feeble light; for +secondary sexual characters proper to one sex are often +developed in a slight degree in the other sex. It is a +more valid objection that the larvæ shine, and in some +species brilliantly: Fritz Müller informs me that the +most luminous insect which he ever beheld in Brazil, +was the larva of some beetle. Both sexes of certain +luminous species of Elater emit light. Kirby and +Spence suspect that the phosphorescence serves to +frighten and drive away enemies.</p> + +<p class="tb"><i>Difference in Size between the Sexes.</i>—With insects +of all kinds the males are commonly smaller than the +females;<a name="FNanchor_438" id="FNanchor_438"></a><a href="#Footnote_438" class="fnanchor">438</a> and this difference can often be detected +even in the larval state. So considerable is the difference +<span class='pagenum'><a name="Page_346" id="Page_346">346</a></span>between the male and female cocoons of the silk-moth +(<i>Bombyx mori</i>), that in France they are separated by +a particular mode of weighing.<a name="FNanchor_439" id="FNanchor_439"></a><a href="#Footnote_439" class="fnanchor">439</a> In the lower classes +of the animal kingdom, the greater size of the females +seems generally to depend on their developing an enormous +number of ova; and this may to a certain extent +hold good with insects. But Dr. Wallace has suggested +a much more probable explanation. He finds, after +carefully attending to the development of the caterpillars +of <i>Bombyx cynthia</i> and <i>Yamamai</i>, and especially +of some dwarfed caterpillars reared from a second brood +on unnatural food, “that in proportion as the individual +moth is finer, so is the time required for its +metamorphosis longer; and for this reason the female, +which is the larger and heavier insect, from having to +carry her numerous eggs, will be preceded by the +male, which is smaller and has less to mature.”<a name="FNanchor_440" id="FNanchor_440"></a><a href="#Footnote_440" class="fnanchor">440</a> Now +as most insects are short-lived, and as they are exposed +to many dangers, it would manifestly be advantageous +to the female to be impregnated as soon as possible. +This end would be gained by the males being first +matured in large numbers ready for the advent of the +females; and this again would naturally follow, as +Mr. A. E. Wallace has remarked,<a name="FNanchor_441" id="FNanchor_441"></a><a href="#Footnote_441" class="fnanchor">441</a> through natural +selection; for the smaller males would be first matured, +and thus would procreate a large number of offspring +which would inherit the reduced size of their male +parents, whilst the larger males from being matured +later would leave fewer offspring.</p> + +<p>There are, however, exceptions to the rule of male +insects being smaller than the females; and some of +<span class='pagenum'><a name="Page_347" id="Page_347">347</a></span>these exceptions are intelligible. Size and strength +would be an advantage to the males, which fight for the +possession of the female; and in these cases the males, +as with the stag-beetle (Lucanus), are larger than the +females. There are, however, other beetles which are +not known to fight together, of which the males exceed +the females in size; and the meaning of this fact is not +known; but in some of these cases, as with the huge +Dynastes and Megasoma, we can at least see that there +would be no necessity for the males to be smaller than +the females, in order to be matured before them, for +these beetles are not short-lived, and there would be +ample time for the pairing of the sexes. So, again, +male dragon-flies (Libellulidæ) are sometimes sensibly +larger, and never smaller, than the females;<a name="FNanchor_442" id="FNanchor_442"></a><a href="#Footnote_442" class="fnanchor">442</a> and +they do not, as Mr. MacLachlan believes, generally +pair with the females, until a week or fortnight has +elapsed, and until they have assumed their proper +masculine colours. But the most curious case, shewing +on what complex and easily-overlooked relations, so +trifling a character as a difference in size between the +sexes may depend, is that of the aculeate Hymenoptera; +for Mr. F. Smith informs me that throughout nearly +the whole of this large group the males, in accordance +with the general rule, are smaller than the +females and emerge about a week before them; but +amongst the Bees, the males of <i>Apis mellifica</i>, <i>Anthidium +manicatum</i> and <i>Anthophora acervorum</i>, and amongst the +Fossores, the males of the <i>Methoca ichneumonides</i>, are +larger than the females. The explanation of this anomaly +is that a marriage-flight is absolutely necessary +<span class='pagenum'><a name="Page_348" id="Page_348">348</a></span>with these species, and the males require great strength +and size in order to carry the females through the air. +Increased size has here been acquired in opposition to +the usual relation between size and the period of development, +for the males, though larger, emerge before +the smaller females.</p> + +<p class="tb">We will now review the several Orders, selecting such +facts as more particularly concern us. The Lepidoptera +(Butterflies and Moths) will be retained for a separate +chapter.</p> + +<p class="tb">Order, <i>Thysanura</i>.—The members of this Order are +lowly organised for their class. They are wingless, +dull-coloured, minute insects, with ugly, almost misshapen +heads and bodies. The sexes do not differ; but +they offer one interesting fact, by showing that the males +pay sedulous court to their females even low down in the +animal scale. Sir J. Lubbock<a name="FNanchor_443" id="FNanchor_443"></a><a href="#Footnote_443" class="fnanchor">443</a> in describing the <i>Smynthurus +luteus</i>, says: “it is very amusing to see these +little creatures coquetting together. The male, which +is much smaller than the female, runs round her, and +they butt one another, standing face to face, and +moving backward and forward like two playful lambs. +Then the female pretends to run away and the male +runs after her with a queer appearance of anger, gets +in front and stands facing her again; then she turns +coyly round, but he, quicker and more active, scuttles +round too, and seems to whip her with his antennæ; +then for a bit they stand face to face, play with their +antennæ, and seem to be all in all to one another.”</p> + +<p class="tb">Order, <i>Diptera</i> (Flies).—The sexes differ little in +colour. The greatest difference, known to Mr. F. Walker, +<span class='pagenum'><a name="Page_349" id="Page_349">349</a></span>is in the genus Bibio, in which the males are blackish +or quite black, and the females obscure brownish-orange. +The genus Elaphomyia, discovered by Mr. Wallace<a name="FNanchor_444" id="FNanchor_444"></a><a href="#Footnote_444" class="fnanchor">444</a> in +New Guinea, is highly remarkable, as the males are +furnished with horns, of which the females are quite +destitute. The horns spring from beneath the eyes, and +curiously resemble those of stags, being either branched +or palmated. They equal in length the whole of the +body in one of the species. They might be thought +to serve for fighting, but as in one species they are +of a beautiful pink colour, edged with black, with a +pale central stripe, and as these insects have altogether +a very elegant appearance, it is perhaps more probable +that the horns serve as ornaments. That the +males of some Diptera fight together is certain; for +Prof. Westwood<a name="FNanchor_445" id="FNanchor_445"></a><a href="#Footnote_445" class="fnanchor">445</a> has several times seen this with some +species of Tipula or Harry-long-legs. Many observers +believe that when gnats (Culicidæ) dance in the air in +a body, alternately rising and falling, the males are +courting the females. The mental faculties of the +Diptera are probably fairly well developed, for their +nervous system is more highly developed than in most +other Orders of insects.<a name="FNanchor_446" id="FNanchor_446"></a><a href="#Footnote_446" class="fnanchor">446</a></p> + +<p>Order, <i>Hemiptera</i> (Field-Bugs).—Mr. J. W. Douglas, +who has particularly attended to the British species, has +kindly given me an account of their sexual differences. +The males of some species are furnished with wings, +whilst the females are wingless; the sexes differ in the +form of the body and elytra; in the second joints of +their antennæ and in their tarsi; but as the signification +<span class='pagenum'><a name="Page_350" id="Page_350">350</a></span>of these differences is quite unknown, they may be here +passed over. The females are generally larger and more +robust than the males. With British, and, as far as +Mr. Douglas knows, with exotic species, the sexes do +not commonly differ much in colour; but in about six +British species the male is considerably darker than the +female, and in about four other species the female is +darker than the male. Both sexes of some species are +beautifully marked with vermilion and black. It is +doubtful whether these colours serve as a protection. +If in any species the males had differed from the females +in an analogous manner, we might have been justified +in attributing such conspicuous colours to sexual selection +with transference to both sexes.</p> + +<p class="tb">Some species of Reduvidæ make a stridulating noise; +and, in the case of <i>Pirates stridulus</i>, this is said<a name="FNanchor_447" id="FNanchor_447"></a><a href="#Footnote_447" class="fnanchor">447</a> to +be effected by the movement of the neck within the +pro-thoracic cavity. According to Westring, <i>Reduvius +personatus</i> also stridulates. But I have not been able +to learn any particulars about these insects; nor have I +any reason to suppose that they differ sexually in this +respect.</p> + +<p>Order, <i>Homoptera.</i>—Every one who has wandered in +a tropical forest must have been astonished at the din +made by the male Cicadæ. The females are mute; +as the Grecian poet Xenarchus says, “Happy the +Cicadas live, since they all have voiceless wives.” +The noise thus made could be plainly heard on board +the “Beagle,” when anchored at a quarter of a mile +from the shore of Brazil; and Captain Hancock says +it can be heard at the distance of a mile. The Greeks +formerly kept, and the Chinese now keep, these insects +<span class='pagenum'><a name="Page_351" id="Page_351">351</a></span>in cages for the sake of their song, so that it must be +pleasing to the ears of some men.<a name="FNanchor_448" id="FNanchor_448"></a><a href="#Footnote_448" class="fnanchor">448</a> The Cicadidæ +usually sing during the day; whilst the Fulgoridæ +appear to be night-songsters. The sound, according +to Landois,<a name="FNanchor_449" id="FNanchor_449"></a><a href="#Footnote_449" class="fnanchor">449</a> who has recently studied the subject, is +produced by the vibration of the lips of the spiracles, +which are set into motion by a current of air emitted +from the tracheæ. It is increased by a wonderfully +complex resounding apparatus, consisting of two cavities +covered by scales. Hence the sound may truly +be called a voice. In the female the musical apparatus +is present, but very much less developed than in the +male, and is never used for producing sound.</p> + +<p>With respect to the object of the music, Dr. Hartman +in speaking of the <i>Cicada septemdecim</i> of the United +States, says,<a name="FNanchor_450" id="FNanchor_450"></a><a href="#Footnote_450" class="fnanchor">450</a> “the drums are now (June 6th and 7th, +1851) heard in all directions. This I believe to be the +marital summons from the males. Standing in thick +chestnut sprouts about as high as my head, where +hundreds were around me, I observed the females +coming around the drumming males.” He adds, “this +season (Aug. 1868) a dwarf pear-tree in my garden +produced about fifty larvæ of <i>Cic. pruinosa</i>; and I +several times noticed the females to alight near a +male while he was uttering his clanging notes.” Fritz +Müller writes to me from S. Brazil that he has often +listened to a musical contest between two or three +males of a Cicada, having a particularly loud voice, and +seated at a considerable distance from each other. As +<span class='pagenum'><a name="Page_352" id="Page_352">352</a></span>soon as the first had finished his song, a second immediately +began; and after he had concluded, another +began, and so on. As there is so much rivalry between +the males, it is probable that the females not only discover +them by the sounds emitted, but that, like female +birds, they are excited or allured by the male with the +most attractive voice.</p> + +<p class="tb">I have not found any well-marked cases of ornamental +differences between the sexes of the Homoptera. +Mr. Douglas informs me that there are three British +species, in which the male is black or marked with black +bands, whilst the females are pale-coloured or obscure.</p> + +<p>Order, <i>Orthoptera</i>.—The males in the three saltatorial +families belonging to this Order are remarkable +for their musical powers, namely the Achetidæ or +crickets, the Locustidæ for which there is no exact +equivalent name in English, and the Acridiidæ or grasshoppers. +The stridulation produced by some of the +Locustidæ is so loud that it can be heard during the +night at the distance of a mile;<a name="FNanchor_451" id="FNanchor_451"></a><a href="#Footnote_451" class="fnanchor">451</a> and that made by +certain species is not unmusical even to the human +ear, so that the Indians on the Amazons keep them +in wicker cages. All observers agree that the sounds +serve either to call or excite the mute females. But it +has been noticed<a name="FNanchor_452" id="FNanchor_452"></a><a href="#Footnote_452" class="fnanchor">452</a> that the male migratory locust of +Russia (one of the Acridiidæ) whilst coupled with the +female, stridulates from anger or jealousy when approached +by another male. The house-cricket when +surprised at night uses its voice to warn its fellows.<a name="FNanchor_453" id="FNanchor_453"></a><a href="#Footnote_453" class="fnanchor">453</a> In +North America the Katy-did (<i>Platyphyllum concavum</i>, +<span class='pagenum'><a name="Page_353" id="Page_353">353</a></span>one of the Locustidæ) is described<a name="FNanchor_454" id="FNanchor_454"></a><a href="#Footnote_454" class="fnanchor">454</a> as mounting on the +upper branches of a tree, and in the evening beginning +“his noisy babble, while rival notes issue from the neighbouring +trees, and the groves resound with the call of +<i>Katy-did-she-did</i>, the live-long night.” Mr. Bates, in +speaking of the European field-cricket (one of the Achetidæ), +says, “the male has been observed to place itself +in the evening at the entrance of its burrow, and +stridulate until a female approaches, when the louder +notes are succeeded by a more subdued tone, whilst +the successful musician caresses with his antennæ + +<span class="figright2" style="width: 250px;"><a name="f10" id="f10"></a><img src="images/fig10.png" width="250" height="250" +alt="Fig. 10. Gryllus campestris (from Landois)." title="" /> + +<span class="indent2">Fig. 10. Gryllus campestris (from Landois).<br /><br /></span> + +<span class="indent2">Right-hand figure, under side of part of the +wing-nervure, much magnified, showing +the teeth, <i>st</i>.<br /></span> + +<span class="indent2">Left-hand figure, upper surface of wing-cover, +with the projecting, smooth nervure, <i>r.</i>, +across which the teeth (<i>st</i>) are scraped.</span></span> + +the mate he has won.”<a name="FNanchor_455" id="FNanchor_455"></a><a href="#Footnote_455" class="fnanchor">455</a> +Dr. Scudder was able to +excite one of these insects +to answer him, by rubbing +on a file with a quill.<a name="FNanchor_456" id="FNanchor_456"></a><a href="#Footnote_456" class="fnanchor">456</a> +In both sexes a remarkable +auditory apparatus +has been discovered by +Von Siebold, situated in +the front legs.<a name="FNanchor_457" id="FNanchor_457"></a><a href="#Footnote_457" class="fnanchor">457</a></p> + +<p>In the three Families +the sounds are differently +produced. In the males Of +the Achetidæ both wing-covers +have the same +structure; and this in the +field-cricket (<i>Gryllus campestris</i>, fig. 10) consists, as de<span class='pagenum'><a name="Page_354" id="Page_354">354</a></span>scribed +by Landois,<a name="FNanchor_458" id="FNanchor_458"></a><a href="#Footnote_458" class="fnanchor">458</a> of from 131 to 138 sharp, transverse +ridges or teeth (<i>st</i>) on the under side of one of the +nervures of the wing-cover. This toothed nervure is +rapidly scraped across a projecting, smooth, hard nervure +(<i>r</i>) on the upper surface of the opposite wing. First + +<span class="figleft2" style="width: 150px;"><a name="f11" id="f11"></a><img src="images/fig11.png" width="150" height="269" +alt="Fig. 11. Teeth of Nervure +of Gryllus domesticus +(from Landois)." title="" /> + +<span class="indent2">Fig. 11. Teeth of Nervure +of Gryllus domesticus +(from Landois).</span></span> + +one wing is rubbed over the other, +and then the movement is reversed. +Both wings are raised a little at the +same time, so as to increase the resonance. +In some species the wing-covers +of the males are furnished at +the base with a talc-like plate.<a name="FNanchor_459" id="FNanchor_459"></a><a href="#Footnote_459" class="fnanchor">459</a> I +have here given a drawing (fig. 11) +of the teeth on the under side of the +nervure of another species of Gryllus, +viz. <i>G. domesticus</i>.</p> + +<p>In the Locustidæ the opposite wing-covers differ in +structure (fig. 12), and cannot, as in the last family, +be indifferently used in a reversed manner. The left +wing, which acts as the bow of the fiddle, lies over the +right wing which serves as the fiddle itself. One of +the nervures (<i>a</i>) on the under surface of the former is +finely serrated, and is scraped across the prominent +nervures on the upper surface of the opposite or right +wing. In our British <i>Phasgonura viridissima</i> it appeared +to me that the serrated nervure is rubbed +against the rounded hind corner of the opposite wing, +the edge of which is thickened, coloured brown, and +very sharp. In the right wing, but not in the left, +there is a little plate, as transparent as talc, surrounded +by nervures, and called the speculum. In <i>Ephippiger +vitium</i>, a member of this same family, we have a curious +<span class='pagenum'><a name="Page_355" id="Page_355">355</a></span>subordinate modification; for the wing-covers are greatly +reduced in size, but “the posterior part of the pro-thorax +is elevated into a kind of dome over the wing-covers, +and which has probably the effect of increasing the +sound.”<a name="FNanchor_460" id="FNanchor_460"></a><a href="#Footnote_460" class="fnanchor">460</a></p> + +<div class="figcenter" style="width: 550px;"><a name="f12" id="f12"></a><img src="images/fig12.png" width="550" height="454" +alt="Fig. 12. Chlorocœlus Tanana (from Bates)." title="" /> + +<p class="center2">Fig. 12. Chlorocœlus Tanana (from Bates), <i>a</i>, <i>b</i>. Lobes of opposite wing-covers.</p></div> + +<p>We thus see that the musical apparatus is more +differentiated or specialised in the Locustidæ, which +includes I believe the most powerful performers in +the Order, than in the Achetidæ, in which both wing-covers +have the same structure and the same function.<a name="FNanchor_461" id="FNanchor_461"></a><a href="#Footnote_461" class="fnanchor">461</a> +Landois, however, detected in one of the Locustidæ, +namely in Decticus, a short and narrow row of small +<span class='pagenum'><a name="Page_356" id="Page_356">356</a></span>teeth, mere rudiments, on the inferior surface of the +right wing-cover, which underlies the other and is +never used as the bow. I observed the same rudimentary +structure on the under side of the right wing-cover +in <i>Phasgonura viridissima</i>. Hence we may with +confidence infer that the Locustidæ are descended from +a form, in which, as in the existing Achetidæ, both +wing-covers had serrated nervures on the under surface, +and could be indifferently used as the bow; but that +in the Locustidæ the two wing-covers gradually became +differentiated and perfected, on the principle of the division +of labour, the one to act exclusively as the bow and +the other as the fiddle. By what steps the more simple +apparatus in the Achetidæ originated, we do not know, +but it is probable that the basal portions of the wing-covers +overlapped each other formerly as at present, and +that the friction of the nervures produced a grating +sound, as I find is now the case with the wing-covers +of the females.<a name="FNanchor_462" id="FNanchor_462"></a><a href="#Footnote_462" class="fnanchor">462</a> A grating sound thus occasionally +and accidentally made by the males, if it served them +ever so little as a love-call to the females, might readily +have been intensified through sexual selection by fitting +variations in the roughness of the nervures having been +continually preserved.</p> + +<p>In the last and third Family, namely the Acridiidæ +or grasshoppers, the stridulation is produced in a very +different manner, and is not so shrill, according to Dr. +Scudder, as in the preceding Families. The inner surface +of the femur (fig. 13, <i>r</i>) is furnished with a longitudinal +row of minute, elegant, lancet-shaped, elastic +teeth, from 85 to 93 in number;<a name="FNanchor_463" id="FNanchor_463"></a><a href="#Footnote_463" class="fnanchor">463</a> and these are scraped +<span class='pagenum'><a name="Page_357" id="Page_357">357</a></span>across the sharp, projecting nervures on the wing-covers, +which, are thus made to vibrate and resound. Harris<a name="FNanchor_464" id="FNanchor_464"></a><a href="#Footnote_464" class="fnanchor">464</a> + +<span class="figright" style="width: 300px;"><a name="f13" id="f13"></a><img src="images/fig13.png" width="300" height="289" +alt="Fig. 13, Hind-leg of Stenobothrus pratorum." title="" /> + +<span class="indent2">Fig. 13, Hind-leg of Stenobothrus pratorum: +<i>r</i>, the stridulating ridge; lower figure, the +teeth, forming the ridge, much magnified +(from Landois).</span></span> + +says that when one of +the males begins to play, +he first “bends the shank +of the hind-leg beneath, +the thigh, where it is +lodged in a furrow designed +to receive it, +and then draws the leg +briskly up and down. +He does not play both +fiddles together, but alternately +first upon one +and then on the other.” +In many species, the base +of the abdomen is hollowed out into a great cavity +which is believed to act as a resounding board. In +Pneumora (fig. 14), a S. African genus belonging to +this same family, we meet with a new and remarkable +modification: in the males a small notched ridge projects +obliquely from each side of the abdomen, against +which the hind femora are rubbed.<a name="FNanchor_465" id="FNanchor_465"></a><a href="#Footnote_465" class="fnanchor">465</a> As the male is +furnished with wings, the female being wingless, it is +remarkable that the thighs are not rubbed in the usual +manner against the wing-covers; but this may perhaps +be accounted for by the unusually small size of the hind-legs. +I have not been able to examine the inner +surface of the thighs, which, judging from analogy, +would be finely serrated. The species of Pneumora +have been more profoundly modified for the sake of +stridulation than any other orthopterous insect; for +<span class='pagenum'><a name="Page_358" id="Page_358">358</a></span>in the male the whole body has been converted into a +musical instrument, being distended with air, like a +great pellucid bladder, so as to increase the resonance. +Mr. Trimen informs me that at the Cape of Good Hope +these insects make a wonderful noise during the night +There is one exception to the rule that the females +in these three Families are destitute of an efficient +musical apparatus; for both sexes of Ephippiger (Locustidæ) +are said<a name="FNanchor_466" id="FNanchor_466"></a><a href="#Footnote_466" class="fnanchor">466</a> to be thus provided. This case may +<span class='pagenum'><a name="Page_359" id="Page_359">359</a></span>be compared with that of the reindeer, in which species +alone both sexes possess horns. Although the female +orthoptera are thus almost invariably mute, yet Landois<a name="FNanchor_467" id="FNanchor_467"></a><a href="#Footnote_467" class="fnanchor">467</a> +found rudiments of the stridulating organs on the femora +of the female Acridiidæ, and similar rudiments on the +under surface of the wing-covers of the female Achetidæ; +but he failed to find any rudiments in the females +of Decticus, one of the Locustidæ. In the Homoptera +the mute females of Cicada, have the proper musical +apparatus in an undeveloped state; and we shall hereafter +meet in other divisions of the animal kingdom with +innumerable instances of structures proper to the male +being present in a rudimentary condition in the female. +Such cases appear at first sight to indicate that both +sexes were primordially constructed in the same manner, +but that certain organs were subsequently lost by the +females. It is, however, a more probable view, as previously +explained, that the organs in question were +acquired by the males and partially transferred to the +females.</p> + +<div class="figcenter" style="width: 450px;"><a name="f14" id="f14"></a><img src="images/fig14.png" width="450" height="492" +alt="Fig. 14. Pneumora (from specimens in the British Museum)." title="" /> + +<p class="center2">Fig. 14. Pneumora (from specimens in the British Museum). Upper figure, male; +lower figure, female.</p></div> + +<p>Landois has observed another interesting fact, namely +that in the females of the Acridiidæ, the stridulating +teeth on the femora remain throughout life in the same +condition in which they first appear in both sexes +during the larval state. In the males, on the other +hand, they become fully developed and acquire their +perfect structure at the last moult, when the insect is +mature and ready to breed.</p> + +<p>From the facts now given, we see that the means +by which the males produce their sounds are extremely +diversified in the Orthoptera, and are altogether different +from those employed by the Homoptera. But +throughout the animal kingdom we incessantly find the +<span class='pagenum'><a name="Page_360" id="Page_360">360</a></span>same object gained by the most diversified means; this +being due to the whole organisation undergoing in +the course of ages multifarious changes; and as part +after part varies, different variations are taken advantage +of for the same general purpose. The diversification of +the means for producing sound in the three families +of the Orthoptera and in the Homoptera, impresses the +mind with the high importance of these structures to +the males, for the sake of calling or alluring the females. +We need feel no surprise at the amount of modification +which the Orthoptera have undergone in this respect, as +we now know, from Dr. Scudder’s remarkable discovery,<a name="FNanchor_468" id="FNanchor_468"></a><a href="#Footnote_468" class="fnanchor">468</a> +that there has been more than ample time. This +naturalist has lately found a fossil insect in the Devonian +formation of New Brunswick, which is furnished with +“the well-known tympanum or stridulating apparatus +of the male Locustidæ.” This insect, though in most +respects related to the Neuroptera, appears to connect, +as is so often the case with very ancient forms, the +two Orders of the Neuroptera and Orthoptera which are +now generally ranked as quite distinct.</p> + +<p class="tb">I have but little more to say on the Orthoptera. +Some of the species are very pugnacious: when two +male field-crickets (<i>Gryllus campestris</i>) are confined +together, they fight till one kills the other; and the +species of Mantis are described as manœuvring with +their sword-like front-limbs, like hussars with their +sabres. The Chinese keep these insects in little bamboo +cages and match them like game-cocks.<a name="FNanchor_469" id="FNanchor_469"></a><a href="#Footnote_469" class="fnanchor">469</a> With respect +to colour, some exotic locusts are beautifully ornamented; +the posterior wings being marked with red, +<span class='pagenum'><a name="Page_361" id="Page_361">361</a></span>blue, and black; but as throughout the Order the +two sexes rarely differ much in colour, it is doubtful +whether they owe these bright tints to sexual selection. +Conspicuous colours may be of use to these insects +as a protection, on the principle to be explained in the +next chapter, by giving notice to their enemies that +they are unpalatable. Thus it has been observed<a name="FNanchor_470" id="FNanchor_470"></a><a href="#Footnote_470" class="fnanchor">470</a> +that an Indian brightly-coloured locust was invariably +rejected when offered to birds and lizards. Some cases, +however, of sexual differences in colour in this Order +are known. The male of an American cricket<a name="FNanchor_471" id="FNanchor_471"></a><a href="#Footnote_471" class="fnanchor">471</a> is described +as being as white as ivory, whilst the female +varies from almost white to greenish-yellow or dusky. +Mr. Walsh informs me that the adult male of <i>Spectrum +femoratum</i> (one of the Phasmidæ) “is of a shining +brownish-yellow colour; the adult female being of +a dull, opaque, cinereous-brown; the young of both +sexes being green.” Lastly, I may mention that the +male of one curious kind of cricket<a name="FNanchor_472" id="FNanchor_472"></a><a href="#Footnote_472" class="fnanchor">472</a> is furnished with +“a long membranous appendage, which falls over the +face like a veil;” but whether this serves as an ornament +is not known.</p> + +<p>Order, <i>Neuroptera</i>.—Little need here be said, except +in regard to colour. In the Ephemeridæ the sexes +often differ slightly in their obscure tints;<a name="FNanchor_473" id="FNanchor_473"></a><a href="#Footnote_473" class="fnanchor">473</a> but it is +not probable that the males are thus rendered attractive +to the females. The Libellulidæ or dragon-flies +are ornamented with splendid green, blue, yellow, and +<span class='pagenum'><a name="Page_362" id="Page_362">362</a></span>vermilion metallic tints; and the sexes often differ. +Thus, the males of some of the Agrionidæ, as Prof. +Westwood remarks<a name="FNanchor_474" id="FNanchor_474"></a><a href="#Footnote_474" class="fnanchor">474</a> “are of a rich blue with black +wings, whilst the females are fine green with colourless +wings.” But in <i>Agrion Ramburii</i> these colours are +exactly reversed in the two sexes.<a name="FNanchor_475" id="FNanchor_475"></a><a href="#Footnote_475" class="fnanchor">475</a> In the extensive +N. American genus of Hetærina, the males alone have +a beautiful carmine spot at the base of each wing. In +<i>Anax junius</i> the basal part of the abdomen in the male +is a vivid ultra-marine blue, and in the female grass-green. +In the allied genus Gomphus, on the other +hand, and in some other genera, the sexes differ but +little in colour. Throughout the animal kingdom, +similar cases of the sexes of closely-allied forms either +differing greatly, or very little, or not at all, are of +frequent occurrence. Although with many Libellulidæ +there is so wide a difference in colour between the sexes, +it is often difficult to say which is the most brilliant; +and the ordinary coloration of the two sexes is exactly +reversed, as we have just seen, in one species of Agrion. +It is not probable that their colours in any case have +been gained as a protection. As Mr. MacLachlan, who +has closely attended to this family, writes to me, dragon-flies—the +tyrants of the insect-world—are the least +liable of any insect to be attacked by birds or other +enemies. He believes that their bright colours serve +as a sexual attraction. It deserves notice, as bearing +on this subject, that certain dragon-flies appear to be +attracted by particular colours: Mr. Patterson observed<a name="FNanchor_476" id="FNanchor_476"></a><a href="#Footnote_476" class="fnanchor">476</a> +that the species of Agrionidæ, of which the males are +blue, settled in numbers on the blue float of a fishing +<span class='pagenum'><a name="Page_363" id="Page_363">363</a></span>line; whilst two other species were attracted by shining +white colours.</p> + +<p>It is an interesting fact, first observed by Schelver, +that the males, in several genera belonging to two sub-families, +when they first emerge from the pupal state +are coloured exactly like the females; but that their +bodies in a short time assume a conspicuous milky-blue +tint, owing to the exudation of a kind of oil, soluble in +ether and alcohol. Mr. MacLachlan believes that in the +male of <i>Libellula depressa</i> this change of colour does not +occur until nearly a fortnight after the metamorphosis, +when the sexes are ready to pair.</p> + +<p>Certain species of Neurothemis present, according +to Brauer<a name="FNanchor_477" id="FNanchor_477"></a><a href="#Footnote_477" class="fnanchor">477</a> a curious case of dimorphism, some of the +females having their wings netted in the usual manner; +whilst other females have them “very richly netted as in +the males of the same species.” Brauer “explains +the phenomenon on Darwinian principles by the +supposition that the close netting of the veins is a +secondary sexual character in the males.” This +latter character is generally developed in the males alone, +but being, like every other masculine character, +latent in the female, is occasionally developed in them. +We have here an illustration of the manner in which +the two sexes of many animals have probably come to +resemble each other, namely by variations first appearing +in the males, being preserved in them, and then +transmitted to and developed in the females; but in +this particular genus a complete transference is occasionally +and abruptly effected. Mr. MacLachlan informs +me of another case of dimorphism occurring in +several species of Agrion in which a certain number of +individuals are found of an orange colour, and these are +<span class='pagenum'><a name="Page_364" id="Page_364">364</a></span>invariably females. This is probably a case of reversion, +for in the true Libellulæ, when the sexes differ in +colour, the females are always orange or yellow, so +that supposing Agrion to be descended from some primordial +form having the characteristic sexual colours +of the typical Libellulæ, it would not be surprising that +a tendency to vary in this manner should occur in the +females alone.</p> + +<p class="tb">Although many dragon-flies are such large, powerful, +and fierce insects, the males have not been observed +by Mr. MacLachlan to fight together, except, as he +believes, in the case of some of the smaller species of +Agrion. In another very distinct group in this Order, +namely in the Termites or white ants, both sexes at +the time of swarming may be seen running about, “the +male after the female, sometimes two chasing one +female, and contending with great eagerness who shall +win the prize.”<a name="FNanchor_478" id="FNanchor_478"></a><a href="#Footnote_478" class="fnanchor">478</a></p> + +<p>Order, <i>Hymenoptera</i>.—That inimitable observer, M. +Fabre,<a name="FNanchor_479" id="FNanchor_479"></a><a href="#Footnote_479" class="fnanchor">479</a> in describing the habits of Cerceris, a wasp-like +insect, remarks that “fights frequently ensue +between the males for the possession of some particular +female, who sits an apparently unconcerned +beholder of the struggle for supremacy, and when the +victory is decided, quietly flies away in company +with the conqueror.” Westwood<a name="FNanchor_480" id="FNanchor_480"></a><a href="#Footnote_480" class="fnanchor">480</a> says that the +males of one of the saw-flies (Tenthredinæ) “have been +found fighting together, with their mandibles locked.” +As M. Fabre speaks of the males of Cerceris striving +to obtain a particular female, it may be well to bear in +<span class='pagenum'><a name="Page_365" id="Page_365">365</a></span>mind that insects belonging to this Order have the +power of recognising each other after long intervals of +time, and are deeply attached. For instance, Pierre +Huber, whose accuracy no one doubts, separated some +ants, and when after an interval of four months they +met others which had formerly belonged to the same +community, they mutually recognised and caressed each +other with their antennæ. Had they been strangers +they would have fought together. Again, when two +communities engage in a battle, the ants on the same +side in the general confusion sometimes attack each +other, but they soon perceive their mistake, and the +one ant soothes the other.<a name="FNanchor_481" id="FNanchor_481"></a><a href="#Footnote_481" class="fnanchor">481</a></p> + +<p>In this Order slight differences in colour, according +to sex, are common, but conspicuous differences are +rare except in the family of Bees; yet both sexes of +certain groups are so brilliantly coloured—for instance +in Chrysis, in which vermilion and metallic greens +prevail—that we are tempted to attribute the result +to sexual selection. In the Ichneumonidæ, according to +Mr. Walsh,<a name="FNanchor_482" id="FNanchor_482"></a><a href="#Footnote_482" class="fnanchor">482</a> the males are almost universally lighter +coloured than the females. On the other hand, in the +Tenthredinidæ the males are generally darker than the +females. In the Siricidæ the sexes frequently differ; +thus the male of <i>Sirex juvencus</i> is banded with orange, +whilst the female is dark purple; but it is difficult to +say which sex is the most ornamented. In <i>Tremex +columbæ</i> the female is much brighter coloured than the +male. With ants, as I am informed by Mr. F. Smith, +the males of several species are black, the females +being testaceous. In the family of Bees, especially in +<span class='pagenum'><a name="Page_366" id="Page_366">366</a></span>the solitary species, as I hear from the same distinguished +entomologist, the sexes often differ in colour. +The males are generally the brightest, and in Bombus +as well as in Apathus, much more variable in colour +than the females. In <i>Anthophora retusa</i> the male is +of a rich fulvous-brown, whilst the female is quite +black: so are the females of several species of Xylocopa, +the males being bright yellow. In an Australian bee +(<i>Lestis bombylans</i>), the female is of an extremely brilliant +steel-blue, sometimes tinted with vivid green; the male +being of a bright brassy colour clothed with rich fulvous +pubescence. As in this group the females are provided +with excellent defensive weapons in their stings, it is +not probable that they have come to differ in colour +from the males for the sake of protection.</p> + +<p class="tb"><i>Mutilla Europæa</i> emits a stridulating noise; and according +to Goureau<a name="FNanchor_483" id="FNanchor_483"></a><a href="#Footnote_483" class="fnanchor">483</a> both sexes have this power. He +attributes the sound to the friction of the third and +preceding abdominal segments; and I find that these +surfaces are marked with very fine concentric ridges, +but so is the projecting thoracic collar, on which the +head articulates; and this collar, when scratched with +the point of a needle, emits the proper sound. It is +rather surprising that both sexes should have the +power of stridulating, as the male is winged and the +female wingless. It is notorious that Bees express +certain emotions, as of anger, by the tone of their +humming, as do some dipterous insects; but I have +not referred to these sounds, as they are not known to +be in any way connected with the act of courtship.</p> + +<p>Order, <i>Coleoptera</i> (Beetles).—Many beetles are +coloured so as to resemble the surfaces which they +<span class='pagenum'><a name="Page_367" id="Page_367">367</a></span>habitually frequent. Other species are ornamented +with gorgeous metallic tints,—for instance, many Carabidæ, +which live on the ground and have the power +of defending themselves by an intensely acrid secretion,—the +splendid diamond-beetles which are protected by +an extremely hard covering,—many species of Chrysomela, +such as <i>C. cerealis</i>, a large species beautifully +striped with various colours, and in Britain confined +to the bare summit of Snowdon,—and a host of other +species. These splendid colours, which are often +arranged in stripes, spots, crosses and other elegant +patterns, can hardly be beneficial, as a protection, except +in the case of some flower-feeding species; and we +cannot believe that they are purposeless. Hence the +suspicion arises, that they serve as a sexual attraction; +but we have no evidence on this head, for the sexes +rarely differ in colour. Blind beetles, which cannot of +course behold each other’s beauty, never exhibit, as I +hear from Mr. Waterhouse, jun., bright colours, though +they often have polished coats: but the explanation of +their obscurity may be that blind insects inhabit caves +and other obscure stations.</p> + +<p>Some Longicorns, however, especially certain Prionidæ, +offer an exception to the common rule that the +sexes of beetles do not differ in colour. Most of these +insects are large and splendidly coloured. The males in +the genus Pyrodes,<a name="FNanchor_484" id="FNanchor_484"></a><a href="#Footnote_484" class="fnanchor">484</a> as I saw in Mr. Bates’ collection, are +<span class='pagenum'><a name="Page_368" id="Page_368">368</a></span>generally redder but rather duller than the females, the +latter being coloured of a more or less splendid golden +green. On the other hand, in one species the male is +golden-green, the female being richly tinted with red +and purple. In the genus Esmeralda the sexes differ so +greatly in colour that they have been ranked as distinct +species: in one species both are of a beautiful shining +green, but the male has a red thorax. On the whole, +as far as I could judge, the females of those Prionidæ, +in which the sexes differ, are coloured more richly +than the males; and this does not accord with the +common rule in regard to colour when acquired through +sexual selection.</p> + +<div class="figcenter" style="width: 450px;"><a name="f15" id="f15"></a><img src="images/fig15.png" width="450" height="363" +alt="Fig. 15. Chalcosoma atlas. Upper figure, male (reduced); lower figure, female +(nat. size)." title="" /> + +<p class="center2">Fig. 15. Chalcosoma atlas. Upper figure, male (reduced); lower figure, female +(nat. size).</p></div> + +<p><span class='pagenum'><a name="Page_369" id="Page_369">369</a></span></p> + +<div class="figcenter" style="width: 500px;"><a name="f16" id="f16"></a><img src="images/fig16.png" width="500" height="186" +alt="Fig. 16. Copris isidis. (Left-hand figures, males.)" title="" /> + +<p class="center2">Fig. 16. Copris isidis. (Left-hand figures, males.)</p></div> + +<div class="figcenter" style="width: 500px;"><a name="f17" id="f17"></a><img src="images/fig17.png" width="500" height="175" +alt="Fig. 17. Phanæus faunus." title="" /> + +<p class="center2">Fig. 17. Phanæus faunus.</p></div> + +<div class="figcenter" style="width: 500px;"><a name="f18" id="f18"></a><img src="images/fig18.png" width="500" height="168" +alt="Fig. 18. Dipelicus cantori." title="" /> + +<p class="center2">Fig. 18. Dipelicus cantori.</p></div> + +<div class="figcenter" style="width: 500px;"><a name="f19" id="f19"></a><img src="images/fig19.png" width="500" height="162" +alt="Fig. 19. Onthophagus rangifer, enlarged." title="" /> + +<p class="center2">Fig. 19. Onthophagus rangifer, enlarged.</p></div> + +<p><span class='pagenum'><a name="Page_370" id="Page_370">370</a></span></p> + +<p>A most remarkable distinction between the sexes of +many beetles is presented by the great horns which +rise from the head, thorax, or clypeus of the males; +and in some few cases from the under surface of the +body. These horns, in the great family of the Lamellicorns, +resemble those of various quadrupeds, such as +stags, rhinoceroses, &c., and are wonderful both from +their size and diversified shapes. Instead of describing +them, I have given figures of the males and females of +some of the more remarkable forms. (Figs. 15 to 19.) +The females generally exhibit rudiments of the horns +in the form of small knobs or ridges; but some are +destitute of even a rudiment. On the other hand, the +horns are nearly as well developed in the female as in +the male of <i>Phanæus lancifer</i>; and only a little less +well developed in the females of some other species of +the same genus and of Copris. In the several subdivisions +of the family, the differences in structure of +the horns do not run parallel, as I am informed by +Mr. Bates, with their more important and characteristic +differences; thus within the same natural section of the +genus Onthophagus, there are species which have either +a single cephalic horn, or two distinct horns.</p> + +<p>In almost all cases, the horns are remarkable from +their excessive variability; so that a graduated series +can be formed, from the most highly developed males +to others so degenerate that they can barely be distinguished +from the females. Mr. Walsh<a name="FNanchor_485" id="FNanchor_485"></a><a href="#Footnote_485" class="fnanchor">485</a> found that in +<i>Phanæus carnifex</i> the horns were thrice as long in some +males as in others. Mr. Bates, after examining above +a hundred males of <i>Onthophagus rangifer</i> (fig. 19), +thought that he had at last discovered a species in +<span class='pagenum'><a name="Page_371" id="Page_371">371</a></span>which the horns did not vary; but further research +proved the contrary.</p> + +<p>The extraordinary size of the horns, and their widely +different structure in closely-allied forms, indicate that +they have been formed for some important purpose; +but their excessive variability in the males of the same +species leads to the inference that this purpose cannot +be of a definite nature. The horns do not show marks +of friction, as if used for any ordinary work. Some +authors suppose<a name="FNanchor_486" id="FNanchor_486"></a><a href="#Footnote_486" class="fnanchor">486</a> that as the males wander much more +than the females, they require horns as a defence +against their enemies; but in many cases the horns +do not seem well adapted for defence, as they are not +sharp. The most obvious conjecture is that they are +used by the males for fighting together; but they +have never been observed to fight; nor could Mr. Bates, +after a careful examination of numerous species, find +any sufficient evidence in their mutilated or broken +condition of their having been thus used. If the males +had been habitual fighters, their size would probably +have been increased through sexual selection, so as to +have exceeded that of the female; but Mr. Bates, after +comparing the two sexes in above a hundred species of +the Copridæ, does not find in well-developed individuals +any marked difference in this respect. There is, moreover, +one beetle, belonging to the same great division +of the Lamellicorns, namely Lethrus, the males of which +are known to fight, but they are not provided with +horns, though their mandibles are much larger than +those of the female.</p> + +<p>The conclusion, which best agrees with the fact of +the horns having been so immensely yet not fixedly +developed,—as shewn by their extreme variability in +<span class='pagenum'><a name="Page_372" id="Page_372">372</a></span>the same species and by their extreme diversity in +closely-allied species—is that they have been acquired +as ornaments. This view will at first appear extremely +improbable; but we shall hereafter find with many +animals, standing much higher in the scale, namely +fishes, amphibians, reptiles and birds, that various +kinds of crests, knobs, horns and combs have been +developed apparently for this sole purpose.</p> + +<p>The males of <i>Onitis furcifer</i> (fig. 20) are furnished +with singular projections on their anterior femora, and + +<span class="figleft2" style="width: 150px;"><a name="f20" id="f20"></a><img src="images/fig20.png" width="150" height="156" +alt="Fig. 20. Onitis furcifer, male, +viewed from beneath." title="" /> + +<span class="indent2">Fig. 20. Onitis furcifer, male, +viewed from beneath.</span></span> + +with a great fork or pair of horns on +the lower surface of the thorax. This +situation seems extremely ill adapted +for the display of these projections, +and they may be of some real service; +but no use can at present be +assigned to them. It is a highly +remarkable fact, that although the +males do not exhibit even a trace of +horns on the upper surface of the +body, yet in the females a rudiment of a single horn on +the head (fig. 21, <i>a</i>), and of a crest (<i>b</i>) on the thorax, +are plainly visible. That the slight thoracic crest in the +female is a rudiment of a projection proper to the male, +though entirely absent in the male of this particular +species, is clear: for the female of <i>Bubas bison</i> (a form<span class='pagenum'><a name="Page_373" id="Page_373">373</a></span> +which comes next to <i>Onitis</i>) has a similar slight crest +on the thorax, and the male has in the same situation a +great projection. So again there can be no doubt that +the little point (<i>a</i>) on the head of the female <i>Onitis +furcifer</i>, as well of the females of two or three allied +species, is a rudimentary representative of the cephalic +horn, which is common to the males of so many lamellicorn +beetles, as in Phanæus, fig. 17. The males indeed +of some unnamed beetles in the British Museum, which +are believed actually to belong to the genus Onitis, are +furnished with a similar horn. The remarkable nature +of this case will be best perceived by an illustration: +the Ruminant quadrupeds run parallel with the lamellicorn +beetles, in some females possessing horns as large +as those of the male, in others having them much +smaller, or existing as mere rudiments (though this is +as rare with ruminants as it is common with Lamellicorns), +or in having none at all. Now if a new species +of deer or sheep were discovered with the female +bearing distinct rudiments of horns, whilst the head +of the male was absolutely smooth, we should have a +case like that of <i>Onitis furcifer</i>.</p> + +<div class="figcenter" style="width: 450px;"><a name="f21" id="f21"></a><img src="images/fig21.png" width="450" height="146" +alt="Fig. 21. Left-hand figure, male of Onitis furcifer." title="" /></div> + +<p class="indent2">Fig. 21. Left-hand figure, male of Onitis furcifer, viewed laterally. Right-hand figure, +female. <i>a.</i> Rudiment of cephalic horn. <i>b.</i> Trace of thoracic horn or crest.</p> + +<p>In this case the old belief of rudiments having been +created to complete the scheme of nature is so far from +holding good, that all ordinary rules are completely +broken through. The view which seems the most probable +is that some early progenitor of Onitis acquired, +like other Lamellicorns, horns on the head and thorax, +and then transferred them, in a rudimentary condition, +as with so many existing species, to the female, by whom +they have ever since been retained. The subsequent +loss of the horns by the male may have resulted through +the principle of compensation from the development of +the projections on the lower surface, whilst the female +has not been thus affected, as she is not furnished with<span class='pagenum'><a name="Page_374" id="Page_374">374</a></span> +these projections, and consequently has retained the +rudiments of the horns on the upper surface. Although +this view is supported by the case of Bledius immediately +to be given, yet the projections on the lower +surface differ greatly in structure and development in +the males of the several species of Onitis, and are even +rudimentary in some; nevertheless the upper surface +in all these species is quite destitute of horns. As +secondary sexual characters are so eminently variable, it +is possible that the projections on the lower surface may +have been first acquired by some progenitor of Onitis and +produced their effect through compensation, and then +have been in certain cases almost completely lost.</p> + +<div class="figcenter" style="width: 500px;"><a name="f22" id="f22"></a><img src="images/fig22.png" width="500" height="98" +alt="Fig. 22. Bledius taurus, magnified. Left-hand figure, male; right-hand figure, female." title="" /></div> + +<p class="center2">Fig. 22. Bledius taurus, magnified. Left-hand figure, male; right-hand figure, female.</p> + +<p class="tb">All the cases hitherto given refer to the Lamellicorns, +but the males of some few other beetles, belonging +to two widely distinct groups, namely, the +Curculionidæ and Staphylinidæ, are furnished with +horns,—in the former on the lower surface of the body,<a name="FNanchor_487" id="FNanchor_487"></a><a href="#Footnote_487" class="fnanchor">487</a> +in the latter on the upper surface of the head and +thorax. In the Staphylinidæ the horns of the males +in the same species are extraordinarily variable, just +as we have seen with the Lamellicorns. In Siagonium +we have a case of dimorphism, for the males can be +divided into two sets, differing greatly in the size of +their bodies, and in the development of their horns, +without any intermediate gradations. In a species of +Bledius (fig. 22), also belonging to the Staphylinidæ, +male specimens can be found in the same locality, as +<span class='pagenum'><a name="Page_375" id="Page_375">375</a></span> +Professor Westwood states, “in which the central horn +of the thorax is very large, but the horns of the head +quite rudimental; and others, in which the thoracic +horn is much shorter, whilst the protuberances on +the head are long.”<a name="FNanchor_488" id="FNanchor_488"></a><a href="#Footnote_488" class="fnanchor">488</a> Here, then, we apparently +have an instance of compensation of growth, which +throws light on the curious case just given of the loss of +the upper horns by the males of <i>Onitis furcifer.</i></p> + +<p><i>Law of Battle.</i>—Some male beetles, which seem ill +fitted for fighting, nevertheless engage in conflicts for +the possession of the females. Mr. Wallace<a name="FNanchor_489" id="FNanchor_489"></a><a href="#Footnote_489" class="fnanchor">489</a> saw two +males of <i>Leptorhynchus angustatus</i>, a linear beetle with +a much elongated rostrum, “fighting for a female, who +stood close by busy at her boring. They pushed at +each other with their rostra, and clawed and thumped, +apparently in the greatest rage.” The smaller male, +however, “soon ran away, acknowledging himself vanquished.” +In some few cases the males are well +adapted for fighting, by possessing great toothed mandibles, +much larger than those of the females. This +is the case with the common stag-beetle (<i>Lucanus +cervus</i>), the males of which emerge from the pupal state +about a week before the other sex, so that several may +often be seen pursuing the same female. At this period +they engage in fierce conflicts. When Mr. A. H. +Davis<a name="FNanchor_490" id="FNanchor_490"></a><a href="#Footnote_490" class="fnanchor">490</a> enclosed two males with one female in a box, +the larger male severely pinched the smaller one, until +he resigned his pretensions. A friend informs me +<span class='pagenum'><a name="Page_376" id="Page_376">376</a></span>that when a boy he often put the males together to see +them fight, and he noticed that they were much bolder +and fiercer than the females, as is well known to be the +case with the higher animals. The males would seize +hold of his finger, if held in front, but not so the females. +With many of the Lucanidæ, as well as with the above-mentioned +Leptorhynchus, the males are larger and +more powerful insects than the females. The two sexes +of <i>Lethrus cephalotes</i> (one of the Lamellicorns) inhabit the +same burrow; and the male has larger mandibles than +the female. If, during the breeding-season, a strange +male attempts to enter the burrow, he is attacked; the +female does not remain passive, but closes the mouth of +the burrow, and encourages her mate by continually +pushing him on from behind. The action does not +cease until the aggressor is killed or runs away.<a name="FNanchor_491" id="FNanchor_491"></a><a href="#Footnote_491" class="fnanchor">491</a> The +two sexes of another lamellicorn beetle, the <i>Ateuchus +cicatricosus</i> live in pairs, and seem much attached to +each other; the male excites the female to roll the +balls of dung in which the ova are deposited; and if +she is removed, he becomes much agitated. If the +male is removed, the female ceases all work, and as +M. Brulerie<a name="FNanchor_492" id="FNanchor_492"></a><a href="#Footnote_492" class="fnanchor">492</a> believes, would remain on the spot until +she died.</p> + +<p>The great mandibles of the male Lucanidæ are extremely +variable both in size and structure, and in this +respect resemble the horns on the head and thorax +of many male Lamellicorns and Staphylinidæ. A perfect +series can be formed from the best-provided to the +worst-provided or degenerate males. Although the +mandibles of the common stag-beetle, and probably of +<span class='pagenum'><a name="Page_377" id="Page_377">377</a></span>many other species, are used as efficient weapons for +fighting, it is doubtful whether their great size can + +<span class="figright2" style="width: 200px;"><a name="f23" id="f23"></a><img src="images/fig23.png" width="200" height="682" +alt="Fig. 23. Chiasognathus grantii, +reduced. Upper figure, male; +lower figure, female." title="" /> + +<span class="indent2">Fig. 23. Chiasognathus grantii, +reduced. Upper figure, male; +lower figure, female.</span></span> + +thus be accounted for. We have +seen that with the <i>Lucanus elaphus</i> +of N. America they are used +for seizing the female. As they +are so conspicuous and so elegantly +branched, the suspicion +has sometimes crossed my mind +that they may be serviceable to +the males as an ornament, in the +same manner as the horns on the +head and thorax of the various +above described species. The +male <i>Chiasognathus grantii</i> of S. +Chile—a splendid beetle belonging +to the same family—has enormously-developed +mandibles (fig. +23); he is bold and pugnacious; +when threatened on any side he +faces round, opening his great +jaws, and at the same time stridulating +loudly; but the mandibles +were not strong enough to pinch +my finger so as to cause actual +pain.</p> + +<p class="tb">Sexual selection, which implies +the possession of considerable perceptive +powers and of strong passions, +seems to have been more +effective with the Lamellicorns +than with any other family of the +Coleoptera or beetles. With some +species the males are provided with weapons for fighting; +some live in pairs and show mutual affection;<span class='pagenum'><a name="Page_378" id="Page_378">378</a></span> +many have the power of stridulating when excited; many +are furnished with the most extraordinary horns, apparently +for the sake of ornament; some which are diurnal +in their habits are gorgeously coloured; and, lastly, +several of the largest beetles in the world belong to this +family, which was placed by Linnæus and Fabricius at +the head of the Order of the Coleoptera.<a name="FNanchor_493" id="FNanchor_493"></a><a href="#Footnote_493" class="fnanchor">493</a></p> + +<p><i>Stridulating organs.</i>—Beetles belonging to many +and widely distinct families possess these organs. The +sound can sometimes be heard at the distance of several +feet or even yards,<a name="FNanchor_494" id="FNanchor_494"></a><a href="#Footnote_494" class="fnanchor">494</a> but is not comparable with that +produced by the Orthoptera. The part which may be +called the rasp generally consists of a narrow slightly-raised +surface, crossed by very fine, parallel ribs, sometimes +so fine as to cause iridescent colours, and having +a very elegant appearance under the microscope. In +some cases, for instance, with Typhæus, it could be +plainly seen that extremely minute, bristly, scale-like +prominences, which cover the whole surrounding surface +in approximately parallel lines, give rise to the +ribs of the rasp by becoming confluent and straight, and +at the same time more prominent and smooth. A hard +ridge on any adjoining part of the body, which in some +cases is specially modified for the purpose, serves as the +scraper for the rasp. The scraper is rapidly moved across +the rasp, or conversely the rasp across the scraper.</p> + +<div class="figcenter" style="width: 450px;"><a name="f24" id="f24"></a><img src="images/fig24.png" width="450" height="166" +alt="Fig. 24. Necrophorus (from Landois). r. The two rasps. Left-hand figure, part of +the rasp highly magnified." title="" /></div> + +<p class="center2">Fig. 24. Necrophorus (from Landois). <i>r.</i> The two rasps. Left-hand figure, part of +the rasp highly magnified.</p> + +<p>These organs are situated in widely different positions. +In the carrion-beetles (Necrophorus) two parallel +rasps (<i>r</i>, fig. 24) stand on the dorsal surface of the fifth +abdominal segment, each rasp being crossed, as described +by Landois,<a name="FNanchor_495" id="FNanchor_495"></a><a href="#Footnote_495" class="fnanchor">495</a> by from 126 to 140 fine ribs. These +<span class='pagenum'><a name="Page_379" id="Page_379">379</a></span>ribs are scraped by the posterior margins of the elytra, a +small portion of which projects beyond the general outline. +In many Crioceridæ, and in <i>Clythra 4-punctata</i> +(one of the Chrysomelidæ), and in some Tenebrionidæ, +&c.,<a name="FNanchor_496" id="FNanchor_496"></a><a href="#Footnote_496" class="fnanchor">496</a> the rasp is seated on the dorsal apex of the abdomen, +on the pygidium or pro-pygidium, and is scraped +as above by the elytra. In Heterocerus, which belongs +to another family, the rasps are placed on the sides of +the first abdominal segment, and are scraped by ridges +on the femora.<a name="FNanchor_497" id="FNanchor_497"></a><a href="#Footnote_497" class="fnanchor">497</a> In certain Curculionidæ and Carabidæ,<a name="FNanchor_498" id="FNanchor_498"></a><a href="#Footnote_498" class="fnanchor">498</a> +the parts are completely reversed in position, +<span class='pagenum'><a name="Page_380" id="Page_380">380</a></span>for the rasps are seated on the inferior surface of the +elytra, near their apices, or along their outer margins, +and the edges of the abdominal segments serve as the +scrapers. In <i>Pelobius hermanni</i> (one of Dytiscidæ or +water-beetles) a strong ridge runs parallel and near to +the sutural margin of the elytra, and is crossed by ribs, +coarse in the middle part, but becoming gradually finer +at both ends, especially at the upper end; when this +insect is held under water or in the air, a stridulating +noise is produced by scraping the extreme horny margin +of the abdomen against the rasp. In a great number +of long-horned beetles (Longicornia) the organs are altogether +differently situated, the rasp being on the meso-thorax, +which is rubbed against the pro-thorax; Landois +counted 238 very fine ribs on the rasp of <i>Cerambyx +heros</i>.</p> + +<p>Many Lamellicorns have the power of stridulating, +and the organs differ greatly in position. Some species + +<span class="figleft2" style="width: 200px;"><a name="f25" id="f25"></a><img src="images/fig25.png" width="200" height="289" +alt="Fig. 25. Hind-leg of Geotrupes +stercorarius (from Landois)." title="" /> + +<span class="indent2">Fig. 25. Hind-leg of Geotrupes +stercorarius (from Landois).<br /><br /></span> +<span class="indent2"><i>r.</i> Rasp. <i>c.</i> Coxa. <i>f.</i> Femur. +<i>t.</i> Tibia. <i>tr.</i> Tarsi.</span></span> + +stridulate very loudly, so that when +Mr. F. Smith caught a <i>Trox sabulosus</i>, +a gamekeeper who stood by +thought that he had caught a +mouse; but I failed to discover the +proper organs in this beetle. In +Geotrupes and Typhæus a narrow +ridge runs obliquely across (<i>r</i>, fig. +25) the coxa of each hind-leg, +having in <i>G. stercorarius</i> 84 ribs, +which are scraped by a specially-projecting +part of one of the abdominal +segments. In the nearly +allied <i>Copris lunaris</i>, an excessively +narrow fine rasp runs along the +sutural margin of the elytra, with another short rasp +near the basal outer margin; but in some other Coprini<span class='pagenum'><a name="Page_381" id="Page_381">381</a></span> +the rasp is seated, according to Leconte,<a name="FNanchor_499" id="FNanchor_499"></a><a href="#Footnote_499" class="fnanchor">499</a> on the dorsal +surface of the abdomen. In Oryctes it is seated on the +pro-pygidium, and in some other Dynastini, according to +the same entomologist, on the under surface of the +elytra. Lastly, Westring states that in <i>Omaloplia brunnea</i> +the rasp is placed on the pro-sternum, and the scraper +on the meta-sternum, the parts thus occupying the under +surface of the body, instead of the upper surface as in +the Longicorns.</p> + +<p>We thus see that the stridulating organs in the different +coleopterous families are wonderfully diversified +in position, but not much in structure. Within the +same family some species are provided with these +organs, and some are quite destitute of them. This +diversity is intelligible, if we suppose that originally +various species made a shuffling or hissing noise by the +rubbing together of the hard and rough parts of their +bodies which were in contact; and that from the noise +thus produced being in some way useful, the rough +surfaces were gradually developed into regular stridulating +organs. Some beetles as they move, now +produce, either intentionally or unintentionally, a shuffling +noise, without possessing any proper organs for the +purpose. Mr. Wallace informs me that the <i>Euchirus +longimanus</i> (a Lamellicorn, with the anterior legs wonderfully +elongated in the male) “makes, whilst moving, +a low hissing sound by the protrusion and contraction +of the abdomen; and when seized it produces a +grating sound by rubbing its hind-legs against the +edges of the elytra.” The hissing sound is clearly +due to a narrow rasp running along the sutural margin +of each elytron; and I could likewise make the grating +<span class='pagenum'><a name="Page_382" id="Page_382">382</a></span>sound by rubbing the shagreened surface of the femur +against the granulated margin of the corresponding +elytron; but I could not here detect any proper rasp; +nor is it likely that I could have overlooked it in so +large an insect. After examining Cychrus and reading +what Westring has written in his two papers about this +beetle, it seems very doubtful whether it possesses any +true rasp, though it has the power of emitting a sound.</p> + +<p>From the analogy of the Orthoptera and Homoptera, +I expected to find that the stridulating organs in the +Coleoptera differed according to sex; but Landois, who +has carefully examined several species, observed no +such difference; nor did Westring; nor did Mr. G. R. +Crotch in preparing the numerous specimens which +he had the kindness to send me for examination. Any +slight sexual difference, however, would be difficult to +detect, on account of the great variability of these organs. +Thus in the first pair of the <i>Necrophorus humator</i> and of +the <i>Pelobius</i> which I examined, the rasp was considerably +larger in the male than in the female; but not so +with succeeding specimens. In <i>Geotrupes stercorarius</i> +the rasp appeared to me thicker, opaquer, and more +prominent in three males than in the same number of +females; consequently my son, Mr. F. Darwin, in order +to discover whether the sexes differed in their power of +stridulating, collected 57 living specimens, which he +separated into two lots, according as they made, when +held in the same manner, a greater or lesser noise. He +then examined their sexes, but found that the males +were very nearly in the same proportion to the females +in both lots. Mr. F. Smith has kept alive numerous +specimens of <i>Mononychus pseudacori</i> (Curculionidæ), and +is satisfied that both sexes stridulate, and apparently in +an equal degree.</p> + +<p>Nevertheless the power of stridulating is certainly a<span class='pagenum'><a name="Page_383" id="Page_383">383</a></span> +sexual character in some few Coleoptera. Mr. Crotch +has discovered that the males alone of two species of +Heliopathes (Tenebrionidæ) possess stridulating organs. +I examined five males of <i>H. gibbus</i>, and in all these +there was a well-developed rasp, partially divided into +two, on the dorsal surface of the terminal abdominal +segment; whilst in the same number of females there +was not even a rudiment of the rasp, the membrane of +this segment being transparent and much thinner than +in the male. In <i>H. cribratostriatus</i> the male has a +similar rasp, excepting that it is not partially divided +into two portions, and the female is completely destitute +of this organ; but in addition the male has on +the apical margins of the elytra, on each side of the +suture, three or four short longitudinal ridges, which +are crossed by extremely fine ribs, parallel to and resembling +those on the abdominal rasp; whether these +ridges serve as an independent rasp, or as a scraper +for the abdominal rasp, I could not decide: the female +exhibits no trace of this latter structure.</p> + +<p>Again, in three species of the Lamellicorn genus +Oryctes, we have a nearly parallel case. In the females +of <i>O. gryphus</i> and <i>nasicornis</i> the ribs on the rasp of the +pro-pygidium are less continuous and less distinct than +in the males; but the chief difference is that the whole +upper surface of this segment, when held in the proper +light, is seen to be clothed with hairs, which are absent +or are represented by excessively fine down in the males. +It should be noticed that in all Coleoptera the effective +part of the rasp is destitute of hairs. In <i>O. senegalensis</i> +the difference between the sexes is more strongly +marked, and this is best seen when the proper segment +is cleaned and viewed as a transparent object. In the +female the whole surface is covered with little separate +crests, bearing spines; whilst in the male these crests<span class='pagenum'><a name="Page_384" id="Page_384">384</a></span> +become, in proceeding towards the apex, more and more +confluent, regular, and naked; so that three-fourths of +the segment is covered with extremely fine parallel +ribs, which are quite absent in the female. In the +females, however, of all three species of Oryctes, when +the abdomen of a softened specimen is pushed backwards +and forwards, a slight grating or stridulating +sound can be produced.</p> + +<p>In the case of the Heliopathes and Oryctes there can +hardly be a doubt that the males stridulate in order to +call or to excite the females; but with most beetles the +stridulation apparently serves both sexes as a mutual +call. This view is not rendered improbable from beetles +stridulating under various emotions; we know that birds +use their voices for many purposes besides singing to +their mates. The great Chiasognathus stridulates in +anger or defiance; many species do the same from distress +or fear, when held so that they cannot escape; +Messrs. Wollaston and Crotch were able, by striking +the hollow stems of trees in the Canary Islands, to discover +the presence of beetles belonging to the genus +Acalles by their stridulation. Lastly the male Ateuchus +stridulates to encourage the female in her work, +and from distress when she is removed.<a name="FNanchor_500" id="FNanchor_500"></a><a href="#Footnote_500" class="fnanchor">500</a> Some naturalists +believe that beetles make this noise to frighten +away their enemies; but I cannot think that the quadrupeds +and birds which are able to devour the larger +beetles with their extremely hard coats, would be frightened +by so slight a grating sound. The belief that +the stridulation serves as a sexual call is supported +by the fact that death-ticks (<i>Anobium tesselatum</i>) are +well known to answer each other’s ticking, or, as I have +<span class='pagenum'><a name="Page_385" id="Page_385">385</a></span>myself observed, a tapping noise artificially made; and +Mr. Doubleday informs me that he has twice or thrice +observed a female ticking,<a name="FNanchor_501" id="FNanchor_501"></a><a href="#Footnote_501" class="fnanchor">501</a> and in the course of an hour +or two has found her united with a male, and on one +occasion surrounded by several males. Finally, it seems +probable that the two sexes of many kinds of beetles +were at first enabled to find each other by the slight +shuffling noise produced by the rubbing together of the +adjoining parts of their hard bodies; and that as the +males or females which made the greatest noise succeeded +best in finding partners, the rugosities on various +parts of their bodies were gradually developed by means +of sexual selection into true stridulating organs.</p> + +<hr /> +<p><span class='pagenum'><a name="Page_386" id="Page_386">386</a></span></p> + +<h3>CHAPTER XI.</h3> + +<h4><span class="smcap">Insects</span>, <i>continued</i>.—<span class="smcap">Order Lepidoptera.</span></h4> + +<div class="blockquot"><p>Courtship of butterflies—Battles—Ticking noise—Colours common +to both sexes, or more brilliant in the males—Examples—Not +due to the direct action of the conditions of life—Colours +adapted for protection—Colours of moths—Display—Perceptive +powers of the Lepidoptera—Variability—Causes of the +difference in colour between the males and females—Mimickry, +female butterflies more brilliantly coloured than the males—Bright +colours of caterpillars—Summary and concluding remarks +on the secondary sexual characters of insects—Birds and +insects compared.</p></div> + +<p>In this great Order the most interesting point for us is +the difference in colour between the sexes of the same +species, and between the distinct species of the same +genus. Nearly the whole of the following chapter will +be devoted to this subject; but I will first make a few +remarks on one or two other points. Several males may +often be seen pursuing and crowding round the same +female. Their courtship appears to be a prolonged affair, +for I have frequently watched one or more males pirouetting +round a female until I became tired, without seeing +the end of the courtship. Although butterflies are such +weak and fragile creatures, they are pugnacious, and an +Emperor butterfly<a name="FNanchor_502" id="FNanchor_502"></a><a href="#Footnote_502" class="fnanchor">502</a> has been captured with the tips of +its wings broken from a conflict with another male. +Mr. Collingwood in speaking of the frequent battles +<span class='pagenum'><a name="Page_387" id="Page_387">387</a></span>between the butterflies of Borneo says, “They whirl +round each other with the greatest rapidity, and appear +to be incited by the greatest ferocity.” One case is +known of a butterfly, namely the <i>Ageronia feronia</i>, +which makes a noise like that produced by a toothed +wheel passing under a spring catch, and which could be +heard at the distance of several yards. At Rio de Janeiro +this sound was noticed by me, only when two were +chasing each other in an irregular course, so that it is +probably made during the courtship of the sexes; but I +neglected to attend to this point.<a name="FNanchor_503" id="FNanchor_503"></a><a href="#Footnote_503" class="fnanchor">503</a></p> + +<p>Every one has admired the extreme beauty of many +butterflies and of some moths; and we are led to ask, +how has this beauty been acquired? Have their colours +and diversified patterns simply resulted from the direct +action of the physical conditions to which these insects +have been exposed, without any benefit being thus derived? +Or have successive variations been accumulated +and determined either as a protection or for some unknown +purpose, or that one sex might be rendered +attractive to the other? And, again, what is the meaning +of the colours being widely different in the males +and females of certain species, and alike in the two +sexes of other species? Before attempting to answer +these questions a body of facts must be given.</p> + +<p>With most of our English butterflies, both those which +are beautiful, such as the admiral, peacock, and painted +lady (Vanessæ), and those which are plain-coloured, +such as the meadow-browns (Hipparchiæ), the sexes +are alike. This is also the case with the magnificent +Heliconidæ and Danaidæ of the tropics. But in certain +<span class='pagenum'><a name="Page_388" id="Page_388">388</a></span>other tropical groups, and with some of our English +butterflies, as the purple emperor, orange-tip, &c. (<i>Apatura +Iris</i> and <i>Anthocharis cardamines</i>), the sexes differ +either greatly or slightly in colour. No language suffices +to describe the splendour of the males of some tropical +species. Even within the same genus we often find species +presenting an extraordinary difference between the +sexes, whilst others have their sexes closely alike. Thus +in the South American genus Epicalia, Mr. Bates, to +whom I am much indebted for most of the following +facts and for looking over this whole discussion, informs +me that he knows twelve species, the two sexes of which +haunt the same stations (and this is not always the case +with butterflies), and therefore cannot have been differently +affected by external conditions<a name="FNanchor_504" id="FNanchor_504"></a><a href="#Footnote_504" class="fnanchor">504</a>. In nine of +these species the males rank amongst the most brilliant +of all butterflies, and differ so greatly from the comparatively +plain females that they were formerly placed +in distinct genera. The females of these nine species +resemble each other in their general type of coloration, +and likewise resemble both sexes in several allied genera, +found in various parts of the world. Hence in accordance +with the descent-theory we may infer that these +nine species, and probably all the others of the genus, +are descended from an ancestral form which was coloured +in nearly the same manner. In the tenth species the +female still retains the same general colouring, but the +male resembles her, so that he is coloured in a much +less gaudy and contrasted manner than the males of the +previous species. In the eleventh and twelfth species, +the females depart from the type of colouring which +<span class='pagenum'><a name="Page_389" id="Page_389">389</a></span>is usual with their sex in this genus, for they are gaily +decorated in nearly the same manner as the males, but +in a somewhat less degree. Hence in these two species +the bright colours of the males seem to have been transferred +to the females; whilst the male of the tenth +species has either retained or recovered the plain +colours of the female as well as of the parent-form of +the genus; the two sexes being thus rendered in both +cases, though in an opposite manner, nearly alike. In +the allied genus Eubagis, both sexes of some of the +species are plain-coloured and nearly alike; whilst +with the greater number the males are decorated with +beautiful metallic tints, in a diversified manner, and +differ much from their females. The females throughout +the genus retain the same general style of colouring, +so that they commonly resemble each other much more +closely than they resemble their own proper males.</p> + +<p>In the genus Papilio, all the species of the Æneas +group are remarkable for their conspicuous and strongly +contrasted colours, and they illustrate the frequent tendency +to gradation in the amount of difference between +the sexes. In a few species, for instance in <i>P. ascanius</i>, +the males and females are alike; in others the males +are a little or very much more superbly coloured than +the females. The genus Junonia allied to our Vanessæ +offers a nearly parallel case, for although the sexes of +most of the species resemble each other and are destitute +of rich colours, yet in certain species, as in <i>J. œnone</i>, +the male is rather more brightly coloured than the +female, and in a few (for instance <i>J. andremiaja</i>) the +male is so different from the female that he might be +mistaken for an entirely distinct species.</p> + +<p>Another striking case was pointed out to me in the +British museum by Mr. A. Butler, namely one of +the Tropical American Theclæ, in which both sexes<span class='pagenum'><a name="Page_390" id="Page_390">390</a></span> +are nearly alike and wonderfully splendid; in another, +the male is coloured in a similarly gorgeous manner, +whilst the whole upper surface of the female is of a dull +uniform brown. Our common little English blue butterflies +of the genus Lycæna, illustrate the various differences +in colour between the sexes, almost as well, +though not in so striking a manner, as the above exotic +genera. In <i>Lycæna agestis</i> both sexes have wings of a +brown colour, bordered with small ocellated orange +spots, and are consequently alike. In <i>L. œgon</i> the +wings of the male are of a fine blue, bordered with +black; whilst the wings of the female are brown, with +a similar border, and closely resemble those of <i>L. agestis</i>. +Lastly, in <i>L. arion</i> both sexes are of a blue colour and +nearly alike, though in the female the edges of the +wings are rather duskier, with the black spots plainer; +and in a bright blue Indian species both sexes are still +more closely alike.</p> + +<p>I have given the foregoing cases in some detail in +order to shew, in the first place, that when the sexes of +butterflies differ, the male as a general rule is the most +beautiful, and departs most from the usual type of colouring +of the group to which the species belongs. Hence in +most groups the females of the several species resemble +each other much more closely than do the males. In +some exceptional cases, however, to which I shall hereafter +allude, the females are coloured more splendidly +than the males. In the second place these cases have +been given to bring clearly before the mind that within +the same genus, the two sexes frequently present every +gradation from no difference in colour to so great a difference +that it was long before the two were placed by +entomologists in the same genus. In the third place, +we have seen that when the sexes nearly resemble +each other, this apparently may be due either to the<span class='pagenum'><a name="Page_391" id="Page_391">391</a></span> +male having transferred his colours to the female, +or to the male having retained, or perhaps recovered, +the primordial colours of the genus to which the +species belongs. It also deserves notice that in those +groups in which the sexes present any difference of +colour, the females usually resemble the males to a certain +extent, so that when the males are beautiful to an +extraordinary degree, the females almost invariably exhibit +some degree of beauty. From the numerous cases +of gradation in the amount of difference between the +sexes, and from the prevalence of the same general type +of coloration throughout the whole of the same group, +we may conclude that the causes, whatever they may +be, which have determined the brilliant colouring of the +males alone of some species, and of both sexes in a more +or less equal degree of other species, have generally +been the same.</p> + +<p>As so many gorgeous butterflies inhabit the tropics, it +has often been supposed that they owe their colours to +the great heat and moisture of these zones; but Mr. +Bates<a name="FNanchor_505" id="FNanchor_505"></a><a href="#Footnote_505" class="fnanchor">505</a> has shewn by the comparison of various closely-allied +groups of insects from the temperate and tropical +regions, that this view cannot be maintained; and the +evidence becomes conclusive when brilliantly-coloured +males and plain-coloured females of the same species +inhabit the same district, feed on the same food, and +follow exactly the same habits of life. Even when +the sexes resemble each other, we can hardly believe +that their brilliant and beautifully-arranged colours are +the purposeless result of the nature of the tissues, and +the action of the surrounding conditions.</p> + +<p>With animals of all kinds, whenever colour has been +modified for some special purpose, this has been, as far +<span class='pagenum'><a name="Page_392" id="Page_392">392</a></span>as we can judge, either for protection or as an attraction +between the sexes. With many species of butterflies +the upper surfaces of the wings are obscurely coloured, +and this in all probability leads to their escaping observation +and danger. But butterflies when at rest +would be particularly liable to be attacked by their +enemies; and almost all the kinds when resting raise +their wings vertically over their backs, so that the lower +sides alone are exposed to view. Hence it is this side +which in many cases is obviously coloured so as to +imitate the surfaces on which these insects commonly +rest. Dr. Rössler, I believe, first noticed the similarity +of the closed wings of certain Vanessæ and other butterflies +to the bark of trees. Many analogous and striking +facts could be given. The most interesting one is that +recorded by Mr. Wallace<a name="FNanchor_506" id="FNanchor_506"></a><a href="#Footnote_506" class="fnanchor">506</a> of a common Indian and +Sumatran butterfly (Kallima), which disappears like +magic when it settles in a bush; for it hides its head +and antennæ between its closed wings, and these in +form, colour, and veining cannot be distinguished from +a withered leaf together with the footstalk. In some +other cases the lower surfaces of the wings are brilliantly +coloured, and yet are protective; thus in <i>Thecla rubi</i> +the wings when closed are of an emerald green and resemble +the young leaves of the bramble, on which this +butterfly in the spring may often be seen seated.</p> + +<p>Although the obscure tints of the upper or under +surface of many butterflies no doubt serve to conceal +them, yet we cannot possibly extend this view to +the brilliant and conspicuous colours of many kinds, +such as our admiral and peacock Vanessæ, our white +<span class='pagenum'><a name="Page_393" id="Page_393">393</a></span>cabbage-butterflies (Pieris), or the great swallow-tail +Papilio which haunts the open fens—for these butterflies +are thus rendered visible to every living creature. +With these species both sexes are alike; but in the +common brimstone butterfly (<i>Gonepteryx rhamni</i>), the +male is of an intense yellow, whilst the female is much +paler; and in the orange-tip (<i>Anthocharis cardamines</i>) +the males alone have the bright orange tips to their +wings. In these cases the males and females are +equally conspicuous, and it is not credible that their +difference in colour stands in any relation to ordinary +protection. Nevertheless it is possible that the conspicuous +colours of many species may be in an indirect +manner beneficial, as will hereafter be explained, by +leading their enemies at once to recognise them as +unpalatable. Even in this case it does not certainly +follow that their bright colours and beautiful patterns +were acquired for this special purpose. In some other +remarkable cases, beauty has been gained for the sake +of protection, through the imitation of other beautiful +species, which inhabit the same district and enjoy an +immunity from attack by being in some way offensive +to their enemies.</p> + +<p class="tb">The female of our orange-tip butterfly, above referred +to, and of an American species (<i>Anth. genutia</i>) +probably shew us, as Mr. Walsh has remarked to me, +the primordial colours of the parent-species of the +genus; for both sexes of four or five widely-distributed +species are coloured in nearly the same manner. We +may infer here, as in several previous cases, that it is +the males of <i>Anth. cardamines</i> and <i>genutia</i> which have +departed from the usual type of colouring of their genus. +In the <i>Anth. sara</i> from California, the orange-tips have +become partially developed in the female; for her wings +are tipped with reddish-orange, but paler than in the<span class='pagenum'><a name="Page_394" id="Page_394">394</a></span> +male, and slightly different in some other respects. In +an allied Indian form, the <i>Iphias glaucippe</i>, the orange-tips +are fully developed in both sexes. In this Iphias +the under surface of the wings marvellously resembles, +as pointed out to me by Mr. A. Butler, a pale-coloured +leaf; and in our English orange-tip, the under surface +resembles the flower-head of the wild parsley, on which +it may be seen going to rest at night.<a name="FNanchor_507" id="FNanchor_507"></a><a href="#Footnote_507" class="fnanchor">507</a> The same +reasoning power which compels us to believe that the +lower surfaces have here been coloured for the sake of +protection, leads us to deny that the wings have been +tipped, especially when this character is confined to the +males, with bright orange for the same purpose.</p> + +<p class="tb">Turning now to Moths: most of these rest motionless +with their wings depressed during the whole or +greater part of the day; and the upper surfaces of their +wings are often shaded and coloured in an admirable +manner, as Mr. Wallace has remarked, for escaping +detection. With most of the Bombycidæ and Noctuidæ,<a name="FNanchor_508" id="FNanchor_508"></a><a href="#Footnote_508" class="fnanchor">508</a> +when at rest, the front-wings overlap and +conceal the hind-wings; so that the latter might be +brightly coloured without much risk; and they +are thus coloured in many species of both families. +During the act of flight, moths would often be able +to escape from their enemies; nevertheless, as the +hind-wings are then fully exposed to view, their bright +colours must generally have been acquired at the +cost of some little risk. But the following fact shews +us how cautious we ought to be in drawing conclusions +on this head. The common yellow under-wings +<span class='pagenum'><a name="Page_395" id="Page_395">395</a></span>(Triphaena) often fly about during the day or early +evening, and are then conspicuous from the colour of +their hind-wings. It would naturally be thought that +this would be a source of danger; but Mr. J. Jenner +Weir believes that it actually serves them as a means +of escape, for birds strike at these brightly coloured and +fragile surfaces, instead of at the body. For instance, +Mr. Weir turned into his aviary a vigorous specimen of +<i>Triphaena pronuba</i>, which was instantly pursued by +a robin; but the bird’s attention being caught by the +coloured wings, the moth was not captured until after +about fifty attempts, and small portions of the wings +were repeatedly broken off. He tried the same experiment, +in the open air, with a <i>T. fimbria</i> and swallow; +but the large size of this moth probably interfered +with its capture.<a name="FNanchor_509" id="FNanchor_509"></a><a href="#Footnote_509" class="fnanchor">509</a> We are thus reminded of a statement +made by Mr. Wallace,<a name="FNanchor_510" id="FNanchor_510"></a><a href="#Footnote_510" class="fnanchor">510</a> namely, that in the Brazilian +forests and Malayan islands, many common and +highly-decorated butterflies are weak flyers, though furnished +with a broad expanse of wings; and they “are +often captured with pierced and broken wings, as if +they had been seized by birds, from which they had +escaped: if the wings had been much smaller in proportion +to the body, it seems probable that the insect +would more frequently have been struck or pierced in +a vital part, and thus the increased expanse of the +wings may have been indirectly beneficial.”</p> + +<p class="tb"><i>Display.</i>—The bright colours of butterflies and of +some moths are specially arranged for display, whether +or not they serve in addition as a protection. Bright +<span class='pagenum'><a name="Page_396" id="Page_396">396</a></span>colours would not be visible during the night; and +there can be no doubt that moths, taken as a body, are +much less gaily decorated than butterflies, all of which +are diurnal in their habits. But the moths in certain +families, such as the Zygænidæ, various Sphingidæ, +Uraniidæ, some Arctiidæ and Saturniidæ, fly about +during the day or early evening, and many of these +are extremely beautiful, being far more brightly +coloured than the strictly nocturnal kinds. A few +exceptional cases, however, of brightly-coloured nocturnal +species have been recorded.<a name="FNanchor_511" id="FNanchor_511"></a><a href="#Footnote_511" class="fnanchor">511</a></p> + +<p>There is evidence of another kind in regard to display. +Butterflies, as before remarked, elevate their wings +when at rest, and whilst basking in the sunshine often +alternately raise and depress them, thus exposing to full +view both surfaces; and although the lower surface is +often coloured in an obscure manner as a protection, +yet in many species it is as highly coloured as the +upper surface, and sometimes in a very different manner. +In some tropical species the lower surface is even +more brilliantly coloured than the upper.<a name="FNanchor_512" id="FNanchor_512"></a><a href="#Footnote_512" class="fnanchor">512</a> In one +English fritillary, the <i>Argynnis aglaia</i>, the lower surface +alone is ornamented with shining silver discs. +Nevertheless, as a general rule, the upper surface, +which is probably the most fully exposed, is coloured +more brightly and in a more diversified manner than +the lower. Hence the lower surface generally affords +<span class='pagenum'><a name="Page_397" id="Page_397">397</a></span>to entomologists the most useful character for detecting +the affinities of the various species.</p> + +<p>Now if we turn to the enormous group of moths, +which do not habitually expose to full view the under +surface of their wings, this side is very rarely, as I hear +from Mr. Stainton, coloured more brightly than the +upper side, or even with equal brightness. Some exceptions +to the rule, either real or apparent, must be +noticed, as that of Hypopira, specified by Mr. Wormald.<a name="FNanchor_513" id="FNanchor_513"></a><a href="#Footnote_513" class="fnanchor">513</a> +Mr. R. Trimen informs me that in Guenée’s great work, +three moths are figured, in which the under surface is +much the most brilliant. For instance, in the Australian +Gastrophora the upper surface of the fore-wing is pale +greyish-ochreous, while the lower surface is magnificently +ornamented by an ocellus of cobalt-blue, placed in the +midst of a black mark, surrounded by orange-yellow, +and this by bluish-white. But the habits of these +three moths are unknown; so that no explanation +can be given of their unusual style of colouring. Mr. +Trimen also informs me that the lower surface of the +wings in certain other Geometræ<a name="FNanchor_514" id="FNanchor_514"></a><a href="#Footnote_514" class="fnanchor">514</a> and quadrifid Noctuæ +are either more variegated or more brightly-coloured +than the upper surface; but some of these species have +the habit of “holding their wings quite erect over their +backs, retaining them in this position for a considerable +time,” and thus exposing to view the under surface. +Other species when settled on the ground or herbage +have the habit of now and then suddenly and slightly +lifting up their wings. Hence the lower surface of the +wings being more brightly-coloured than the upper sur<span class='pagenum'><a name="Page_398" id="Page_398">398</a></span>face +in certain moths is not so anomalous a circumstance +as it at first appears. The Saturniidæ include +some of the most beautiful of all moths, their wings +being decorated, as in our British Emperor moth, with +fine ocelli; and Mr. T. W. Wood<a name="FNanchor_515" id="FNanchor_515"></a><a href="#Footnote_515" class="fnanchor">515</a> observes that they +resemble butterflies in some of their movements; “for +instance, in the gentle waving up and down of the +wings, as if for display, which is more characteristic +of diurnal than of nocturnal Lepidoptera.”</p> + +<p>It is a singular fact that no British moths, nor as +far as I can discover hardly any foreign species, which +are brilliantly coloured, differ much in colour according +to sex; though this is the case with many brilliant butterflies. +The male, however, of one American moth, the +<i>Saturnia Io</i>, is described as having its fore-wings deep +yellow, curiously marked with purplish-red spots; whilst +the wings of the female are purple-brown, marked with +grey lines.<a name="FNanchor_516" id="FNanchor_516"></a><a href="#Footnote_516" class="fnanchor">516</a> The British moths which differ sexually in +colour are all brown, or various tints of dull yellow, or +nearly white. In several species the males are much +darker than the females,<a name="FNanchor_517" id="FNanchor_517"></a><a href="#Footnote_517" class="fnanchor">517</a> and these belong to groups +which generally fly about during the afternoon. On the +other hand, in many genera, as Mr. Stainton informs me, +<span class='pagenum'><a name="Page_399" id="Page_399">399</a></span>the males have the hind-wings whiter than those of +the female—of which, fact <i>Agrotis exclamationis</i> offers a +good instance. The males are thus rendered more +conspicuous than the females, whilst flying about in +the dusk. In the Ghost Moth (<i>Hepialus humuli</i>) the +difference is more strongly marked; the males being +white and the females yellow with darker markings. +It is difficult to conjecture what the meaning can be +of these differences between the sexes in the shades of +darkness or lightness; but we can hardly suppose that +they are the result of mere variability with sexually-limited +inheritance, independently of any benefit thus +derived.</p> + +<p>From the foregoing statements it is impossible to +admit that the brilliant colours of butterflies and of +some few moths, have commonly been acquired for the +sake of protection. We have seen that their colours +and elegant patterns are arranged and exhibited as +if for display. Hence I am led to suppose that the +females generally prefer, or are most excited by the +more brilliant males; for on any other supposition +the males would be ornamented, as far as we can +see, for no purpose. We know that ants and certain +lamellicorn beetles are capable of feeling an attachment +for each other, and that ants recognise their fellows +after an interval of several months. Hence there is no +abstract improbability in the Lepidoptera, which probably +stand nearly or quite as high in the scale as these +insects, having sufficient mental capacity to admire +bright colours. They certainly discover flowers by +colour, and, as I have elsewhere shewn, the plants +which are fertilised exclusively by the wind never have +a conspicuously-coloured corolla. The Humming-bird +Sphinx may often be seen to swoop down from a distance +on a bunch of flowers in the midst of green foliage;<span class='pagenum'><a name="Page_400" id="Page_400">400</a></span> +and I have been assured by a friend, that these moths +repeatedly visited flowers painted on the walls of a room +in the South of France. The common white butterfly, +as I hear from Mr. Doubleday, often flies down to a bit +of paper on the ground, no doubt mistaking it for one of +its own species. Mr. Collingwood<a name="FNanchor_518" id="FNanchor_518"></a><a href="#Footnote_518" class="fnanchor">518</a> in speaking of the +difficulty of collecting certain butterflies in the Malay +Archipelago, states that “a dead specimen pinned upon +a conspicuous twig will often arrest an insect of the +same species in its headlong flight, and bring it down +within easy reach of the net, especially if it be of the +opposite sex.”</p> + +<p>The courtship of butterflies is a prolonged affair. The +males sometimes fight together in rivalry; and many +may be seen pursuing or crowding round the same +female. If, then, the females do not prefer one male to +another, the pairing must be left to mere chance, and +this does not appear to me a probable event. If, on the +other hand, the females habitually, or even occasionally, +prefer the more beautiful males, the colours of the latter +will have been rendered brighter by degrees, and will +have been transmitted to both sexes or to one sex, +according to which law of inheritance prevailed. The +process of sexual selection will have been much facilitated, +if the conclusions arrived at from various kinds of +evidence in the supplement to the ninth chapter can be +trusted; namely that the males of many Lepidoptera, +at least in the imago state, greatly exceed in number +the females.</p> + +<p>Some facts, however, are opposed to the belief that +female butterflies prefer the more beautiful males; thus, +as I have been assured by several observers, fresh females +may frequently be seen paired with battered, faded or +<span class='pagenum'><a name="Page_401" id="Page_401">401</a></span>dingy males; but this is a circumstance which could +hardly fail often to follow from the males emerging +from their cocoons earlier than the females. With moths +of the family of the Bombycidæ, the sexes pair immediately +after assuming the imago state; for they cannot +feed, owing to the rudimentary condition of their mouths. +The females, as several entomologists have remarked +to me, lie in an almost torpid state, and appear not to +evince the least choice in regard to their partners, +This is the case with the common silk-moth (<i>B. mori</i>), +as I have been told by some continental and English +breeders. Dr. Wallace, who has had such immense +experience in breeding <i>Bombyx cynthia</i>, is convinced +that the females evince no choice or preference. He +has kept above 300 of these moths living together, and +has often found the most vigorous females mated with +stunted males. The reverse apparently seldom occurs; +for, as he believes, the more vigorous males pass over the +weakly females, being attracted by those endowed with +most vitality. Although we have been indirectly induced +to believe that the females of many species prefer +the more beautiful males, I have no reason to suspect, +either with moths or butterflies, that the males are +attracted by the beauty of the females. If the more +beautiful females had been continually preferred, it is +almost certain, from the colours of butterflies being so +frequently transmitted to one sex alone, that the females +would often have been rendered more beautiful than +their male partners. But this does not occur except in +a few instances; and these can be explained, as we +shall presently see, on the principle of mimickry and +protection.</p> + +<p class="tb">As sexual selection primarily depends on variability, +a few words must be added on this subject. In respect<span class='pagenum'><a name="Page_402" id="Page_402">402</a></span> +to colour there is no difficulty, as any number of highly +variable Lepidoptera could be named. One good instance +will suffice. Mr. Bates shewed me a whole series +of specimens of <i>Papilio sesostris</i> and <i>childrenæ</i>; in the +latter the males varied much in the extent of the beautifully +enamelled green patch on the fore-wings, and +in the size of the white mark, as well as of the splendid +crimson stripe on the hind-wings; so that there was +a great contrast between the most and least gaudy +males. The male of <i>Papilio sesostris</i>, though a beautiful +insect, is much less so than <i>P. childrenæ</i>. It likewise +varies a little in the size of the green patch on the fore-wings, +and in the occasional appearance of a small +crimson stripe on the hind-wings, borrowed, as it would +seem, from its own female; for the females of this and +of many other species in the Æneas group possess this +crimson stripe. Hence between the brightest specimens +of <i>P. sesostris</i> and the least bright of <i>P. childrenæ</i>, there +was but a small interval; and it was evident that as far +as mere variability is concerned, there would be no +difficulty in permanently increasing by means of selection +the beauty of either species. The variability is +here almost confined to the male sex; but Mr. Wallace +and Mr. Bates have shewn<a name="FNanchor_519" id="FNanchor_519"></a><a href="#Footnote_519" class="fnanchor">519</a> that the females of some +other species are extremely variable, the males being +nearly constant. As I have before mentioned the Ghost +Moth (<i>Hepialus humuli</i>) as one of the best instances in +Britain of a difference in colour between the sexes of +moths, it may be worth adding<a name="FNanchor_520" id="FNanchor_520"></a><a href="#Footnote_520" class="fnanchor">520</a> that in the Shetland +<span class='pagenum'><a name="Page_403" id="Page_403">403</a></span>Islands, males are frequently found which closely +resemble the females. In a future chapter I shall have +occasion to shew that the beautiful eye-like spots or +ocelli, so common on the wings of many Lepidoptera, +are eminently variable.</p> + +<p>On the whole, although many serious objections may +be urged, it seems probable that most of the species of +Lepidoptera which are brilliantly coloured, owe their +colours to sexual selection, excepting in certain cases, +presently to be mentioned, in which conspicuous colours +are beneficial as a protection. From the ardour of the +male throughout the animal kingdom, he is generally +willing to accept any female; and it is the female which +usually exerts a choice. Hence if sexual selection has +here acted, the male, when the sexes differ, ought to be +the most brilliantly coloured; and this undoubtedly is +the ordinary rule. When the sexes are brilliantly coloured +and resemble each other, the characters acquired +by the males appear to have been transmitted to both +sexes. But will this explanation of the similarity and +dissimilarity in colour between the sexes suffice?</p> + +<p>The males and females of the same species of butterfly +are known<a name="FNanchor_521" id="FNanchor_521"></a><a href="#Footnote_521" class="fnanchor">521</a> in several cases to inhabit different stations, +the former commonly basking in the sunshine, the latter +haunting gloomy forests. It is therefore possible that +different conditions of life may have acted directly on +the two sexes; but this is not probable,<a name="FNanchor_522" id="FNanchor_522"></a><a href="#Footnote_522" class="fnanchor">522</a> as in the adult +state they are exposed during a very short period to +different conditions; and the larvæ of both are exposed +to the same conditions. Mr. Wallace believes +<span class='pagenum'><a name="Page_404" id="Page_404">404</a></span>that the less brilliant colours of the female have been +specially gained in all or almost all cases for the sake +of protection. On the contrary it seems to me more +probable that the males alone, in the large majority of +cases, have acquired their bright colours through sexual +selection, the females having been but little modified. +Consequently the females of distinct but allied species +ought to resemble each other much more closely than +do the males of the same species; and this is the general +rule. The females thus approximately show us the primordial +colouring of the parent-species of the group to +which they belong. They have, however, almost always +been modified to a certain extent by some of the successive +steps of variation, through the accumulation of +which the males were rendered beautiful, having been +transferred to them. The males and females of allied +though distinct species will also generally have been +exposed during their prolonged larval state to different +conditions, and may have been thus indirectly affected; +though with the males any slight change of colour thus +caused will often have been completely masked by the +brilliant tints gained through sexual selection. When +we treat of Birds, I shall have to discuss the whole +question whether the differences in colour between the +males and females have been in part specially gained +by the latter as a protection; so that I will here only +give unavoidable details.</p> + +<p>In all cases when the more common form of equal +inheritance by both sexes has prevailed, the selection of +bright-coloured males would tend to make the females +bright-coloured; and the selection of dull-coloured females +would tend to make the males dull. If both processes +were carried on simultaneously, they would tend +to neutralise each other. As far as I can see, it would +be extremely difficult to change through selection the<span class='pagenum'><a name="Page_405" id="Page_405">405</a></span> +one form of inheritance into the other. But by the +selection of successive variations, which were from the +first sexually limited in their transmission, there would +not be the slightest difficulty in giving bright colours to +the males alone, and at the same time or subsequently, +dull colours to the females alone. In this latter manner +female butterflies and moths may, as I fully admit, have +been rendered inconspicuous for the sake of protection, +and widely different from their males.</p> + +<p>Mr. Wallace<a name="FNanchor_523" id="FNanchor_523"></a><a href="#Footnote_523" class="fnanchor">523</a> has argued with much force in favour +of his view that when the sexes differ, the female has +been specially modified for the sake of protection; and +that this has been effected by one form of inheritance, +namely, the transmission of characters to both sexes, +having been changed through the agency of natural +selection into the other form, namely, transmission to +one sex. I was at first strongly inclined to accept this +view; but the more I have studied the various classes +throughout the animal kingdom, the less probable it +has appeared. Mr. Wallace urges that both sexes of +the <i>Heliconidæ</i>, <i>Danaidæ</i>, <i>Acroeidæ</i> are equally brilliant +because both are protected from the attacks of birds +and other enemies, by their offensive odour; but that +in other groups, which do not possess this immunity, +the females have been rendered inconspicuous, from +having more need of protection than the males. This +supposed difference in the “need of protection by the +two sexes” is rather deceptive, and requires some +discussion. It is obvious that brightly-coloured individuals, +whether males or females, would equally attract, +and obscurely-coloured individuals equally escape, the +<span class='pagenum'><a name="Page_406" id="Page_406">406</a></span>attention of their enemies. But we are concerned +with the effects of the destruction or preservation of +certain individuals of either sex, on the character of +the race. With insects, after the male has fertilised +the female, and after the latter has laid her eggs, +the greater or less immunity from danger of either sex +could not possibly have any effect on the offspring. +Before the sexes have performed their proper functions, +if they existed in equal numbers and if they strictly +paired (all other circumstances being the same), the +preservation of the males and females would be equally +important for the existence of the species and for the +character of the offspring. But with most animals, as +is known to be the case with the domestic silk-moth, +the male can fertilise two or three females; so that the +destruction of the males would not be so injurious to +the species as that of the females. On the other hand, +Dr. Wallace believes that with moths the progeny from +a second or third fertilisation is apt to be weakly, and +therefore would not have so good chance of surviving. +When the males exist in much greater numbers than the +females, no doubt many males might be destroyed with +impunity to the species; but I cannot see that the +results of ordinary selection for the sake of protection +would be influenced by the sexes existing in unequal +numbers; for the same proportion of the more conspicuous +individuals, whether males or females, would +probably be destroyed. If indeed the males presented +a greater range of variation in colour, the result would +be different; but we need not here follow out such complex +details. On the whole I cannot perceive that an +inequality in the numbers of the two sexes would influence +in any marked manner the effects of ordinary +selection on the character of the offspring.</p> + +<p>Female Lepidoptera require, as Mr. Wallace insists,<span class='pagenum'><a name="Page_407" id="Page_407">407</a></span> +some days to deposit their fertilised ova and to search +for a proper place; during this period (whilst the life +of the male was of no importance) the brighter-coloured +females would be exposed to danger and would be +liable to be destroyed. The duller-coloured females on +the other hand would survive, and thus would influence, +it might be thought, in a marked manner the +character of the species,—either of both sexes or of +one sex, according to which form of inheritance prevailed. +But it must not be forgotten that the males +emerge from the cocoon-state some days before the +females, and during this period, whilst the unborn +females were safe, the brighter-coloured males would be +exposed to danger; so that ultimately both sexes would +probably be exposed during a nearly equal length of +time to danger, and the elimination of conspicuous +colours would not be much more effective in the one +than the other sex.</p> + +<p>It is a more important consideration that female +Lepidoptera, as Mr. Wallace remarks, and as is known +to every collector, are generally slower flyers than +the males. Consequently the latter, if exposed to +greater danger from being conspicuously coloured, +might be able to escape from their enemies, whilst the +similarly-coloured females would be destroyed; and thus +the females would have the most influence in modifying +the colour of their progeny.</p> + +<p>There is one other consideration: bright colours, as +far as sexual selection is concerned, are commonly of +no service to the females; so that if the latter varied +in brightness, and the variations were sexually limited +in their transmission, it would depend on mere chance +whether the females had their bright colours increased; +and this would tend throughout the Order to diminish +the number of species with brightly-coloured females<span class='pagenum'><a name="Page_408" id="Page_408">408</a></span> +in comparison with the species having brightly-coloured +males. On the other hand, as bright colours are supposed +to be highly serviceable to the males in their +love-struggles, the brighter males (as we shall see +in the chapter on Birds) although exposed to rather +greater danger, would on an average procreate a greater +number of offspring than the duller males. In this +case, if the variations were limited in their transmission +to the male sex, the males alone would be rendered +more brilliantly coloured; but if the variations were +not thus limited, the preservation and augmentation of +such variations would depend on whether more evil was +caused to the species by the females being rendered +conspicuous, than good to the males by certain individuals +being successful over their rivals.</p> + +<p>As there can hardly be a doubt that both sexes of +many butterflies and moths have been rendered dull-coloured +for the sake of protection, so it may have +been with the females alone of some species in which +successive variations towards dullness first appeared +in the female sex and were from the first limited in +their transmission to the same sex. If not thus limited, +both sexes would become dull-coloured. We shall +immediately see, when we treat of mimickry, that +the females alone of certain butterflies have been rendered +extremely beautiful for the sake of protection, +without any of the successive protective variations +having been transferred to the male, to whom they +could not possibly have been in the least degree injurious, +and therefore could not have been eliminated +through natural selection. Whether in each particular +species, in which the sexes differ in colour, it is the +female which has been specially modified for the sake +of protection; or whether it is the male which has been +specially modified for the sake of sexual attraction, the<span class='pagenum'><a name="Page_409" id="Page_409">409</a></span> +female having retained her primordial colouring only +slightly changed through the agencies before alluded +to; or whether again both sexes have been modified, +the female for protection and the male for sexual attraction, +can only be definitely decided when we know the +life-history of each species.</p> + +<p>Without distinct evidence, I am unwilling to admit +that a double process of selection has long been going +on with a multitude of species,—the males having been +rendered more brilliant by beating their rivals; and the +females more dull-coloured by having escaped from their +enemies. We may take as an instance the common brimstone +butterfly (Gonepteryx), which appears early in the +spring before any other kind. The male of this species +is of a far more intense yellow than the female, though +she is almost equally conspicuous; and in this case it +does not seem probable that she specially acquired +her pale tints as a protection, though it is probable +that the male acquired his bright colours as a sexual +attraction. The female of <i>Anthocharis cardamines</i> does +not possess the beautiful orange tips to her wings with +which the male is ornamented; consequently she closely +resembles the white butterflies (Pieris) so common in +our gardens; but we have no evidence that this resemblance +is beneficial. On the contrary, as she resembles +both sexes of several species of the same genus inhabiting +various quarters of the world, it is more probable +that she has simply retained to a large extent her +primordial colours.</p> + +<p class="tb">Various facts support the conclusion that with the +greater number of brilliantly-coloured Lepidoptera, it +is the male which has been modified; the two sexes +having come to differ from each other, or to resemble +each other, according to which form of inheritance has +prevailed. Inheritance is governed by so many un<span class='pagenum'><a name="Page_410" id="Page_410">410</a></span>known +laws or conditions, that they seem to us to be +most capricious in their action;<a name="FNanchor_524" id="FNanchor_524"></a><a href="#Footnote_524" class="fnanchor">524</a> and we can so far +understand how it is that with closely-allied species the +sexes of some differ to an astonishing degree, whilst +the sexes of others are identical in colour. As the +successive steps in the process of variation are necessarily +all transmitted through the female, a greater +or less number of such steps might readily become +developed in her; and thus we can understand the +frequent gradations from an extreme difference to no +difference at all between the sexes of the species within +the same group. These cases of gradation are much +too common to favour the supposition that we here see +females actually undergoing the process of transition +and losing their brightness for the sake of protection; +for we have every reason to conclude that at any +one time the greater number of species are in a fixed +condition. With respect to the differences between the +females of the species in the same genus or family, we +can perceive that they depend, at least in part, on the +females partaking of the colours of their respective +males. This is well illustrated in those groups in which +the males are ornamented to an extraordinary degree; +for the females in these groups generally partake to a +certain extent of the splendour of their male partners. +Lastly, we continually find, as already remarked, that +the females of almost all the species in the same genus, +or even family, resemble each other much more closely +in colour than do the males; and this indicates that +the males have undergone a greater amount of modification +than the females.</p> + +<p class="tb"><span class='pagenum'><a name="Page_411" id="Page_411">411</a></span><i>Mimickry.</i>—This principle was first made clear in an +admirable paper by Mr. Bates,<a name="FNanchor_525" id="FNanchor_525"></a><a href="#Footnote_525" class="fnanchor">525</a> who thus threw a flood +of light on many obscure problems. It had previously +been observed that certain butterflies in S. America +belonging to quite distinct families, resembled the Heliconidæ +so closely in every stripe and shade of colour +that they could not be distinguished except by an +experienced entomologist. As the Heliconidæ are +coloured in their usual manner, whilst the others depart +from the usual colouring of the groups to which they +belong, it is clear that the latter are the imitators, and +the Heliconidæ the imitated. Mr. Bates further observed +that the imitating species are comparatively rare, whilst +the imitated swarm in large numbers; the two sets +living mingled together. From the fact of the Heliconidæ +being conspicuous and beautiful insects, yet +so numerous in individuals and species, he concluded +that they must be protected from the attacks of birds +by some secretion or odour; and this hypothesis has +now been confirmed by a considerable body of curious +evidence.<a name="FNanchor_526" id="FNanchor_526"></a><a href="#Footnote_526" class="fnanchor">526</a> From these considerations Mr. Bates inferred +that the butterflies which imitate the protected +species had acquired their present marvellously deceptive +appearance, through variation and natural selection, +in order to be mistaken for the protected kinds and +thus to escape being devoured. No explanation is +here attempted of the brilliant colours of the imitated, +but only of the imitating butterflies. We must +account for the colours of the former in the same +general manner, as in the cases previously discussed in +this chapter. Since the publication of Mr. Bates’ paper, +similar and equally striking facts have been observed +<span class='pagenum'><a name="Page_412" id="Page_412">412</a></span>by Mr. Wallace<a name="FNanchor_527" id="FNanchor_527"></a><a href="#Footnote_527" class="fnanchor">527</a> in the Malayan region, and by +Mr. Trimen in South Africa.</p> + +<p>As some writers<a name="FNanchor_528" id="FNanchor_528"></a><a href="#Footnote_528" class="fnanchor">528</a> have felt much difficulty in understanding +how the first steps in the process of mimickry +could have been effected through natural selection, it +may be well to remark that the process probably has +never commenced with forms widely dissimilar in colour. +But with two species moderately like each other, the +closest resemblance if beneficial to either form could +readily be thus gained; and if the imitated form was +subsequently and gradually modified through sexual +selection or any other means, the imitating form would +be led along the same track, and thus be modified to +almost any extent, so that it might ultimately assume +an appearance or colouring wholly unlike that of the +other members of the group to which it belonged. As +extremely slight variations in colour would not in many +cases suffice to render a species so like another protected +species as to lead to its preservation, it should +be remembered that many species of Lepidoptera are +liable to considerable and abrupt variations in colour. +A few instances have been given in this chapter; but +under this point of view Mr. Bates’ original paper on +mimickry, as well as Mr. Wallace’s papers, should be +consulted.</p> + +<p>In the foregoing cases both sexes of the imitating +species resemble the imitated; but occasionally the +<span class='pagenum'><a name="Page_413" id="Page_413">413</a></span>female alone mocks a brilliantly-coloured and protected +species inhabiting the same district. Consequently the +female differs in colour from her own male, and, which +is a rare and anomalous circumstance, is the more +brightly-coloured of the two. In all the few species of +Pieridæ, in which the female is more conspicuously +coloured than the male, she imitates, as I am informed +by Mr. Wallace, some protected species inhabiting the +same region. The female of <i>Diadema anomala</i> is rich +purple-brown with almost the whole surface glossed with +satiny blue, and she closely imitates the <i>Euplœa midamus</i>, +“one of the commonest butterflies of the East;” +whilst the male is bronzy or olive-brown, with only a +slight blue gloss on the outer parts of the wings.<a name="FNanchor_529" id="FNanchor_529"></a><a href="#Footnote_529" class="fnanchor">529</a> +Both sexes of this Diadema and of <i>D. bolina</i> follow +the same habits of life, so that the differences in colour +between the sexes cannot be accounted for by exposure +to different conditions;<a name="FNanchor_530" id="FNanchor_530"></a><a href="#Footnote_530" class="fnanchor">530</a> even if this explanation were +admissible in other instances.<a name="FNanchor_531" id="FNanchor_531"></a><a href="#Footnote_531" class="fnanchor">531</a></p> + +<p>The above cases of female butterflies which are more +brightly-coloured than the males, shew us, firstly, that +variations have arisen in a state of nature in the female +sex, and have been transmitted exclusively, or almost exclusively, +to the same sex; and, secondly, that this form +of inheritance has not been determined through natural +selection. For if we assume that the females, before +they became brightly coloured in imitation of some protected +kind, were exposed during each season for a longer +period to danger than the males; or if we assume that +<span class='pagenum'><a name="Page_414" id="Page_414">414</a></span>they could not escape so swiftly from their enemies, +we can understand how they alone might originally +have acquired through natural selection and sexually-limited +inheritance their present protective colours. +But except on the principle of these variations having +been transmitted exclusively to the female offspring, +we cannot understand why the males should have remained +dull-coloured; for it would surely not have +been in any way injurious to each individual male to +have partaken by inheritance of the protective colours +of the female, and thus to have had a better chance +of escaping destruction. In a group in which brilliant +colours are so common as with butterflies, it cannot be +supposed that the males have been kept dull-coloured +through sexual selection by the females rejecting the +individuals which were rendered as beautiful as themselves. +We may, therefore, conclude that in these cases +inheritance by one sex is not due to the modification +through natural selection of a tendency to equal inheritance +by both sexes.</p> + +<p class="tb">It may be well here to give an analogous case in +another Order, of characters acquired only by the female, +though not in the least injurious, as far as we can judge, +to the male. Amongst the Phasmidæ, or spectre-insects, +Mr. Wallace states that “it is often the females alone +that so strikingly resemble leaves, while the males show +only a rude approximation.” Now, whatever may be +the habits of these insects, it is highly improbable that +it could be disadvantageous to the males to escape detection +by resembling leaves.<a name="FNanchor_532" id="FNanchor_532"></a><a href="#Footnote_532" class="fnanchor">532</a> Hence we may conclude +<span class='pagenum'><a name="Page_415" id="Page_415">415</a></span>that the females alone in this latter as in the previous +cases originally varied in certain characters; these characters +having been preserved and augmented through +ordinary selection for the sake of protection and from +the first transmitted to the female offspring alone.</p> + +<p><i>Bright Colours of Caterpillars.</i>—Whilst reflecting on +the beauty of many butterflies, it occurred to me that +some caterpillars were splendidly coloured, and as +sexual selection could not possibly have here acted, +it appeared rash to attribute the beauty of the mature +insect to this agency, unless the bright colours of their +larvæ could be in some manner explained. In the first +place it may be observed that the colours of caterpillars +do not stand in any close correlation with those of the +mature insect. Secondly, their bright colours do not +<span class='pagenum'><a name="Page_416" id="Page_416">416</a></span>serve in any ordinary manner as a protection. As an +instance of this, Mr. Bates informs me that the most +conspicuous caterpillar which he ever beheld (that of a +Sphinx) lived on the large green leaves of a tree on the +open llanos of South America; it was about four inches +in length, transversely banded with black and yellow, +and with its head, legs, and tail of a bright red. Hence +it caught the eye of any man who passed by at the +distance of many yards, and no doubt of every passing +bird.</p> + +<p>I then applied to Mr. Wallace, who has an innate +genius for solving difficulties. After some consideration +he replied: “Most caterpillars require protection, as +may be inferred from some kinds being furnished +with spines or irritating hairs, and from many being +coloured green like the leaves on which they feed, +or curiously like the twigs of the trees on which they +live.” I may add as another instance of protection, +that there is a caterpillar of a moth, as I am informed +by Mr. J. Mansel Weale, which lives on the mimosas in +South Africa, and fabricates for itself a case, quite undistinguishable +from the surrounding thorns. From +such considerations Mr. Wallace thought it probable +that conspicuously-coloured caterpillars were protected +by having a nauseous taste; but as their skin is extremely +tender, and as their intestines readily protrude +from a wound, a slight peck from the beak of a bird +would be as fatal to them as if they had been devoured. +Hence, as Mr. Wallace remarks, “distastefulness alone +would be insufficient to protect a caterpillar unless +some outward sign indicated to its would-be destroyer +that its prey was a disgusting morsel.” Under these +circumstances it would be highly advantageous to a +caterpillar to be instantaneously and certainly recognised +as unpalatable by all birds and other animals.<span class='pagenum'><a name="Page_417" id="Page_417">417</a></span> +Thus the most gaudy colours would be serviceable, and +might have been gained by variation and the survival +of the most easily-recognised individuals.</p> + +<p class="tb">This hypothesis appears at first sight very bold; but +when it was brought before the Entomological Society<a name="FNanchor_533" id="FNanchor_533"></a><a href="#Footnote_533" class="fnanchor">533</a> +it was supported by various statements; and Mr. J. +Jenner Weir, who keeps a large number of birds in an +aviary, has made, as he informs me, numerous trials, +and finds no exception to the rule, that all caterpillars +of nocturnal and retiring habits with smooth skins, +all of a green colour, and all which imitate twigs, are +greedily devoured by his birds. The hairy and spinose +kinds are invariably rejected, as were four conspicuously-coloured +species. When the birds rejected a caterpillar, +they plainly shewed, by shaking their heads and cleansing +their beaks, that they were disgusted by the taste.<a name="FNanchor_534" id="FNanchor_534"></a><a href="#Footnote_534" class="fnanchor">534</a> +Three conspicuous kinds of caterpillars and moths were +also given by Mr. A. Butler to some lizards and frogs, +and were rejected; though other kinds were eagerly +eaten. Thus the probable truth of Mr. Wallace’s view +is confirmed, namely, that certain caterpillars have been +made conspicuous for their own good, so as to be easily +recognised by their enemies, on nearly the same principle +that certain poisons are coloured by druggists for +the good of man. This view will, it is probable, be +hereafter extended to many animals, which are coloured +in a conspicuous manner.</p> + +<p><i>Summary and Concluding Remarks on Insects.</i>—Looking +back to the several Orders, we have seen that +the sexes often differ in various characters, the meaning +<span class='pagenum'><a name="Page_418" id="Page_418">418</a></span>of which is not understood. The sexes, also, often differ +in their organs of sense or locomotion, so that the males +may quickly discover or reach the females, and still +oftener in the males possessing diversified contrivances +for retaining the females when found. But we are not +here much concerned with sexual differences of these +kinds.</p> + +<p>In almost all the Orders, the males of some species, +even of weak and delicate kinds, are known to be highly +pugnacious; and some few are furnished with special +weapons for fighting with their rivals. But the law of +battle does not prevail nearly so widely with insects as +with the higher animals. Hence probably it is that the +males have not often been rendered larger and stronger +than the females. On the contrary they are usually +smaller, in order that they may be developed within a +shorter time, so as to be ready in large numbers for the +emergence of the females.</p> + +<p>In two families of the Homoptera the males alone +possess, in an efficient state, organs which may be called +vocal; and in three families of the Orthoptera the males +alone possess stridulating organs. In both cases these +organs are incessantly used during the breeding-season, +not only for calling the females, but for charming or +exciting them in rivalry with other males. No one +who admits the agency of natural selection, will dispute +that these musical instruments have been acquired +through sexual selection. In four other Orders the +members of one sex, or more commonly of both sexes, +are provided with organs for producing various sounds, +which apparently serve merely as call-notes. Even +when both sexes are thus provided, the individuals +which were able to make the loudest or most continuous +noise would gain partners before those which were less +noisy, so that their organs have probably been gained<span class='pagenum'><a name="Page_419" id="Page_419">419</a></span> +through sexual selection. It is instructive to reflect +on the wonderful diversity of the means for producing +sound, possessed by the males alone or by both sexes +in no less than six Orders, and which were possessed +by at least one insect at an extremely remote geological +epoch. We thus learn how effectual sexual selection +has been in leading to modifications of structure, +which sometimes, as with the Homoptera, are of an important +nature.</p> + +<p>From the reasons assigned in the last chapter, it is +probable that the great horns of the males of many +lamellicorn, and some other beetles, have been acquired +as ornaments. So perhaps it may be with certain +other peculiarities confined to the male sex. From +the small size of insects, we are apt to undervalue their +appearance. If we could imagine a male Chalcosoma +(fig. <a href="#f15">15</a>) with its polished, bronzed coat of mail, and +vast complex horns, magnified to the size of a horse or +even of a dog, it would be one of the most imposing +animals in the world.</p> + +<p>The colouring of insects is a complex and obscure +subject. When the male differs slightly from the female, +and neither are brilliantly coloured, it is probable that +the two sexes have varied in a slightly different manner, +with the variations transmitted to the same sex, without +any benefit having been thus derived or evil suffered. +When the male is brilliantly coloured and differs conspicuously +from the female, as with some dragon-flies +and many butterflies, it is probable that he alone has +been modified, and that he owes his colours to sexual +selection; whilst the female has retained a primordial +or very ancient type of colouring, slightly modified by +the agencies before explained, and has therefore not +been rendered obscure, at least in most cases, for the +sake of protection. But the female alone has some<span class='pagenum'><a name="Page_420" id="Page_420">420</a></span>times +been coloured brilliantly so as to imitate other +protected species inhabiting the same district. When +the sexes resemble each other and both are obscurely +coloured, there is no doubt that they have been in a +multitude of cases coloured for the sake of protection. +So it is in some instances when both are brightly +coloured, causing them to resemble surrounding objects +such as flowers, or other protected species, or indirectly +by giving notice to their enemies that they are of an +unpalatable nature. In many other cases in which the +sexes resemble each other and are brilliantly coloured, +especially when the colours are arranged for display, we +may conclude that they have been gained by the male +sex as an attraction, and have been transferred to both +sexes. We are more especially led to this conclusion +whenever the same type of coloration prevails throughout +a group, and we find that the males of some species +differ widely in colour from the females, whilst both +sexes of other species are quite alike, with intermediate +gradations connecting these extreme states.</p> + +<p>In the same manner as bright colours have often +been partially transferred from the males to the females, +so it has been with the extraordinary horns of many +lamellicorn and some other beetles. So, again, the +vocal or instrumental organs proper to the males of +the Homoptera and Orthoptera have generally been +transferred in a rudimentary, or even in a nearly perfect +condition to the females; yet not sufficiently perfect to +be used for producing sound. It is also an interesting +fact, as bearing on sexual selection, that the stridulating +organs of certain male Orthoptera are not fully developed +until the last moult; and that the colours of certain +male dragon-flies are not fully developed until +some little time after their emergence from the pupal +state, and when they are ready to breed.</p> + +<p><span class='pagenum'><a name="Page_421" id="Page_421">421</a></span>Sexual selection implies that the more attractive +individuals are preferred by the opposite sex; and as +with insects, when the sexes differ, it is the male which, +with rare exceptions, is the most ornamented and +departs most from the type to which the species +belongs;—and as it is the male which searches eagerly +for the female, we must suppose that the females habitually +or occasionally prefer the more beautiful males, +and that these have thus acquired their beauty. That +in most or all the orders the females have the power +of rejecting any particular male, we may safely infer +from the many singular contrivances possessed by the +males, such as great jaws, adhesive cushions, spines, +elongated legs, &c., for seizing the female; for these +contrivances shew that there is some difficulty in the +act. In the case of unions between distinct species, +of which many instances have been recorded, the +female must have been a consenting party. Judging +from what we know of the perceptive powers and +affections of various insects, there is no antecedent improbability +in sexual selection having come largely into +action; but we have as yet no direct evidence on this +head, and some facts are opposed to the belief. Nevertheless, +when we see many males pursuing the same +female, we can hardly believe that the pairing is left to +blind chance—that the female exerts no choice, and +is not influenced by the gorgeous colours or other +ornaments, with which the male alone is decorated.</p> + +<p>If we admit that the females of the Homoptera and +Orthoptera appreciate the musical tones emitted by their +male partners, and that the various instruments for this +purpose have been perfected through sexual selection, +there is little improbability in the females of other +insects appreciating beauty in form or colour, and consequently +in such characters having been thus gained<span class='pagenum'><a name="Page_422" id="Page_422">422</a></span> +by the males. But from the circumstance of colour +being so variable, and from its having been so often +modified for the sake of protection, it is extremely +difficult to decide in how large a proportion of cases +sexual selection has come into play. This is more +especially difficult in those Orders, such as the Orthoptera, +Hymenoptera, and Coleoptera, in which the two +sexes rarely differ much in colour; for we are thus cut +off from our best evidence of some relation between the +reproduction of the species and colour. With the +Coleoptera, however, as before remarked, it is in the +great lamellicorn group, placed by some authors at +the head of the Order, and in which we sometimes +see a mutual attachment between the sexes, that we +find the males of some species possessing weapons for +sexual strife, others furnished with wonderful horns, +many with stridulating organs, and others ornamented +with splendid metallic tints. Hence it seems probable +that all these characters have been gained through +the same means, namely sexual selection.</p> + +<p>When we treat of Birds, we shall see that they present +in their secondary sexual characters the closest +analogy with insects. Thus, many male birds are +highly pugnacious, and some are furnished with special +weapons for fighting with their rivals. They possess +organs which are used during the breeding-season for +producing vocal and instrumental music. They are +frequently ornamented with combs, horns, wattles and +plumes of the most diversified kinds, and are decorated +with beautiful colours, all evidently for the sake of display. +We shall find that, as with insects, both sexes, +in certain groups, are equally beautiful, and are equally +provided with ornaments which are usually confined to +the male sex. In other groups both sexes are equally +plain-coloured and unornamented. Lastly, in some few<span class='pagenum'><a name="Page_423" id="Page_423">423</a></span> +anomalous cases, the females are more beautiful than +the males. We shall often find, in the same group of +birds, every gradation from no difference between the +sexes, to an extreme difference. In the latter case we +shall see that the females, like female insects, often +possess more or less plain traces of the characters which +properly belong to the males. The analogy, indeed, in +all these respects between birds and insects, is curiously +close. Whatever explanation applies to the one class +probably applies to the other; and this explanation, as +we shall hereafter attempt to shew, is almost certainly +sexual selection.</p> + +<h3>FOOTNOTES:</h3> + +<div class="footnote"><p><a name="Footnote_1" id="Footnote_1"></a><a href="#FNanchor_1"><span class="label">1</span></a> As the works of the first-named authors are so well known, I need +not give the titles; but as those of the latter are less well known in +England, I will give them:—‘Sechs Vorlesungen über die Darwin’sche +Theorie:’ zweite Auflage, 1868, von Dr. L. Büchner; translated +into French under the title ‘Conférences sur la Théorie Darwinienne,’ +1869. ‘Der Mensch, im Lichte der Darwin’sche Lehre,’ 1865, von +Dr. F. Rolle. I will not attempt to give references to all the authors +who have taken the same side of the question. Thus G. Canestrini +has published (‘Annuario della Soc. d. Nat.,’ Modena, 1867, p. 81) a +very curious paper on rudimentary characters, as bearing on the origin +of man. Another work has (1869) been published by Dr. Barrago +Francesco, bearing in Italian the title of “Man, made in the image of +God, was also made in the image of the ape.”</p></div> + +<div class="footnote"><p><a name="Footnote_2" id="Footnote_2"></a><a href="#FNanchor_2"><span class="label">2</span></a> Prof. Häckel is the sole author who, since the publication of the +‘Origin,’ has discussed, in his various works, in a very able manner, +the subject of sexual selection, and has seen its full importance.</p></div> + +<div class="footnote"><p><a name="Footnote_3" id="Footnote_3"></a><a href="#FNanchor_3"><span class="label">3</span></a> ‘Grosshirnwindungen des Menschen,’ 1868, s. 96.</p></div> + +<div class="footnote"><p><a name="Footnote_4" id="Footnote_4"></a><a href="#FNanchor_4"><span class="label">4</span></a> ‘Leç. sur la Phys.’ 1866, p. 890, as quoted by M. Dally, ‘L’Ordre +des Primates et le Transformisme,’ 1868, p. 29.</p></div> + +<div class="footnote"><p><a name="Footnote_5" id="Footnote_5"></a><a href="#FNanchor_5"><span class="label">5</span></a> ‘Naturgeschichte der Säugethiere von Paraguay,’ 1830, s. 50.</p></div> + +<div class="footnote"><p><a name="Footnote_6" id="Footnote_6"></a><a href="#FNanchor_6"><span class="label">6</span></a> Brehm, ‘Thierleben,’ B. i. 1864, s. 75, 86. On the Ateles, s. 105. +For other analogous statements, see s. 25, 107.</p></div> + +<div class="footnote"><p><a name="Footnote_7" id="Footnote_7"></a><a href="#FNanchor_7"><span class="label">7</span></a> With respect to insects see Dr. Laycock ‘On a General Law of +Vital Periodicity,’ British Association, 1842. Dr. Macculloch, ‘Silliman’s +North American Journal of Science,’ vol. xvii. p. 305, has seen +a dog suffering from tertian ague.</p></div> + +<div class="footnote"><p><a name="Footnote_8" id="Footnote_8"></a><a href="#FNanchor_8"><span class="label">8</span></a> I have given the evidence on this head in my ‘Variation of +Animals and Plants under Domestication,’ vol. ii. p. 15.</p></div> + +<div class="footnote"><p><a name="Footnote_9" id="Footnote_9"></a><a href="#FNanchor_9"><span class="label">9</span></a> “Mares e diversis generibus Quadrumanorum sine dubio dignoscunt +feminas humanas a maribus. Primum, credo, odoratu, postea aspectu. +Mr. Youatt, qui diu in Hortis Zoologicis (Bestiariis) medicus animalium +erat, vir in rebus observandis cautus et sagax, hoc mihi certissime +probavit, et curatores ejusdem loci et alii e ministris confirmaverunt. +Sir Andrew Smith et Brehm notabant idem in Cynocephalo. Illustrissimus +Cuvier etiam narrat multa de hac re quâ ut opinor nihil +turpius potest indicari inter omnia hominibus et Quadrumanis communia. +Narrat enim Cynocephalum quendam in furorem incidere +aspectu feminarum aliquarum, sed nequaquam accendi tanto furore +ab omnibus. Semper eligebat juniores, et dignoscebat in turba, et +advocabat voce gestuque.”</p></div> + +<div class="footnote"><p><a name="Footnote_10" id="Footnote_10"></a><a href="#FNanchor_10"><span class="label">10</span></a> This remark is made with respect to Cynocephalus and the anthropomorphous +apes by Geoffroy Saint-Hilaire and F. Cuvier, ‘Hist. +Nat. des Mammifères,’ tom. i. 1824.</p></div> + +<div class="footnote"><p><a name="Footnote_11" id="Footnote_11"></a><a href="#FNanchor_11"><span class="label">11</span></a> Huxley, ‘Man’s Place in Nature,’ 1863, p. 34.</p></div> + +<div class="footnote"><p><a name="Footnote_12" id="Footnote_12"></a><a href="#FNanchor_12"><span class="label">12</span></a> ‘Man’s Place in Nature,’ 1863, p. 67.</p></div> + +<div class="footnote"><p><a name="Footnote_13" id="Footnote_13"></a><a href="#FNanchor_13"><span class="label">13</span></a> The human embryo (upper fig.) is from Ecker, ‘Icones Phys.,’ +1851-1859, tab. xxx. fig. 2. This embryo was ten lines in length, so +that the drawing is much magnified. The embryo of the dog is from +Bischoff, ‘Entwicklungsgeschichte des Hunde-Eies,’ 1845, tab. xi. fig. +42 <span class="smcap">B</span>. This drawing is five times magnified, the embryo being 25 days +old. The internal viscera have been omitted, and the uterine appendages +in both drawings removed. I was directed to these figures by +Prof. Huxley, from whose work, ‘Man’s Place in Nature.’ the idea of +giving them was taken. Häckel has also given analogous drawings in +his ‘Schöpfungsgeschichte.’</p></div> + +<div class="footnote"><p><a name="Footnote_14" id="Footnote_14"></a><a href="#FNanchor_14"><span class="label">14</span></a> Prof. Wyman in ‘Proc. of American Acad. of Sciences,’ vol. iv. +1860, p. 17.</p></div> + +<div class="footnote"><p><a name="Footnote_15" id="Footnote_15"></a><a href="#FNanchor_15"><span class="label">15</span></a> Owen, ‘Anatomy of Vertebrates,’ vol. i. p. 533.</p></div> + +<div class="footnote"><p><a name="Footnote_16" id="Footnote_16"></a><a href="#FNanchor_16"><span class="label">16</span></a> ‘Die Grosshirnwindungen des Menschen,’ 1868, s. 95.</p></div> + +<div class="footnote"><p><a name="Footnote_17" id="Footnote_17"></a><a href="#FNanchor_17"><span class="label">17</span></a> ‘Anatomy of Vertebrates,’ vol. ii. p. 553.</p></div> + +<div class="footnote"><p><a name="Footnote_18" id="Footnote_18"></a><a href="#FNanchor_18"><span class="label">18</span></a> ‘Proc. Soc. Nat. Hist.’ Boston, 1863, vol. ix. p. 185.</p></div> + +<div class="footnote"><p><a name="Footnote_19" id="Footnote_19"></a><a href="#FNanchor_19"><span class="label">19</span></a> ‘Man’s Place in Nature,’ p. 65.</p></div> + +<div class="footnote"><p><a name="Footnote_20" id="Footnote_20"></a><a href="#FNanchor_20"><span class="label">20</span></a> I had written a rough copy of this chapter before reading a valuable +paper, “Caratteri rudimentali in ordine all’origine del uomo” +(‘Annuario della Soc. d. Nat.,’ Modena, 1867, p. 81), by G. Canestrini, +to which paper I am considerably indebted. Häckel has given admirable +discussions on this whole subject, under the title of Dysteleology, +in his ‘Generelle Morphologie’ and ‘Schöpfungsgeschichte.’</p></div> + +<div class="footnote"><p><a name="Footnote_21" id="Footnote_21"></a><a href="#FNanchor_21"><span class="label">21</span></a> Some good criticisms on this subject have been given by Messrs. +Murie and Mivart, in ‘Transact. Zoolog. Soc.’ 1869, vol. vii. p. 92.</p></div> + +<div class="footnote"><p><a name="Footnote_22" id="Footnote_22"></a><a href="#FNanchor_22"><span class="label">22</span></a> ‘Variation of Animals and Plants under Domestication,’ vol. ii. +pp. 317 and 397. See also ‘Origin of Species,’ 5th edit. p. 535.</p></div> + +<div class="footnote"><p><a name="Footnote_23" id="Footnote_23"></a><a href="#FNanchor_23"><span class="label">23</span></a> For instance M. Richard (‘Annales des Sciences Nat.’ 3rd series, +Zoolog. 1852, tom. xviii. p. 13) describes and figures rudiments of what +he calls the “muscle pédieux de la main,” which he says is sometimes +“infiniment petit.” Another muscle, called “le tibial postérieur,” is +generally quite absent in the hand, but appears from time to time in a +more or less rudimentary condition.</p></div> + +<div class="footnote"><p><a name="Footnote_24" id="Footnote_24"></a><a href="#FNanchor_24"><span class="label">24</span></a> Prof. W. Turner, ‘Proc. Royal Soc. Edinburgh,’ 1866-67, p. 65.</p></div> + +<div class="footnote"><p><a name="Footnote_25" id="Footnote_25"></a><a href="#FNanchor_25"><span class="label">25</span></a> Canestrini quotes Hyrt. (‘Annuario della Soc. dei Naturalisti,’ +Modena, 1867, p. 97) to the same effect.</p></div> + +<div class="footnote"><p><a name="Footnote_26" id="Footnote_26"></a><a href="#FNanchor_26"><span class="label">26</span></a> ‘The Diseases of the Ear,’ by J. Toynbee, F.R.S., 1860, p. 12.</p></div> + +<div class="footnote"><p><a name="Footnote_27" id="Footnote_27"></a><a href="#FNanchor_27"><span class="label">27</span></a> See also some remarks, and the drawings of the ears of the Lemuroidea, +in Messrs. Murie and Mivart’s excellent paper in ‘Transact. +Zoolog. Soc.’ vol. vii. 1869, pp. 6 and 90.</p></div> + +<div class="footnote"><p><a name="Footnote_28" id="Footnote_28"></a><a href="#FNanchor_28"><span class="label">28</span></a> Müller’s ‘Elements of Physiology,’ Eng. translat., 1842, vol. ii. p. +1117. Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 260; ibid. on the +Walrus, ‘Proc. Zoolog. Soc.’ November 8th, 1854. See also R. Knox, +'Great Artists and Anatomists,’ p. 106. This rudiment apparently is +somewhat larger in Negroes and Australians than in Europeans, see Carl +Vogt, ‘Lectures on Man,’ Eng. translat. p. 129.</p></div> + +<div class="footnote"><p><a name="Footnote_29" id="Footnote_29"></a><a href="#FNanchor_29"><span class="label">29</span></a> ‘The Physiology and Pathology of Mind,’ 2nd edit. 1868, p. 134.</p></div> + +<div class="footnote"><p><a name="Footnote_30" id="Footnote_30"></a><a href="#FNanchor_30"><span class="label">30</span></a> Eschricht, Ueber die Richtung der Haare am menschlichen Körper +'Müllers Archiv für Anat. und Phys.’ 1837, s. 47. I shall often have +to refer to this very curious paper.</p></div> + +<div class="footnote"><p><a name="Footnote_31" id="Footnote_31"></a><a href="#FNanchor_31"><span class="label">31</span></a> Paget, ‘Lectures on Surgical Pathology,’ 1853, vol. i. p. 71.</p></div> + +<div class="footnote"><p><a name="Footnote_32" id="Footnote_32"></a><a href="#FNanchor_32"><span class="label">32</span></a> Eschricht, ibid. s. 40, 47.</p></div> + +<div class="footnote"><p><a name="Footnote_33" id="Footnote_33"></a><a href="#FNanchor_33"><span class="label">33</span></a> Dr. Webb, ‘Teeth in Man and the Anthropoid Apes,’ as quoted by +Dr. C. Carter Blake in ‘Anthropological Review,’ July, 1867, p. 299.</p></div> + +<div class="footnote"><p><a name="Footnote_34" id="Footnote_34"></a><a href="#FNanchor_34"><span class="label">34</span></a> Owen, ‘Anatomy of Vertebrates,’ vol. iii. pp. 320, 321, and 325.</p></div> + +<div class="footnote"><p><a name="Footnote_35" id="Footnote_35"></a><a href="#FNanchor_35"><span class="label">35</span></a> ‘On the Primitive Form of the Skull,’ Eng. translat. in ‘Anthropological +Review,’ Oct. 1868, p. 426.</p></div> + +<div class="footnote"><p><a name="Footnote_36" id="Footnote_36"></a><a href="#FNanchor_36"><span class="label">36</span></a> Owen, ‘Anatomy of Vertebrates,’ vol. iii. pp. 416, 434, 441.</p></div> + +<div class="footnote"><p><a name="Footnote_37" id="Footnote_37"></a><a href="#FNanchor_37"><span class="label">37</span></a> ‘Annuario della Soc. d. Nat.’ Modena, 1867, p. 94.</p></div> + +<div class="footnote"><p><a name="Footnote_38" id="Footnote_38"></a><a href="#FNanchor_38"><span class="label">38</span></a> M. C. Martins (“De l’Unité Organique,” in ‘Revue des Deux +Mondes,’ June 15, 1862, p. 16), and Häckel (‘Generelle Morphologie,’ +B. ii. s. 278), have both remarked on the singular fact of this rudiment +sometimes causing death.</p></div> + +<div class="footnote"><p><a name="Footnote_39" id="Footnote_39"></a><a href="#FNanchor_39"><span class="label">39</span></a> ‘The Lancet,’ Jan. 24, 1863, p. 83. Dr. Knox, ‘Great Artists and +Anatomists,’ p. 63. See also an important memoir on this process by +Dr. Grube, in the ‘Bulletin de l’Acad. Imp. de St. Pétersbourg,’ tom. +xii. 1867, p. 448.</p></div> + +<div class="footnote"><p><a name="Footnote_40" id="Footnote_40"></a><a href="#FNanchor_40"><span class="label">40</span></a> “On the Caves of Gibraltar,” ‘Transact. Internat. Congress of +Prehist. Arch.’ Third Session, 1869, p. 54.</p></div> + +<div class="footnote"><p><a name="Footnote_41" id="Footnote_41"></a><a href="#FNanchor_41"><span class="label">41</span></a> Quatrefages has lately collected the evidence on this subject. +'Revue des Cours Scientifiques,’ 1867-1868, p. 625.</p></div> + +<div class="footnote"><p><a name="Footnote_42" id="Footnote_42"></a><a href="#FNanchor_42"><span class="label">42</span></a> Owen, ‘On the Nature of Limbs,’ 1849, p. 114.</p></div> + +<div class="footnote"><p><a name="Footnote_43" id="Footnote_43"></a><a href="#FNanchor_43"><span class="label">43</span></a> Leuckart, in Todd’s ‘Cyclop. of Anat.’ 1849-52, vol. iv. p. 1415. +In man this organ is only from three to six lines in length, but, like +so many other rudimentary parts, it is variable in development as well +as in other characters.</p></div> + +<div class="footnote"><p><a name="Footnote_44" id="Footnote_44"></a><a href="#FNanchor_44"><span class="label">44</span></a> See, on this subject, Owen, ‘Anatomy of Vertebrates,’ vol. iii. pp. +675, 676, 706.</p></div> + +<div class="footnote"><p><a name="Footnote_45" id="Footnote_45"></a><a href="#FNanchor_45"><span class="label">45</span></a> See the evidence on these points, as given by Lubbock, ‘Prehistoric +Times,’ p. 354, &c.</p></div> + +<div class="footnote"><p><a name="Footnote_46" id="Footnote_46"></a><a href="#FNanchor_46"><span class="label">46</span></a> ‘L’Instinct chez les Insectes.’ ‘Revue des Deux Mondes,’ Feb. +1870, p. 690.</p></div> + +<div class="footnote"><p><a name="Footnote_47" id="Footnote_47"></a><a href="#FNanchor_47"><span class="label">47</span></a> ‘The American Beaver and his Works,’ 1868.</p></div> + +<div class="footnote"><p><a name="Footnote_48" id="Footnote_48"></a><a href="#FNanchor_48"><span class="label">48</span></a> ‘The Principles of Psychology,’ 2nd edit. 1870, pp. 418-443.</p></div> + +<div class="footnote"><p><a name="Footnote_49" id="Footnote_49"></a><a href="#FNanchor_49"><span class="label">49</span></a> ‘Contributions to the Theory of Natural Selection,’ 1870, p. 212</p></div> + +<div class="footnote"><p><a name="Footnote_50" id="Footnote_50"></a><a href="#FNanchor_50"><span class="label">50</span></a> ‘Recherches sur les Mœurs des Fourmis,’ 1810, p. 173.</p></div> + +<div class="footnote"><p><a name="Footnote_51" id="Footnote_51"></a><a href="#FNanchor_51"><span class="label">51</span></a> All the following statements, given on the authority of these two +naturalists, are taken from Rengger’s ‘Naturges. der Säugethiere +von Paraguay,’ 1830, s. 41-57, and from Brehm’s ‘Thierleben,’ B. i. +s. 10-87.</p></div> + +<div class="footnote"><p><a name="Footnote_52" id="Footnote_52"></a><a href="#FNanchor_52"><span class="label">52</span></a> ‘Bridgewater Treatise,’ p. 263.</p></div> + +<div class="footnote"><p><a name="Footnote_53" id="Footnote_53"></a><a href="#FNanchor_53"><span class="label">53</span></a> W. C. L. Martin, ‘Nat. Hist. of Mammalia,’ 1841, p. 405.</p></div> + +<div class="footnote"><p><a name="Footnote_54" id="Footnote_54"></a><a href="#FNanchor_54"><span class="label">54</span></a> Quoted by Vogt, ‘Mémoire sur les Microcéphales,’ 1867, p. 168.</p></div> + +<div class="footnote"><p><a name="Footnote_55" id="Footnote_55"></a><a href="#FNanchor_55"><span class="label">55</span></a> ‘The Variation of Animals and Plants under Domestication,’ vol. +i. p. 27.</p></div> + +<div class="footnote"><p><a name="Footnote_56" id="Footnote_56"></a><a href="#FNanchor_56"><span class="label">56</span></a> ‘Les Mœurs des Fourmis,’ 1810, p. 150.</p></div> + +<div class="footnote"><p><a name="Footnote_57" id="Footnote_57"></a><a href="#FNanchor_57"><span class="label">57</span></a> Quoted in Dr. Maudsley’s ‘Physiology and Pathology of Mind,’ +1868, pp. 19, 220.</p></div> + +<div class="footnote"><p><a name="Footnote_58" id="Footnote_58"></a><a href="#FNanchor_58"><span class="label">58</span></a> Dr. Jerdon, ‘Birds of India,’ vol. i. 1862, p. xxi.</p></div> + +<div class="footnote"><p><a name="Footnote_59" id="Footnote_59"></a><a href="#FNanchor_59"><span class="label">59</span></a> Mr. L. H. Morgan’s work on ‘The American Beaver,’ 1868, offers +a good illustration of this remark. I cannot, however, avoid thinking +that he goes too far in underrating the power of Instinct.</p></div> + +<div class="footnote"><p><a name="Footnote_60" id="Footnote_60"></a><a href="#FNanchor_60"><span class="label">60</span></a> ‘The Moor and the Loch,’ p. 45. Col. Hutchinson on ‘Dog +Breaking,’ 1850, p. 46.</p></div> + +<div class="footnote"><p><a name="Footnote_61" id="Footnote_61"></a><a href="#FNanchor_61"><span class="label">61</span></a> ‘Personal Narrative,’ Eng. translat., vol. iii. p. 106.</p></div> + +<div class="footnote"><p><a name="Footnote_62" id="Footnote_62"></a><a href="#FNanchor_62"><span class="label">62</span></a> Quoted by Sir C. Lyell, ‘Antiquity of Man,’ p. 497.</p></div> + +<div class="footnote"><p><a name="Footnote_63" id="Footnote_63"></a><a href="#FNanchor_63"><span class="label">63</span></a> ‘Journal of Researches during the Voyage of the “Beagle,”’ 1845, +p. 398. ‘Origin of Species,’ 5th edit. p. 260.</p></div> + +<div class="footnote"><p><a name="Footnote_64" id="Footnote_64"></a><a href="#FNanchor_64"><span class="label">64</span></a> ‘Lettres Phil. sur l’Intelligence des Animaux,’ nouvelle edit. +1802, p. 86.</p></div> + +<div class="footnote"><p><a name="Footnote_65" id="Footnote_65"></a><a href="#FNanchor_65"><span class="label">65</span></a> See the evidence on this head in chap. i. vol. i. ‘On the Variation +of Animals and Plants under Domestication.’</p></div> + +<div class="footnote"><p><a name="Footnote_66" id="Footnote_66"></a><a href="#FNanchor_66"><span class="label">66</span></a> ‘Proc. Zoolog. Soc.’ 1864, p. 186.</p></div> + +<div class="footnote"><p><a name="Footnote_67" id="Footnote_67"></a><a href="#FNanchor_67"><span class="label">67</span></a> Savage and Wyman in ‘Boston Journal of Nat. Hist.’ vol. iv. +1843-44, p. 383.</p></div> + +<div class="footnote"><p><a name="Footnote_68" id="Footnote_68"></a><a href="#FNanchor_68"><span class="label">68</span></a> ‘Säugethiere von Paraguay,’ 1830, s. 51-56.</p></div> + +<div class="footnote"><p><a name="Footnote_69" id="Footnote_69"></a><a href="#FNanchor_69"><span class="label">69</span></a> ‘Thierleben,’ B. i. s. 79, 82.</p></div> + +<div class="footnote"><p><a name="Footnote_70" id="Footnote_70"></a><a href="#FNanchor_70"><span class="label">70</span></a> ‘The Malay Archipelago,’ vol. i. 1869, p. 87.</p></div> + +<div class="footnote"><p><a name="Footnote_71" id="Footnote_71"></a><a href="#FNanchor_71"><span class="label">71</span></a> ‘Primeval Man,’ 1869, pp. 145, 147.</p></div> + +<div class="footnote"><p><a name="Footnote_72" id="Footnote_72"></a><a href="#FNanchor_72"><span class="label">72</span></a> ‘Prehistoric Times,’ 1865, p. 473, &c.</p></div> + +<div class="footnote"><p><a name="Footnote_73" id="Footnote_73"></a><a href="#FNanchor_73"><span class="label">73</span></a> Quoted in ‘Anthropological Review,’ 1864, p. 158.</p></div> + +<div class="footnote"><p><a name="Footnote_74" id="Footnote_74"></a><a href="#FNanchor_74"><span class="label">74</span></a> Rengger, ibid. s. 45.</p></div> + +<div class="footnote"><p><a name="Footnote_75" id="Footnote_75"></a><a href="#FNanchor_75"><span class="label">75</span></a> See my ‘Variation of Animals and Plants under Domestication,’ +vol. i. p. 27.</p></div> + +<div class="footnote"><p><a name="Footnote_76" id="Footnote_76"></a><a href="#FNanchor_76"><span class="label">76</span></a> See a discussion on this subject in Mr. E. B. Tylor’s very interesting +work, ‘Researches into the Early History of Mankind,’ 1865, chaps. +ii. to iv.</p></div> + +<div class="footnote"><p><a name="Footnote_77" id="Footnote_77"></a><a href="#FNanchor_77"><span class="label">77</span></a> Hon. Daines Barrington in ‘Philosoph. Transactions,’ 1773, p. +262. See also Dureau de la Malle, in ‘Ann. des Sc. Nat.’ 3rd series, +Zoolog. tom. x. p. 119.</p></div> + +<div class="footnote"><p><a name="Footnote_78" id="Footnote_78"></a><a href="#FNanchor_78"><span class="label">78</span></a> ‘On the Origin of Language,’ by H. Wedgwood, 1866. ‘Chapters +on Language,’ by the Rev. F. W. Farrar, 1865. These works are most +interesting. See also ‘De la Phys. et de Parole,’ par Albert Lemoine, +1865, p. 190. The work on this subject, by the late Prof. Aug. Schleicher, +has been translated by Dr. Bikkers into English, under the title +of ‘Darwinism tested by the Science of Language,’ 1869.</p></div> + +<div class="footnote"><p><a name="Footnote_79" id="Footnote_79"></a><a href="#FNanchor_79"><span class="label">79</span></a> Vogt, ‘Mémoire sur les Microcéphales,’ 1867, p. 169. With +respect to savages, I have given some facts in my ‘Journal of +Researches,’ &c., 1845, p. 206.</p></div> + +<div class="footnote"><p><a name="Footnote_80" id="Footnote_80"></a><a href="#FNanchor_80"><span class="label">80</span></a> See clear evidence on this head in the two works so often quoted, +by Brehm and Rengger.</p></div> + +<div class="footnote"><p><a name="Footnote_81" id="Footnote_81"></a><a href="#FNanchor_81"><span class="label">81</span></a> See remarks on this head by Dr. Maudsley, ‘The Physiology +and Pathology of Mind,’ 2nd edit. 1868, p. 199.</p></div> + +<div class="footnote"><p><a name="Footnote_82" id="Footnote_82"></a><a href="#FNanchor_82"><span class="label">82</span></a> Many curious cases have been recorded. See, for instance, +'Inquiries Concerning the Intellectual Powers,’ by Dr. Abercrombie, +1838, p. 150.</p></div> + +<div class="footnote"><p><a name="Footnote_83" id="Footnote_83"></a><a href="#FNanchor_83"><span class="label">83</span></a> ‘The Variation of Animals and Plants under Domestication,’ vol. +ii. p. 6.</p></div> + +<div class="footnote"><p><a name="Footnote_84" id="Footnote_84"></a><a href="#FNanchor_84"><span class="label">84</span></a> See some good remarks to this effect by Dr. Maudsley, ‘The +Physiology and Pathology of Mind,’ 1808, p. 199.</p></div> + +<div class="footnote"><p><a name="Footnote_85" id="Footnote_85"></a><a href="#FNanchor_85"><span class="label">85</span></a> Macgillivray, ‘Hist. of British Birds,’ vol. ii. 1839, p. 29. An +excellent observer, Mr. Blackwall, remarks that the magpie learns to +pronounce single words, and even short sentences, more readily than +almost any other British bird; yet, as he adds, after long and closely +investigating its habits, he has never known it, in a state of nature, +display any unusual capacity for imitation. ‘Researches in Zoology,’ +1834, p. 158.</p></div> + +<div class="footnote"><p><a name="Footnote_86" id="Footnote_86"></a><a href="#FNanchor_86"><span class="label">86</span></a> See the very interesting parallelism between the development of +speech and languages, given by Sir C. Lyell in ‘The Geolog. Evidences +of the Antiquity of Man,’ 1863, chap. xxiii.</p></div> + +<div class="footnote"><p><a name="Footnote_87" id="Footnote_87"></a><a href="#FNanchor_87"><span class="label">87</span></a> See remarks to this effect by the Rev. F. W. Farrar, in an interesting +article, entitled “Philology and Darwinism” in ‘Nature,’ March +24th, 1870, p. 528.</p></div> + +<div class="footnote"><p><a name="Footnote_88" id="Footnote_88"></a><a href="#FNanchor_88"><span class="label">88</span></a> ‘Nature,’ Jan. 6th, 1870, p. 257.</p></div> + +<div class="footnote"><p><a name="Footnote_89" id="Footnote_89"></a><a href="#FNanchor_89"><span class="label">89</span></a> Quoted by C. S. Wake, ‘Chapters on Man,’ 1868, p. 101.</p></div> + +<div class="footnote"><p><a name="Footnote_90" id="Footnote_90"></a><a href="#FNanchor_90"><span class="label">90</span></a> Buckland, ‘Bridgewater Treatise,’ p. 411.</p></div> + +<div class="footnote"><p><a name="Footnote_91" id="Footnote_91"></a><a href="#FNanchor_91"><span class="label">91</span></a> See some good remarks on the simplification of languages, by Sir +J. Lubbock, ‘Origin of Civilisation,’ 1870, p. 278.</p></div> + +<div class="footnote"><p><a name="Footnote_92" id="Footnote_92"></a><a href="#FNanchor_92"><span class="label">92</span></a> ‘Conférences sur la Théorie Darwinienne,’ French translat., 1869, +p. 132.</p></div> + +<div class="footnote"><p><a name="Footnote_93" id="Footnote_93"></a><a href="#FNanchor_93"><span class="label">93</span></a> The Rev. Dr. J. M’Cann, ‘Anti-Darwinism,’ 1869, p. 13.</p></div> + +<div class="footnote"><p><a name="Footnote_94" id="Footnote_94"></a><a href="#FNanchor_94"><span class="label">94</span></a> ‘The Spectator,’ Dec. 4th, 1869, p. 1430.</p></div> + +<div class="footnote"><p><a name="Footnote_95" id="Footnote_95"></a><a href="#FNanchor_95"><span class="label">95</span></a> See an excellent article on this subject by the Rev. F. W. Farrar, +in the ‘Anthropological Review,’ Aug. 1864, p. ccxvii. For further +facts see Sir J. Lubbock, ‘Prehistoric Times,’ 2nd edit. 1869. p. 564; +and especially the chapters on Religion in his ‘Origin of Civilisation,’ +1870.</p></div> + +<div class="footnote"><p><a name="Footnote_96" id="Footnote_96"></a><a href="#FNanchor_96"><span class="label">96</span></a> The Worship of Animals and Plants, in the ‘Fortnightly Review,’ +Oct. 1, 1869, p. 422.</p></div> + +<div class="footnote"><p><a name="Footnote_97" id="Footnote_97"></a><a href="#FNanchor_97"><span class="label">97</span></a> Tylor, ‘Early History of Mankind,’ 1865, p. 6. See also the three +striking chapters on the Development of Religion, in Lubbock’s ‘Origin +of Civilisation,’ 1870. In a like manner Mr. Herbert Spencer, in his +ingenious essay in the ‘Fortnightly Review’ (May 1st, 1870, p. 535), +accounts for the earliest forms of religious belief throughout the world, +by man being led through dreams, shadows, and other causes, to look +at himself as a double essence, corporeal and spiritual. As the spiritual +being is supposed to exist after death and to be powerful, it is propitiated +by various gifts and ceremonies, and its aid invoked. He then +further shews that names or nicknames given from some animal or +other object to the early progenitors or founders of a tribe, are supposed +after a long interval to represent the real progenitor of the tribe; +and such animal or object is then naturally believed still to exist as +a spirit, is held sacred, and worshipped as a god. Nevertheless I cannot +but suspect that there is a still earlier and ruder stage, when anything +which manifests power or movement is thought to be endowed with +some form of life, and with mental faculties analogous to our own.</p></div> + +<div class="footnote"><p><a name="Footnote_98" id="Footnote_98"></a><a href="#FNanchor_98"><span class="label">98</span></a> See an able article on the Psychical Elements of Religion, by +Mr. L. Owen Pike, in ‘Anthropolog. Review,’ April, 1870, p. lxiii.</p></div> + +<div class="footnote"><p><a name="Footnote_99" id="Footnote_99"></a><a href="#FNanchor_99"><span class="label">99</span></a> ‘Religion, Moral, &c., der Darwin’schen Art-Lehre,’ 1869, s. 53.</p></div> + +<div class="footnote"><p><a name="Footnote_100" id="Footnote_100"></a><a href="#FNanchor_100"><span class="label">100</span></a> ‘Prehistoric Times,’ 2nd edit. p. 571. In this work (at p. 553) +there will be found an excellent account of the many strange and +capricious customs of savages.</p></div> + +<div class="footnote"><p><a name="Footnote_101" id="Footnote_101"></a><a href="#FNanchor_101"><span class="label">101</span></a> See, for instance, on this subject, Quatrefages, ‘Unité de l’Espèce +Humaine,’ 1861, p. 21, &c.</p></div> + +<div class="footnote"><p><a name="Footnote_102" id="Footnote_102"></a><a href="#FNanchor_102"><span class="label">102</span></a> ‘Dissertation on Ethical Philosophy,’ 1837, p. 231, &c.</p></div> + +<div class="footnote"><p><a name="Footnote_103" id="Footnote_103"></a><a href="#FNanchor_103"><span class="label">103</span></a> ‘Metaphysics of Ethics,’ translated by J. W. Semple, Edinburgh, +1836, p. 136.</p></div> + +<div class="footnote"><p><a name="Footnote_104" id="Footnote_104"></a><a href="#FNanchor_104"><span class="label">104</span></a> Mr. Bain gives a list (‘Mental and Moral Science,’ 1868, p. 543-725) +of twenty-six British authors who have written on this subject, +and whose names are familiar to every reader; to these, Mr. Bain’s own +name, and those of Mr. Lecky, Mr. Shadworth Hodgson, and Sir J. +Lubbock, as well as of others, may be added.</p></div> + +<div class="footnote"><p><a name="Footnote_105" id="Footnote_105"></a><a href="#FNanchor_105"><span class="label">105</span></a> Sir B. Brodie, after observing that man is a social animal (‘Psychological +Enquiries,’ 1854, p. 192), asks the pregnant question, +“ought not this to settle the disputed question as to the existence of a +moral sense?” Similar ideas have probably occurred to many persons, +as they did long ago to Marcus Aurelius. Mr. J. S. Mill speaks, in +his celebrated work, ‘Utilitarianism,’ (1864, p. 46), of the social feelings +as a “powerful natural sentiment,” and as “the natural basis of +sentiment for utilitarian morality;” but on the previous page he +says, “if, as is my own belief, the moral feelings are not innate, but +acquired, they are not for that reason less natural.” It is with hesitation +that I venture to differ from so profound a thinker, but it can +hardly be disputed that the social feelings are instinctive or innate +in the lower animals; and why should they not be so in man? +Mr. Bain (see, for instance, ‘The Emotions and the Will,’ 1865, p. 481) +and others believe that the moral sense is acquired by each individual +during his lifetime. On the general theory of evolution this is at +least extremely improbable.</p></div> + +<div class="footnote"><p><a name="Footnote_106" id="Footnote_106"></a><a href="#FNanchor_106"><span class="label">106</span></a> ‘Die Darwin’sche Theorie,’ s. 101.</p></div> + +<div class="footnote"><p><a name="Footnote_107" id="Footnote_107"></a><a href="#FNanchor_107"><span class="label">107</span></a> Mr. R. Browne in ‘Proc. Zoolog. Soc.’ 1868, p. 409.</p></div> + +<div class="footnote"><p><a name="Footnote_108" id="Footnote_108"></a><a href="#FNanchor_108"><span class="label">108</span></a> Brehm, ‘Thierleben,’ B. i. 1864, s. 52, 79. For the case of the +monkeys extracting thorns from each other, see s. 54. With respect to +the Hamadryas turning over stones, the fact is given (s. 76) on the +evidence of Alvarez, whose observations Brehm thinks quite trustworthy. +For the cases of the old male baboons attacking the dogs, see +s. 79; and with respect to the eagle, s. 56.</p></div> + +<div class="footnote"><p><a name="Footnote_109" id="Footnote_109"></a><a href="#FNanchor_109"><span class="label">109</span></a> ‘Annals and Mag. of Nat. Hist.’ November, 1868, p. 382.</p></div> + +<div class="footnote"><p><a name="Footnote_110" id="Footnote_110"></a><a href="#FNanchor_110"><span class="label">110</span></a> Sir J. Lubbock, ‘Prehistoric Times,’ 2nd edit. p. 446.</p></div> + +<div class="footnote"><p><a name="Footnote_111" id="Footnote_111"></a><a href="#FNanchor_111"><span class="label">111</span></a> As quoted by Mr. L. H. Morgan, ‘The American Beaver,’ 1868, +p. 272. Capt. Stansbury also gives an interesting account of the manner +in which a very young pelican, carried away by a strong stream, +was guided and encouraged in its attempts to reach the shore by half a +dozen old birds.</p></div> + +<div class="footnote"><p><a name="Footnote_112" id="Footnote_112"></a><a href="#FNanchor_112"><span class="label">112</span></a> As Mr. Bain states, “effective aid to a sufferer springs from sympathy +proper:” ‘Mental and Moral Science,’ 1868, p. 245.</p></div> + +<div class="footnote"><p><a name="Footnote_113" id="Footnote_113"></a><a href="#FNanchor_113"><span class="label">113</span></a> ‘Thierleben,’ B. i. s. 85.</p></div> + +<div class="footnote"><p><a name="Footnote_114" id="Footnote_114"></a><a href="#FNanchor_114"><span class="label">114</span></a> ‘De l’Espèce et de la Class.’ 1869, p. 97.</p></div> + +<div class="footnote"><p><a name="Footnote_115" id="Footnote_115"></a><a href="#FNanchor_115"><span class="label">115</span></a> ‘Der Darwin’schen Art-Lehre,’ 1869, s. 54.</p></div> + +<div class="footnote"><p><a name="Footnote_116" id="Footnote_116"></a><a href="#FNanchor_116"><span class="label">116</span></a> Brehm, ‘Thierleben,’ B. i. s. 76.</p></div> + +<div class="footnote"><p><a name="Footnote_117" id="Footnote_117"></a><a href="#FNanchor_117"><span class="label">117</span></a> See the first and striking chapter in Adam Smith’s ‘Theory of +Moral Sentiments.’ Also Mr. Bain’s ‘Mental and Moral Science,’ +1868, p. 244, and 275-282. Mr. Bain states, that “sympathy is, +indirectly, a source of pleasure to the sympathiser;” and he accounts +for this through reciprocity. He remarks that “the person benefited, +or others in his stead, may make up, by sympathy and good offices +returned, for all the sacrifice.” But if, as appears to be the case, +sympathy is strictly an instinct, its exercise would give direct pleasure, +in the same manner as the exercise, as before remarked, of almost every +other instinct.</p></div> + +<div class="footnote"><p><a name="Footnote_118" id="Footnote_118"></a><a href="#FNanchor_118"><span class="label">118</span></a> This fact, the Rev. L. Jenyns states (see his edition of ‘White’s +Nat. Hist. of Selborne,’ 1853, p. 204) was first recorded by the illustrious +Jenner, in ‘Phil. Transact.’ 1824, and has since been confirmed +by several observers, especially by Mr. Blackwall. This latter careful +observer examined, late in the autumn, during two years, thirty-six +nests; he found that twelve contained young dead birds, five contained +eggs on the point of being hatched, and three eggs not nearly hatched. +Many birds not yet old enough for a prolonged flight are likewise +deserted and left behind. See Blackwall, ‘Researches in Zoology,’ +1834, pp. 108, 118. For some additional evidence, although this is not +wanted, see Leroy, ‘Lettres Phil.’ 1802, p. 217.</p></div> + +<div class="footnote"><p><a name="Footnote_119" id="Footnote_119"></a><a href="#FNanchor_119"><span class="label">119</span></a> Hume remarks (‘An Enquiry Concerning the Principles of Morals,’ +edit. of 1751, p. 132), “there seems a necessity for confessing that the +happiness and misery of others are not spectacles altogether indifferent +to us, but that the view of the former ... communicates a +secret joy; the appearance of the latter ... throws a melancholy +damp over the imagination.”</p></div> + +<div class="footnote"><p><a name="Footnote_120" id="Footnote_120"></a><a href="#FNanchor_120"><span class="label">120</span></a> ‘Mental and Moral Science,’ 1868, p. 254.</p></div> + +<div class="footnote"><p><a name="Footnote_121" id="Footnote_121"></a><a href="#FNanchor_121"><span class="label">121</span></a> I have given one such case, namely of three Patagonian Indians +who preferred being shot, one after the other, to betraying the plans of +their companions in war (‘Journal of Researches,’ 1845, p. 103).</p></div> + +<div class="footnote"><p><a name="Footnote_122" id="Footnote_122"></a><a href="#FNanchor_122"><span class="label">122</span></a> Dr. Prosper Despine, in his ‘Psychologie Naturelle,’ 1868 (tom. +i. p. 243; tom ii. p. 169) gives many curious cases of the worst criminals, +who apparently have been entirely destitute of conscience.</p></div> + +<div class="footnote"><p><a name="Footnote_123" id="Footnote_123"></a><a href="#FNanchor_123"><span class="label">123</span></a> See an able article in the ‘North British Review,’ 1867, p. 395. +See also Mr. W. Bagehot’s articles on the Importance of Obedience +and Coherence to Primitive Man, in the ‘Fortnightly Review,’ 1867, +p. 529, and 1868, p. 457, &c.</p></div> + +<div class="footnote"><p><a name="Footnote_124" id="Footnote_124"></a><a href="#FNanchor_124"><span class="label">124</span></a> The fullest account which I have met with is by Dr. Gerland, in +his ‘Ueber das Aussterben der Naturvölker,’ 1868; but I shall have to +recur to the subject of infanticide in a future chapter.</p></div> + +<div class="footnote"><p><a name="Footnote_125" id="Footnote_125"></a><a href="#FNanchor_125"><span class="label">125</span></a> See the very interesting discussion on Suicide in Lecky’s ‘History +of European Morals,’ vol. i. 1869, p. 223.</p></div> + +<div class="footnote"><p><a name="Footnote_126" id="Footnote_126"></a><a href="#FNanchor_126"><span class="label">126</span></a> See, for instance, Mr. Hamilton’s account of the Kaffirs, ‘Anthropological +Review,’ 1870, p. xv.</p></div> + +<div class="footnote"><p><a name="Footnote_127" id="Footnote_127"></a><a href="#FNanchor_127"><span class="label">127</span></a> Mr. M’Lennan has given ‘Primitive Marriage,’ 1865, p. 176, a +good collection of facts on this head.</p></div> + +<div class="footnote"><p><a name="Footnote_128" id="Footnote_128"></a><a href="#FNanchor_128"><span class="label">128</span></a> Lecky, ‘History of European Morals,’ vol. i. 1869, p. 109.</p></div> + +<div class="footnote"><p><a name="Footnote_129" id="Footnote_129"></a><a href="#FNanchor_129"><span class="label">129</span></a> ‘Embassy to China,’ vol. ii. p. 348.</p></div> + +<div class="footnote"><p><a name="Footnote_130" id="Footnote_130"></a><a href="#FNanchor_130"><span class="label">130</span></a> See on this subject copious evidence in Chap. vii. of Sir J. +Lubbock, ‘Origin of Civilisation,’ 1870.</p></div> + +<div class="footnote"><p><a name="Footnote_131" id="Footnote_131"></a><a href="#FNanchor_131"><span class="label">131</span></a> For instance Lecky, ‘Hist. European Morals,’ vol. i. p. 124.</p></div> + +<div class="footnote"><p><a name="Footnote_132" id="Footnote_132"></a><a href="#FNanchor_132"><span class="label">132</span></a> This term is used in an able article in the ‘Westminster Review,’ +Oct. 1869, p. 498. For the Greatest Happiness principle, see J. S. +Mill, ‘Utilitarianism,’ p. 17.</p></div> + +<div class="footnote"><p><a name="Footnote_133" id="Footnote_133"></a><a href="#FNanchor_133"><span class="label">133</span></a> Good instances are given by Mr. Wallace in ‘Scientific Opinion,’ +Sept. 15, 1869; and more fully in his ‘Contributions to the Theory of +Natural Selection,’ 1870, p. 353.</p></div> + +<div class="footnote"><p><a name="Footnote_134" id="Footnote_134"></a><a href="#FNanchor_134"><span class="label">134</span></a> Tennyson, ‘Idylls of the King,’ p. 244.</p></div> + +<div class="footnote"><p><a name="Footnote_135" id="Footnote_135"></a><a href="#FNanchor_135"><span class="label">135</span></a> ‘The Thoughts of the Emperor M. Aurelius Antoninus,’ Eng. +translat., 2nd edit., 1869, p. 112. Marcus Aurelius was born <span class="smcap">A.D.</span> 121.</p></div> + +<div class="footnote"><p><a name="Footnote_136" id="Footnote_136"></a><a href="#FNanchor_136"><span class="label">136</span></a> Letter to Mr. Mill in Bain’s ‘Mental and Moral Science,’ 1868, +p. 722.</p></div> + +<div class="footnote"><p><a name="Footnote_137" id="Footnote_137"></a><a href="#FNanchor_137"><span class="label">137</span></a> A writer in the ‘North British Review’ (July, 1869, p. 531), well +capable of forming a sound judgment, expresses himself strongly to this +effect. Mr. Lecky (‘Hist. of Morals,’ vol. i. p. 143) seems to a certain +extent to coincide.</p></div> + +<div class="footnote"><p><a name="Footnote_138" id="Footnote_138"></a><a href="#FNanchor_138"><span class="label">138</span></a> See his remarkable work on ‘Hereditary Genius,’ 1869, p. 349. +The Duke of Argyll (‘Primeval Man,’ 1869, p. 188) has some good +remarks on the contest in man’s nature between right and wrong.</p></div> + +<div class="footnote"><p><a name="Footnote_139" id="Footnote_139"></a><a href="#FNanchor_139"><span class="label">139</span></a> ‘The Thoughts of Marcus Aurelius,’ &c., p. 139.</p></div> + +<div class="footnote"><p><a name="Footnote_140" id="Footnote_140"></a><a href="#FNanchor_140"><span class="label">140</span></a> ‘Investigations in Military and Anthropolog. Statistics of American +Soldiers,’ by B. A. Gould, 1869, p. 256.</p></div> + +<div class="footnote"><p><a name="Footnote_141" id="Footnote_141"></a><a href="#FNanchor_141"><span class="label">141</span></a> With respect to the “Cranial forms of the American aborigines,” +see Dr. Aitken Meigs in ‘Proc. Acad. Nat. Sci.’ Philadelphia, May, +1866. On the Australians, see Huxley, in Lyell’s ‘Antiquity of Man,’ +1863, p. 87. On the Sandwich Islanders, Prof. J. Wyman, ‘Observations +on Crania,’ Boston, 1868, p. 18.</p></div> + +<div class="footnote"><p><a name="Footnote_142" id="Footnote_142"></a><a href="#FNanchor_142"><span class="label">142</span></a> ‘Anatomy of the Arteries,’ by R. Quain.</p></div> + +<div class="footnote"><p><a name="Footnote_143" id="Footnote_143"></a><a href="#FNanchor_143"><span class="label">143</span></a> ‘Transact. Royal Soc.’ Edinburgh, vol. xxiv. p. 175, 189.</p></div> + +<div class="footnote"><p><a name="Footnote_144" id="Footnote_144"></a><a href="#FNanchor_144"><span class="label">144</span></a> ‘Proc. Royal Soc.’ 1867, p. 544; also 1868, p. 483, 524. There is +a previous paper, 1866, p. 229.</p></div> + +<div class="footnote"><p><a name="Footnote_145" id="Footnote_145"></a><a href="#FNanchor_145"><span class="label">145</span></a> ‘Proc. R. Irish Academy,’ vol. x. 1868, p. 141.</p></div> + +<div class="footnote"><p><a name="Footnote_146" id="Footnote_146"></a><a href="#FNanchor_146"><span class="label">146</span></a> ‘Act. Acad.,’ St. Petersburg, 1778, part ii. p. 217.</p></div> + +<div class="footnote"><p><a name="Footnote_147" id="Footnote_147"></a><a href="#FNanchor_147"><span class="label">147</span></a> Brehm, ‘Thierleben,’ B. i. s. 58, 87. Rengger, ‘Säugethiere von +Paraguay,’ s. 57.</p></div> + +<div class="footnote"><p><a name="Footnote_148" id="Footnote_148"></a><a href="#FNanchor_148"><span class="label">148</span></a> ‘Variation of Animals and Plants under Domestication,’ vol. ii. +chap. xii.</p></div> + +<div class="footnote"><p><a name="Footnote_149" id="Footnote_149"></a><a href="#FNanchor_149"><span class="label">149</span></a> ‘Hereditary Genius: an Inquiry into its Laws and Consequences,’ +1869.</p></div> + +<div class="footnote"><p><a name="Footnote_150" id="Footnote_150"></a><a href="#FNanchor_150"><span class="label">150</span></a> Mr. Bates remarks (‘The Naturalist on the Amazons,’ 1863, vol. ii. +p. 159), with respect to the Indians of the same S. American tribe, +“no two of them were at all similar in the shape of the head; one +man had an oval visage with fine features, and another was quite +Mongolian in breadth and prominence of cheek, spread of nostrils, +and obliquity of eyes.”</p></div> + +<div class="footnote"><p><a name="Footnote_151" id="Footnote_151"></a><a href="#FNanchor_151"><span class="label">151</span></a> Blumenbach, ‘Treatises on Anthropolog.’ Eng. translat., 1865, +p. 205.</p></div> + +<div class="footnote"><p><a name="Footnote_152" id="Footnote_152"></a><a href="#FNanchor_152"><span class="label">152</span></a> Godron, ‘De l’Espèce,’ 1859, tom. ii. livre 3. Quatrefages, ‘Unité +de l’Espèce Humaine,’ 1861. Also Lectures on Anthropology, given +in the ‘Revue des Cours Scientifiques,’ 1866-1868.</p></div> + +<div class="footnote"><p><a name="Footnote_153" id="Footnote_153"></a><a href="#FNanchor_153"><span class="label">153</span></a> ‘Hist. Gen. et Part. des Anomalies de l’Organisation,’ in three +volumes, tom. i. 1832.</p></div> + +<div class="footnote"><p><a name="Footnote_154" id="Footnote_154"></a><a href="#FNanchor_154"><span class="label">154</span></a> I have fully discussed these laws in my ‘Variation of Animals +and Plants under Domestication,’ vol. ii. chap. xxii. and xxiii. M. J. +P. Durand has lately 1868; published a valuable essay ‘De l’Influence +des Milieux, &c.’ He lays much stress on the nature of the soil.</p></div> + +<div class="footnote"><p><a name="Footnote_155" id="Footnote_155"></a><a href="#FNanchor_155"><span class="label">155</span></a> ‘Investigations in Military and Anthrop. Statistics,’ &c. 1869, by +B. A. Gould, p. 93, 107, 126, 131, 134.</p></div> + +<div class="footnote"><p><a name="Footnote_156" id="Footnote_156"></a><a href="#FNanchor_156"><span class="label">156</span></a> For the Polynesians, see Prichard’s ‘Physical Hist. of Mankind,’ +vol. v. 1847, p. 145, 283. Also Godron, ‘De l’Espèce,’ tom. ii. p. 289. +There is also a remarkable difference in appearance between the closely-allied +Hindoos inhabiting the Upper Ganges and Bengal; see Elphinstone’s +'History of India,’ vol. i. p. 324.</p></div> + +<div class="footnote"><p><a name="Footnote_157" id="Footnote_157"></a><a href="#FNanchor_157"><span class="label">157</span></a> ‘Memoirs, Anthropolog. Soc.’ vol. iii. 1867-69, p. 561, 565, 567.</p></div> + +<div class="footnote"><p><a name="Footnote_158" id="Footnote_158"></a><a href="#FNanchor_158"><span class="label">158</span></a> Dr. Brakenridge, ‘Theory of Diathesis,’ ‘Medical Times,’ June 19 +and July 17, 1869.</p></div> + +<div class="footnote"><p><a name="Footnote_159" id="Footnote_159"></a><a href="#FNanchor_159"><span class="label">159</span></a> I have given authorities for these several statements in my ‘Variation +of Animals under Domestication,’ vol. ii. p. 297-300. Dr. Jaeger, +“Ueber das Längenwachsthum der Knochen,” ‘Jenaischen Zeitschrift,’ +B. v. Heft i.</p></div> + +<div class="footnote"><p><a name="Footnote_160" id="Footnote_160"></a><a href="#FNanchor_160"><span class="label">160</span></a> ‘Investigations,’ &c. By B. A. Gould, 1869, p. 288.</p></div> + +<div class="footnote"><p><a name="Footnote_161" id="Footnote_161"></a><a href="#FNanchor_161"><span class="label">161</span></a> ‘Säugethiere von Paraguay,’ 1830, s. 4.</p></div> + +<div class="footnote"><p><a name="Footnote_162" id="Footnote_162"></a><a href="#FNanchor_162"><span class="label">162</span></a> ‘History of Greenland,’ Eng. translat. 1767, vol. i. p. 230.</p></div> + +<div class="footnote"><p><a name="Footnote_163" id="Footnote_163"></a><a href="#FNanchor_163"><span class="label">163</span></a> ‘Intermarriage.’ By Alex. Walker, 1838. p. 377.</p></div> + +<div class="footnote"><p><a name="Footnote_164" id="Footnote_164"></a><a href="#FNanchor_164"><span class="label">164</span></a> ‘The Variation of Animals under Domestication,’ vol. i. p. 173.</p></div> + +<div class="footnote"><p><a name="Footnote_165" id="Footnote_165"></a><a href="#FNanchor_165"><span class="label">165</span></a> ‘Principles of Biology,’ vol. i. p. 455.</p></div> + +<div class="footnote"><p><a name="Footnote_166" id="Footnote_166"></a><a href="#FNanchor_166"><span class="label">166</span></a> Paget, ‘Lectures on Surgical Pathology,’ vol. i. 1853, p. 209.</p></div> + +<div class="footnote"><p><a name="Footnote_167" id="Footnote_167"></a><a href="#FNanchor_167"><span class="label">167</span></a> ‘The Variation of Animals under Domestication,’ vol. i. p. 8.</p></div> + +<div class="footnote"><p><a name="Footnote_168" id="Footnote_168"></a><a href="#FNanchor_168"><span class="label">168</span></a> ‘Säugethiere von Paraguay,’ s. 8, 10. I have had good opportunities +for observing the extraordinary power of eyesight in the +Fuegians.’ See also Lawrence (‘Lectures on Physiology,’ &c., 1822, p. +404) on this same subject. M. Giraud-Teulon has recently collected +(‘Revue des Cours Scientifiques,’ 1870, p. 625) a large and valuable +body of evidence proving that the cause of short-sight, “<i>C’est le travail +assidu, de près.</i>”</p></div> + +<div class="footnote"><p><a name="Footnote_169" id="Footnote_169"></a><a href="#FNanchor_169"><span class="label">169</span></a> Prichard, ‘Phys. Hist. of Mankind,’ on the authority of Blumenbach, +vol. i. 1851, p. 311; for the statement by Pallas, vol. iv. 1844, p. +407.</p></div> + +<div class="footnote"><p><a name="Footnote_170" id="Footnote_170"></a><a href="#FNanchor_170"><span class="label">170</span></a> Quoted by Prichard, ‘Researches into the Phys. Hist. of Mankind,’ +vol. v. p. 463.</p></div> + +<div class="footnote"><p><a name="Footnote_171" id="Footnote_171"></a><a href="#FNanchor_171"><span class="label">171</span></a> Mr. Forbes’ valuable paper is now published in the ‘Journal of +the Ethnological Soc. of London,’ new series, vol. ii. 1870, p. 193.</p></div> + +<div class="footnote"><p><a name="Footnote_172" id="Footnote_172"></a><a href="#FNanchor_172"><span class="label">172</span></a> Dr. Wilckens (‘Landwirthschaft. Wochenblatt,’ No. 10, 1869) +has lately published an interesting essay shewing how domestic +animals, which live in mountainous regions, have their frames +modified.</p></div> + +<div class="footnote"><p><a name="Footnote_173" id="Footnote_173"></a><a href="#FNanchor_173"><span class="label">173</span></a> ‘Mémoire sur les Microcéphales,’ 1867, p. 50, 125, 169, 171, 184-198.</p></div> + +<div class="footnote"><p><a name="Footnote_174" id="Footnote_174"></a><a href="#FNanchor_174"><span class="label">174</span></a> See Dr. A. Farre’s well-known article in the ‘Cyclop. of Anat. +and Phys.’ vol. v. 1859, p. 642. Owen ‘Anatomy of Vertebrates,’ vol. +iii. 1868, p. 687. Prof. Turner in ‘Edinburgh Medical Journal,’ Feb. +1865.</p></div> + +<div class="footnote"><p><a name="Footnote_175" id="Footnote_175"></a><a href="#FNanchor_175"><span class="label">175</span></a> ‘Annuario della Soc. dei Naturalisti in Modena,’ 1867, p. 83. +Prof. Canestrini gives extracts on this subject from various authorities. +Laurillard remarks, that as he has found a complete similarity in the +form, proportions, and connexion of the two malar bones in several +human subjects and in certain apes, he cannot consider this disposition +of the parts as simply accidental.</p></div> + +<div class="footnote"><p><a name="Footnote_176" id="Footnote_176"></a><a href="#FNanchor_176"><span class="label">176</span></a> A whole series of cases is given by Isid. Geoffroy St.-Hilaire, +'Hist. des Anomalies,’ tom. iii. p. 437.</p></div> + +<div class="footnote"><p><a name="Footnote_177" id="Footnote_177"></a><a href="#FNanchor_177"><span class="label">177</span></a> In my ‘Variation of Animals under Domestication’ (vol. ii. p. 57) +I attributed the not very rare cases of supernumerary mammæ in +women to reversion. I was led to this as a <i>probable</i> conclusion, by the +additional mammæ being generally placed symmetrically on the breast, +and more especially from one case, in which a single efficient mamma +occurred in the inguinal region of a woman, the daughter of another +woman with supernumerary mammæ. But Prof. Preyer (‘Der Kampf +um das Dasein,’ 1869, s. 45) states that <i>mammæ erraticæ</i> have been +known to occur in other situations, even on the back; so that the force +of my argument is greatly weakened or perhaps quite destroyed. +</p><p> +With much hesitation I, in the same work (vol. ii. p. 12), attributed +the frequent cases of polydactylism in men to reversion. I was partly +led to this through Prof. Owen’s statement, that some of the Ichthyopterygia +possess more than five digits, and therefore, as I supposed, had +retained a primordial condition; but after reading Prof. Gegenbaur’s +paper (‘Jenaischen Zeitschrift,’ B. v. Heft 3, s. 341), who is the highest +authority in Europe on such a point, and who disputes Owen’s conclusion, +I see that it is extremely doubtful whether supernumerary +digits can thus be accounted for. It was the fact that such digits not +only frequently occur and are strongly inherited, but have the power +of regrowth after amputation, like the normal digits of the lower vertebrata, +that chiefly led me to the above conclusion. This extraordinary +fact of their regrowth remains inexplicable, if the belief in reversion to +some extremely remote progenitor must be rejected. I cannot, however, +follow Prof. Gegenbaur in supposing that additional digits could +not reappear through reversion, without at the same time other parts of +the skeleton being simultaneously and similarly modified; for single +characters often reappear through reversion.</p></div> + +<div class="footnote"><p><a name="Footnote_178" id="Footnote_178"></a><a href="#FNanchor_178"><span class="label">178</span></a> ‘Anatomy of Vertebrates,’ vol. iii. 1868, p. 323.</p></div> + +<div class="footnote"><p><a name="Footnote_179" id="Footnote_179"></a><a href="#FNanchor_179"><span class="label">179</span></a> ‘Generelle Morphologie,’ 1866, B. ii. s. clv.</p></div> + +<div class="footnote"><p><a name="Footnote_180" id="Footnote_180"></a><a href="#FNanchor_180"><span class="label">180</span></a> Carl Vogt’s ‘Lectures on Man,’ Eng. translat. 1864, p. 151.</p></div> + +<div class="footnote"><p><a name="Footnote_181" id="Footnote_181"></a><a href="#FNanchor_181"><span class="label">181</span></a> C. Carter Blake, on a jaw from La Naulette, ‘Anthropolog. +Review,’ 1867, p. 295. Schaaffhausen, ibid. 1868, p. 426.</p></div> + +<div class="footnote"><p><a name="Footnote_182" id="Footnote_182"></a><a href="#FNanchor_182"><span class="label">182</span></a> ‘The Anatomy of Expression,’ 1844, p. 110, 131.</p></div> + +<div class="footnote"><p><a name="Footnote_183" id="Footnote_183"></a><a href="#FNanchor_183"><span class="label">183</span></a> Quoted by Prof. Canestrini in the ‘Annuario,’ &c., 1867, p. 90.</p></div> + +<div class="footnote"><p><a name="Footnote_184" id="Footnote_184"></a><a href="#FNanchor_184"><span class="label">184</span></a> These papers deserve careful study by any one who desires to +learn how frequently our muscles vary, and in varying come to resemble +those of the Quadrumana. The following references relate to +the few points touched on in my text: vol. xiv. 1865, p. 379-384; vol. +xv. 1866, p. 241, 242; vol. xv. 1867, p. 544; vol. xvi. 1868, p. 524. I +may here add that Dr. Murie and Mr. St. George Mivart have shewn +in their Memoir on the Lemuroidea (‘Transact. Zoolog. Soc.’ vol. vii. +1869, p. 96), how extraordinarily variable some of the muscles are in +these animals, the lowest members of the Primates. Gradations, also, +in the muscles leading to structures found in animals still lower in +the scale, are numerous in the Lemuroidea.</p></div> + +<div class="footnote"><p><a name="Footnote_185" id="Footnote_185"></a><a href="#FNanchor_185"><span class="label">185</span></a> Prof. Macalister in ‘Proc. R. Irish Academy,’ vol. x. 1868, p. 124.</p></div> + +<div class="footnote"><p><a name="Footnote_186" id="Footnote_186"></a><a href="#FNanchor_186"><span class="label">186</span></a> Prof. Macalister (ibid. p. 121) has tabulated his observations, and +finds that muscular abnormalities are most frequent in the fore-arms, +secondly in the face, thirdly in the foot, &c.</p></div> + +<div class="footnote"><p><a name="Footnote_187" id="Footnote_187"></a><a href="#FNanchor_187"><span class="label">187</span></a> The Rev. Dr. Haughton, after giving (‘Proc. R. Irish Academy,’ +June 27, 1864, p. 715) a remarkable case of variation in the human +<i>flexor pollicis longus</i>, adds, “This remarkable example shews that man +may sometimes possess the arrangement of tendons of thumb and +fingers characteristic of the macaque; but whether such a case should +be regarded as a macaque passing upwards into a man, or a man +passing downwards into a macaque, or as a congenital freak of +nature, I cannot undertake to say.” It is satisfactory to hear so +capable an anatomist, and so embittered an opponent of evolutionism, +admitting even the possibility of either of his first propositions. Prof. +Macalister has also described (‘Proc. R. Irish Acad.’ vol. x. 1864, p. +138) variations in the <i>flexor pollicis longus</i>, remarkable from their relations +to the same muscle in the Quadrumana.</p></div> + +<div class="footnote"><p><a name="Footnote_188" id="Footnote_188"></a><a href="#FNanchor_188"><span class="label">188</span></a> The authorities for these several statements are given in my +'Variation of Animals under Domestication,’ vol. ii. p. 320-335.</p></div> + +<div class="footnote"><p><a name="Footnote_189" id="Footnote_189"></a><a href="#FNanchor_189"><span class="label">189</span></a> This whole subject has been discussed in chap. xxiii. vol. ii. of +my ‘Variation of Animals and Plants under Domestication.’</p></div> + +<div class="footnote"><p><a name="Footnote_190" id="Footnote_190"></a><a href="#FNanchor_190"><span class="label">190</span></a> See the ever memorable ‘Essay on the Principle of Population,’ +by the Rev. T. Malthus, vol. i. 1826, p. 6, 517.</p></div> + +<div class="footnote"><p><a name="Footnote_191" id="Footnote_191"></a><a href="#FNanchor_191"><span class="label">191</span></a> ‘Variation of Animals and Plants under Domestication,’ vol. ii. +p. 111-113, 163.</p></div> + +<div class="footnote"><p><a name="Footnote_192" id="Footnote_192"></a><a href="#FNanchor_192"><span class="label">192</span></a> Mr. Sedgwick, ‘British and Foreign Medico-Chirurg. Review,’ +July, 1863, p. 170.</p></div> + +<div class="footnote"><p><a name="Footnote_193" id="Footnote_193"></a><a href="#FNanchor_193"><span class="label">193</span></a> ‘The Annals of Rural Bengal,’ by W. W. Hunter, 1868, p. 259.</p></div> + +<div class="footnote"><p><a name="Footnote_194" id="Footnote_194"></a><a href="#FNanchor_194"><span class="label">194</span></a> ‘Primitive Marriage,’ 1865.</p></div> + +<div class="footnote"><p><a name="Footnote_195" id="Footnote_195"></a><a href="#FNanchor_195"><span class="label">195</span></a> See some good remarks to this effect by W. Stanley Jevons, “A +Deduction from Darwin’s Theory,” ‘Nature,’ 1869, p. 231.</p></div> + +<div class="footnote"><p><a name="Footnote_196" id="Footnote_196"></a><a href="#FNanchor_196"><span class="label">196</span></a> Latham, ‘Man and his Migrations,’ 1851, p. 135.</p></div> + +<div class="footnote"><p><a name="Footnote_197" id="Footnote_197"></a><a href="#FNanchor_197"><span class="label">197</span></a> Messrs. Murie and Mivart in their “Anatomy of the Lemuroidea” +(‘Transact. Zoolog. Soc.’ vol. vii. 1869, p. 96-98) say, “some muscles +are so irregular in their distribution that they cannot be well classed +in any of the above groups.” These muscles differ even on the opposite +sides of the same individual.</p></div> + +<div class="footnote"><p><a name="Footnote_198" id="Footnote_198"></a><a href="#FNanchor_198"><span class="label">198</span></a> ‘Quarterly Review,’ April, 1869, p. 392. This subject is more +fully discussed in Mr. Wallace’s ‘Contributions to the Theory of Natural +Selection,’ 1870, in which all the essays referred to in this work are +republished. The ‘Essay on Man’ has been ably criticised by Prof. +Claparède, one of the most distinguished zoologists in Europe, in an +article published in the ‘Bibliothèque Universelle,’ June, 1870. The +remark quoted in my text will surprise every one who has read +Mr. Wallace’s celebrated paper on ‘The Origin of Human Races +deduced from the Theory of Natural Selection,’ originally published +in the ‘Anthropological Review,’ May, 1864, p. clviii. I cannot here +resist quoting a most just remark by Sir J. Lubbock (‘Prehistoric +Times,’ 1865, p. 479) in reference to this paper, namely, that Mr. +Wallace, “with characteristic unselfishness, ascribes it (<i>i.e.</i> the idea of +natural selection) unreservedly to Mr. Darwin, although, as is well +known, he struck out the idea independently, and published it, +though not with the same elaboration, at the same time.”</p></div> + +<div class="footnote"><p><a name="Footnote_199" id="Footnote_199"></a><a href="#FNanchor_199"><span class="label">199</span></a> Quoted by Mr. Lawson Tait in his “Law of Natural Selection,”—‘Dublin +Quarterly Journal of Medical Science,’ Feb. 1869. Dr. +Keller is likewise quoted to the same effect.</p></div> + +<div class="footnote"><p><a name="Footnote_200" id="Footnote_200"></a><a href="#FNanchor_200"><span class="label">200</span></a> Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 71.</p></div> + +<div class="footnote"><p><a name="Footnote_201" id="Footnote_201"></a><a href="#FNanchor_201"><span class="label">201</span></a> ‘Quarterly Review,’ April, 1869, p. 392.</p></div> + +<div class="footnote"><p><a name="Footnote_202" id="Footnote_202"></a><a href="#FNanchor_202"><span class="label">202</span></a> In <i>Hylobates syndactylus</i>, as the name expresses, two of the digits +regularly cohere; and this, as Mr. Blyth informs me, is occasionally +the case with the digits of <i>H. agilis</i>, <i>lar</i>, and <i>leuciscus</i>.</p></div> + +<div class="footnote"><p><a name="Footnote_203" id="Footnote_203"></a><a href="#FNanchor_203"><span class="label">203</span></a> Brehm, ‘Thierleben,’ B. i. s. 80.</p></div> + +<div class="footnote"><p><a name="Footnote_204" id="Footnote_204"></a><a href="#FNanchor_204"><span class="label">204</span></a> “The Hand, its mechanism,” &c. ‘Bridgewater Treatise,’ 1833, +p. 38.</p></div> + +<div class="footnote"><p><a name="Footnote_205" id="Footnote_205"></a><a href="#FNanchor_205"><span class="label">205</span></a> Häckel has an excellent discussion on the steps by which man +became a biped: ‘Natürliche Schöpfungsgeschichte,’ 1868, s. 507. Dr. +Büchner (‘Conférences sur la Théorie Darwinienne,’ 1869, p. 135) has +given good cases of the use of the foot as a prehensile organ by man; +also on the manner of progression of the higher apes to which I allude +in the following paragraph: see also Owen (‘Anatomy of Vertebrates,’ +vol. iii. p. 71) on this latter subject.</p></div> + +<div class="footnote"><p><a name="Footnote_206" id="Footnote_206"></a><a href="#FNanchor_206"><span class="label">206</span></a> “On the Primitive Form of the Skull,” translated in ‘Anthropological +Review,’ Oct. 1868, p. 428. Owen (‘Anatomy of Vertebrates,’ +vol. ii. 1866, p. 551) on the mastoid processes in the higher apes.</p></div> + +<div class="footnote"><p><a name="Footnote_207" id="Footnote_207"></a><a href="#FNanchor_207"><span class="label">207</span></a> ‘Die Grenzen der Thierwelt, eine Betrachtung zu Darwin’s +Lehre,’ 1868, s. 51.</p></div> + +<div class="footnote"><p><a name="Footnote_208" id="Footnote_208"></a><a href="#FNanchor_208"><span class="label">208</span></a> Dujardin, ‘Annales des Sc. Nat.’ 3rd series, Zoolog. tom. xiv. +1850, p. 203. See also Mr. Lowne, ‘Anatomy and Phys. of the <i>Musca +vomitoria</i>,’ 1870, p. 14. My son, Mr. F. Darwin, dissected for me the +cerebral ganglia of the <i>Formica rufa</i>.</p></div> + +<div class="footnote"><p><a name="Footnote_209" id="Footnote_209"></a><a href="#FNanchor_209"><span class="label">209</span></a> ‘Philosophical Transactions,’ 1869, p. 513.</p></div> + +<div class="footnote"><p><a name="Footnote_210" id="Footnote_210"></a><a href="#FNanchor_210"><span class="label">210</span></a> Quoted in C. Vogt’s ‘Lectures on Man,’ Eng. translat. 1864, p. +88, 90. Prichard, ‘Phys. Hist. of Mankind,’ vol. i. 1838, p. 305.</p></div> + +<div class="footnote"><p><a name="Footnote_211" id="Footnote_211"></a><a href="#FNanchor_211"><span class="label">211</span></a> ‘Comptes Rendus des Séances,’ &c. June 1, 1868.</p></div> + +<div class="footnote"><p><a name="Footnote_212" id="Footnote_212"></a><a href="#FNanchor_212"><span class="label">212</span></a> ‘The Variation of Animals and Plants under Domestication,’ vol. +ii. p. 124-129.</p></div> + +<div class="footnote"><p><a name="Footnote_213" id="Footnote_213"></a><a href="#FNanchor_213"><span class="label">213</span></a> Schaaffhausen gives from Blumenbach and Busch, the cases of the +spasms and cicatrix, in ‘Anthropolog. Review,’ Oct. 1868, p. 420. Dr. +Jarrold (‘Anthropologia,’ 1808, p. 115, 116) adduces from Camper and +from his own observations, cases of the modification of the skull from +the head being fixed in an unnatural position. He believes that certain +trades, such as that of a shoemaker, by causing the head to be +habitually held forward, makes the forehead more rounded and +prominent.</p></div> + +<div class="footnote"><p><a name="Footnote_214" id="Footnote_214"></a><a href="#FNanchor_214"><span class="label">214</span></a> ‘Variation of Animals,’ &c., vol. i. p. 117 on the elongation of the +skull; p. 119, on the effect of the lopping of one ear.</p></div> + +<div class="footnote"><p><a name="Footnote_215" id="Footnote_215"></a><a href="#FNanchor_215"><span class="label">215</span></a> Quoted by Schaaffhausen, in ‘Anthropolog. Review,’ Oct. 1868, +p. 419.</p></div> + +<div class="footnote"><p><a name="Footnote_216" id="Footnote_216"></a><a href="#FNanchor_216"><span class="label">216</span></a> Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 619.</p></div> + +<div class="footnote"><p><a name="Footnote_217" id="Footnote_217"></a><a href="#FNanchor_217"><span class="label">217</span></a> Isidore Geoffroy St.-Hilaire remarks (‘Hist. Nat. Générale,’ tom. +ii. 1859, p. 215-217) on the head of man being covered with long hair; +also on the upper surfaces of monkeys and of other mammals being +more thickly clothed than the lower surfaces. This has likewise been +observed by various authors. Prof. P. Gervais (‘Hist. Nat. des Mammifères,’ +tom. i. 1854, p. 28), however, states that in the Gorilla the +hair is thinner on the back, where it is partly rubbed off, than on the +lower surface.</p></div> + +<div class="footnote"><p><a name="Footnote_218" id="Footnote_218"></a><a href="#FNanchor_218"><span class="label">218</span></a> Mr. St. George Mivart, ‘Proc. Zoolog. Soc.’ 1865, p. 562, 583. +Dr. J. E. Gray, ‘Cat. Brit. Mus.: Skeletons.’ Owen, ‘Anatomy of +Vertebrates,’ vol. ii. p. 517. Isidore Geoffroy, ‘Hist. Nat. Gén.’ tom. +ii. p. 244.</p></div> + +<div class="footnote"><p><a name="Footnote_219" id="Footnote_219"></a><a href="#FNanchor_219"><span class="label">219</span></a> ‘The Variation of Animals and Plants under Domestication,’ vol. +ii. p. 280, 282.</p></div> + +<div class="footnote"><p><a name="Footnote_220" id="Footnote_220"></a><a href="#FNanchor_220"><span class="label">220</span></a> ‘Primeval Man,’ 1869, p. 66.</p></div> + +<div class="footnote"><p><a name="Footnote_221" id="Footnote_221"></a><a href="#FNanchor_221"><span class="label">221</span></a> ‘Anthropological Review,’ May, 1864, p. clviii.</p></div> + +<div class="footnote"><p><a name="Footnote_222" id="Footnote_222"></a><a href="#FNanchor_222"><span class="label">222</span></a> After a time the members or tribes which are absorbed into another +tribe assume, as Mr. Maine remarks (‘Ancient Law,’ 1861, p. 131), that +they are the co-descendants of the same ancestors.</p></div> + +<div class="footnote"><p><a name="Footnote_223" id="Footnote_223"></a><a href="#FNanchor_223"><span class="label">223</span></a> Morlot, ‘Soc. Vaud. Sc. Nat.’ 1860, p. 294.</p></div> + +<div class="footnote"><p><a name="Footnote_224" id="Footnote_224"></a><a href="#FNanchor_224"><span class="label">224</span></a> I have given instances in my ‘Variation of Animals under Domestication,’ +vol. ii. p. 196.</p></div> + +<div class="footnote"><p><a name="Footnote_225" id="Footnote_225"></a><a href="#FNanchor_225"><span class="label">225</span></a> See a remarkable series of articles on Physics and Politics in the +'Fortnightly Review,’ Nov. 1867; April 1, 1868; July 1, 1869.</p></div> + +<div class="footnote"><p><a name="Footnote_226" id="Footnote_226"></a><a href="#FNanchor_226"><span class="label">226</span></a> ‘Origin of Civilisation,’ 1870, p. 265.</p></div> + +<div class="footnote"><p><a name="Footnote_227" id="Footnote_227"></a><a href="#FNanchor_227"><span class="label">227</span></a> Mr. Wallace gives cases in his ‘Contributions to the Theory of +Natural Selection,’ 1870, p. 354.</p></div> + +<div class="footnote"><p><a name="Footnote_228" id="Footnote_228"></a><a href="#FNanchor_228"><span class="label">228</span></a> ‘Ancient Law,’ 1861, p. 22. For Mr. Bagehot’s remarks, ‘Fortnightly +Review,’ April 1, 1868, p. 452.</p></div> + +<div class="footnote"><p><a name="Footnote_229" id="Footnote_229"></a><a href="#FNanchor_229"><span class="label">229</span></a> ‘The Variation of Animals and Plants under Domestication,’ vol. i. +p. 309.</p></div> + +<div class="footnote"><p><a name="Footnote_230" id="Footnote_230"></a><a href="#FNanchor_230"><span class="label">230</span></a> ‘Fraser’s Magazine,’ Sept. 1868, p. 353. This article seems to +have struck many persons, and has given rise to two remarkable essays +and a rejoinder in the ‘Spectator,’ Oct. 3rd and 17th 1868. It has +also been discussed in the ‘Q. Journal of Science,’ 1869, p. 152, and by +Mr. Lawson Tait in the ‘Dublin Q. Journal of Medical Science,’ Feb. +1869, and by Mr. E. Ray Lankester in his ‘Comparative Longevity,’ +1870, p. 128. Similar views appeared previously in the ‘Australasian,’ +July 13, 1867. I have borrowed ideas from several of these writers.</p></div> + +<div class="footnote"><p><a name="Footnote_231" id="Footnote_231"></a><a href="#FNanchor_231"><span class="label">231</span></a> For Mr. Wallace, see ‘Anthropolog. Review,’ as before cited. +Mr. Galton in ‘Macmillan’s Magazine,’ Aug. 1865, p. 318; also his +great work, ‘Hereditary Genius,’ 1870.</p></div> + +<div class="footnote"><p><a name="Footnote_232" id="Footnote_232"></a><a href="#FNanchor_232"><span class="label">232</span></a> ‘Hereditary Genius,’ 1870, p. 132-140.</p></div> + +<div class="footnote"><p><a name="Footnote_233" id="Footnote_233"></a><a href="#FNanchor_233"><span class="label">233</span></a> See the fifth and sixth columns, compiled from good authorities, +in the table given in Mr. E. R. Lankester’s ‘Comparative Longevity,’ +1870, p. 115.</p></div> + +<div class="footnote"><p><a name="Footnote_234" id="Footnote_234"></a><a href="#FNanchor_234"><span class="label">234</span></a> ‘Hereditary Genius,’ 1870, p. 330.</p></div> + +<div class="footnote"><p><a name="Footnote_235" id="Footnote_235"></a><a href="#FNanchor_235"><span class="label">235</span></a> ‘Origin of Species’ (fifth edition, 1869), p. 104.</p></div> + +<div class="footnote"><p><a name="Footnote_236" id="Footnote_236"></a><a href="#FNanchor_236"><span class="label">236</span></a> ‘Hereditary Genius,’ 1870, p. 347.</p></div> + +<div class="footnote"><p><a name="Footnote_237" id="Footnote_237"></a><a href="#FNanchor_237"><span class="label">237</span></a> E. Ray Lankester, ‘Comparative Longevity,’ 1870, p. 115. The +table of the intemperate is from Nelson’s ‘Vital Statistics.’ In regard +to profligacy, see Dr. Farr, “Influence of Marriage on Mortality,” ‘Nat. +Assoc. for the Promotion of Social Science,’ 1858.</p></div> + +<div class="footnote"><p><a name="Footnote_238" id="Footnote_238"></a><a href="#FNanchor_238"><span class="label">238</span></a> ‘Fraser’s Magazine,’ Sept. 1868, p. 353. ‘Macmillan’s Magazine,’ +Aug. 1865, p. 318. The Rev. F. W. Farrar (‘Fraser’s Mag.,’ Aug. 1870, +p. 264) takes a different view.</p></div> + +<div class="footnote"><p><a name="Footnote_239" id="Footnote_239"></a><a href="#FNanchor_239"><span class="label">239</span></a> “On the Laws of the Fertility of Women,” in ‘Transact. Royal +Soc.’ Edinburgh, vol. xxiv. p. 287. See, also, Mr. Galton, ‘Hereditary +Genius,’ p. 352-357, for observations to the above effect.</p></div> + +<div class="footnote"><p><a name="Footnote_240" id="Footnote_240"></a><a href="#FNanchor_240"><span class="label">240</span></a> ‘Tenth Annual Report of Births, Deaths, &c., in Scotland,’ 1867, +p. xxix.</p></div> + +<div class="footnote"><p><a name="Footnote_241" id="Footnote_241"></a><a href="#FNanchor_241"><span class="label">241</span></a> These quotations are taken from our highest authority on such +questions, namely, Dr. Farr, in his paper “On the Influence of Marriage +on the Mortality of the French People,” read before the Nat. Assoc. +for the Promotion of Social Science, 1858.</p></div> + +<div class="footnote"><p><a name="Footnote_242" id="Footnote_242"></a><a href="#FNanchor_242"><span class="label">242</span></a> Dr. Farr, ibid. The quotations given below are extracted from +the same striking paper.</p></div> + +<div class="footnote"><p><a name="Footnote_243" id="Footnote_243"></a><a href="#FNanchor_243"><span class="label">243</span></a> I have taken the mean of the quinquennial means, given in ‘The +Tenth Annual Report of Births, Deaths, &c., in Scotland,’ 1867. The +quotation from Dr. Stark is copied from an article in the ‘Daily News,’ +Oct. 17th, 1868, which Dr. Farr considers very carefully written.</p></div> + +<div class="footnote"><p><a name="Footnote_244" id="Footnote_244"></a><a href="#FNanchor_244"><span class="label">244</span></a> See the ingenious and original argument on this subject by Mr. +Galton, ‘Hereditary Genius,’ p. 340-342.</p></div> + +<div class="footnote"><p><a name="Footnote_245" id="Footnote_245"></a><a href="#FNanchor_245"><span class="label">245</span></a> Mr. Greg, ‘Fraser’s Magazine,’ Sept. 1868, p. 357.</p></div> + +<div class="footnote"><p><a name="Footnote_246" id="Footnote_246"></a><a href="#FNanchor_246"><span class="label">246</span></a> ‘Hereditary Genius,’ 1870, p. 357-359. The Rev. F. H. Farrar +(‘Fraser’s Mag.’, Aug. 1870, p. 257) advances arguments on the other +side. Sir C. Lyell had already (‘Principles of Geology,’ vol. ii. 1868, +p. 489) called attention, in a striking passage, to the evil influence of +the Holy Inquisition in having lowered, through selection, the general +standard of intelligence in Europe.</p></div> + +<div class="footnote"><p><a name="Footnote_247" id="Footnote_247"></a><a href="#FNanchor_247"><span class="label">247</span></a> Mr. Galton, ‘Macmillan’s Magazine,’ August, 1865, p. 325. See, +also, ‘Nature,’ “On Darwinism and National Life,” Dec. 1869, p. 184.</p></div> + +<div class="footnote"><p><a name="Footnote_248" id="Footnote_248"></a><a href="#FNanchor_248"><span class="label">248</span></a> ‘Last Winter in the United States,’ 1868, p. 29.</p></div> + +<div class="footnote"><p><a name="Footnote_249" id="Footnote_249"></a><a href="#FNanchor_249"><span class="label">249</span></a> ‘On the Origin of Civilisation,’ ‘Proc. Ethnological Soc.’ Nov. +26, 1867.</p></div> + +<div class="footnote"><p><a name="Footnote_250" id="Footnote_250"></a><a href="#FNanchor_250"><span class="label">250</span></a> ‘Primeval Man,’ 1869.</p></div> + +<div class="footnote"><p><a name="Footnote_251" id="Footnote_251"></a><a href="#FNanchor_251"><span class="label">251</span></a> ‘Royal Institution of Great Britain,’ March 15, 1867. Also, +'Researches into the Early History of Mankind,’ 1865.</p></div> + +<div class="footnote"><p><a name="Footnote_252" id="Footnote_252"></a><a href="#FNanchor_252"><span class="label">252</span></a> ‘Primitive Marriage,’ 1865. See, likewise, an excellent article, +evidently by the same author, in the ‘North British Review,’ July, +1869. Also, Mr. L. H. Morgan, “A Conjectural Solution of the Origin +of the Class. System of Relationship,” in ‘Proc. American Acad. of +Sciences,’ vol. vii. Feb. 1868. Prof. Schaaffhausen (‘Anthropolog. +Review,’ Oct. 1869, p. 373) remarks on “the vestiges of human sacrifices +found both in Homer and the Old Testament.”</p></div> + +<div class="footnote"><p><a name="Footnote_253" id="Footnote_253"></a><a href="#FNanchor_253"><span class="label">253</span></a> Sir J. Lubbock, ‘Prehistoric Times,’ 2nd edit. 1869, chap. xv. and +xvi. <i>et passim</i>.</p></div> + +<div class="footnote"><p><a name="Footnote_254" id="Footnote_254"></a><a href="#FNanchor_254"><span class="label">254</span></a> Dr. F. Müller has made some good remarks to this effect in the +'Reise der Novara: Anthropolog. Theil,’ Abtheil. iii. 1868, s. 127.</p></div> + +<div class="footnote"><p><a name="Footnote_255" id="Footnote_255"></a><a href="#FNanchor_255"><span class="label">255</span></a> Isidore Geoffroy St.-Hilaire gives a detailed account of the position +assigned to man by various naturalists in their classifications: ‘Hist. +Nat. Gén.’ tom. ii. 1859, p. 170-189.</p></div> + +<div class="footnote"><p><a name="Footnote_256" id="Footnote_256"></a><a href="#FNanchor_256"><span class="label">256</span></a> See the very interesting article, “L’Instinct chez les Insectes,” by +M. George Pouchet, ‘Revue des Deux Mondes,’ Feb. 1870, p. 682.</p></div> + +<div class="footnote"><p><a name="Footnote_257" id="Footnote_257"></a><a href="#FNanchor_257"><span class="label">257</span></a> Westwood, ‘Modern Class. of Insects,’ vol. ii. 1840, p. 87.</p></div> + +<div class="footnote"><p><a name="Footnote_258" id="Footnote_258"></a><a href="#FNanchor_258"><span class="label">258</span></a> ‘Proc. Zoolog. Soc.’ 1869, p. 4.</p></div> + +<div class="footnote"><p><a name="Footnote_259" id="Footnote_259"></a><a href="#FNanchor_259"><span class="label">259</span></a> ‘Evidence as to Man’s Place in Nature,’ 1863, p. 70, <i>et passim</i>.</p></div> + +<div class="footnote"><p><a name="Footnote_260" id="Footnote_260"></a><a href="#FNanchor_260"><span class="label">260</span></a> Isid. Geoffroy, ‘Hist. Nat. Gén.’ tom. ii. 1859, p. 217.</p></div> + +<div class="footnote"><p><a name="Footnote_261" id="Footnote_261"></a><a href="#FNanchor_261"><span class="label">261</span></a> “Ueber die Richtung der Haare,” &c., Müller’s ‘Archiv für Anat. +und Phys.’ 1837, s. 51.</p></div> + +<div class="footnote"><p><a name="Footnote_262" id="Footnote_262"></a><a href="#FNanchor_262"><span class="label">262</span></a> On the hair in Hylobates, see ‘Nat. Hist. of Mammals,’ by C. L. +Martin, 1841, p. 415. Also, Isid. Geoffroy on the American monkeys +and other kinds, ‘Hist. Nat. Gén.’ vol. ii. 1859, p. 216, 243. Eschricht, +ibid. s. 46, 55, 61. Owen, ‘Anat. of Vertebrates,’ vol. iii. p. 619. +Wallace, ‘Contributions to the Theory of Natural Selection,’ 1870. p. +344.</p></div> + +<div class="footnote"><p><a name="Footnote_263" id="Footnote_263"></a><a href="#FNanchor_263"><span class="label">263</span></a> ‘Origin of Species,’ 5th edit. 1869, p. 194. ‘The Variation of +Animals and Plants under Domestication,’ vol. ii. 1868, p. 348.</p></div> + +<div class="footnote"><p><a name="Footnote_264" id="Footnote_264"></a><a href="#FNanchor_264"><span class="label">264</span></a> ‘An Introduction to the Classification of Animals,’ 1869, p. 99.</p></div> + +<div class="footnote"><p><a name="Footnote_265" id="Footnote_265"></a><a href="#FNanchor_265"><span class="label">265</span></a> This is nearly the same classification as that provisionally adopted +by Mr. St. George Mivart (‘Transact. Philosoph. Soc.’ 1867, p. 300), +who, after separating the Lemuridæ, divides the remainder of the +Primates into the Hominidæ, the Simiadæ answering to the Catarhines, +the Cebidæ, and the Hapalidæ,—these two latter groups answering to +the Platyrhines.</p></div> + +<div class="footnote"><p><a name="Footnote_266" id="Footnote_266"></a><a href="#FNanchor_266"><span class="label">266</span></a> ‘Transact. Zoolog. Soc.’ vol. vi. 1867, p. 214.</p></div> + +<div class="footnote"><p><a name="Footnote_267" id="Footnote_267"></a><a href="#FNanchor_267"><span class="label">267</span></a> Mr. St. G. Mivart, ‘Transact. Phil. Soc.’ 1867, p. 410.</p></div> + +<div class="footnote"><p><a name="Footnote_268" id="Footnote_268"></a><a href="#FNanchor_268"><span class="label">268</span></a> Messrs. Murie and Mivart on the Lemuroidea. ‘Transact. Zoolog. +Soc.’ vol. vii. 1869, p. 5.</p></div> + +<div class="footnote"><p><a name="Footnote_269" id="Footnote_269"></a><a href="#FNanchor_269"><span class="label">269</span></a> Häckel has come to this same conclusion. See ‘Ueber die Entstehung +des Menschengeschlechts,’ in Virchow’s ‘Sammlung. gemein. +wissen. Vorträge,’ 1868, s. 61. Also his ‘Natürliche Schöpfungsgeschichte,’ +1868, in which he gives in detail his views on the genealogy +of man.</p></div> + +<div class="footnote"><p><a name="Footnote_270" id="Footnote_270"></a><a href="#FNanchor_270"><span class="label">270</span></a> ‘Anthropological Review,’ April, 1867, p. 236.</p></div> + +<div class="footnote"><p><a name="Footnote_271" id="Footnote_271"></a><a href="#FNanchor_271"><span class="label">271</span></a> ‘Elements of Geology,’ 1865, p. 583-585. ‘Antiquity of Man’, +1863; p. 145.</p></div> + +<div class="footnote"><p><a name="Footnote_272" id="Footnote_272"></a><a href="#FNanchor_272"><span class="label">272</span></a> ‘Man’s Place in Nature,’ p. 105.</p></div> + +<div class="footnote"><p><a name="Footnote_273" id="Footnote_273"></a><a href="#FNanchor_273"><span class="label">273</span></a> Elaborate tables are given in his ‘Generelle Morphologie’ (B. ii. +s. cliii. and s. 425); and with more especial reference to man in his +'Natürliche Schöpfungsgeschichte,’ 1868. Prof. Huxley, in reviewing +this latter work (‘The Academy,’ 1869, p. 42) says, that he considers +the phylum or lines of descent of the Vertebrata to be admirably discussed +by Häckel, although he differs on some points. He expresses, +also, his high estimate of the value of the general tenor and spirit of +the whole work.</p></div> + +<div class="footnote"><p><a name="Footnote_274" id="Footnote_274"></a><a href="#FNanchor_274"><span class="label">274</span></a> ‘Palæontology,’ 1860, p. 199.</p></div> + +<div class="footnote"><p><a name="Footnote_275" id="Footnote_275"></a><a href="#FNanchor_275"><span class="label">275</span></a> I had the satisfaction of seeing, at the Falkland Islands, in April, +1833, and therefore some years before any other naturalist, the locomotive +larvæ of a compound Ascidian, closely allied to, but apparently +generically distinct from, Synoicum. The tail was about five times as +long as the oblong head, and terminated in a very fine filament. It +was plainly divided, as sketched by me under a simple microscope, by +transverse opaque partitions, which I presume represent the great cells +figured by Kowalevsky. At an early stage of development the tail was +closely coiled round the head of the larva.</p></div> + +<div class="footnote"><p><a name="Footnote_276" id="Footnote_276"></a><a href="#FNanchor_276"><span class="label">276</span></a> ‘Mémoires de l’Acad. des Sciences de St. Pétersbourg,’ tom. x. +No. 15, 1866.</p></div> + +<div class="footnote"><p><a name="Footnote_277" id="Footnote_277"></a><a href="#FNanchor_277"><span class="label">277</span></a> This is the conclusion of one of the highest authorities in comparative +anatomy, namely, Prof. Gegenbaur: ‘Grundzüge der vergleich. +Anat.’ 1870, s. 876. The result has been arrived at chiefly from the +study of the Amphibia; but it appears from the researches of Waldeyer +(as quoted in Humphry’s ‘Journal of Anat. and Phys.’ 1869, p. 161), +that the sexual organs of even “the higher vertebrata are, in their early +condition, hermaphrodite.” Similar views have long been held by +some authors, though until recently not well based.</p></div> + +<div class="footnote"><p><a name="Footnote_278" id="Footnote_278"></a><a href="#FNanchor_278"><span class="label">278</span></a> The male Thylacinus offers the best instance. Owen, ‘Anatomy +of Vertebrates,’ vol. iii. p. 771.</p></div> + +<div class="footnote"><p><a name="Footnote_279" id="Footnote_279"></a><a href="#FNanchor_279"><span class="label">279</span></a> Serranus is well known often to be in an hermaphrodite condition; +but Dr. Günther informs me that he is convinced that this is not its +normal state. Descent from an ancient androgynous prototype would, +however, naturally favour and explain, to a certain extent, the recurrence +of this condition in these fishes.</p></div> + +<div class="footnote"><p><a name="Footnote_280" id="Footnote_280"></a><a href="#FNanchor_280"><span class="label">280</span></a> Mr. Lockwood believes (as quoted in ‘Quart. Journal of Science,’ +April, 1868, p. 269), from what he has observed of the development of +Hippocampus, that the walls of the abdominal pouch of the male +in some way afford nourishment. On male fishes hatching the ova in +their mouths, see a very interesting paper by Prof. Wyman, in ‘Proc. +Boston Soc. of Nat. Hist.’ Sept. 15, 1857; also Prof. Turner, in ‘Journal +of Anat. and Phys.’ Nov. 1, 1866, p. 78. Dr. Günther has likewise +described similar cases.</p></div> + +<div class="footnote"><p><a name="Footnote_281" id="Footnote_281"></a><a href="#FNanchor_281"><span class="label">281</span></a> All vital functions tend to run their course in fixed and recurrent +periods, and with tidal animals the periods would probably be lunar; +for such animals must have been left dry or covered deep with water,—supplied +with copious food or stinted,—during endless generations, at +regular lunar intervals. If then the Vertebrata are descended from an +animal allied to the existing tidal Ascidians, the mysterious fact, that +with the higher and now terrestrial Vertebrata, not to mention other +classes, many normal and abnormal vital processes run their course +according to lunar periods, is rendered intelligible. A recurrent period, +if approximately of the right duration, when once gained, would not, as +far as we can judge, be liable to be changed; consequently it might +be thus transmitted during almost any number of generations. This +conclusion, if it could be proved sound, would be curious; for we should +then see that the period of gestation in each mammal, and the hatching +of each bird’s eggs, and many other vital processes, still betrayed the +primordial birthplace of these animals.</p></div> + +<div class="footnote"><p><a name="Footnote_282" id="Footnote_282"></a><a href="#FNanchor_282"><span class="label">282</span></a> ‘History of India,’ 1841, vol. i. p. 323. Father Ripa makes exactly +the same remark with respect to the Chinese.</p></div> + +<div class="footnote"><p><a name="Footnote_283" id="Footnote_283"></a><a href="#FNanchor_283"><span class="label">283</span></a> A vast number of measurements of Whites, Blacks, and Indians, are +given in the ‘Investigations in the Military and Anthropolog. Statistics +of American Soldiers,’ by B. A. Gould, 1869, p. 298-358; on the +capacity of the lungs, p. 471. See also the numerous and valuable +tables, by Dr. Weisbach, from the observations of Dr. Scherzer and +Dr. Schwarz, in the ‘Reise der Novara: Anthropolog. Theil,’ 1867.</p></div> + +<div class="footnote"><p><a name="Footnote_284" id="Footnote_284"></a><a href="#FNanchor_284"><span class="label">284</span></a> See, for instance, Mr. Marshall’s account of the brain of a Bush-woman, +in ‘Phil. Transact.’ 1864, p. 519.</p></div> + +<div class="footnote"><p><a name="Footnote_285" id="Footnote_285"></a><a href="#FNanchor_285"><span class="label">285</span></a> Wallace, ‘The Malay Archipelago,’ vol. ii. 1869, p. 178.</p></div> + +<div class="footnote"><p><a name="Footnote_286" id="Footnote_286"></a><a href="#FNanchor_286"><span class="label">286</span></a> With respect to the figures in the famous Egyptian caves of Abou-Simbel, +M. Pouchet says (‘The Plurality of the Human Races,’ Eng. +translat. 1864, p. 50), that he was far from finding recognisable representations +of the dozen or more nations which some authors believe +that they can recognise. Even some of the most strongly-marked +races cannot be identified with that degree of unanimity which might +have been expected from what has been written on the subject. +Thus Messrs. Nott and Gliddon (‘Types of Mankind,’ p. 148) state +that Rameses II., or the Great, has features superbly European; whereas +Knox, another firm believer in the specific distinction of the races +of man (‘Races of Man,’ 1850, p. 201), speaking of young Memnon +(the same person with Rameses II., as I am informed by Mr. Birch) insists +in the strongest manner that he is identical in character with the +Jews of Antwerp. Again, whilst looking in the British Museum with +two competent judges, officers of the establishment, at the statue of +Amunoph III., we agreed that he had a strongly negro cast of features; +but Messrs. Nott and Gliddon (ibid. p. 146, fig. 53) describe him as +“a hybrid, but not of negro intermixture.”</p></div> + +<div class="footnote"><p><a name="Footnote_287" id="Footnote_287"></a><a href="#FNanchor_287"><span class="label">287</span></a> As quoted by Nott and Gliddon, ‘Types of Mankind,’ 1854, p. 439. +They give also corroborative evidence; but C. Vogt thinks that the +subject requires further investigation.</p></div> + +<div class="footnote"><p><a name="Footnote_288" id="Footnote_288"></a><a href="#FNanchor_288"><span class="label">288</span></a> “Diversity of Origin of the Human Races,” in the ‘Christian +Examiner,’ July, 1850.</p></div> + +<div class="footnote"><p><a name="Footnote_289" id="Footnote_289"></a><a href="#FNanchor_289"><span class="label">289</span></a> ‘Transact. B. Soc. of Edinburgh,’ vol. xxii. 1861, p. 567.</p></div> + +<div class="footnote"><p><a name="Footnote_290" id="Footnote_290"></a><a href="#FNanchor_290"><span class="label">290</span></a> ‘On the Phenomena of Hybridity in the Genus Homo,’ Eng. +translat. 1864.</p></div> + +<div class="footnote"><p><a name="Footnote_291" id="Footnote_291"></a><a href="#FNanchor_291"><span class="label">291</span></a> See the interesting letter by Mr. T. A. Murray, in the ‘Anthropolog. +Review,’ April, 1868, p. liii. In this letter Count Strzelecki’s +statement, that Australian women who have borne children to a white +man are afterwards sterile with their own race, is disproved. M. A. de +Quatrefages has also collected (‘Revue des Cours Scientifiques,’ March, +1869, p. 239) much evidence that Australians and Europeans are not +sterile when crossed.</p></div> + +<div class="footnote"><p><a name="Footnote_292" id="Footnote_292"></a><a href="#FNanchor_292"><span class="label">292</span></a> ‘An Examination of Prof. Agassiz’s Sketch of the Nat. Provinces +of the Animal World,’ Charleston, 1855, p. 44.</p></div> + +<div class="footnote"><p><a name="Footnote_293" id="Footnote_293"></a><a href="#FNanchor_293"><span class="label">293</span></a> ‘Military and Anthropolog. Statistics of American Soldiers,’ by +B. A. Gould, 1869, p. 319.</p></div> + +<div class="footnote"><p><a name="Footnote_294" id="Footnote_294"></a><a href="#FNanchor_294"><span class="label">294</span></a> ‘The Variation of Animals and Plants under Domestication,’ vol. ii. +p. 109. I may here remind the reader that the sterility of species +when crossed is not a specially-acquired quality; but, like the incapacity +of certain trees to be grafted together, is incidental on other +acquired differences. The nature of these differences is unknown, but +they relate more especially to the reproductive system, and much less +to external structure or to ordinary differences in constitution. One +important element in the sterility of crossed species apparently lies in +one or both having been long habituated to fixed conditions; for we +know that changed conditions have a special influence on the reproductive +system, and we have good reason to believe (as before remarked) +that the fluctuating conditions of domestication tend to eliminate +that sterility which is so general with species in a natural state +when crossed. It has elsewhere been shewn by me (ibid. vol. ii. p. 185, +and ‘Origin of Species,’ 5th edit. p. 317) that the sterility of crossed +species has not been acquired through natural selection: we can see that +when two forms have already been rendered very sterile, it is scarcely +possible that their sterility should be augmented by the preservation or +survival of the more and more sterile individuals; for as the sterility +increases fewer and fewer offspring will be produced from which to +breed, and at last only single individuals will be produced, at the rarest +intervals. But there is even a higher grade of sterility than this. Both +Gärtner and Kölreuter have proved that in genera of plants including +numerous species, a series can be formed from species which when +crossed yield fewer and fewer seeds, to species which never produce a +single seed, but yet are affected by the pollen of the other species, for +the germen swells. It is here manifestly impossible to select the more +sterile individuals, which have already ceased to yield seeds; so that +the acme of sterility, when the germen alone is affected, cannot be +gained through selection. This acme, and no doubt the other grades +of sterility, are the incidental results of certain unknown differences +in the constitution of the reproductive system of the species which are +crossed.</p></div> + +<div class="footnote"><p><a name="Footnote_295" id="Footnote_295"></a><a href="#FNanchor_295"><span class="label">295</span></a> ‘The Variation of Animals,’ &c., vol. ii. p. 92.</p></div> + +<div class="footnote"><p><a name="Footnote_296" id="Footnote_296"></a><a href="#FNanchor_296"><span class="label">296</span></a> M. de Quatrefages has given (‘Anthropolog. Review,’ Jan. 1869, +p. 22) an interesting account of the success and energy of the Paulistas +in Brazil, who are a much crossed race of Portuguese and Indians, with +a mixture of the blood of other races.</p></div> + +<div class="footnote"><p><a name="Footnote_297" id="Footnote_297"></a><a href="#FNanchor_297"><span class="label">297</span></a> For instance with the aborigines of America and Australia. Prof. +Huxley says (‘Transact. Internat. Congress of Prehist. Arch.’ 1868. p. +105) that the skulls of many South Germans and Swiss are “as short +and as broad as those of the Tartars,” &c.</p></div> + +<div class="footnote"><p><a name="Footnote_298" id="Footnote_298"></a><a href="#FNanchor_298"><span class="label">298</span></a> See a good discussion on this subject in Waitz, ‘Introduct. to +Anthropology,’ Eng. translat. 1863, p. 198-208, 227. I have taken +some of the above statements from H. Tuttle’s ‘Origin and Antiquity +of Physical Man,’ Boston, 1866, p. 35.</p></div> + +<div class="footnote"><p><a name="Footnote_299" id="Footnote_299"></a><a href="#FNanchor_299"><span class="label">299</span></a> Prof. Nägeli has carefully described several striking cases in his +'Botanische Mittheilungen,’ B. ii. 1866, s. 294-369. Prof. Asa Gray +has made analogous remarks on some intermediate forms in the Compositæ +of N. America.</p></div> + +<div class="footnote"><p><a name="Footnote_300" id="Footnote_300"></a><a href="#FNanchor_300"><span class="label">300</span></a> ‘Origin of Species,’ 5th edit. p. 68.</p></div> + +<div class="footnote"><p><a name="Footnote_301" id="Footnote_301"></a><a href="#FNanchor_301"><span class="label">301</span></a> See Prof. Huxley to this effect in the ‘Fortnightly Review,’ 1865, +p. 275.</p></div> + +<div class="footnote"><p><a name="Footnote_302" id="Footnote_302"></a><a href="#FNanchor_302"><span class="label">302</span></a> ‘Lectures on Man,’ Eng. translat. 1864, p. 468.</p></div> + +<div class="footnote"><p><a name="Footnote_303" id="Footnote_303"></a><a href="#FNanchor_303"><span class="label">303</span></a> ‘Die Racen des Schweines,’ 1860, s. 46. ‘Vorstudien für Geschichte, +&c., Schweineschädel,’ 1864, s. 104. With respect to cattle, +see M. de Quatrefages, ‘Unité de l’Espèce Humaine,’ 1861, p. 119.</p></div> + +<div class="footnote"><p><a name="Footnote_304" id="Footnote_304"></a><a href="#FNanchor_304"><span class="label">304</span></a> Tylor’s ‘Early History of Mankind,’ 1865; for the evidence with +respect to gesture-language, see p. 54. Lubbock’s ‘Prehistoric Times,’ +2nd edit. 1869.</p></div> + +<div class="footnote"><p><a name="Footnote_305" id="Footnote_305"></a><a href="#FNanchor_305"><span class="label">305</span></a> ‘The Primitive Inhabitants of Scandinavia,’ Eng. translat. edited +by Sir J. Lubbock, 1868, p. 104.</p></div> + +<div class="footnote"><p><a name="Footnote_306" id="Footnote_306"></a><a href="#FNanchor_306"><span class="label">306</span></a> Hodder M. Westropp, on Cromlechs, &c., ‘Journal of Ethnological +Soc.’ as given in ‘Scientific Opinion,’ June 2nd. 1869, p. 3.</p></div> + +<div class="footnote"><p><a name="Footnote_307" id="Footnote_307"></a><a href="#FNanchor_307"><span class="label">307</span></a> ‘Journal of Researches: Voyage of the “Beagle,”’ p. 46.</p></div> + +<div class="footnote"><p><a name="Footnote_308" id="Footnote_308"></a><a href="#FNanchor_308"><span class="label">308</span></a> ‘Prehistoric Times,’ 1869, p. 574.</p></div> + +<div class="footnote"><p><a name="Footnote_309" id="Footnote_309"></a><a href="#FNanchor_309"><span class="label">309</span></a> Translation in ‘Anthropological Review,’ Oct. 1868, p. 431.</p></div> + +<div class="footnote"><p><a name="Footnote_310" id="Footnote_310"></a><a href="#FNanchor_310"><span class="label">310</span></a> ‘Transact. Internat. Congress of Prehistoric Arch.’ 1868, p. 172-175. +See also Broca (translation) in ‘Anthropological Review,’ Oct. +1868, p. 410.</p></div> + +<div class="footnote"><p><a name="Footnote_311" id="Footnote_311"></a><a href="#FNanchor_311"><span class="label">311</span></a> Dr. Gerland, ‘Ueber das Aussterben der Naturvölker,’ 1868, s. 82.</p></div> + +<div class="footnote"><p><a name="Footnote_312" id="Footnote_312"></a><a href="#FNanchor_312"><span class="label">312</span></a> Gerland (ibid. s. 12) gives facts in support of this statement.</p></div> + +<div class="footnote"><p><a name="Footnote_313" id="Footnote_313"></a><a href="#FNanchor_313"><span class="label">313</span></a> See remarks to this effect in Sir H. Holland’s ‘Medical Notes and +Reflections,’ 1839, p. 390.</p></div> + +<div class="footnote"><p><a name="Footnote_314" id="Footnote_314"></a><a href="#FNanchor_314"><span class="label">314</span></a> I have collected (‘Journal of Researches, Voyage of the “Beagle,”’ +p. 435) a good many cases bearing on this subject: see also Gerland, +ibid. s. 8. Poeppig speaks of the “breath of civilisation as poisonous +to savages.”</p></div> + +<div class="footnote"><p><a name="Footnote_315" id="Footnote_315"></a><a href="#FNanchor_315"><span class="label">315</span></a> Sproat, ‘Scenes and Studies of Savage Life,’ 1868, p. 284.</p></div> + +<div class="footnote"><p><a name="Footnote_316" id="Footnote_316"></a><a href="#FNanchor_316"><span class="label">316</span></a> Bagehot, “Physics and Politics,” ‘Fortnightly Review,’ April 1, +1868, p. 455.</p></div> + +<div class="footnote"><p><a name="Footnote_317" id="Footnote_317"></a><a href="#FNanchor_317"><span class="label">317</span></a> “On Anthropology,” translation, ‘Anthropolog. Review,’ Jan. +1868, p. 38.</p></div> + +<div class="footnote"><p><a name="Footnote_318" id="Footnote_318"></a><a href="#FNanchor_318"><span class="label">318</span></a> ‘The Annals of Rural Bengal,’ 1868, p. 134.</p></div> + +<div class="footnote"><p><a name="Footnote_319" id="Footnote_319"></a><a href="#FNanchor_319"><span class="label">319</span></a> ‘The Variation of Animals and Plants under Domestication,’ vol. +ii. p. 95.</p></div> + +<div class="footnote"><p><a name="Footnote_320" id="Footnote_320"></a><a href="#FNanchor_320"><span class="label">320</span></a> Pallas, ‘Act. Acad. St. Petersburgh,’ 1780, part ii. p. 69. He +was followed by Rudolphi, in his ‘Beyträge zur Anthropologie,’ 1812. +An excellent summary of the evidence is given by Godron, ‘De +l’Espèce,’ 1859, vol. ii. p. 246, &c.</p></div> + +<div class="footnote"><p><a name="Footnote_321" id="Footnote_321"></a><a href="#FNanchor_321"><span class="label">321</span></a> Sir Andrew Smith, as quoted by Knox, ‘Races of Man,’ 1850, +p. 473.</p></div> + +<div class="footnote"><p><a name="Footnote_322" id="Footnote_322"></a><a href="#FNanchor_322"><span class="label">322</span></a> See De Quatrefages on this head, ‘Revue des Cours Scientifiques,’ +Oct. 17, 1868, p. 731.</p></div> + +<div class="footnote"><p><a name="Footnote_323" id="Footnote_323"></a><a href="#FNanchor_323"><span class="label">323</span></a> Livingstone’s ‘Travels and Researches in S. Africa,’ 1857, p. 338, +329. D’Orbigny, as quoted by Godron, ‘De l’Espèce,’ vol. ii. p. 266.</p></div> + +<div class="footnote"><p><a name="Footnote_324" id="Footnote_324"></a><a href="#FNanchor_324"><span class="label">324</span></a> See a paper read before the Royal Soc. in 1813, and published in +his Essays in 1818. I have given an account of Dr. Wells’ views in the +Historical Sketch (p. xvi) to my ‘Origin of Species.’ Various cases of +colour correlated with constitutional peculiarities are given in my +'Variation of Animals under Domestication,’ vol. ii. p. 227, 335.</p></div> + +<div class="footnote"><p><a name="Footnote_325" id="Footnote_325"></a><a href="#FNanchor_325"><span class="label">325</span></a> See, for instance, Nott and Gliddon, ‘Types of Mankind,’ p. 68.</p></div> + +<div class="footnote"><p><a name="Footnote_326" id="Footnote_326"></a><a href="#FNanchor_326"><span class="label">326</span></a> Major Tulloch, in a paper read before the Statistical Society, +April 20th, 1840, and given in the ‘Athenæum,’ 1840, p. 353.</p></div> + +<div class="footnote"><p><a name="Footnote_327" id="Footnote_327"></a><a href="#FNanchor_327"><span class="label">327</span></a> ‘The Plurality of the Human Race’ (translat.), 1864, p. 60.</p></div> + +<div class="footnote"><p><a name="Footnote_328" id="Footnote_328"></a><a href="#FNanchor_328"><span class="label">328</span></a> Quatrefages, ‘Unité de l’Espèce Humaine,’ 1861, p. 205. Waitz, +'Introduct. to Anthropology,’ translat. vol. i. 1863, p. 124. Livingstone +gives analogous cases in his ‘Travels.’</p></div> + +<div class="footnote"><p><a name="Footnote_329" id="Footnote_329"></a><a href="#FNanchor_329"><span class="label">329</span></a> In the spring of 1862 I obtained permission from the Director-General +of the Medical department of the Army, to transmit to the +surgeons of the various regiments on foreign service a blank table, +with the following appended remarks, but I have received no returns. +“As several well-marked cases have been recorded with our domestic +animals of a relation between the colour of the dermal appendages +and the constitution; and it being notorious that there is some limited +degree of relation between the colour of the races of man and the +climate inhabited by them; the following investigation seems worth +consideration. Namely, whether there is any relation in Europeans +between the colour of their hair, and their liability to the diseases of +tropical countries. If the surgeons of the several regiments, when +stationed in unhealthy tropical districts, would be so good as first to +count, as a standard of comparison, how many men, in the force +whence the sick are drawn, have dark and light-coloured hair, and +hair of intermediate or doubtful tints; and if a similar account were +kept by the same medical gentlemen, of all the men who suffered +from malarious and yellow fevers, or from dysentery, it would soon +be apparent, after some thousand cases had been tabulated, whether +there exists any relation between the colour of the hair and constitutional +liability to tropical diseases. Perhaps no such relation would +be discovered, but the investigation is well worth making. In case +any positive result were obtained, it might be of some practical use +in selecting men for any particular service. Theoretically the result +would be of high interest, as indicating one means by which a race +of men inhabiting from a remote period an unhealthy tropical climate, +might have become dark-coloured by the better preservation of dark-haired +or dark-complexioned individuals during a long succession of +generations.”</p></div> + +<div class="footnote"><p><a name="Footnote_330" id="Footnote_330"></a><a href="#FNanchor_330"><span class="label">330</span></a> ‘Anthropological Review,’ Jan. 1866, p. xxi.</p></div> + +<div class="footnote"><p><a name="Footnote_331" id="Footnote_331"></a><a href="#FNanchor_331"><span class="label">331</span></a> See, for instance, Quatrefages (‘Revue des Cours Scientifiques,’ +Oct. 10, 1868, p. 724) on the effects of residence in Abyssinia and +Arabia, and other analogous cases. Dr. Rolle (‘Der Mensch, seine +Abstammung,’ &c., 1865, s. 99) states, on the authority of Khanikof, +that the greater number of German families settled in Georgia, have +acquired in the course of two generations dark hair and eyes. Mr. D. +Forbes informs me that the Quichuas in the Andes vary greatly in +colour, according to the position of the valleys inhabited by them.</p></div> + +<div class="footnote"><p><a name="Footnote_332" id="Footnote_332"></a><a href="#FNanchor_332"><span class="label">332</span></a> Harlan, ‘Medical Researches,’ p. 532. Quatrefages (‘Unité de +l’Espèce Humaine,’ 1861, p. 128) has collected much evidence on this +head.</p></div> + +<div class="footnote"><p><a name="Footnote_333" id="Footnote_333"></a><a href="#FNanchor_333"><span class="label">333</span></a> See Prof. Schaaffhausen, translat. in ‘Anthropological Review,’ +Oct. 1868, p. 429.</p></div> + +<div class="footnote"><p><a name="Footnote_334" id="Footnote_334"></a><a href="#FNanchor_334"><span class="label">334</span></a> Mr. Catlin states (‘N. American Indians,’ 3rd edit. 1842, vol. i. p. +49) that in the whole tribe of the Mandans, about one in ten or twelve +of the members of all ages and both sexes have bright silvery grey hair, +which is hereditary. Now this hair is as coarse and harsh as that of +a horse’s mane, whilst the hair of other colours is fine and soft.</p></div> + +<div class="footnote"><p><a name="Footnote_335" id="Footnote_335"></a><a href="#FNanchor_335"><span class="label">335</span></a> On the odour of the skin, Godron, ‘Sur l’Espèce,’ tom. ii. p. 217. +On the pores in the skin, Dr. Wilckens, ‘Die Aufgaben der landwirth. +Zootechnik,’ 1869, s. 7.</p></div> + +<div class="footnote"><p><a name="Footnote_336" id="Footnote_336"></a><a href="#FNanchor_336"><span class="label">336</span></a> Westwood, ‘Modern Class. of Insects,’ vol. ii. 1810, p. 541. In +regard to the statement about Tanais, mentioned below, I am indebted +to Fritz Müller.</p></div> + +<div class="footnote"><p><a name="Footnote_337" id="Footnote_337"></a><a href="#FNanchor_337"><span class="label">337</span></a> Kirby and Spence, ‘Introduction to Entomology,’ vol. iii. 1826, +p. 309.</p></div> + +<div class="footnote"><p><a name="Footnote_338" id="Footnote_338"></a><a href="#FNanchor_338"><span class="label">338</span></a> Even with those of plants in which the sexes are separate, the male +flowers are generally mature before the female. Many hermaphrodite +plants are, as first shewn by C. K. Sprengel, dichogamous; that is, +their male and female organs are not ready at the same time, so that +they cannot be self-fertilised. Now with such plants the pollen is +generally mature in the same flower before the stigma, though there +are some exceptional species in which the female organs are mature +before the male.</p></div> + +<div class="footnote"><p><a name="Footnote_339" id="Footnote_339"></a><a href="#FNanchor_339"><span class="label">339</span></a> I have received information, hereafter to be given, to this effect +with respect to poultry. Even with birds, such as pigeons, which +pair for life, the female, as I hear from Mr. Jenner Weir, will desert +her mate if he is injured or grows weak.</p></div> + +<div class="footnote"><p><a name="Footnote_340" id="Footnote_340"></a><a href="#FNanchor_340"><span class="label">340</span></a> On the Gorilla, Savage and Wyman, ‘Boston Journal of Nat. Hist.’ +vol. v. 1845-47, p. 423. On Cynocephalus, Brehm, ‘Illust. Thierleben,’ +B. i. 1864, s. 77. On Mycetes, Rengger, ‘Naturgesch.: Säugethiere +von Paraguay,’ 1830, s. 14, 20. On Cebus, Brehm, ibid. s. 108.</p></div> + +<div class="footnote"><p><a name="Footnote_341" id="Footnote_341"></a><a href="#FNanchor_341"><span class="label">341</span></a> Pallas, ‘Spicilegia Zoolog.’ Fasc. xii. 1777, p. 29. Sir Andrew +Smith, ‘Illustrations of the Zoology of S. Africa,’ 1849, pl. 29, on the +Kobus. Owen, in his ‘Anatomy of Vertebrates’ (vol. iii. 1868, p. 633) +gives a table incidentally showing which species of Antelopes pair and +which are gregarious.</p></div> + +<div class="footnote"><p><a name="Footnote_342" id="Footnote_342"></a><a href="#FNanchor_342"><span class="label">342</span></a> Dr. Campbell, in ‘Proc. Zoolog. Soc.’ 1869, p. 138. See also an +interesting paper, by Lieut. Johnstone, in ‘Proc. Asiatic Soc. of Bengal,’ +May, 1868.</p></div> + +<div class="footnote"><p><a name="Footnote_343" id="Footnote_343"></a><a href="#FNanchor_343"><span class="label">343</span></a> ‘The Ibis,’ vol. iii. 1861, p. 133, on the Progne Widow-bird. See +also on the Vidua axillaris, ibid. vol. ii. 1860, p. 211. On the polygamy +of the Capercailzie and Great Bustard, see L. Lloyd, ‘Game Birds +of Sweden,’ 1867, p. 19, and 182. Montagu and Selby speak of the +Black Grouse as polygamous and of the Red Grouse as monogamous.</p></div> + +<div class="footnote"><p><a name="Footnote_344" id="Footnote_344"></a><a href="#FNanchor_344"><span class="label">344</span></a> The Rev. E. S. Dixon, however, speaks positively (‘Ornamental +Poultry,’ 1848, p. 76) about the eggs of the guinea-fowl being infertile +when more than one female is kept with the same male.</p></div> + +<div class="footnote"><p><a name="Footnote_345" id="Footnote_345"></a><a href="#FNanchor_345"><span class="label">345</span></a> Noel Humphreys, ‘River Gardens,’ 1857.</p></div> + +<div class="footnote"><p><a name="Footnote_346" id="Footnote_346"></a><a href="#FNanchor_346"><span class="label">346</span></a> Kirby and Spence, ‘Introduction to Entomology,’ vol. iii. 1826, +p. 342.</p></div> + +<div class="footnote"><p><a name="Footnote_347" id="Footnote_347"></a><a href="#FNanchor_347"><span class="label">347</span></a> One parasitic Hymenopterous insect (Westwood, ‘Modern Class. +of Insects,’ vol. ii, p. 160) forms an exception to the rule, as the male +has rudimentary wings, and never quits the cell in which it is born, +whilst the female has well-developed wings. Audouin believes that +the females are impregnated by the males which are born in the same +cells with them; but it is much more probable that the females visit +other cells, and thus avoid close inter-breeding. We shall hereafter +meet with a few exceptional cases, in various classes, in which the +female, instead of the male, is the seeker and wooer.</p></div> + +<div class="footnote"><p><a name="Footnote_348" id="Footnote_348"></a><a href="#FNanchor_348"><span class="label">348</span></a> ‘Essays and Observations,’ edited by Owen, vol. i. 1861, p. 194.</p></div> + +<div class="footnote"><p><a name="Footnote_349" id="Footnote_349"></a><a href="#FNanchor_349"><span class="label">349</span></a> Prof. Sachs (‘Lehrbuch der Botanik,’ 1870, s. 633) in speaking of +the male and female reproductive cells, remarks, “verhält sich die eine +bei der Vereinigung activ, ... die andere erscheint bei der Vereinigung +passiv.”</p></div> + +<div class="footnote"><p><a name="Footnote_350" id="Footnote_350"></a><a href="#FNanchor_350"><span class="label">350</span></a> ‘Reise der Novara: Anthropolog. Theil,’ 1867, s. 216-269. The +results were calculated by Dr. Weisbach from measurements made by +Drs. K. Scherzer and Schwarz. On the greater variability of the males +of domesticated animals, see my ‘Variation of Animals and Plants +under Domestication,’ vol. ii. 1868, p. 75.</p></div> + +<div class="footnote"><p><a name="Footnote_351" id="Footnote_351"></a><a href="#FNanchor_351"><span class="label">351</span></a> ‘Proceedings Royal Soc.’ vol. xvi. July, 1868, p. 519 and 524.</p></div> + +<div class="footnote"><p><a name="Footnote_352" id="Footnote_352"></a><a href="#FNanchor_352"><span class="label">352</span></a> ‘Proc. Royal Irish Academy,’ vol. x. 1868, p. 123.</p></div> + +<div class="footnote"><p><a name="Footnote_353" id="Footnote_353"></a><a href="#FNanchor_353"><span class="label">353</span></a> ‘Massachusetts Medical Soc.’ vol. ii. No. 3, 1808, p. 9.</p></div> + +<div class="footnote"><p><a name="Footnote_354" id="Footnote_354"></a><a href="#FNanchor_354"><span class="label">354</span></a> ‘The Variation of Animals and Plants under Domestication,’ vol. +ii. 1868, p. 75. In the last chapter but one, the provisional hypothesis +of pangenesis, above alluded to, is fully explained.</p></div> + +<div class="footnote"><p><a name="Footnote_355" id="Footnote_355"></a><a href="#FNanchor_355"><span class="label">355</span></a> These facts are given on the high authority of a great breeder, +Mr. Teebay, in Tegetmeier’s ‘Poultry Book,’ 1868, p. 158. On the +characters of chickens of different breeds, and on the breeds of the +pigeon, alluded to in the above paragraph, see ‘Variation of Animals,’ +&c., vol. i. p. 160, 249; vol. ii. p. 77.</p></div> + +<div class="footnote"><p><a name="Footnote_356" id="Footnote_356"></a><a href="#FNanchor_356"><span class="label">356</span></a> ‘Novæ species Quadrupedum e Glirium ordine,’ 1778, p. 7. On +the transmission of colour by the horse, see ‘Variation of Animals, &c. +under Domestication,’ vol. i. p. 21. Also vol. ii. p. 71, for a general +discussion on Inheritance as limited by Sex.</p></div> + +<div class="footnote"><p><a name="Footnote_357" id="Footnote_357"></a><a href="#FNanchor_357"><span class="label">357</span></a> Dr. Chapuis, ‘Le Pigeon Voyageur Belge,’ 1865, p. 87. Boitard +et Corbié, ‘Les Pigeons de Volière,’ &c., 1824, p. 173.</p></div> + +<div class="footnote"><p><a name="Footnote_358" id="Footnote_358"></a><a href="#FNanchor_358"><span class="label">358</span></a> References are given in my ‘Variation of Animals under Domestication,’ +vol. ii. p. 72.</p></div> + +<div class="footnote"><p><a name="Footnote_359" id="Footnote_359"></a><a href="#FNanchor_359"><span class="label">359</span></a> I am much obliged to Mr. Cupples for having made enquiries for +me in regard to the Roebuck and Red Deer of Scotland from Mr. +Robertson, the experienced head-forester to the Marquis of Breadalbane. +In regard to Fallow-deer, I am obliged to Mr. Eyton and others for +information. For the <i>Cervus alces</i> of N. America, see ‘Land and Water,’ +1868, p. 221 and 254; and for the <i>C. Virginianus</i> and <i>strongyloceros</i> of +the same continent, see J. D. Caton, in ‘Ottawa Acad. of Nat. Sc.’ +1868, p. 13. For <i>Cervus Eldi</i> of Pegu, see Lieut. Beavan, ‘Proc. +Zoolog. Soc.’ 1867, p. 762.</p></div> + +<div class="footnote"><p><a name="Footnote_360" id="Footnote_360"></a><a href="#FNanchor_360"><span class="label">360</span></a> <i>Antilocapra Americana.</i> Owen, ‘Anatomy of Vertebrates,’ vol. +iii. p. 627.</p></div> + +<div class="footnote"><p><a name="Footnote_361" id="Footnote_361"></a><a href="#FNanchor_361"><span class="label">361</span></a> I have been assured that the horns of the sheep in North Wales +can always be felt, and are sometimes even an inch in length, at birth. +With cattle Youatt says (‘Cattle,’ 1834, p. 277) that the prominence of +the frontal bone penetrates the cutis at birth, and that the horny +matter is soon formed over it.</p></div> + +<div class="footnote"><p><a name="Footnote_362" id="Footnote_362"></a><a href="#FNanchor_362"><span class="label">362</span></a> I am greatly indebted to Prof. Victor Carus for having made +inquiries for me, from the highest authorities, with respect to the +merino sheep of Saxony. On the Guinea coast of Africa there is a +breed of sheep in which, as with merinos, the rams alone bear horns; +and Mr. Winwood Reade informs me that in the one case observed, a +young ram born on Feb. 10th first showed horns on March 6th, so +that in this instance the development of the horns occurred at a later +period of life, conformably with our rule, than in the Welsh sheep, in +which both sexes are horned.</p></div> + +<div class="footnote"><p><a name="Footnote_363" id="Footnote_363"></a><a href="#FNanchor_363"><span class="label">363</span></a> In the common peacock (<i>Pavo cristatus</i>) the male alone possesses +spurs, whilst both sexes of the Java peacock (<i>P. muticus</i>) offer the +unusual case of being furnished with spurs. Hence I fully expected +that in the latter species they would have been developed earlier in life +than in the common peacock; but M. Hegt of Amsterdam informs me, +that with young birds of the previous year, belonging to both species, +compared on April 23rd, 1869, there was no difference in the development +of the spurs. The spurs, however, were as yet represented merely +by slight knobs or elevations. I presume that I should have been informed +if any difference in the rate of development had subsequently +been observed.</p></div> + +<div class="footnote"><p><a name="Footnote_364" id="Footnote_364"></a><a href="#FNanchor_364"><span class="label">364</span></a> In some other species of the Duck Family the speculum in the +two sexes differs in a greater degree; but I have not been able to discover +whether its full development occurs later in life in the males of +such species, than in the male of the common duck, as ought to be the +case according to our rule. With the allied <i>Mergus cucullatus</i> we have, +however, a case of this kind: the two sexes differ conspicuously in +general plumage, and to a considerable degree in the speculum, which +is pure white in the male and greyish-white in the female. Now the +young males at first resemble, in all respects, the female, and have a +greyish-white speculum, but this becomes pure white at an earlier age +than that at which the adult male acquires his other more strongly-marked +sexual differences in plumage: see Audubon, ‘Ornithological +Biography,’ vol. iii. 1835, p. 249-250.</p></div> + +<div class="footnote"><p><a name="Footnote_365" id="Footnote_365"></a><a href="#FNanchor_365"><span class="label">365</span></a> ‘Das Ganze der Taubenzucht,’ 1837, s. 21, 24. For the case of +the streaked pigeons, see Dr. Chapuis, ‘Le Pigeon Voyageur Belge.’ +1865, p. 87.</p></div> + +<div class="footnote"><p><a name="Footnote_366" id="Footnote_366"></a><a href="#FNanchor_366"><span class="label">366</span></a> For full particulars and references on all these points respecting +the several breeds of the Fowl, see ‘Variation of Animals and Plants +under Domestication,’ vol. i. p. 250, 256. In regard to the higher +animals, the sexual differences which have arisen under domestication +are described in the same work under the head of each species.</p></div> + +<div class="footnote"><p><a name="Footnote_367" id="Footnote_367"></a><a href="#FNanchor_367"><span class="label">367</span></a> ‘Twenty-ninth Annual Report of the Registrar-General for 1866.’ +In this report (p. xii) a special decennial table is given.</p></div> + +<div class="footnote"><p><a name="Footnote_368" id="Footnote_368"></a><a href="#FNanchor_368"><span class="label">368</span></a> For Norway and Russia, see abstract of Prof. Faye’s researches, +in ‘British and Foreign Medico-Chirurg. Review,’ April, 1867, p. 343, +345. For France, the ‘Annuaire pour l’An 1867.’ p. 213.</p></div> + +<div class="footnote"><p><a name="Footnote_369" id="Footnote_369"></a><a href="#FNanchor_369"><span class="label">369</span></a> In regard to the Jews, see M. Thury, ‘La Loi de Production des +Sexes,’ 1863, p. 25.</p></div> + +<div class="footnote"><p><a name="Footnote_370" id="Footnote_370"></a><a href="#FNanchor_370"><span class="label">370</span></a> Babbage, ‘Edinburgh Journal of Science,’ 1829, vol. i. p. 88; also +p. 90, on still-born children. On illegitimate children in England, +see ‘Report of Registrar-General for 1866,’ p. xv.</p></div> + +<div class="footnote"><p><a name="Footnote_371" id="Footnote_371"></a><a href="#FNanchor_371"><span class="label">371</span></a> ‘British and Foreign Medico-Chirurg. Review,’ April, 1867, p. +343. Dr. Stark also remarks (‘Tenth Annual Report of Births, Deaths, +&c., in Scotland,’ 1867, p. xxviii) that “These examples may suffice +to shew that, at almost every stage of life, the males in Scotland +have a greater liability to death and a higher death-rate than the +females. The fact, however, of this peculiarity being most strongly +developed at that infantile period of life when the dress, food, and +general treatment of both sexes are alike, seems to prove that the +higher male death-rate is an impressed, natural, and constitutional +peculiarity due to sex alone.”</p></div> + +<div class="footnote"><p><a name="Footnote_372" id="Footnote_372"></a><a href="#FNanchor_372"><span class="label">372</span></a> With the savage Guaranys of Paraguay, according to the accurate +Azara (‘Voyages dans l’Amérique mérid.’ tom. ii. 1809, p. 60, 179), +the women in proportion to the men are as 14 to 13.</p></div> + +<div class="footnote"><p><a name="Footnote_373" id="Footnote_373"></a><a href="#FNanchor_373"><span class="label">373</span></a> Leuckart in Wagner, ‘Handwörterbuch der Phys.’ B. iv. 1853, +s. 774.</p></div> + +<div class="footnote"><p><a name="Footnote_374" id="Footnote_374"></a><a href="#FNanchor_374"><span class="label">374</span></a> Anthropological Review, April, 1870, p. cviii.</p></div> + +<div class="footnote"><p><a name="Footnote_375" id="Footnote_375"></a><a href="#FNanchor_375"><span class="label">375</span></a> During the last eleven years a record has been kept of the number of +mares which have proved barren or prematurely slipped their foals; and it +deserves notice, as shewing how infertile these highly-nurtured and rather +closely-interbred animals have become, that not far from one-third of the +mares failed to produce living foals. Thus during 1866, 809 male colts +and 816 female colts were born, and 743 mares failed to produce offspring. +During 1867, 836 males and 902 females were born, and 794 mares failed.</p></div> + +<div class="footnote"><p><a name="Footnote_376" id="Footnote_376"></a><a href="#FNanchor_376"><span class="label">376</span></a> I am much indebted to Mr. Cupples for having procured for me the +above returns from Scotland, as well as some of the following returns on +cattle. Mr. R. Elliot, of Laighwood, first called my attention to the premature +deaths of the males,—a statement subsequently confirmed by Mr. +Aitchison and others. To this latter gentleman, and to Mr. Payan, I owe +my thanks for the larger returns on sheep.</p></div> + +<div class="footnote"><p><a name="Footnote_377" id="Footnote_377"></a><a href="#FNanchor_377"><span class="label">377</span></a> Bell, ‘History of British Quadrupeds,’ p. 100.</p></div> + +<div class="footnote"><p><a name="Footnote_378" id="Footnote_378"></a><a href="#FNanchor_378"><span class="label">378</span></a> ‘Illustrations of the Zoology of S. Africa,’ 1849, pl. 29.</p></div> + +<div class="footnote"><p><a name="Footnote_379" id="Footnote_379"></a><a href="#FNanchor_379"><span class="label">379</span></a> Brehm (‘Illust. Thierleben,’ B. iv. s. 990) comes to the same conclusion.</p></div> + +<div class="footnote"><p><a name="Footnote_380" id="Footnote_380"></a><a href="#FNanchor_380"><span class="label">380</span></a> On the authority of L. Lloyd, ‘Game Birds of Sweden,’ 1867, p. 12, +132.</p></div> + +<div class="footnote"><p><a name="Footnote_381" id="Footnote_381"></a><a href="#FNanchor_381"><span class="label">381</span></a> ‘Nat. Hist. of Selbourne,’ letter xxix. edit. of 1825, vol. i. p. 139.</p></div> + +<div class="footnote"><p><a name="Footnote_382" id="Footnote_382"></a><a href="#FNanchor_382"><span class="label">382</span></a> Mr. Jenner Weir received similar information, on making enquiries during +the following year. To shew the number of chaffinches caught, I +may mention that in 1869 there was a match between two experts; and +one man caught in a day 62, and another 40, male chaffinches. The greatest +number ever caught by one man in a single day was 70.</p></div> + +<div class="footnote"><p><a name="Footnote_383" id="Footnote_383"></a><a href="#FNanchor_383"><span class="label">383</span></a> ‘Ibis,’ vol. ii. p. 260, as quoted in Gould’s ‘Trochilidæ,’ 1861, p. 52. +For the foregoing proportions, I am indebted to Mr. Salvin for a table of +his results.</p></div> + +<div class="footnote"><p><a name="Footnote_384" id="Footnote_384"></a><a href="#FNanchor_384"><span class="label">384</span></a> ‘Ibis,’ 1860, p. 137; and 1867, p. 369.</p></div> + +<div class="footnote"><p><a name="Footnote_385" id="Footnote_385"></a><a href="#FNanchor_385"><span class="label">385</span></a> ‘Ibis,’ 1862, p. 137.</p></div> + +<div class="footnote"><p><a name="Footnote_386" id="Footnote_386"></a><a href="#FNanchor_386"><span class="label">386</span></a> Leuckart quotes Bloch (Wagner, ‘Handwörterbuch der Phys.’ B. iv. +1853, s. 775), that with fish there are twice as many males as females.</p></div> + +<div class="footnote"><p><a name="Footnote_387" id="Footnote_387"></a><a href="#FNanchor_387"><span class="label">387</span></a> Quoted in the ‘Farmer,’ March 18, 1869, p. 369.</p></div> + +<div class="footnote"><p><a name="Footnote_388" id="Footnote_388"></a><a href="#FNanchor_388"><span class="label">388</span></a> ‘The Stormontfield Piscicultural Experiments,’ 1866, p. 23. The +'Field’ newspaper, June 29th, 1867.</p></div> + +<div class="footnote"><p><a name="Footnote_389" id="Footnote_389"></a><a href="#FNanchor_389"><span class="label">389</span></a> ‘Land and Water,’ 1868, p. 41.</p></div> + +<div class="footnote"><p><a name="Footnote_390" id="Footnote_390"></a><a href="#FNanchor_390"><span class="label">390</span></a> Yarrell, ‘Hist. British Fishes,’ vol. i. 1836, p. 307; on the <i>Cyprinus +carpio</i>, p. 331; on the <i>Tinca vulgaris</i>, p. 331; on the <i>Abramis brama</i>, p. +336. See, for the minnow (<i>Leuciscus phoxinus</i>), ‘Loudon’s Mag. of Nat. +Hist.’ vol. v. 1832, p. 682.</p></div> + +<div class="footnote"><p><a name="Footnote_391" id="Footnote_391"></a><a href="#FNanchor_391"><span class="label">391</span></a> Leuckart quotes Meinecke (Wagner, ‘Handwörterbuch der Phys.’ +B. iv. 1853, s. 775) that with Butterflies the males are three or four times +as numerous as the females.</p></div> + +<div class="footnote"><p><a name="Footnote_392" id="Footnote_392"></a><a href="#FNanchor_392"><span class="label">392</span></a> ‘The Naturalist on the Amazons,’ vol. ii. 1863, p. 228, 347.</p></div> + +<div class="footnote"><p><a name="Footnote_393" id="Footnote_393"></a><a href="#FNanchor_393"><span class="label">393</span></a> Four of these cases are given by Mr. Trimen in his ‘Rhopalocera +Africæ Australis.’</p></div> + +<div class="footnote"><p><a name="Footnote_394" id="Footnote_394"></a><a href="#FNanchor_394"><span class="label">394</span></a> Quoted by Trimen, ‘Transact. Ent. Soc.’ vol. v. part iv. 1866, p. 330.</p></div> + +<div class="footnote"><p><a name="Footnote_395" id="Footnote_395"></a><a href="#FNanchor_395"><span class="label">395</span></a> ‘Transact. Linn. Soc.’ vol. xxv. p. 37.</p></div> + +<div class="footnote"><p><a name="Footnote_396" id="Footnote_396"></a><a href="#FNanchor_396"><span class="label">396</span></a> ‘Proc. Entomolog. Soc.’ Feb. 17th, 1868.</p></div> + +<div class="footnote"><p><a name="Footnote_397" id="Footnote_397"></a><a href="#FNanchor_397"><span class="label">397</span></a> Quoted by Dr. Wallace in ‘Proc. Ent. Soc.’ 3rd series, vol. v. 1867, +p. 487.</p></div> + +<div class="footnote"><p><a name="Footnote_398" id="Footnote_398"></a><a href="#FNanchor_398"><span class="label">398</span></a> Blanchard, ‘Metamorphoses, Mœurs des Insectes,’ 1868, p. 225-226.</p></div> + +<div class="footnote"><p><a name="Footnote_399" id="Footnote_399"></a><a href="#FNanchor_399"><span class="label">399</span></a> ‘Lepidopteren-Doubblettren Liste,’ Berlin, No. x. 1866.</p></div> + +<div class="footnote"><p><a name="Footnote_400" id="Footnote_400"></a><a href="#FNanchor_400"><span class="label">400</span></a> This naturalist has been so kind as to send me some results from +former years, in which the females seemed to preponderate; but so many +of the figures were estimates, that I found it impossible to tabulate them.</p></div> + +<div class="footnote"><p><a name="Footnote_401" id="Footnote_401"></a><a href="#FNanchor_401"><span class="label">401</span></a> Günther’s ‘Record of Zoological Literature,’ 1867, p. 260. On the +excess of female Lucanus, ibid. p. 250. On the males of Lucanus in England, +Westwood, ‘Modern Class. of Insects,’ vol. i. p. 187. On the Siagonium, +ibid. p. 172.</p></div> + +<div class="footnote"><p><a name="Footnote_402" id="Footnote_402"></a><a href="#FNanchor_402"><span class="label">402</span></a> Walsh, in ‘The American Entomologist,’ vol. i. 1869, p. 103. F. Smith, +'Record of Zoological Literature,’ 1867, p. 328.</p></div> + +<div class="footnote"><p><a name="Footnote_403" id="Footnote_403"></a><a href="#FNanchor_403"><span class="label">403</span></a> ‘Farm Insects,’ p. 45-46.</p></div> + +<div class="footnote"><p><a name="Footnote_404" id="Footnote_404"></a><a href="#FNanchor_404"><span class="label">404</span></a> ‘Observations on N. American Neuroptera,’ by H. Hagen and B. D. +Walsh, ‘Proc. Ent. Soc. Philadelphia,’ Oct. 1863, p. 168, 223, 239.</p></div> + +<div class="footnote"><p><a name="Footnote_405" id="Footnote_405"></a><a href="#FNanchor_405"><span class="label">405</span></a> ‘Proc. Ent. Soc. London,’ Feb. 17, 1868.</p></div> + +<div class="footnote"><p><a name="Footnote_406" id="Footnote_406"></a><a href="#FNanchor_406"><span class="label">406</span></a> Another great authority in this class, Prof. Thorell of Upsala (‘On +European Spiders,’ 1869-70, part i. p. 205) speaks as if female spiders were +generally commoner than the males.</p></div> + +<div class="footnote"><p><a name="Footnote_407" id="Footnote_407"></a><a href="#FNanchor_407"><span class="label">407</span></a> See, on this subject, Mr. Pickard-Cambridge, as quoted in ‘Quarterly +Journal of Science,’ 1868, p. 429.</p></div> + +<div class="footnote"><p><a name="Footnote_408" id="Footnote_408"></a><a href="#FNanchor_408"><span class="label">408</span></a> I have often been struck with the fact, that in several species of +Primula the seeds in the capsules which contained only a few were +very much larger than the numerous seeds in the more productive +capsules.</p></div> + +<div class="footnote"><p><a name="Footnote_409" id="Footnote_409"></a><a href="#FNanchor_409"><span class="label">409</span></a> ‘Principles of Biology,’ vol. ii. 1867, chaps. ii.-xi.</p></div> + +<div class="footnote"><p><a name="Footnote_410" id="Footnote_410"></a><a href="#FNanchor_410"><span class="label">410</span></a> De l’Espèce et de la Class.’ &c., 1869, p. 106.</p></div> + +<div class="footnote"><p><a name="Footnote_411" id="Footnote_411"></a><a href="#FNanchor_411"><span class="label">411</span></a> See, for instance, the account which I have given in my ‘Journal +of Researches,’ 1845, p. 7.</p></div> + +<div class="footnote"><p><a name="Footnote_412" id="Footnote_412"></a><a href="#FNanchor_412"><span class="label">412</span></a> I have given (‘Geolog. Observations on Volcanic Islands,’ 1844, +p. 53) a curious instance of the influence of light on the colours of +a frondescent incrustation, deposited by the surf on the coast-rocks of +Ascension, and formed by the solution of triturated sea-shells.</p></div> + +<div class="footnote"><p><a name="Footnote_413" id="Footnote_413"></a><a href="#FNanchor_413"><span class="label">413</span></a> ‘Facts and Arguments for Darwin,’ English translat. 1869, p. 20. +See the previous discussion on the olfactory threads. Sars has described +a somewhat analogous case (as quoted in ‘Nature,’ 1870, +p. 455) in a Norwegian crustacean, the <i>Pontoporeia affinis</i>.</p></div> + +<div class="footnote"><p><a name="Footnote_414" id="Footnote_414"></a><a href="#FNanchor_414"><span class="label">414</span></a> See Sir J. Lubbock in ‘Annals. and Mag. of Nat. Hist.’ vol. xi. +1853, pl. i. and x.; and vol. xii. (1853) pl. vii. See also Lubbock in +'Transact. Ent. Soc.’ vol. iv. new series, 1856-1858, p. 8. With respect +to the zigzagged antennæ mentioned below, see Fritz Müller, ‘Facts +and Arguments for Darwin’ 1869, p. 40, foot-note.</p></div> + +<div class="footnote"><p><a name="Footnote_415" id="Footnote_415"></a><a href="#FNanchor_415"><span class="label">415</span></a> See a paper by Mr. C. Spence Bate, with figures, in ‘Proc. Zoolog. +Soc.’ 1868, p. 363; and on the nomenclature of the genus, ibid. p. 585. +I am greatly indebted to Mr. Spence Bate for nearly all the above +statements with respect to the chelæ of the higher crustaceans.</p></div> + +<div class="footnote"><p><a name="Footnote_416" id="Footnote_416"></a><a href="#FNanchor_416"><span class="label">416</span></a> ‘Hist. Nat. des Crust.’ tom. ii. 1837, p. 50.</p></div> + +<div class="footnote"><p><a name="Footnote_417" id="Footnote_417"></a><a href="#FNanchor_417"><span class="label">417</span></a> Fritz Müller, ‘Facts and Arguments for Darwin,’ 1869, p. 25-28.</p></div> + +<div class="footnote"><p><a name="Footnote_418" id="Footnote_418"></a><a href="#FNanchor_418"><span class="label">418</span></a> ‘Travels in the Interior of Brazil,’ 1846, p. 111. I have given, in +my ‘Journal of Researches,’ p. 463, an account of the habits of the +Birgos.</p></div> + +<div class="footnote"><p><a name="Footnote_419" id="Footnote_419"></a><a href="#FNanchor_419"><span class="label">419</span></a> Mr. Ch. Fraser, in ‘Proc. Zoolog. Soc.’ 1869, p. 3. I am indebted +to Mr. Bate for the statement from Dr. Power.</p></div> + +<div class="footnote"><p><a name="Footnote_420" id="Footnote_420"></a><a href="#FNanchor_420"><span class="label">420</span></a> Claus, ‘Die freilebenden Copepoden,’ 1863, s. 35.</p></div> + +<div class="footnote"><p><a name="Footnote_421" id="Footnote_421"></a><a href="#FNanchor_421"><span class="label">421</span></a> ‘Facts and Arguments,’ &c., p. 79.</p></div> + +<div class="footnote"><p><a name="Footnote_422" id="Footnote_422"></a><a href="#FNanchor_422"><span class="label">422</span></a> ‘A History of the Spiders of Great Britain,’ 1861-64. For the +following facts, see p. 102, 77, 88.</p></div> + +<div class="footnote"><p><a name="Footnote_423" id="Footnote_423"></a><a href="#FNanchor_423"><span class="label">423</span></a> Aug. Vinson (‘Aranéides des Iles de la Réunion,’ pl. vi. figs. 1 +and 2) gives a good instance of the small size of the male in <i>Epeira +nigra</i>. In this species, as I may add, the male is testaceous and the +female black with legs banded with red. Other even more striking +cases of inequality in size between the sexes have been recorded +(‘Quarterly Journal of Science,’ 1868, July, p. 429); but I have not seen +the original accounts.</p></div> + +<div class="footnote"><p><a name="Footnote_424" id="Footnote_424"></a><a href="#FNanchor_424"><span class="label">424</span></a> Kirby and Spence, ‘Introduction to Entomology,’ vol. i. 1818, +p. 280.</p></div> + +<div class="footnote"><p><a name="Footnote_425" id="Footnote_425"></a><a href="#FNanchor_425"><span class="label">425</span></a> Theridion (Asagena, Sund.) serratipes, 4-punctatum et guttatum; +see Westring, in Kroyer, ‘Naturhist. Tidskrift,’ vol. iv. 1842-1843, +p. 349; and vol. ii. 1846-1849, p. 342. See, also, for other species, +'Araneæ Svecicæ,’ p. 184.</p></div> + +<div class="footnote"><p><a name="Footnote_426" id="Footnote_426"></a><a href="#FNanchor_426"><span class="label">426</span></a> Walckenaer et P. Gervais, ‘Hist. Nat. des Insectes: Aptères,’ +tom. iv. 1847, p. 17, 19, 68.</p></div> + +<div class="footnote"><p><a name="Footnote_427" id="Footnote_427"></a><a href="#FNanchor_427"><span class="label">427</span></a> Sir J. Lubbock, ‘Transact. Linnean Soc.’ vol. xxv. 1866, p. 484. +With respect to the Mutillidæ see Westwood, ‘Modern Class. of Insects,’ +vol. ii. p. 213.</p></div> + +<div class="footnote"><p><a name="Footnote_428" id="Footnote_428"></a><a href="#FNanchor_428"><span class="label">428</span></a> These organs in the male often differ in closely-allied species, and +afford excellent specific characters. But their importance, under a +functional point of view, as Mr. E. MacLachlan has remarked to me, +has probably been overrated. It has been suggested, that slight differences +in these organs would suffice to prevent the intercrossing of +well-marked varieties or incipient species, and would thus aid in their +development. That this can hardly be the case, we may infer from the +many recorded cases (see for instance, Bronn, ‘Geschichte der Natur,’ +B. ii. 1843, s. 164; and Westwood, ‘Transact. Ent. Soc.’ vol. iii. 1842, +p. 195) of distinct species having been observed in union. Mr. +MacLachlan informs me (vide ‘Stett. Ent. Zeitung,’ 1867, s. 155) that +when several species of Phryganidæ, which present strongly-pronounced +differences of this kind, were confined together by Dr. Aug. Meyer, +<i>they coupled</i>, and one pair produced fertile ova.</p></div> + +<div class="footnote"><p><a name="Footnote_429" id="Footnote_429"></a><a href="#FNanchor_429"><span class="label">429</span></a> ‘The Practical Entomologist,’ Philadelphia, vol. ii. May, 1867, p. +88.</p></div> + +<div class="footnote"><p><a name="Footnote_430" id="Footnote_430"></a><a href="#FNanchor_430"><span class="label">430</span></a> Mr. Walsh, ibid. p. 107.</p></div> + +<div class="footnote"><p><a name="Footnote_431" id="Footnote_431"></a><a href="#FNanchor_431"><span class="label">431</span></a> ‘Modern Classification of Insects,’ vol. ii. 1840, p. 206, 205. Mr. +Walsh, who called my attention to this double use of the jaws, says +that he has repeatedly observed this fact.</p></div> + +<div class="footnote"><p><a name="Footnote_432" id="Footnote_432"></a><a href="#FNanchor_432"><span class="label">432</span></a> We have here a curious and inexplicable case of dimorphism, for +some of the females of four European species of Dytiscus, and of certain +species of Hydroporus, have their elytra smooth; and no intermediate +gradations between sulcated or punctured and quite smooth elytra +have been observed. See Dr. H. Schaum, as quoted in the ‘Zoologist,’ +vol. v.-vi. 1847-48, p. 1896. Also Kirby and Spence, ‘Introduction to +Entomology,’ vol. iii. 1826, p. 305.</p></div> + +<div class="footnote"><p><a name="Footnote_433" id="Footnote_433"></a><a href="#FNanchor_433"><span class="label">433</span></a> Westwood, ‘Modern Class.’ vol. ii. p. 193. The following statement +about Penthe, and others in inverted commas, are taken from +Mr. Walsh, ‘Practical Entomologist,’ Philadelphia, vol. ii. p. 88.</p></div> + +<div class="footnote"><p><a name="Footnote_434" id="Footnote_434"></a><a href="#FNanchor_434"><span class="label">434</span></a> Kirby and Spence, ‘Introduct.’ &c., vol. iii. p. 332-336.</p></div> + +<div class="footnote"><p><a name="Footnote_435" id="Footnote_435"></a><a href="#FNanchor_435"><span class="label">435</span></a> ‘Insecta Maderensia,’ 1854, p. 20.</p></div> + +<div class="footnote"><p><a name="Footnote_436" id="Footnote_436"></a><a href="#FNanchor_436"><span class="label">436</span></a> E. Doubleday, ‘Annals and Mag. of Nat. Hist.’ vol. i. 1848, p. 379. +I may add that the wings in certain Hymenoptera (see Shuckard, +'Fossorial Hymenop.’ 1837, p. 39-43) differ in neuration according to +sex.</p></div> + +<div class="footnote"><p><a name="Footnote_437" id="Footnote_437"></a><a href="#FNanchor_437"><span class="label">437</span></a> H. W. Bates, in ‘Journal of Proc. Linn. Soc.’ vol. vi. 1862, p. 74. +Mr. Wonfor’s observations are quoted in ‘Popular Science Review,’ +1868, p. 343.</p></div> + +<div class="footnote"><p><a name="Footnote_438" id="Footnote_438"></a><a href="#FNanchor_438"><span class="label">438</span></a> Kirby and Spence, ‘Introduction to Entomology,’ vol. iii. p. 299.</p></div> + +<div class="footnote"><p><a name="Footnote_439" id="Footnote_439"></a><a href="#FNanchor_439"><span class="label">439</span></a> Robinet, ‘Vers à Soie,’ 1848, p. 207.</p></div> + +<div class="footnote"><p><a name="Footnote_440" id="Footnote_440"></a><a href="#FNanchor_440"><span class="label">440</span></a> ‘Transact. Ent. Soc.’ 3rd series, vol. v. p. 486.</p></div> + +<div class="footnote"><p><a name="Footnote_441" id="Footnote_441"></a><a href="#FNanchor_441"><span class="label">441</span></a> ‘Journal of Proc. Ent. Soc.’ Feb. 4th, 1867, p. lxxi.</p></div> + +<div class="footnote"><p><a name="Footnote_442" id="Footnote_442"></a><a href="#FNanchor_442"><span class="label">442</span></a> For this and other statements on the size of the sexes, see Kirby +and Spence, ibid. vol. iii. p. 300; on the duration of life in insects, +see p. 344.</p></div> + +<div class="footnote"><p><a name="Footnote_443" id="Footnote_443"></a><a href="#FNanchor_443"><span class="label">443</span></a> ‘Transact. Linnean Soc.’ vol. xxvi. 1868, p. 296.</p></div> + +<div class="footnote"><p><a name="Footnote_444" id="Footnote_444"></a><a href="#FNanchor_444"><span class="label">444</span></a> ‘The Malay Archipelago,’ vol. ii. 1869, p. 313.</p></div> + +<div class="footnote"><p><a name="Footnote_445" id="Footnote_445"></a><a href="#FNanchor_445"><span class="label">445</span></a> ‘Modern Classification of Insects,’ vol. ii. 1840, p. 526.</p></div> + +<div class="footnote"><p><a name="Footnote_446" id="Footnote_446"></a><a href="#FNanchor_446"><span class="label">446</span></a> See Mr. B. T. Lowne’s very interesting work, ‘On the Anatomy of +the Blow-Fly, Musca vomitoria,’ 1870, p. 14.</p></div> + +<div class="footnote"><p><a name="Footnote_447" id="Footnote_447"></a><a href="#FNanchor_447"><span class="label">447</span></a> Westwood, ‘Modern Class. of Insects,’ vol. ii. p. 473.</p></div> + +<div class="footnote"><p><a name="Footnote_448" id="Footnote_448"></a><a href="#FNanchor_448"><span class="label">448</span></a> These particulars are taken from Westwood’s ‘Modern Class. of +Insects,’ vol. ii. 1840, p. 422. See, also, on the Fulgoridæ, Kirby and +Spence, ‘Introduct.’ vol. ii. p. 401.</p></div> + +<div class="footnote"><p><a name="Footnote_449" id="Footnote_449"></a><a href="#FNanchor_449"><span class="label">449</span></a> ‘Zeitschrift für wissenschaft. Zoolog.’ B. xvii. 1867, s. 152-158.</p></div> + +<div class="footnote"><p><a name="Footnote_450" id="Footnote_450"></a><a href="#FNanchor_450"><span class="label">450</span></a> I am indebted to Mr. Walsh for having sent me this extract from +a ‘Journal of the Doings of Cicada septemdecim,’ by Dr. Hartman.</p></div> + +<div class="footnote"><p><a name="Footnote_451" id="Footnote_451"></a><a href="#FNanchor_451"><span class="label">451</span></a> L. Guilding, ‘Transact. Linn. Soc.’ vol. xv. p. 154.</p></div> + +<div class="footnote"><p><a name="Footnote_452" id="Footnote_452"></a><a href="#FNanchor_452"><span class="label">452</span></a> Köppen, as quoted in the ‘Zoological Record,’ for 1867, p. 460.</p></div> + +<div class="footnote"><p><a name="Footnote_453" id="Footnote_453"></a><a href="#FNanchor_453"><span class="label">453</span></a> Gilbert White, ‘Nat. Hist. of Selborne,’ vol. ii. 1825, p. 262.</p></div> + +<div class="footnote"><p><a name="Footnote_454" id="Footnote_454"></a><a href="#FNanchor_454"><span class="label">454</span></a> Harris, ‘Insects of New England,’ 1842, p. 128.</p></div> + +<div class="footnote"><p><a name="Footnote_455" id="Footnote_455"></a><a href="#FNanchor_455"><span class="label">455</span></a> ‘The Naturalist on the Amazons,’ vol. i. 1863, p. 252. Mr. Bates +gives a very interesting discussion on the gradations in the musical +apparatus of the three families. See also Westwood, ‘Modern Class.’ +vol. ii. p. 445 and 453.</p></div> + +<div class="footnote"><p><a name="Footnote_456" id="Footnote_456"></a><a href="#FNanchor_456"><span class="label">456</span></a> ‘Proc. Boston Soc. of Nat. Hist.’ vol. xi. April, 1868.</p></div> + +<div class="footnote"><p><a name="Footnote_457" id="Footnote_457"></a><a href="#FNanchor_457"><span class="label">457</span></a> ‘Nouveau Manuel d’Anat. Comp.’ (French translat.), tom. i. 1850 +p. 567.</p></div> + +<div class="footnote"><p><a name="Footnote_458" id="Footnote_458"></a><a href="#FNanchor_458"><span class="label">458</span></a> ‘Zeitschrift für wissenschaft. Zoolog.’ B. xvii. 1867, s. 117.</p></div> + +<div class="footnote"><p><a name="Footnote_459" id="Footnote_459"></a><a href="#FNanchor_459"><span class="label">459</span></a> Westwood, ‘Modern Class. of Insects,’ vol. i. p. 440.</p></div> + +<div class="footnote"><p><a name="Footnote_460" id="Footnote_460"></a><a href="#FNanchor_460"><span class="label">460</span></a> Westwood, ‘Modern Class. of Insects,’ vol. i. p. 453.</p></div> + +<div class="footnote"><p><a name="Footnote_461" id="Footnote_461"></a><a href="#FNanchor_461"><span class="label">461</span></a> Landois, ibid. s. 121, 122.</p></div> + +<div class="footnote"><p><a name="Footnote_462" id="Footnote_462"></a><a href="#FNanchor_462"><span class="label">462</span></a> Mr. Walsh also informs me that he has noticed that the female of +the <i>Platyphyllum concavum</i>, “when captured makes a feeble grating +noise by shuffling her wing-covers together.”</p></div> + +<div class="footnote"><p><a name="Footnote_463" id="Footnote_463"></a><a href="#FNanchor_463"><span class="label">463</span></a> Landois, ibid. s. 113.</p></div> + +<div class="footnote"><p><a name="Footnote_464" id="Footnote_464"></a><a href="#FNanchor_464"><span class="label">464</span></a> ‘Insects of New England,’ 1842, p. 133.</p></div> + +<div class="footnote"><p><a name="Footnote_465" id="Footnote_465"></a><a href="#FNanchor_465"><span class="label">465</span></a> Westwood, ‘Modern Classification,’ vol. i. p. 462.</p></div> + +<div class="footnote"><p><a name="Footnote_466" id="Footnote_466"></a><a href="#FNanchor_466"><span class="label">466</span></a> Westwood, ibid. vol. i. p. 453.</p></div> + +<div class="footnote"><p><a name="Footnote_467" id="Footnote_467"></a><a href="#FNanchor_467"><span class="label">467</span></a> Landois, ibid. s. 115, 116, 120, 122.</p></div> + +<div class="footnote"><p><a name="Footnote_468" id="Footnote_468"></a><a href="#FNanchor_468"><span class="label">468</span></a> ‘Transact. Ent. Soc.’ 3rd series, vol. ii. (‘Journal of Proceedings, +p. 117.)</p></div> + +<div class="footnote"><p><a name="Footnote_469" id="Footnote_469"></a><a href="#FNanchor_469"><span class="label">469</span></a> Westwood, ‘Modern Class. of Insects,’ vol. i. p. 427; for crickets, +p. 445.</p></div> + +<div class="footnote"><p><a name="Footnote_470" id="Footnote_470"></a><a href="#FNanchor_470"><span class="label">470</span></a> Mr. Ch. Horne, in ‘Proc. Ent. Soc.’ May 3, 1869, p. xii.</p></div> + +<div class="footnote"><p><a name="Footnote_471" id="Footnote_471"></a><a href="#FNanchor_471"><span class="label">471</span></a> The <i>Oecanthus nivalis</i>, Harris, ‘Insects of New England,’ 1842, +p. 124.</p></div> + +<div class="footnote"><p><a name="Footnote_472" id="Footnote_472"></a><a href="#FNanchor_472"><span class="label">472</span></a> Platyblemnus: Westwood, ‘Modern. Class.’ vol. i. p. 447.</p></div> + +<div class="footnote"><p><a name="Footnote_473" id="Footnote_473"></a><a href="#FNanchor_473"><span class="label">473</span></a> B. D. Walsh, the Pseudo-neuroptera of Illinois, +in ‘Proc. Ent. Soc. +of Philadelphia,’ 1862, p. 361.</p></div> + +<div class="footnote"><p><a name="Footnote_474" id="Footnote_474"></a><a href="#FNanchor_474"><span class="label">474</span></a> ‘Modern Class.’ vol. ii. p. 37.</p></div> + +<div class="footnote"><p><a name="Footnote_475" id="Footnote_475"></a><a href="#FNanchor_475"><span class="label">475</span></a> Walsh, ibid. p. 381. I am indebted to this naturalist for the +following facts on Hetærina, Anax, and Gomphus.</p></div> + +<div class="footnote"><p><a name="Footnote_476" id="Footnote_476"></a><a href="#FNanchor_476"><span class="label">476</span></a> ‘Transact. Ent. Soc’ vol. i. 1836, p. lxxxi.</p></div> + +<div class="footnote"><p><a name="Footnote_477" id="Footnote_477"></a><a href="#FNanchor_477"><span class="label">477</span></a> See abstract in the ‘Zoological Record’ for 1867, p. 450.</p></div> + +<div class="footnote"><p><a name="Footnote_478" id="Footnote_478"></a><a href="#FNanchor_478"><span class="label">478</span></a> Kirby and Spence, ‘Introduct. to Entomology,’ vol. ii. 1818, p. 35.</p></div> + +<div class="footnote"><p><a name="Footnote_479" id="Footnote_479"></a><a href="#FNanchor_479"><span class="label">479</span></a> See an interesting article, “The Writings of Fabre,” in ‘Nat. Hist. +Review,’ April, 1862, p. 122.</p></div> + +<div class="footnote"><p><a name="Footnote_480" id="Footnote_480"></a><a href="#FNanchor_480"><span class="label">480</span></a> ‘Journal of Proc. of Entomolog. Soc.’ Sept. 7th, 1863, p. 169.</p></div> + +<div class="footnote"><p><a name="Footnote_481" id="Footnote_481"></a><a href="#FNanchor_481"><span class="label">481</span></a> P. Huber, ‘Recherches sur les Mœurs des Fourmis,’ 1810, p. 150, +165.</p></div> + +<div class="footnote"><p><a name="Footnote_482" id="Footnote_482"></a><a href="#FNanchor_482"><span class="label">482</span></a> ‘Proc. Entomolog. Soc. of Philadelphia,’ 1866, p. 238-239.</p></div> + +<div class="footnote"><p><a name="Footnote_483" id="Footnote_483"></a><a href="#FNanchor_483"><span class="label">483</span></a> Quoted by Westwood, ‘Modern Class. of Insects,’ vol. ii. p. 214.</p></div> + +<div class="footnote"><p><a name="Footnote_484" id="Footnote_484"></a><a href="#FNanchor_484"><span class="label">484</span></a> Pyrodes pulcherrimus, in which the sexes differ conspicuously, has +been described by Mr. Bates in ‘Transact. Ent. Soc.’ 1869, p. 50. I +will specify the few other cases in which I have heard of a difference +in colour between the sexes of beetles. Kirby and Spence (‘Introduct. +to Entomology,’ vol. iii. p. 301) mention a Cantharis, Meloe, Rhagium, +and the <i>Leptura testacea</i>; the male of the latter being testaceous, with +a black thorax, and the female of a dull red all over. These two +latter beetles belong to the Order of Longicorns. Messrs. R. Trimen +and Waterhouse, junr., inform me of two Lamellicorns, viz., a Peritrichia +and Trichius, the male of the latter being more obscurely +coloured than the female. In <i>Tillus elongatus</i> the male is black, and +the female always, as it is believed, of a dark blue colour with a red +thorax. The male, also, of <i>Orsodacna atra</i>, as I hear from Mr. Walsh, +is black, the female (the so-called <i>O. ruficollis</i>) having a rufous thorax.</p></div> + +<div class="footnote"><p><a name="Footnote_485" id="Footnote_485"></a><a href="#FNanchor_485"><span class="label">485</span></a> ‘Proc. Entomolog. Soc. of Philadelphia,’ 1864, p. 228.</p></div> + +<div class="footnote"><p><a name="Footnote_486" id="Footnote_486"></a><a href="#FNanchor_486"><span class="label">486</span></a> Kirby and Spence, ‘Introduct. Entomolog.’ vol. iii. p. 300.</p></div> + +<div class="footnote"><p><a name="Footnote_487" id="Footnote_487"></a><a href="#FNanchor_487"><span class="label">487</span></a> Kirby and Spence, ibid. vol. iii. p. 329.</p></div> + +<div class="footnote"><p><a name="Footnote_488" id="Footnote_488"></a><a href="#FNanchor_488"><span class="label">488</span></a> ‘Modern Classification of Insects,’ vol. i. p. 172. On the same +page there is an account of Siagonium. In the British Museum I +noticed one male specimen of Siagonium in an intermediate condition, +so that the dimorphism is not strict.</p></div> + +<div class="footnote"><p><a name="Footnote_489" id="Footnote_489"></a><a href="#FNanchor_489"><span class="label">489</span></a> ‘The Malay Archipelago,’ vol. ii. 1869, p. 276.</p></div> + +<div class="footnote"><p><a name="Footnote_490" id="Footnote_490"></a><a href="#FNanchor_490"><span class="label">490</span></a> ‘Entomological Magazine,’ vol. i. 1833, p. 82. See also on the +conflicts of this species, Kirby and Spence, ibid. vol. iii. p. 314; and +Westwood, ibid. vol. i. p. 187.</p></div> + +<div class="footnote"><p><a name="Footnote_491" id="Footnote_491"></a><a href="#FNanchor_491"><span class="label">491</span></a> Quoted from Fischer, in ‘Dict. Class. d’Hist. Nat.’ tom. x. p. 324.</p></div> + +<div class="footnote"><p><a name="Footnote_492" id="Footnote_492"></a><a href="#FNanchor_492"><span class="label">492</span></a> ‘Ann. Soc. Entomolog. France,’ 1866, as quoted in ‘Journal of +Travel,’ by A. Murray, 1868, p. 135.</p></div> + +<div class="footnote"><p><a name="Footnote_493" id="Footnote_493"></a><a href="#FNanchor_493"><span class="label">493</span></a> Westwood, ‘Modern Class.’ vol. i. p. 184.</p></div> + +<div class="footnote"><p><a name="Footnote_494" id="Footnote_494"></a><a href="#FNanchor_494"><span class="label">494</span></a> Wollaston, On certain musical Curculionidæ, ‘Annals and Mag. of +Nat. Hist.’ vol. vi. 1860, p. 14.</p></div> + +<div class="footnote"><p><a name="Footnote_495" id="Footnote_495"></a><a href="#FNanchor_495"><span class="label">495</span></a> ‘Zeitschrift für wiss. Zoolog.’ B. xvii. 1867, s. 127.</p></div> + +<div class="footnote"><p><a name="Footnote_496" id="Footnote_496"></a><a href="#FNanchor_496"><span class="label">496</span></a> I am greatly indebted to Mr. G. R. Crotch for having sent me +numerous prepared specimens of various beetles belonging to these three +families and others, as well as for valuable information of all kinds. He +believes that the power of stridulation in the Clythra has not been previously +observed. I am also much indebted to Mr. E. W. Janson, for +information and specimens. I may add that my son, Mr. F. Darwin, +finds that <i>Dermestes murinus</i> stridulates, but he searched in vain for the +apparatus. Scolytus has lately been described by Mr. Algen as a stridulator, +in the ‘Edinburgh Monthly Magazine,’ 1869, Nov., p. 130.</p></div> + +<div class="footnote"><p><a name="Footnote_497" id="Footnote_497"></a><a href="#FNanchor_497"><span class="label">497</span></a> Schiödte, translated in ‘Annals and Mag. of Nat. Hist.’ vol. xx. +1867, p. 37.</p></div> + +<div class="footnote"><p><a name="Footnote_498" id="Footnote_498"></a><a href="#FNanchor_498"><span class="label">498</span></a> Westring has described (Kroyer, ‘Naturhist. Tidskrift,’ B. ii. 1848-49, +p. 334) the stridulating organs in these two, as well as in other +families. In the Carabidæ I have examined <i>Elaphrus uliginosus</i> and +<i>Blethisa multipunctata</i>, sent to me by Mr. Crotch. In Blethisa the +transverse ridges on the furrowed border of the abdominal segment do +not come into play, as far as I could judge, in scraping the rasps on the +elytra.</p></div> + +<div class="footnote"><p><a name="Footnote_499" id="Footnote_499"></a><a href="#FNanchor_499"><span class="label">499</span></a> I am indebted to Mr. Walsh, of Illinois, for having sent me +extracts from Leconte’s ‘Introduction to Entomology,’ p. 101, 143.</p></div> + +<div class="footnote"><p><a name="Footnote_500" id="Footnote_500"></a><a href="#FNanchor_500"><span class="label">500</span></a> M. P. de la Brulerie, as quoted in ‘Journal of Travel,’ A. Murray, +vol. i. 1868, p. 135.</p></div> + +<div class="footnote"><p><a name="Footnote_501" id="Footnote_501"></a><a href="#FNanchor_501"><span class="label">501</span></a> Mr. Doubleday informs me that “the noise is produced by the +insect raising itself on its legs as high as it can, and then sinking its +thorax five or six times, in rapid succession, against the substance +upon which it is sitting.” For references on this subject see Landois, +'Zeitschrift für wissen. Zoolog.’ B. xvii. s. 131. Olivier says (as quoted +by Kirby and Spence, ‘Introduct.’ vol. ii. p. 395) that the female of +<i>Pimelia striata</i> produces a rather loud sound by striking her abdomen +against any hard substance, “and that the male, obedient to this call, +soon attends her and they pair.”</p></div> + +<div class="footnote"><p><a name="Footnote_502" id="Footnote_502"></a><a href="#FNanchor_502"><span class="label">502</span></a> Apatura Iris: ‘The Entomologist’s Weekly Intelligencer,’ 1859, +p. 139. For the Bornean Butterflies see C. Collingwood, ‘Rambles of +a Naturalist,’ 1868, p. 183.</p></div> + +<div class="footnote"><p><a name="Footnote_503" id="Footnote_503"></a><a href="#FNanchor_503"><span class="label">503</span></a> See my ‘Journal of Researches,’ 1845, p. 33. Mr. Doubleday has +detected (‘Proc. Ent. Soc.’ March 3rd, 1845, p. 123) a peculiar membranous +sac at the base of the front wings, which is probably connected +with the production of the sound.</p></div> + +<div class="footnote"><p><a name="Footnote_504" id="Footnote_504"></a><a href="#FNanchor_504"><span class="label">504</span></a> See also Mr. Bates’ paper in ‘Proc. Ent. Soc. of Philadelphia,’ +1865, p. 206. Also Mr. Wallace on the same subject, in regard to +Diadema, in ‘Transact. Entomolog. Soc. of London,’ 1869, p. 278.</p></div> + +<div class="footnote"><p><a name="Footnote_505" id="Footnote_505"></a><a href="#FNanchor_505"><span class="label">505</span></a> ‘The Naturalist on the Amazons,’ vol. i. 1863, p. 19.</p></div> + +<div class="footnote"><p><a name="Footnote_506" id="Footnote_506"></a><a href="#FNanchor_506"><span class="label">506</span></a> See the interesting article in the ‘Westminster Review,’ July, 1867, +p. 10. A woodcut of the Kallima is given by Mr. Wallace in Hardwicke’s +'Science Gossip,’ Sept. 1867, p. 196.</p></div> + +<div class="footnote"><p><a name="Footnote_507" id="Footnote_507"></a><a href="#FNanchor_507"><span class="label">507</span></a> See the interesting observations by Mr. T. W. Wood, ‘The Student,’ +Sept. 1868, p. 81.</p></div> + +<div class="footnote"><p><a name="Footnote_508" id="Footnote_508"></a><a href="#FNanchor_508"><span class="label">508</span></a> Mr. Wallace in ‘Hardwicke’s Science Gossip,’ Sept. 1867, p. 193.</p></div> + +<div class="footnote"><p><a name="Footnote_509" id="Footnote_509"></a><a href="#FNanchor_509"><span class="label">509</span></a> See also, on this subject, Mr. Weir’s paper in ‘Transact. Ent. Soc.’ +1869, p. 23.</p></div> + +<div class="footnote"><p><a name="Footnote_510" id="Footnote_510"></a><a href="#FNanchor_510"><span class="label">510</span></a> ‘Westminster Review,’ July, 1867, p. 16.</p></div> + +<div class="footnote"><p><a name="Footnote_511" id="Footnote_511"></a><a href="#FNanchor_511"><span class="label">511</span></a> For instance, Lithosia; but Prof. Westwood (‘Modern Class. of +Insects,’ vol. ii. p. 390) seems surprised at this case. On the relative +colours of diurnal and nocturnal Lepidoptera, see ibid. p. 333 and 392; +also Harris, ‘Treatise on the Insects of New England,’ 1842, p. 315.</p></div> + +<div class="footnote"><p><a name="Footnote_512" id="Footnote_512"></a><a href="#FNanchor_512"><span class="label">512</span></a> Such differences between the upper and lower surfaces of the +wings of several species of Papilio, may be seen in the beautiful plates +to Mr. Wallace’s Memoir on the Papilionidæ of the Malayan Region, +in ‘Transact. Linn. Soc.’ vol. xxv. part i. 1865.</p></div> + +<div class="footnote"><p><a name="Footnote_513" id="Footnote_513"></a><a href="#FNanchor_513"><span class="label">513</span></a> ‘Proc. Ent. Soc.’ March 2nd, 1868.</p></div> + +<div class="footnote"><p><a name="Footnote_514" id="Footnote_514"></a><a href="#FNanchor_514"><span class="label">514</span></a> See also an account of the S. American genus Erateina (one of +the Geometræ) in ‘Transact. Ent. Soc.’ new series, vol. v. pl. xv. and +xvi.</p></div> + +<div class="footnote"><p><a name="Footnote_515" id="Footnote_515"></a><a href="#FNanchor_515"><span class="label">515</span></a> ‘Proc. Ent. Soc. of London,’ July 6, 1868, p. xxvii.</p></div> + +<div class="footnote"><p><a name="Footnote_516" id="Footnote_516"></a><a href="#FNanchor_516"><span class="label">516</span></a> Harris, ‘Treatise,’ &c., edited by Flint, 1862, p. 395.</p></div> + +<div class="footnote"><p><a name="Footnote_517" id="Footnote_517"></a><a href="#FNanchor_517"><span class="label">517</span></a> For instance, I observe in my son’s cabinet that the males are +darker than the females in the <i>Lasiocampa quercus</i>, <i>Odonestis potatoria</i>, +<i>Hypogymna dispar</i>, <i>Dasychira pudibunda</i>, and <i>Cycnia mendica</i>. In this +latter species the difference in colour between the two sexes is strongly +marked; and Mr. Wallace informs me that we here have, as he believes, +an instance of protective mimickry confined to one sex, as will hereafter +be more fully explained. The white female of the Cycnia resembles +the very common <i>Spilosoma menthrasti</i>, both sexes of which are white; +and Mr. Stainton observed that this latter moth was rejected with utter +disgust by a whole brood of young turkeys, which were fond of eating +other moths; so that if the Cycnia was commonly mistaken by British +birds for the Spilosoma, it would escape being devoured, and its white +deceptive colour would thus be highly beneficial.</p></div> + +<div class="footnote"><p><a name="Footnote_518" id="Footnote_518"></a><a href="#FNanchor_518"><span class="label">518</span></a> ‘Rambles of a Naturalist in the Chinese Seas,’ 1868, p. 182.</p></div> + +<div class="footnote"><p><a name="Footnote_519" id="Footnote_519"></a><a href="#FNanchor_519"><span class="label">519</span></a> Wallace on the Papilionidæ of the Malayan Region, in ‘Transact. +Linn. Soc. vol. xxv. 1865, p. 8, 36. A striking case of a rare variety, +strictly intermediate between two other well-marked female varieties, +is given by Mr. Wallace. See also Mr. Bates, in ‘Proc. Entomolog. +Soc.’ Nov. 19th, 1866, p. xl.</p></div> + +<div class="footnote"><p><a name="Footnote_520" id="Footnote_520"></a><a href="#FNanchor_520"><span class="label">520</span></a> Mr. R. MacLachlan, ‘Transact. Ent. Soc.’ vol. ii. part 6th, 3rd +series, 1866, p. 459.</p></div> + +<div class="footnote"><p><a name="Footnote_521" id="Footnote_521"></a><a href="#FNanchor_521"><span class="label">521</span></a> H. W. Bates, ‘The Naturalist on the Amazons,’ vol. ii. 1863, +p. 228. A. R. Wallace, in ‘Transact. Linn. Soc.’ vol. xxv. 1865, p. 10.</p></div> + +<div class="footnote"><p><a name="Footnote_522" id="Footnote_522"></a><a href="#FNanchor_522"><span class="label">522</span></a> On this whole subject see ‘The Variation of Animals and Plants +under Domestication,’ vol. ii. 1868, chap. xxiii.</p></div> + +<div class="footnote"><p><a name="Footnote_523" id="Footnote_523"></a><a href="#FNanchor_523"><span class="label">523</span></a> A. R. Wallace, in ‘The Journal of Travel,’ vol. i. 1868, p. 88. +'Westminster Review,’ July, 1857, p. 37. See also Messrs. Wallace +and Bates in ‘Proc. Ent. Soc.’ Nov. 19th, 1866, p. xxxix.</p></div> + +<div class="footnote"><p><a name="Footnote_524" id="Footnote_524"></a><a href="#FNanchor_524"><span class="label">524</span></a> ‘The Variation of Animals and Plants under Domestication,’ +vol. ii. chap. xii. p. 17.</p></div> + +<div class="footnote"><p><a name="Footnote_525" id="Footnote_525"></a><a href="#FNanchor_525"><span class="label">525</span></a> ‘Transact. Linn. Soc.’ vol. xxiii. 1862, p. 495.</p></div> + +<div class="footnote"><p><a name="Footnote_526" id="Footnote_526"></a><a href="#FNanchor_526"><span class="label">526</span></a> ‘Proc. Ent. Soc.’ Dec. 3rd, 1866, p. xlv.</p></div> + +<div class="footnote"><p><a name="Footnote_527" id="Footnote_527"></a><a href="#FNanchor_527"><span class="label">527</span></a> ‘Transact. Linn. Soc.’ vol. xxv. 1865, p. 1; also ‘Transact. Ent. +Soc.’ vol. iv. (3rd series), 1867, p. 301.</p></div> + +<div class="footnote"><p><a name="Footnote_528" id="Footnote_528"></a><a href="#FNanchor_528"><span class="label">528</span></a> See an ingenious article entitled, “Difficulties of the Theory of +Natural Selection,” in the ‘Month,’ 1869. The writer strangely supposes +that I attribute the variations in colour of the Lepidoptera, by +which certain species belonging to distinct families have come to +resemble others, to reversion to a common progenitor; but there is no +more reason to attribute these variations to reversion than in the case +of any ordinary variation.</p></div> + +<div class="footnote"><p><a name="Footnote_529" id="Footnote_529"></a><a href="#FNanchor_529"><span class="label">529</span></a> Wallace, “Notes on Eastern Butterflies,” ‘Transact. Ent. Soc.’ +1869, p. 287.</p></div> + +<div class="footnote"><p><a name="Footnote_530" id="Footnote_530"></a><a href="#FNanchor_530"><span class="label">530</span></a> Wallace, in ‘Westminster Review,’ July, 1867, p. 37; and in +'Journal of Travel and Nat. Hist.’ vol. i. 1868, p. 88.</p></div> + +<div class="footnote"><p><a name="Footnote_531" id="Footnote_531"></a><a href="#FNanchor_531"><span class="label">531</span></a> See remarks by Messrs. Bates and Wallace, in ‘Proc. Ent. Soc.’ +Nov. 19, 1866, p. xxxix.</p></div> + +<div class="footnote"><p><a name="Footnote_532" id="Footnote_532"></a><a href="#FNanchor_532"><span class="label">532</span></a> See Mr. Wallace in ‘Westminster Review,’ July, 1867, p. 11 and +37. The male of no butterfly, as Mr. Wallace informs me, is known to +differ in colour, as a protection, from the female; and he asks me how +I can explain this fact on the principle that one sex alone has varied +and has transmitted its variations exclusively to the same sex, without +the aid of selection to check the variations being inherited by the +other sex. No doubt if it could be shewn that the females of very +many species had been rendered beautiful through protective mimickry, +but that this has never occurred with the males, it would be a serious +difficulty. But the number of cases as yet known hardly suffices for a +fair judgment. We can see that the males, from having the power of +flying more swiftly, and thus escaping danger, would not be so likely +as the females to have had their colours modified for the sake of protection; +but this would not in the least have interfered with their receiving +protective colours through inheritance from the females. In the second +place, it is probable that sexual selection would actually tend to prevent +a beautiful male from becoming obscure, for the less brilliant individuals +would be less attractive to the females. Supposing that the beauty of +the male of any species had been mainly acquired through sexual +selection, yet if this beauty likewise served as a protection, the acquisition +would have been aided by natural selection. But it would be +quite beyond our power to distinguish between the two processes of +sexual and ordinary selection. Hence it is not likely that we should +be able to adduce cases of the males having been rendered brilliant +exclusively through protective mimickry, though this is comparatively +easy with the females, which have rarely or never been rendered beautiful, +as far as we can judge, for the sake of sexual attraction, although +they have often received beauty through inheritance from their male +parents.</p></div> + +<div class="footnote"><p><a name="Footnote_533" id="Footnote_533"></a><a href="#FNanchor_533"><span class="label">533</span></a> ‘Proc. Entomolog. Soc.’ Dec. 3rd, 1866, p. xlv., and March 4th, +1867, p. lxxx.</p></div> + +<div class="footnote"><p><a name="Footnote_534" id="Footnote_534"></a><a href="#FNanchor_534"><span class="label">534</span></a> See Mr. J. Jenner Weir’s paper on insects and insectivorous birds, +in ‘Transact. Ent. Soc.’ 1869, p. 21; also Mr. Butler’s paper, ibid. p. 27.</p></div> + +<h4>END OF VOL. I.</h4> + +<h5>LONDON:</h5> + +<h6>PRINTED BY WILLIAM CLOWES AND SONS, DUKE STREET, STAMFORD STREET, +AND CHARING CROSS.</h6> + + +<p class="quotsig"> +<span class="smcap">50a, Albemarle Street, London</span><br /> + <i>January, 1871.</i></p> + +<hr /> + +<p><span class='pagenum'><a name="Page_1a" id="Page_1a">1</a></span></p> + +<h3>MR. MURRAY’S</h3> + +<h2>LIST OF POPULAR WORKS.</h2> + +<h4><i>One Shilling.</i></h4> + +<p class="indent">THE PRINCIPAL SPEECHES AND ADDRESSES OF +H.R.H. THE PRINCE CONSORT, with an Introduction giving some Outlines +of his Character. With Portrait.</p> + +<p class="indent">MUSIC AND THE ART OF DRESS. Two Essays. By A +LADY.</p> + +<p class="indent">THE FALL OF JERUSALEM. A Dramatic Poem. By DEAN +MILMAN.</p> + +<p class="indent">THEODORE HOOK. A Sketch. By J. G. LOCKHART.</p> + +<p class="indent">HISTORY OF THE GUILLOTINE. By <span class="smcap">Rt. Hon.</span> J. W. +CROKER. Woodcuts.</p> + +<p class="indent">THE CHACE. A Descriptive Essay. By C. J. 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From the Autobiography of HENRY +STEFFENS.</p> + +<p class="indent">STOKERS AND POKERS; or, The North-Western Railway.—The +Electric Telegraph—and Railway Clearing House. By SIR FRANCIS +HEAD.</p> + +<p class="indent">TRAVELS IN EGYPT, NUBIA, SYRIA, AND THE HOLY +LAND, with a Journey round the Dead Sea, and through the Country East of +the Jordan. By IRBY and MANGLES.</p> + +<p class="indent">WESTERN BARBARY; An Account of the Wild Tribes and +Savage Animals. By JOHN H. DRUMMOND HAY.</p> + +<p class="indent">LETTERS FROM THE SHORES OF THE BALTIC. By A +LADY.</p> + +<p class="indent">NOTES AND SKETCHES OF NEW SOUTH WALES +DURING A RESIDENCE OF MANY YEARS IN THAT COLONY. +By MRS. CHARLES MEREDITH.</p> + +<p class="indent">RECOLLECTIONS OF BUSH LIFE IN AUSTRALIA, during +a Residence of Eight Years in the Interior. By REV. H. W. HAYGARTH.</p> + +<p class="indent">JOURNAL OF A RESIDENCE AMONG THE NEGROES +IN THE WEST INDIES. By M. G. LEWIS.</p> + +<p class="indent">MEMOIRS OF FATHER RIPA DURING THIRTEEN YEARS’ +RESIDENCE AT THE COURT OF PEKIN, in the Service of the Emperor +of China, with an Account of the Foundation of the College for the Education +of Young Chinese in Naples. From the Italian. By FORTUNATO PRANDI.</p> + +<p class="indent">PHILIP MUSGRAVE; or, Memoirs of a Church of England +Missionary in the North American Colonies. By REV. J. ABBOTT.</p> + +<p class="indent">A MONTH IN NORWAY. By JOHN G. HOLLWAY.</p> + +<p class="indent">LETTERS FROM MADRAS; or, First Impressions of Life and +Manners in India. By A LADY.</p> + +<p class="indent">ROUGH NOTES TAKEN DURING SOME RAPID RIDES +ACROSS THE PAMPAS AND AMONG THE ANDES. By SIR FRANCIS +HEAD.</p> + +<p class="indent">A VOYAGE UP THE RIVER AMAZON, including a Visit to +PARA. By WILLIAM H. EDWARDS.<span class='pagenum'><a name="Page_4a" id="Page_4a">4</a></span></p> + +<p class="indent">A POPULAR ACCOUNT <span class="smcap">of the MANNERS and CUSTOMS +OF INDIA</span>. By REV. CHARLES ACLAND.</p> + +<p class="indent">ADVENTURES IN THE LIBYAN DESERT AND THE +OASIS OF JUPITER AMMON. By BAYLE ST. JOHN.</p> + +<p class="indent">LIFE OF SIR FOWELL BUXTON. By his Son, CHARLES +BUXTON. With an Inquiry into the Results of Emancipation.</p> + +<h4> +<i>Two Shillings and Sixpence.</i> +</h4> + +<p class="indent">THE CONTINUITY OF SCRIPTURE, as declared by the +Testimony of our Lord and of the Evangelists and Apostles. By LORD +HATHERLEY.</p> + +<p class="indent">LIFE OF LORD BACON. By LORD CAMPBELL.</p> + +<p class="indent">CHILDE HAROLD’S PILGRIMAGE. A Romaunt. By LORD +BYRON.</p> + +<p class="indent">TALES AND POEMS. By LORD BYRON.</p> + +<p class="indent">LITTLE ARTHUR’S HISTORY OF ENGLAND. By LADY +CALLCOTT. With 28 Woodcuts.</p> + +<p class="indent">THE EMIGRANT. By SIR FRANCIS HEAD.</p> + +<p class="indent">THE FABLES OF ÆSOP; <span class="smcap">A New Version</span>, chiefly from +<span class="smcap">Original Sources</span>. By REV. THOMAS JAMES. With 100 Woodcuts.</p> + +<p class="indent">STORIES FOR CHILDREN SELECTED FROM THE HISTORY +OF ENGLAND. By <span class="smcap">Rt. Hon.</span> J. W. CROKER. With 24 Woodcuts.</p> + +<p class="indent">THE COMPLETE POETICAL WORKS OF LORD BYRON.</p> + +<p class="indent">THE PHILOSOPHY OF THE MORAL FEELINGS. By +JOHN ABERCROMBIE.</p> + +<p class="indent">POEMS AND SONGS. By ALLAN CUNNINGHAM. With +Biographical Notice.</p> + +<p class="indent">THE FRANCHISE; a Privilege and not a Right. Proved by the +Political Experience of the Ancients. By H. S. TREMENHEERE.</p> + +<h4> +<i>Three Shillings.</i> +</h4> + +<p class="indent">“THE FORTY-FIVE;” <span class="smcap">A Narrative of the Insurrection of +1745 in Scotland</span>. To which are added Letters of <span class="smcap">Prince Charles Stuart</span>. +By LORD MAHON.<span class='pagenum'><a name="Page_5a" id="Page_5a">5</a></span></p> + +<h4><i>Three Shillings and Sixpence.</i></h4> + +<p class="indent">THE BIBLE IN SPAIN; or, The Adventures and Imprisonment +of an Englishman in attempting to Circulate the Scriptures in the Peninsula. +By GEORGE BORROW.</p> + +<p class="indent">THE GYPSIES OF SPAIN; their Manners, Customs, Religion, +and Language. By GEORGE BORROW.</p> + +<p class="indent">THE BEAUTIES OF LORD BYRON’S WRITINGS; <span class="smcap">Poetry +And Prose.</span> With Portrait.</p> + +<p class="indent">LIFE OF LOUIS PRINCE OF CONDÉ, SURNAMED THE +GREAT. By LORD MAHON.</p> + +<p class="indent">SKETCHES OF GERMAN LIFE AND SCENES FROM THE +WAR OF LIBERATION IN GERMANY. By VARNHAGEN VON +ERESE. Translated by SIR ALEXANDER DUFF GORDON.</p> + +<p class="indent">THE TRUE STORY OF THE BATTLE OF WATERLOO. By +REV. G. R. GLEIG.</p> + +<p class="indent">DEEDS OF NAVAL DARING; or, <span class="smcap">Anecdotes of The +British Navy.</span> By EDWARD GIFFARD.</p> + +<p class="indent">TYPEE; or, The Marquesas Islanders. By HERMAN MELVILLE.</p> + +<p class="indent">OMOO: A Narrative of Adventures in the South Seas; a Sequel +to ‘Typee.’ By HERMAN MELVILLE.</p> + +<p class="indent">THE BIBLE IN THE HOLY LAND; being Extracts from +Dean Stanley’s ‘Sinai and Palestine.’ With Woodcuts.</p> + +<p class="indent">AN ESSAY ON ENGLISH POETRY. With Short Notices of the +British Poets. By THOMAS CAMPBELL.</p> + +<p class="indent">HISTORICAL AND CRITICAL ESSAYS. By LORD MAHON.</p> + +<p class="indent">LIFE OF LORD CLIVE. By REV. G. R. GLEIG.</p> + +<p class="indent">LIFE OF SIR THOMAS MUNRO. With Selections from his +Correspondence. By REV. G. R. GLEIG.</p> + +<p class="indent">THE RISE OF OUR INDIAN EMPIRE: being a History of +British India from its Origin till the Peace of 1783. By LORD MAHON.<span class='pagenum'><a name="Page_6a" id="Page_6a">6</a></span></p> + +<p class="indent">A NARRATIVE OF THE SIEGE OF KARS, and of the Six +Months’ Resistance by the Turkish Garrison under General Williams. With +Travels and Adventures in Armenia, &c. By HUMPHREY SANDWITH, +M.D.</p> + +<p class="indent">SKETCHES OF PERSIA. By SIR JOHN MALCOLM.</p> + +<p class="indent">THE WILD SPORTS AND NATURAL HISTORY OF THE +HIGHLANDS. By CHARLES ST. JOHN.</p> + +<p class="indent">GATHERINGS FROM SPAIN. By RICHARD FORD.</p> + +<p class="indent">TRAVELS IN MEXICO AND THE ROCKY MOUNTAINS. +By G. F. RUXTON.</p> + +<p class="indent">PORTUGAL AND GALLICIA; <span class="smcap">With an Account of the +Social and Political State of the Basque Provinces.</span> By LORD +CARNARVON.</p> + +<p class="indent">A RESIDENCE AT SIERRA LEONE. Described from a +Journal kept on the Spot, and Letters written to Friends at Home. By +A LADY.</p> + +<p class="indent">THE REMAINS IN VERSE AND PROSE OF ARTHUR +HENRY HALLAM. With Preface, Memoir, and Portrait.</p> + +<p class="indent">THE POETICAL WORKS OF BISHOP HEBER; containing +Palestine, Europe, the Red Sea, Hymns, &c. With Portrait.</p> + +<p class="indent">GLEANINGS IN NATURAL HISTORY. By EDWARD +JESSE. With Woodcuts.</p> + +<p class="indent">THE REJECTED ADDRESSES: or, The New Theatrum +Poetarum. By HORACE and JAMES SMITH. With Portrait and +Woodcuts.</p> + +<p class="indent">CONSOLATIONS IN TRAVEL; or, The Last Days of a Philosopher. +By SIR HUMPHRY DAVY. With Woodcuts.</p> + +<p class="indent">SALMONIA; or, Days of Fly-fishing. By SIR HUMPHRY +DAVY. With Woodcuts.</p> + +<p class="indent">THE INTELLECTUAL POWERS AND THE INVESTIGATION +OF TRUTH. By JOHN ABERCROMBIE.</p> + +<p class="indent">SPECIMENS OF THE TABLE-TALK OF THE LATE +SAMUEL TAYLOR COLERIDGE. With Portrait.</p> + +<p class="indent">PRACTICAL INSTRUCTIONS IN GARDENING, for every +Month in the Year. By MRS. LOUDON. With Woodcuts.<span class='pagenum'><a name="Page_7a" id="Page_7a">7</a></span></p> + +<p class="indent">A SMALLER HISTORY OF ENGLAND. Edited by DR. WM. +SMITH. With Woodcuts.</p> + +<p class="indent">A SMALLER HISTORY OF GREECE. Edited by DR. WM. +SMITH. With Woodcuts.</p> + +<p class="indent">A SMALLER HISTORY OF ROME. Edited by DR. WM. +SMITH. With Woodcuts.</p> + +<p class="indent">A SMALLER CLASSICAL MYTHOLOGY. With Translations +from the Ancient Poets. Edited by DR. WM. SMITH. With Woodcuts.</p> + +<p class="indent">A SMALLER ANCIENT HISTORY, from the Earliest Times +to the Conquest of Alexander the Great. Edited by DR. WM. SMITH. +With Woodcuts.</p> + +<p class="indent">A SMALLER HISTORY OF ENGLISH LITERATURE, from +the earliest period to the Georgian Era. Edited by DR. WM. SMITH.</p> + +<p class="indent">SPECIMENS OF THE CHIEF WRITERS IN ENGLISH +LITERATURE. Chronologically arranged. Edited by DR. WM. SMITH.</p> + +<p class="indent">A SMALLER SCRIPTURE HISTORY.—I. Old Testament History; +II. Connection of Old and New Testaments; III. New Testament +History to <span class="smcap">A.D.</span> 70. Edited by DR. WM. SMITH. With Woodcuts.</p> + +<h4><i>Four Shillings.</i></h4> + +<p class="indent">HISTORY OF ENGLAND, from the <span class="smcap">First Invasion</span> by the +<span class="smcap">Romans</span>, continued down to 1865. With <span class="smcap">Conversations</span> at the end of each +<span class="smcap">Chapter</span>. By MRS. MARKHAM. With 100 Woodcuts.</p> + +<p class="indent">HISTORY OF FRANCE, from the <span class="smcap">Conquest</span> by the <span class="smcap">Gauls</span>, continued +down to 1867. With <span class="smcap">Conversations</span> at the end of each <span class="smcap">Chapter</span>. +By MRS. MARKHAM. With 70 Woodcuts.</p> + +<p class="indent">HISTORY OF GERMANY, from the <span class="smcap">Invasion</span> of the <span class="smcap">Kingdom</span> by +the <span class="smcap">Romans</span> under <span class="smcap">Marius</span>, continued down to 1867. On the Plan of MRS. +MARKHAM. With 50 Woodcuts.</p> + +<p class="indent">SHALL AND WILL; or, the Future Auxiliary Verb. By SIR +EDMUND HEAD.</p> + +<h4><i>Four Shillings and Sixpence.</i></h4> + +<p class="indent">CHILDREN OF THE LAKE. A Poem. By EDWARD +SALLESBURY.</p> + +<p class="indent">A LADY’S DIARY OF THE SIEGE OF LUCKNOW.</p> + +<p class="indent">HOUSEHOLD SURGERY; or, Hints on Emergencies. By JOHN +F. SOUTH. With Woodcuts.<span class='pagenum'><a name="Page_8a" id="Page_8a">8</a></span></p> + +<h4><i>Five Shillings.</i></h4> + +<p class="indent">ANCIENT SPANISH BALLADS; <span class="smcap">Historical and Romantic.</span> +Translated with Notes by J. G. LOCKHART. With Portrait and Illustrations.</p> + +<p class="indent">MISCELLANIES. By LORD BYRON. 2 vols.</p> + +<p class="indent">INTRODUCTIONS TO THE STUDY OF THE GREEK +CLASSIC POETS. By HENRY NELSON COLERIDGE.</p> + +<p class="indent">HYMNS IN PROSE FOR CHILDREN. By MRS. BARBAULD. +With 112 Illustrations.</p> + +<p class="indent">RECOLLECTIONS OF THE DRUSES, <span class="smcap">and some Notes on +their Religion</span>. By LORD CARNARVON.</p> + +<p class="indent">THE ORIGIN OF LANGUAGE. BASED ON MODERN RESEARCHES. +By REV. F. W. FARRAR.</p> + +<p class="indent">MODERN DOMESTIC COOKERY. Founded on Principles of +Economy and Practical Knowledge, and adapted for Private Families. +With Woodcuts.</p> + +<p class="indent">DRAMAS AND PLAYS. By LORD BYRON. 2 vols.</p> + +<p class="indent">THE HORSE AND HIS RIDER. By SIR FRANCIS HEAD. +With Woodcuts.</p> + +<p class="indent">HANDBOOK OF FAMILIAR QUOTATIONS, chiefly from +English Authors.</p> + +<p class="indent">THE CHACE—THE TURF—AND THE ROAD. A Series of +Popular Essays. By C. J. APPERLEY (<span class="smcap">Nimrod</span>). With Portrait and +Illustrations.</p> + +<p class="indent">AUNT IDA’S WALKS AND TALKS. A Story Book for +Children. By A LADY.</p> + +<p class="indent">STORIES FOR DARLINGS. A Book for Boys and Girls. With +Illustrations.</p> + +<p class="indent">THE CHARMED ROE. A Story Book for Young People. Illustrated +by OTTO SPECKTER.</p> + +<p class="indent">DON JUAN AND BEPPO. By LORD BYRON. 2 vols.</p> + +<p class="indent">LIFE IN THE LIGHT OF GOD’S WORD. By ARCHBISHOP +THOMSON, D.D.</p> + +<p class="indent">ATHENS AND ATTICA; Notes of a Tour. By BISHOP +WORDSWORTH, D.D. With Illustrations.</p> + +<p class="indent">ANNALS OF THE WARS—XVIIITH CENTURY, 1700-1799. +Compiled from the most Authentic Sources. By SIR EDWARD CUST, D.C.L. +With Maps. 5 vols. Post 8vo. 5s. each.</p> + +<p class="indent">ANNALS OF THE WARS—<span class="smcap">XIXth</span> CENTURY, 1800-15. +Compiled from the most Authentic Sources. By SIR EDWARD CUST. +4 vols. Fcap. 8vo. 5s. each.<span class='pagenum'><a name="Page_9a" id="Page_9a">9</a></span></p> + +<h4><i>Six Shillings.</i></h4> + +<p class="indent">BENEDICITE; or, <span class="smcap">THE SONG of the THREE CHILDREN</span>. +Being <span class="smcap">Illustrations</span> of the <span class="smcap">Power</span>, <span class="smcap">Beneficence</span>, and <span class="smcap">Design</span> manifested by +the <span class="smcap">Creator</span> in <span class="smcap">His Works</span>. By DR. CHAPLIN CHILD.</p> + +<p class="indent">OLD DECCAN DAYS; or, HINDOO FAIRY LEGENDS +current in Southern India. By M. FRERE. With Introduction by <span class="smcap">Sir +Bartle Frere</span>. With Illustrations.</p> + +<p class="indent">THE WILD GARDEN; or, <span class="smcap">Our Groves and Shrubberies +made beautiful by the Naturalization of Hardy Exotic Plants</span>. +By WILLIAM ROBINSON. With Frontispiece.</p> + +<p class="indent">MISSIONARY TRAVELS AND RESEARCHES IN SOUTH +AFRICA. By DAVID LIVINGSTONE, M.D. With Map and Illustrations.</p> + +<p class="indent">FIVE YEARS OF A HUNTER’S LIFE IN SOUTH AFRICA; +By GORDON CUMMING. With Illustrations.</p> + +<p class="indent">THOUGHTS ON ANIMALCULES; or, The Invisible World, as +revealed by the Microscope. By GIDEON A. MANTELL. With Plates.</p> + +<p class="indent">LIVES OF BRINDLEY AND THE EARLY ENGINEERS. +By SAMUEL SMILES. With Woodcuts.</p> + +<p class="indent">LIFE OF TELFORD. With a History of Roads and Travelling +in England. By SAMUEL SMILES. With Woodcuts.</p> + +<p class="indent">LIVES OF GEORGE AND ROBERT STEPHENSON. By +SAMUEL SMILES. With Woodcuts.</p> + +<p class="indent">SELF-HELP. With Illustrations of Character and Conduct. By +SAMUEL SMILES.</p> + +<p class="indent">INDUSTRIAL BIOGRAPHY: Iron-workers and Toolmakers. A +Sequel to ‘Self-help.’ By SAMUEL SMILES.</p> + +<p class="indent">THE HUGUENOTS IN ENGLAND AND IRELAND: their +Settlements, Churches, and Industries. By SAMUEL SMILES.</p> + +<p class="indent">WILD WALES; its People, Language, and Scenery. With Introductory +Remarks. By GEORGE BORROW.</p> + +<p class="indent">A MANUAL OF ETHNOLOGY; or, <span class="smcap">A Popular History</span> of the +<span class="smcap">Races</span> of the <span class="smcap">Old World</span>. By CHARLES L. BRACE.</p> + +<h4><i>Seven Shillings.</i></h4> + +<p class="indent">JOURNALS OF A TOUR IN INDIA. By BISHOP HEBER. +2 vols.</p> + +<p class="indent">ADVENTURES AMONG THE MARQUESAS AND SOUTH +SEA ISLANDERS. By HERMAN MELVILLE. 2 vols.</p> + +<p class="indent">CONSTITUTIONAL PROGRESS. By MONTAGU BURROWS, +M.A.</p> + +<p class="indent">LIFE AND POETICAL WORKS OF REV. GEORGE +CRABBE. Edited by HIS SON. With Notes, Portrait, and Illustrations.<span class='pagenum'><a name="Page_10a" id="Page_10a">10</a></span></p> + +<h4><i>Seven Shillings and Sixpence.</i></h4> + +<p class="indent">THE ART OF TRAVEL; or, Hints on the Shifts and Contrivances +available in Wild Countries. By FRANCIS GALTON. With +Woodcuts.</p> + +<p class="indent">VISITS TO THE MONASTERIES OF THE LEVANT. By +HON. R. CURZON. With Illustrations.</p> + +<p class="indent">LETTERS FROM HIGH LATITUDES; an Account of a Yacht +Voyage to Iceland, Jan Mayen, and Spitzbergen, &c. By LORD DUFFERIN. +With Illustrations.</p> + +<p class="indent">BUBBLES FROM THE BRUNNEN OF NASSAU. By an +Old Man (SIR FRANCIS HEAD). With Illustrations.</p> + +<p class="indent">NINEVEH AND ITS REMAINS; a Narrative of an Expedition +to Assyria in 1845-47. By A. H. LAYARD. With Illustrations.</p> + +<p class="indent">NINEVEH AND BABYLON; a Narrative of a Second Expedition +to Assyria in 1849-51. By A. H. LAYARD. With Illustrations.</p> + +<p class="indent">THREE YEARS’ RESIDENCE IN ABYSSINIA, with Travels +in that Country. By MANSFIELD PARKYNS. With Illustrations.</p> + +<p class="indent">FIVE YEARS IN DAMASCUS, with <span class="smcap">Travels</span> in <span class="smcap">Palmyra</span>, +<span class="smcap">Lebanon</span>, and among the <span class="smcap">Giant Cities of Bashan</span> and <span class="smcap">The Hauran</span>. By +REV. J. L. PORTER. With Illustrations.</p> + +<p class="indent">THE VOYAGE OF THE ‘FOX’ IN THE ARCTIC SEAS, +and the Discovery of the Fate of Sir John Franklin and his Companions. +By SIR LEOPOLD McCLINTOCK. With Illustrations.</p> + +<p class="indent">REMINISCENCES OF ATHENS AND THE MOREA, during +Travels in Greece. By LORD CARNARVON. With Map.</p> + +<p class="indent">PEN AND PENCIL SKETCHES IN INDIA. By GENERAL +MUNDY. With Illustrations.</p> + +<p class="indent">PHILOSOPHY IN SPORT, MADE SCIENCE IN EARNEST: +or, The First Principles of Natural Philosophy explained by the Toys and +Sports of Youth. By DR. PARIS. With Woodcuts.</p> + +<p class="indent">BLIND PEOPLE; their Works and Ways. With Lives of some +famous Blind Men. By REV. B. G. JOHNS. With Illustrations.</p> + +<p class="indent">HORACE: A New Edition of the Text. Edited by DEAN +MILMAN. With 100 Woodcuts.</p> + +<p class="indent">THE BOOK OF THE CHURCH. By ROBERT SOUTHEY.</p> + +<p class="indent">A HANDBOOK FOR YOUNG PAINTERS. By C. R. +LESLIE, R.A. With 24 Illustrations.</p> + +<p class="indent">A GEOGRAPHICAL HANDBOOK OF FERNS, with Tables +to show their Distribution. By K. M. LYELL. With a Frontispiece.</p> + +<p class="indent">THE STORY OF THE LIFE OF LORD BACON. By W. +HEPWORTH DIXON.<span class='pagenum'><a name="Page_11a" id="Page_11a">11</a></span></p> + +<p class="indent">A SMALLER DICTIONARY OF THE BIBLE; Its Antiquities, +Geography, Biography, and Natural History. By DR. WM. SMITH. With +Maps and Illustrations.</p> + +<p class="indent">A SMALLER CLASSICAL DICTIONARY OF MYTHOLOGY, +BIOGRAPHY, AND GEOGRAPHY. By DR. WM. SMITH. With +200 Woodcuts.</p> + +<p class="indent">A SMALLER DICTIONARY OF GREEK AND ROMAN +ANTIQUITIES. By DR. WM. SMITH. With 200 Woodcuts.</p> + +<p class="indent">A SMALLER LATIN-ENGLISH DICTIONARY. With a +Dictionary of Proper Names, and Tables of the Roman Calendar, Measures, +Weights, and Moneys. By DR. WM. SMITH.</p> + +<p class="indent">A SMALLER ENGLISH-LATIN DICTIONARY, By DR. WM. +SMITH.</p> + +<p class="indent">THE STUDENT’S HUME; <span class="smcap">An Epitome</span> of the <span class="smcap">History Of +England</span>. By DAVID HUME. Corrected and continued to 1868. With +Woodcuts.</p> + +<p class="indent">THE STUDENT’S CONSTITUTIONAL HISTORY OF ENGLAND. +By HENRY HALLAM. With the Author’s latest Additions. +Edited by <span class="smcap">Dr. Wm. Smith</span>. [<i>In the Press.</i></p> + +<p class="indent">THE STUDENT’S HISTORY OF THE MIDDLE AGES OF +EUROPE. By HENRY HALLAM. With the Author’s Supplemental +Notes. Edited by <span class="smcap">Dr. Wm. Smith</span>. [<i>In the Press.</i></p> + +<p class="indent">THE STUDENT’S HISTORY OF FRANCE. <span class="smcap">From the +Earliest Times to the Establishment of the Second Empire</span>, 1852. +With Woodcuts.</p> + +<p class="indent">THE STUDENT’S HISTORY OF ROME. <span class="smcap">From the Earliest +Times to the Establishment of the Empire.</span> With Chapters on the +History of Literature and Art. By DEAN LIDDELL. With Woodcuts.</p> + +<p class="indent">THE STUDENT’S GIBBON; <span class="smcap">An Epitome of the History of +the Decline and Fall of the Roman Empire</span>. By EDWARD GIBBON. +With Woodcuts.</p> + +<p class="indent">THE STUDENT’S HISTORY OF GREECE. <span class="smcap">From the +Earliest Times to the Roman Conquest.</span> With Chapters on the +History of Literature and Art. By DR. WM. SMITH. With Woodcuts.</p> + +<p class="indent">THE STUDENT’S ANCIENT HISTORY OF THE EAST. +From the Earliest Times to the Conquests of Alexander the Great, including +Egypt, Assyria, Babylonia, Media, Persia, Asia Minor, and Phœnicia. By +PHILIP SMITH, B.A. With Woodcuts.</p> + +<p class="indent">THE STUDENT’S OLD TESTAMENT HISTORY. <span class="smcap">From the +Creation to the Return of the Jews from Captivity.</span> With Maps and +Woodcuts.</p> + +<p class="indent">THE STUDENT’S NEW TESTAMENT HISTORY. With an +Introduction, containing the connection of the Old and New Testaments. +With Maps and Woodcuts.</p> + +<p class="indent">THE STUDENT’S MANUAL OF THE ENGLISH LANGUAGE. +By GEORGE P. MARSH. Edited, with additional Chapters and +Notes.<span class='pagenum'><a name="Page_12a" id="Page_12a">12</a></span></p> + +<p class="indent">THE STUDENT’S MANUAL OF ENGLISH LITERATURE. +By T. B. SHAW, M.A. Edited, with Notes and Illustrations.</p> + +<p class="indent">THE STUDENT’S SPECIMENS OF ENGLISH LITERATURE. +Selected from the <span class="smcap">Best Writers</span>. By THOS. B. SHAW, M.A. +Edited, with Additions.</p> + +<p class="indent">THE STUDENT’S MANUAL OF ANCIENT GEOGRAPHY. +By REV. W. L. BEVAN. With Woodcuts.</p> + +<p class="indent">THE STUDENT’S MANUAL OF MODERN GEOGRAPHY. +Mathematical, Physical, and Descriptive. By REV. W. L. BEVAN. With +Woodcuts.</p> + +<p class="indent">THE STUDENT’S MANUAL OF MORAL PHILOSOPHY. +With Quotations and References. By WILLIAM FLEMING, D.D.</p> + +<p class="indent">THE STUDENT’S BLACKSTONE. <span class="smcap">A Systematic Abridgment +of the entire Commentaries.</span> By R. MALCOLM KERR, LL.D.</p> + +<p class="indent">A PRACTICAL HEBREW GRAMMAR. With the Hebrew +text of Genesis i.-vi. and Psalms i.-vi., Grammatical Analysis and Vocabulary. +By REV. STANLEY LEATHES.</p> + +<h4><i>Nine Shillings.</i></h4> + +<p class="indent">THE CONNECTION OF THE PHYSICAL SCIENCES. By +MARY SOMERVILLE. With Woodcuts.</p> + +<p class="indent">PHYSICAL GEOGRAPHY. By MARY SOMERVILLE. +Revised by H. W. BATES. With Portrait.</p> + +<p class="indent">THE STUDENT’S ELEMENTS OF GEOLOGY. By SIR +CHARLES LYELL. With 600 Woodcuts.</p> + +<p class="indent">A MANUAL OF SCIENTIFIC INQUIRY. By <span class="smcap">Various Writers</span>. +Edited by SIR J. F. HERSCHEL and REV. ROBERT MAIN.</p> + +<p class="indent">POETICAL WORKS OF LORD BYRON. With Notes, Illustrations, +and Portrait.</p> + +<p class="indent">LIFE OF LORD BYRON; with his Letters and Journals. By +THOMAS MOORE. With Portraits.</p> + +<p class="indent">ARCHBISHOP BECKET; A BIOGRAPHY. By CANON +ROBERTSON, M.A. With Illustrations.</p> + +<p class="indent">PICTURES OF THE CHINESE, DRAWN BY THEMSELVES. +Described by REV. R. H. COBBOLD. With 34 Illustrations.</p> + +<p class="indent">THE ENGLISH BATTLES AND SIEGES OF THE PENINSULAR +WAR. By SIR WILLIAM NAPIER. With Portrait.</p> + +<p class="indent">THE YOUNG OFFICER’S COMPANION; or, <span class="smcap">Essays</span> on +<span class="smcap">Military Duties</span> and <span class="smcap">Qualities</span>: with <span class="smcap">Illustrations</span> from <span class="smcap">History</span>. By +LORD DE ROS.</p> + +<p class="indent">DOG-BREAKING; the most Expeditious, Certain, and Easy +Method, whether great Excellence or only Mediocrity be required. With +a Few Hints for those who Love the Dog and the Gun. By GENERAL +HUTCHINSON. With Woodcuts.<span class='pagenum'><a name="Page_13a" id="Page_13a">13</a></span></p> + +<h4>LIST OF SCHOOL CLASSICS.</h4> + +<h4>By DR. WILLIAM SMITH.</h4> + +<p class="indent">PRINCIPIA LATINA, <span class="smcap">Part I. A First Latin Course</span>. A +Grammar, Delectus, and Exercise Book with Vocabularies. <i>13th Edition.</i> +<i>3s. 6d.</i></p> + +<p class="indent"><sup>*</sup><sub>*</sub><sup>*</sup> This Edition contains the Accidence arranged for the ‘Public School Latin +Primer.’</p> + +<p class="indent">PRINCIPIA LATINA, <span class="smcap">Part II. Latin Reading Book</span>. An +Introduction to Ancient Mythology, Geography, Roman Antiquities, and +History. With Notes and a Dictionary. <i>3s. 6d.</i></p> + +<p class="indent">PRINCIPIA LATINA, <span class="smcap">Part III. Latin Poetry</span>. 1. Easy +Hexameters and Pentameters. 2. Eclogæ Ovidianæ. 3. Prosody and Metre. +4. First Latin Verse Book. <i>3s. 6d.</i></p> + +<p class="indent">PRINCIPIA LATINA, <span class="smcap">Part IV. Latin Prose Composition</span>. +Rules of Syntax, with Examples, Explanations of Synonyms, and Exercises +on the Syntax. <i>3s. 6d.</i></p> + +<p class="indent">PRINCIPIA LATINA, <span class="smcap">Part V. Short Tales and Anecdotes +from Ancient History, for Translation into Latin Prose</span>. <i>3s.</i></p> + +<p class="indent">A LATIN-ENGLISH VOCABULARY, with a Latin-English +Dictionary to Phædrus, Cornelius Nepos, and Cæsar’s ‘Gallic War.’ <i>3s. 6d.</i></p> + +<p class="indent">THE STUDENT’S LATIN GRAMMAR. By WM. SMITH, +D.C.L., and THEOPHILUS D. HALL. <i>6s.</i></p> + +<p class="indent">A SMALLER LATIN GRAMMAR. Abridged from the above +Work. <i>3s. 6d.</i></p> + +<p class="indent">INITIA GRÆCA, <span class="smcap">Part I. A First Greek Course</span>, containing +Grammar, Delectus, Exercise Book, and Vocabularies. By DR. WM. SMITH. +<i>3s. 6d.</i></p> + +<p class="indent">INITIA GRÆCA, <span class="smcap">Part II. A Reading Book</span>; containing short +Tales, Anecdotes, Fables, Mythology, and Grecian History. With a Lexicon. +<i>3s. 6d.</i></p> + +<p class="indent">INITIA GRÆCA, <span class="smcap">Part III. Greek Prose Composition</span>; containing +the Rules of Syntax, with copious Examples and Exercises. <i>3s. 6d.</i></p> + +<p class="indent">THE STUDENT’S GREEK GRAMMAR. By PROFESSOR +CURTIUS, and WM. SMITH, D.C.L. <i>6s.</i></p> + +<p class="indent">A SMALLER GREEK GRAMMAR. Abridged from the above +Work. <i>3s. 6d.</i></p> + +<hr /> + +<p class="indent">PRINCIPIA GRÆCA. <span class="smcap">A First Greek Course.</span> A Grammar, +Delectus, and Exercise Book, with Vocabularies. By H. E. BUTTON, M.A. +<i>3s. 6d.</i></p> + +<p class="indent">MATTHIÆ’S GREEK GRAMMAR. Abridged by BLOMFIELD. +Revised and enlarged, by <span class="smcap">E. S. Crooke</span>, B.A. <i>4s.</i></p> + +<p class="indent">KING EDWARD VI.’S FIRST LATIN BOOK; including +a Short Syntax and Prosody with an English Translation. <i>2s. 6d.</i></p> + +<p class="indent">KING EDWARD VI.’S LATIN GRAMMAR. <i>3s. 6d.</i></p> + +<p class="indent">ENGLISH NOTES FOR LATIN ELEGIACS; designed for +Early Proficients in the Art of Latin Versification. By REV. W. OXENHAM. +<i>3s. 6d.</i><span class='pagenum'><a name="Page_14a" id="Page_14a">14</a></span></p> + +<h4>LIST OF HANDBOOKS FOR TRAVELLERS.</h4> + +<p class="indent">THE CONTINENT, &c.</p> + +<p class="indent">HANDBOOK—TRAVEL TALK,—<span class="smcap">English, French, German, +and Italian</span>. <i>3s. 6d.</i></p> + +<p class="indent">HANDBOOK—NORTH GERMANY, <span class="smcap">Holland, Belgium, +Prussia, and the Rhine to Switzerland</span>. With Map and Plans.</p> + +<p class="indent">HANDBOOK—SOUTH GERMANY, <span class="smcap">The Tyrol, Bavaria, +Austria, Salzburg, Styria, Hungary, and The Danube, from Ulm to the +Black Sea</span>. With Map and Plans. <i>10s.</i></p> + +<p class="indent">HANDBOOK—SWITZERLAND, <span class="smcap">The Alps of Savoy and +Piedmont</span>. With Maps and Plans. <i>10s.</i></p> + +<p class="indent">HANDBOOK—FRANCE, <span class="smcap">Normandy, Brittany, The French +Alps, Dauphine, Provence, and the Pyrenees</span>. With Maps. <i>12s.</i></p> + +<p class="indent">HANDBOOK—PARIS AND ITS ENVIRONS. With Map and +Plans. <i>3s. 6d.</i></p> + +<p class="indent"><sup>*</sup><sub>*</sub><sup>*</sup> <span class="smcap">Murray’s Plan of Paris.</span> <i>3s. 6d.</i></p> + +<p class="indent">HANDBOOK—CORSICA AND SARDINIA. With Map. <i>4s.</i></p> + +<p class="indent">HANDBOOK—SPAIN, <span class="smcap">Madrid, The Castiles, The Basque +Provinces, Leon, The Asturias, Galicia, Estremadura, Andalusia, +Ronda, Granada, Murcia, Valencia, Catalonia, Aragon, Navarre, +The Balearic Islands</span>, &c., &c. With Maps. 2 vols. <i>24s.</i></p> + +<p class="indent">HANDBOOK—PORTUGAL, <span class="smcap">Lisbon, Porto, Cintra, Mafra</span>, +&c. With Map. <i>9s.</i></p> + +<p class="indent">HANDBOOK—NORTH ITALY, <span class="smcap">Piedmont, Nice, Lombardy, +Venice, Parma, Modena, and Romagna</span>. With Map and Plans. <i>12s.</i></p> + +<p class="indent">HANDBOOK—CENTRAL ITALY, <span class="smcap">Tuscany, Florence, Lucca, +Umbria, The Marches, and the Patrimony of St. Peter</span>. With Map. +<i>10s.</i></p> + +<p class="indent">HANDBOOK—ROME AND ITS ENVIRONS. With Map and +Plans. <i>9s.</i></p> + +<p class="indent">HANDBOOK—SOUTH ITALY, <span class="smcap">Two Sicilies, Naples, Pompeii, +Herculaneum, Vesuvius, Abruzzi</span>, &c. With Map. <i>10s.</i></p> + +<p class="indent">HANDBOOK—SICILY, <span class="smcap">Palermo, Messina, Catania, Syracuse, +Etna, and the Ruins of the Greek Temples</span>. With Map. <i>12s.</i></p> + +<p class="indent">HANDBOOK—EGYPT, <span class="smcap">The Nile, Alexandria, Cairo, +Thebes, and the Overland Route to India</span>. With Map. <i>15s.</i></p> + +<p class="indent">HANDBOOK—GREECE, <span class="smcap">The Ionian Islands, Athens, +Albania, Thessaly, and Macedonia</span>. With Map.</p> + +<p class="indent">HANDBOOK—<span class="smcap">Constantinople</span>, <span class="smcap">The Bosphorus</span>, <span class="smcap">Dardanelles</span>, +<span class="smcap">Brousa, and Plain of Troy</span>, with General Hints for Travellers in Turkey, +&c. With Map.<span class='pagenum'><a name="Page_15a" id="Page_15a">15</a></span></p> + +<p class="indent">HANDBOOK—DENMARK, <span class="smcap">Norway, Sweden, and Iceland</span>. +With Map and Plans.</p> + +<p class="indent">HANDBOOK—RUSSIA, <span class="smcap">St. Petersburg, Moscow, Finland</span>, &c. +With Map. <i>15s.</i></p> + +<p class="indent">HANDBOOK—INDIA, <span class="smcap">Bombay and Madras</span>. With Map. 2 vols. +<i>12s.</i> each.</p> + +<p class="indent">HANDBOOK—HOLY LAND, <span class="smcap">Syria, Palestine, Sinai, Edom, +and the Syrian Deserts</span>. With Map. 2 vols. <i>24s.</i></p> + +<h4>KNAPSACK GUIDES FOR TRAVELLERS.</h4> + +<p class="indent">KNAPSACK GUIDE TO SWITZERLAND. With Plans. <i>5s.</i></p> + +<p class="indent">KNAPSACK GUIDE TO NORWAY. With Map. <i>6s.</i></p> + +<p class="indent">KNAPSACK GUIDE TO ITALY. With Plans. <i>6s.</i></p> + +<p class="indent">KNAPSACK GUIDE TO THE TYROL. With Plans. <i>6s.</i></p> + +<p class="indent">ENGLAND AND WALES.</p> + +<p class="indent">HANDBOOK—LONDON AS IT IS. With Map and Plans. +<i>3s. 6d.</i></p> + +<p class="indent">HANDBOOK—ESSEX, CAMBRIDGE, SUFFOLK, AND +NORFOLK—<span class="smcap">Chelmsford, Colchester, Maldon, Cambridge, Ely, Newmarket, +Bury, Ipswich, Woodbridge, Felixstowe, Lowestoft, Norwich, +Yarmouth, Cromer</span>, &c. With Maps and Plans. <i>12s.</i></p> + +<p class="indent">HANDBOOK—KENT AND SUSSEX—<span class="smcap">Canterbury, Dover, +Ramsgate, Rochester, Chatham, Brighton, Chichester, Worthing, Hastings, +Lewes, Arundel</span>. With Map. <i>10s.</i></p> + +<p class="indent">HANDBOOK—SURREY AND HANTS—<span class="smcap">Kingston, Croydon, +Reigate, Guildford, Dorking, Boxhill, Winchester, Southampton, Portsmouth, +and The Isle of Wight</span>. With Map. <i>10s.</i></p> + +<p class="indent">HANDBOOK—BERKS, BUCKS, AND OXON—<span class="smcap">Windsor, +Eton, Reading, Aylesbury, Henley, Oxford, and the Thames</span>. With +Map. <i>7s. 6d.</i></p> + +<p class="indent">HANDBOOK—WILTS, DORSET, AND SOMERSET—<span class="smcap">Salisbury, +Chippenham, Weymouth, Sherborne, Wells, Bath, Bristol, +Taunton</span>, &c. With Map. <i>10s.</i></p> + +<p class="indent">HANDBOOK—DEVON AND CORNWALL—<span class="smcap">Exeter, +Ilfracombe, Linton, Sidmouth, Dawlish, Teignmouth, Plymouth, Devonport, +Torquay, Launceston, Penzance, Falmouth, The Lizard, Land‘s +End</span>, &c. With Map. <i>10s.</i><span class='pagenum'><a name="Page_16a" id="Page_16a">16</a></span></p> + +<p class="indent">HANDBOOK—GLOUCESTER, HEREFORD, AND +WORCESTER—<span class="smcap">Cirencester, Cheltenham, Stroud, Tewkesbury, Leominster, +Ross, Malvern, Kidderminster, Dudley, Bromsgrove, Evesham</span>. +With Map. <i>6s. 6d.</i></p> + +<p class="indent">HANDBOOK—NORTH WALES—<span class="smcap">Bangor, Carnarvon, +Beaumaris, Snowdon, Conway</span>, &c. With Map. <i>6s. 6d.</i></p> + +<p class="indent">HANDBOOK—SOUTH WALES—<span class="smcap">Monmouth, Carmarthen, +Tenby, Swansea, and the Wye</span>, &c. With Map. <i>7s.</i></p> + +<p class="indent">HANDBOOK—DERBY, NOTTS, LEICESTER, AND +STAFFORD—<span class="smcap">Matlock, Bakewell, Chatsworth, The Peak, Buxton, +Hardwick, Dove Dale, Ashborne, Southwell, Mansfield, Retford, +Burton, Belvoir, Melton Mowbray, Wolverhampton, Litchfield, +Walsall, Tamworth</span>. With Map. <i>7s. 6d.</i></p> + +<p class="indent">HANDBOOK—SHROPSHIRE, CHESHIRE, AND LANCASHIRE—<span class="smcap">Shrewsbury, +Ludlow, Bridgnorth, Oswestry, Chester, Crewe, +Alderley, Stockport, Birkenhead, Warrington, Bury, Manchester, +Liverpool, Burnley, Clitheroe, Bolton, Blackburn, Wigan, Preston, +Rochdale, Lancaster, Southport, Blackpool</span>, &c. With Map. <i>10s.</i></p> + +<p class="indent">HANDBOOK—YORKSHIRE—<span class="smcap">Doncaster, Hull, Selby, +Beverley, Scarborough, Whitby, Harrogate, Ripon, Leeds, Wakefield, +Bradford, Halifax, Huddersfield, Sheffield</span>. With Map. <i>12s.</i></p> + +<p class="indent">HANDBOOK—DURHAM AND NORTHUMBERLAND—<span class="smcap">Newcastle, +Darlington, Bishop Auckland, Stockton, Hartlepool, Sunderland, +Shields, Berwick, Tynemouth, Alnwick</span>. With Map. <i>9s.</i></p> + +<p class="indent">HANDBOOK—WESTMORLAND AND CUMBERLAND—<span class="smcap">Lancaster, +Furness Abbey, Ambleside, Kendal, Windermere, Coniston, +Keswick, Grasmere, Carlisle, Cockermouth, Penrith, Appleby</span>. With +Map. <i>6s.</i></p> + +<p class="indent"><sup>*</sup><sub>*</sub><sup>*</sup> <span class="smcap">Murray‘s Map of the Lake District</span>, <i>3s. 6d.</i></p> + +<p class="indent">HANDBOOK—SCOTLAND—<span class="smcap">Edinburgh, Melrose, Kelso, +Glasgow, Dumfries, Ayr, Stirling, Arran, The Clyde, Oban, Inverary, +Loch Lomond, Loch Katrine and Trosachs, Caledonian Canal, Inverness, +Perth, Dundee, Aberdeen, Braemar, Skye, Caithness, Ross, and +Sutherland</span>. With Maps and Plans. <i>9s.</i></p> + +<p class="indent">HANDBOOK—IRELAND—<span class="smcap">Dublin, Belfast, Donegal, Galway, +Wexford, Cork, Limerick, Waterford, Killarney, Munster</span>. +With Map. <i>12s.</i></p> + +<h4> +<span class="smcap">JOHN MURRAY, 50a, ALBEMARLE STREET.</span></h4> +<hr /> +<h6>LONDON: PRINTED BY WILLIAM CLOWES AND SONS, STAMFORD STREET, +AND CHARING CROSS.</h6> + +<div class="bbox"> +<h3>TRANSCRIBERS' NOTES</h3> + +<p>Footnote 292 spells the city Charleston but the text has it as +Charlestown. 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The Descent of Man and Selection in +Relation to Sex, Vol. I (1st editio, by Charles Darwin + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: The Descent of Man and Selection in Relation to Sex, Vol. I (1st edition) + +Author: Charles Darwin + +Release Date: January 15, 2011 [EBook #34967] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK THE DESCENT OF MAN AND *** + + + + +Produced by StevenGibbs, Turgut Dincer and the Online +Distributed Proofreading Team at http://www.pgdp.net + + + + + + + + + +THE DESCENT OF MAN, + +AND + +SELECTION IN RELATION TO SEX. + + + + +BY CHARLES DARWIN, M.A., F.R.S., &c. + +IN TWO VOLUMES.--VOL. I. + +WITH ILLUSTRATIONS. + +LONDON: + +JOHN MURRAY, ALBEMARLE STREET. + +1871. + +[_The right of Translation is reserved._] + +BY THE SAME AUTHOR. + + + ON THE ORIGIN OF SPECIES BY MEANS OF NATURAL SELECTION; or, THE + PRESERVATION OF FAVOURED RACES IN THE STRUGGLE FOR LIFE; Fifth + Edition (_Tenth Thousand_), with Additions and Corrections. + 1869. MURRAY. + + THE VARIATION OF ANIMALS AND PLANTS UNDER DOMESTICATION. In two vols. + With Illustrations. 1868. MURRAY. + + ON THE VARIOUS CONTRIVANCES BY WHICH BRITISH AND FOREIGN ORCHIDS ARE + FERTILISED BY INSECTS; and on the GOOD EFFECTS OF CROSSING. With + numerous Woodcuts. MURRAY. + + A NATURALIST'S VOYAGE ROUND THE WORLD; or, A JOURNAL OF RESEARCHES INTO + THE NATURAL HISTORY AND GEOLOGY OF THE COUNTRIES visited during the + voyage of H.M.S. 'Beagle,' under the command of Captain FITZROY, R.N. + _Eleventh Thousand_. MURRAY. + + ON THE STRUCTURE AND DISTRIBUTION OF CORAL REEFS. SMITH, ELDER, & CO. + + GEOLOGICAL OBSERVATIONS ON VOLCANIC ISLANDS. SMITH, ELDER, & CO. + + GEOLOGICAL OBSERVATIONS ON SOUTH AMERICA. SMITH, ELDER & CO. + + A MONOGRAPH OF THE CIRRIPEDIA. With numerous Illustrations. 2 vols. + 8vo. HARDWICKE. + + ON THE MOVEMENTS AND HABITS OF CLIMBING PLANTS. With Woodcuts. WILLIAMS + & NORGATE. + + +LONDON: PRINTED BY WILLIAM CLOWES AND SONS, STAMFORD STREET, AND CHARING +CROSS. + + + + + CONTENTS. + + + INTRODUCTION Page 1-5 + + + PART I. + + ON THE DESCENT OF MAN. + + + CHAPTER I. + + THE EVIDENCE OF THE DESCENT OF MAN FROM SOME LOWER FORM. + + Nature of the evidence bearing on the origin of man--Homologous + structures in man and the lower animals--Miscellaneous points + of correspondence--Development--Rudimentary structures, + muscles, sense-organs, hair, bones, reproductive organs, + &c.--The bearing of these three great classes of facts on the + origin of man 9-33 + + + CHAPTER II. + + COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER ANIMALS. + + The difference in mental power between the highest ape and the + lowest savage, immense--Certain instincts in common--The + emotions--Curiosity--Imitation--Attention--Memory--Imagination-- + Reason--Progressive improvement--Tools and weapons used by + animals--Language--Self-consciousness--Sense of beauty--Belief + in God, spiritual agencies, superstitions 34-69 + + + CHAPTER III. + + COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER + ANIMALS--_continued_. + + The moral sense--Fundamental proposition--The qualities of + social animals--Origin of sociability--Struggle between opposed + instincts--Man and social animal--The more enduring social + instincts conquer other less persistent instincts--The social + virtues alone regarded by savages--The self-regarding virtues + acquired at a later stage of development--The importance of the + judgment of the members of the same community on + conduct--Transmission of moral tendencies--Summary 70-106 + + + CHAPTER IV. + + ON THE MANNER OF DEVELOPMENT OF MAN FROM SOME LOWER FORM. + + Variability of body and mind in man--Inheritance--Causes of + variability--Laws of variation the same in man as in the lower + animals--Direct action of the conditions of life--Effects of + the increased use and disuse of parts--Arrested + development--Reversion--Correlated variation--Rate of + increase--Checks to increase--Natural selection--Man the most + dominant animal in the world--Importance of his corporeal + structure--The causes which have led to his becoming + erect--Consequent changes of structure--Decrease in size of the + canine teeth--Increased size and altered shape of the + skull--Nakedness--Absence of a tail--Defenceless condition of + man 107-157 + + + CHAPTER V. + + ON THE DEVELOPMENT OF THE INTELLECTUAL AND MORAL FACULTIES + DURING PRIMEVAL AND CIVILISED TIMES. + + The advancement of the intellectual powers through natural + selection--Importance of imitation--Social and moral + faculties--Their development within the limits of the same + tribe--Natural selection as affecting civilised + nations--Evidence that civilised nations were once barbarous 158-184 + + + CHAPTER VI. + + ON THE AFFINITIES AND GENEALOGY OF MAN. + + Position of man in the animal series--The natural system + genealogical--Adaptive characters of slight value--Various + small points of resemblance between man and the + Quadrumana--Rank of man in the natural system--Birthplace and + antiquity of man--Absence of fossil connecting-links--Lower + stages in the genealogy of man, as inferred, firstly from his + affinities and secondly from his structure--Early androgynous + condition of the Vertebrata--Conclusion 185-213 + + + CHAPTER VII. + + ON THE RACES OF MAN. + + The nature and value of specific characters--Application to the + races of man--Arguments in favour of, and opposed to, ranking + the so-called races of man as distinct species--Sub-species-- + Monogenists and polygenists--Convergence of character--Numerous + points of resemblance in body and mind between the most + distinct races of man--The state of man when he first spread + over the earth--Each race not descended from a single + pair--The extinction of races--The formation of races-- + The effects of crossing--Slight influence of the direct + action of the conditions of life--Slight or no influence of + natural selection--Sexual selection. 214-250 + + + PART II. SEXUAL SELECTION. + + CHAPTER VIII. + + PRINCIPLES OF SEXUAL SELECTION. + + Secondary sexual characters--Sexual selection--Manner of + action--Excess of males--Polygamy--The male alone generally + modified through sexual selection--Eagerness of the + male--Variability of the male--Choice exerted by the + female--Sexual compared with natural selection--Inheritance at + corresponding periods of life, at corresponding seasons of the + year, and as limited by sex--Relations between the several + forms of inheritance--Causes why one sex and the young are not + modified through sexual selection--Supplement on the + proportional numbers of the two sexes throughout the animal + kingdom--On the limitation of the numbers of the two sexes + through natural selection 253-320 + + + CHAPTER IX. + + SECONDARY SEXUAL CHARACTERS IN THE LOWER CLASSES OF THE ANIMAL + KINGDOM. + + These characters absent in the lowest classes--Brilliant + colours--Mollusca--Annelids--Crustacea, secondary sexual + characters strongly developed; dimorphism; colour; characters + not acquired before maturity--Spiders, sexual colours of; + stridulation by the males--Myriapoda 321-340 + + + CHAPTER X. + + SECONDARY SEXUAL CHARACTERS OF INSECTS. + + Diversified structures possessed by the males for seizing the + females--Differences between the sexes, of which the meaning is + not understood--Difference in size between the + sexes--Thysanura--Diptera--Hemiptera--Homoptera, musical powers + possessed by the males alone--Orthoptera, musical instruments + of the males, much diversified in structure; pugnacity; + colours--Neuroptera, sexual differences in colour--Hymenoptera, + pugnacity and colours--Coleoptera, colours; furnished with + great horns, apparently as an ornament; battles; stridulating + organs generally common to both sexes 341-385 + + + CHAPTER XI. + + INSECTS, _continued_.--ORDER LEPIDOPTERA. + + Courtship of butterflies--Battles--Ticking noise--Colours + common to both sexes, or more brilliant in the + males--Examples--Not due to the direct action of the conditions + of life--Colours adapted for protection--Colours of + moths--Display--Perceptive powers of the + Lepidoptera--Variability--Causes of the difference in colour + between the males and females--Mimickry, female butterflies + more brilliantly coloured than the males--Bright colours of + caterpillars--Summary and concluding remarks on the secondary + sexual characters of insects--Birds and insects compared 386-423 + + + + +THE DESCENT OF MAN; + +AND ON + +SELECTION IN RELATION TO SEX. + + +INTRODUCTION. + +The nature of the following work will be best understood by a brief +account of how it came to be written. During many years I collected +notes on the origin or descent of man, without any intention of +publishing on the subject, but rather with the determination not to +publish, as I thought that I should thus only add to the prejudices +against my views. It seemed to me sufficient to indicate, in the first +edition of my 'Origin of Species,' that by this work "light would be +thrown on the origin of man and his history;" and this implies that man +must be included with other organic beings in any general conclusion +respecting his manner of appearance on this earth. Now the case wears a +wholly different aspect. When a naturalist like Carl Vogt ventures to +say in his address as President of the National Institution of Geneva +(1869), "personne, en Europe au moins, n'ose plus soutenir la creation +independante et de toutes pieces, des especes," it is manifest that at +least a large number of naturalists must admit that species are the +modified descendants of other species; and this especially holds good +with the younger and rising naturalists. The greater number accept the +agency of natural selection; though some urge, whether with justice the +future must decide, that I have greatly overrated its importance. Of the +older and honoured chiefs in natural science, many unfortunately are +still opposed to evolution in every form. + +In consequence of the views now adopted by most naturalists, and which +will ultimately, as in every other case, be followed by other men, I +have been led to put together my notes, so as to see how far the general +conclusions arrived at in my former works were applicable to man. This +seemed all the more desirable as I had never deliberately applied these +views to a species taken singly. When we confine our attention to any +one form, we are deprived of the weighty arguments derived from the +nature of the affinities which connect together whole groups of +organisms--their geographical distribution in past and present times, +and their geological succession. The homological structure, +embryological development, and rudimentary organs of a species, whether +it be man or any other animal, to which our attention may be directed, +remain to be considered; but these great classes of facts afford, as it +appears to me, ample and conclusive evidence in favour of the principle +of gradual evolution. The strong support derived from the other +arguments should, however, always be kept before the mind. + +The sole object of this work is to consider, firstly, whether man, like +every other species, is descended from some pre-existing form; secondly, +the manner of his development; and thirdly, the value of the +differences between the so-called races of man. As I shall confine +myself to these points, it will not be necessary to describe in detail +the differences between the several races--an enormous subject which has +been fully discussed in many valuable works. The high antiquity of man +has recently been demonstrated by the labours of a host of eminent men, +beginning with M. Boucher de Perthes; and this is the indispensable +basis for understanding his origin. I shall, therefore, take this +conclusion for granted, and may refer my readers to the admirable +treatises of Sir Charles Lyell, Sir John Lubbock, and others. Nor shall +I have occasion to do more than to allude to the amount of difference +between man and the anthropomorphous apes; for Prof. Huxley, in the +opinion of most competent judges, has conclusively shewn that in every +single visible character man differs less from the higher apes than +these do from the lower members of the same order of Primates. + +This work contains hardly any original facts in regard to man; but as +the conclusions at which I arrived, after drawing up a rough draft, +appeared to me interesting, I thought that they might interest others. +It has often and confidently been asserted, that man's origin can never +be known: but ignorance more frequently begets confidence than does +knowledge: it is those who know little, and not those who know much, who +so positively assert that this or that problem will never be solved by +science. The conclusion that man is the co-descendant with other species +of some ancient, lower, and extinct form, is not in any degree new. +Lamarck long ago came to this conclusion, which has lately been +maintained by several eminent naturalists and philosophers; for instance +by Wallace, Huxley, Lyell, Vogt, Lubbock, Buechner, Rolle, &c.,[1] and +especially by Haeckel. This last naturalist, besides his great work, +'Generelle Morphologie' (1866), has recently (1868, with a second edit. +in 1870), published his 'Natuerliche Schoepfungsgeschichte,' in which he +fully discusses the genealogy of man. If this work had appeared before +my essay had been written, I should probably never have completed it. +Almost all the conclusions at which I have arrived I find confirmed by +this naturalist, whose knowledge on many points is much fuller than +mine. Wherever I have added any fact or view from Prof. Haeckel's +writings, I give his authority in the text, other statements I leave as +they originally stood in my manuscript, occasionally giving in the +foot-notes references to his works, as a confirmation of the more +doubtful or interesting points. + +During many years it has seemed to me highly probable that sexual +selection has played an important part in differentiating the races of +man; but in my 'Origin of Species' (first edition, p. 199) I contented +myself by merely alluding to this belief. When I came to apply this view +to man, I found it indispensable to treat the whole subject in full +detail.[2] Consequently the second part of the present work, treating of +sexual selection, has extended to an inordinate length, compared with +the first part; but this could not be avoided. + +I had intended adding to the present volumes an essay on the expression +of the various emotions by man and the lower animals. My attention was +called to this subject many years ago by Sir Charles Bell's admirable +work. This illustrious anatomist maintains that man is endowed with +certain muscles solely for the sake of expressing his emotions. As this +view is obviously opposed to the belief that man is descended from some +other and lower form, it was necessary for me to consider it. I likewise +wished to ascertain how far the emotions are expressed in the same +manner by the different races of man. But owing to the length of the +present work, I have thought it better to reserve my essay, which is +partially completed, for separate publication. + + + + +PART I. + +THE DESCENT OR ORIGIN OF MAN. + +PART I.--THE DESCENT OF MAN. + + + + +CHAPTER I. + +THE EVIDENCE OF THE DESCENT OF MAN FROM SOME LOWER FORM. + + + Nature of the evidence bearing on the origin of man--Homologous + structures in man and the lower animals--Miscellaneous points + of correspondence--Development--Rudimentary structures, + muscles, sense-organs, hair, bones, reproductive organs, + &c.--The bearing of these three great classes of facts on the + origin of man. + + +He who wishes to decide whether man is the modified descendant of some +pre-existing form, would probably first enquire whether man varies, +however slightly, in bodily structure and in mental faculties; and if +so, whether the variations are transmitted to his offspring in +accordance with the laws which prevail with the lower animals; such as +that of the transmission of characters to the same age or sex. Again, +are the variations the result, as far as our ignorance permits us to +judge, of the same general causes, and are they governed by the same +general laws, as in the case of other organisms; for instance by +correlation, the inherited effects of use and disuse, &c.? Is man +subject to similar malconformations, the result of arrested development, +of reduplication of parts, &c., and does he display in any of his +anomalies reversion to some former and ancient type of structure? It +might also naturally be enquired whether man, like so many other +animals, has given rise to varieties and sub-races, differing but +slightly from each other, or to races differing so much that they must +be classed as doubtful species? How are such races distributed over the +world; and how, when crossed, do they react on each other, both in the +first and succeeding generations? And so with many other points. + +The enquirer would next come to the important point, whether man tends +to increase at so rapid a rate, as to lead to occasional severe +struggles for existence, and consequently to beneficial variations, +whether in body or mind, being preserved, and injurious ones eliminated. +Do the races or species of men, whichever term may be applied, encroach +on and replace each other, so that some finally become extinct? We shall +see that all these questions, as indeed is obvious in respect to most of +them, must be answered in the affirmative, in the same manner as with +the lower animals. But the several considerations just referred to may +be conveniently deferred for a time; and we will first see how far the +bodily structure of man shows traces, more or less plain, of his descent +from some lower form. In the two succeeding chapters the mental powers +of man, in comparison with those of the lower animals, will be +considered. + + +_The Bodily Structure of Man._--It is notorious that man is constructed +on the same general type or model with other mammals. All the bones in +his skeleton can be compared with corresponding bones in a monkey, bat, +or seal. So it is with his muscles, nerves, blood-vessels and internal +viscera. The brain, the most important of all the organs, follows the +same law, as shewn by Huxley and other anatomists. Bischoff,[3] who is a +hostile witness, admits that every chief fissure and fold in the brain +of man has its analogy in that of the orang; but he adds that at no +period of development do their brains perfectly agree; nor could this be +expected, for otherwise their mental powers would have been the same. +Vulpian[4] remarks: "Les differences reelles qui existent entre +l'encephale de l'homme et celui des singes superieurs, sont bien +minimes. Il ne faut pas se faire d'illusions a cet egard. L'homme est +bien plus pres des singes anthropomorphes par les caracteres anatomiques +de son cerveau que ceux-ci ne le sont non-seulement des autres +mammiferes, mais memes de certains quadrumanes, des guenons et des +macaques." But it would be superfluous here to give further details on +the correspondence between man and the higher mammals in the structure +of the brain and all other parts of the body. + +It may, however, be worth while to specify a few points, not directly or +obviously connected with structure, by which this correspondence or +relationship is well shewn. + +Man is liable to receive from the lower animals, and to communicate to +them, certain diseases as hydrophobia, variola, the glanders, &c.; and +this fact proves the close similarity of their tissues and blood, both +in minute structure and composition, far more plainly than does their +comparison under the best microscope, or by the aid of the best chemical +analysis. Monkeys are liable to many of the same non-contagious diseases +as we are; thus Rengger,[5] who carefully observed for a long time the +_Cebus Azarae_ in its native land, found it liable to catarrh, with the +usual symptoms, and which when often recurrent led to consumption. +These monkeys suffered also from apoplexy, inflammation of the bowels, +and cataract in the eye. The younger ones when shedding their milk-teeth +often died from fever. Medicines produced the same effect on them as on +us. Many kinds of monkeys have a strong taste for tea, coffee, and +spirituous liquors: they will also, as I have myself seen, smoke tobacco +with pleasure. Brehm asserts that the natives of north-eastern Africa +catch the wild baboons by exposing vessels with strong beer, by which +they are made drunk. He has seen some of these animals, which he kept in +confinement, in this state; and he gives a laughable account of their +behaviour and strange grimaces. On the following morning they were very +cross and dismal; they held their aching heads with both hands and wore +a most pitiable expression: when beer or wine was offered them, they +turned away with disgust, but relished the juice of lemons.[6] An +American monkey, an Ateles, after getting drunk on brandy, would never +touch it again, and thus was wiser than many men. These trifling facts +prove how similar the nerves of taste must be in monkeys and man, and +how similarly their whole nervous system is affected. + +Man is infested with internal parasites, sometimes causing fatal +effects, and is plagued by external parasites, all of which belong to +the same genera or families with those infesting other mammals. Man is +subject like other mammals, birds, and even insects, to that mysterious +law, which causes certain normal processes, such as gestation, as well +as the maturation and duration of various diseases, to follow lunar +periods.[7] His wounds are repaired by the same process of healing; and +the stumps left after the amputation of his limbs occasionally possess, +especially during an early embryonic period, some power of regeneration, +as in the lowest animals.[8] + +The whole process of that most important function, the reproduction of +the species, is strikingly the same in all mammals, from the first act +of courtship by the male[9] to the birth and nurturing of the young. +Monkeys are born in almost as helpless a condition as our own infants; +and in certain genera the young differ fully as much in appearance from +the adults, as do our children from their full-grown parents.[10] It has +been urged by some writers as an important distinction, that with man +the young arrive at maturity at a much later age than with any other +animal; but if we look to the races of mankind which inhabit tropical +countries the difference is not great, for the orang is believed not to +be adult till the age of from ten to fifteen years.[11] Man differs +from woman in size, bodily strength, hairyness, &c., as well as in mind, +in the same manner as do the two sexes of many mammals. It is, in short, +scarcely possible to exaggerate the close correspondence in general +structure, in the minute structure of the tissues, in chemical +composition and in constitution, between man and the higher animals, +especially the anthropomorphous apes. + + +_Embryonic Development._--Man is developed from an ovule, about the +125th of an inch in diameter, which differs in no respect from the +ovules of other animals. The embryo itself at a very early period can +hardly be distinguished from that of other members of the vertebrate +kingdom. At this period the arteries run in arch-like branches, as if to +carry the blood to branchiae which are not present in the higher +vertebrata, though the slits on the sides of the neck still remain (_f_, +_g_, fig. 1), marking their former position. At a somewhat later period, +when the extremities are developed, "the feet of lizards and mammals," +as the illustrious Von Baer remarks, "the wings and feet of birds, no +less than the hands and feet of man, all arise from the same fundamental +form." It is, says Prof. Huxley,[12] "quite in the later stages of +development that the young human being presents marked differences from +the young ape, while the latter departs as much from the dog in its +developments, as the man does. Startling as this last assertion may +appear to be, it is demonstrably true." + +As some of my readers may never have seen a drawing of an embryo, I have +given one of man and another of a dog, at about the same early stage of +development, carefully copied from two works of undoubted accuracy.[13] + +[Illustration: Fig. 1. Upper figure human embryo, from Ecker. Lower +figure that of a dog, from Bischoff. + + _a._ Fore-brain, cerebral hemispheres, &c. + _b._ Mid-brain, corpora quadrigemina. + _c._ Hind-brain, cerebellum, medulla oblongata. + _d._ Eye. + _e._ Ear. + _f._ First visceral arch. + _g._ Second visceral arch. + H. Vertebral columns and muscles in process of development. + _i._ Anterior {~BOX DRAWINGS LIGHT DOWN AND LEFT~} + {~BOX DRAWINGS LIGHT VERTICAL~} extremities. + K. Posterior {~BOX DRAWINGS LIGHT UP AND LEFT~} + L. Tail or os coccyx.] + +After the foregoing statements made by such high authorities, it would +be superfluous on my part to give a number of borrowed details, shewing +that the embryo of man closely resembles that of other mammals. It may, +however, be added that the human embryo likewise resembles in various +points of structure certain low forms when adult. For instance, the +heart at first exists as a simple pulsating vessel; the excreta are +voided through a cloacal passage; and the os coccyx projects like a true +tail, "extending considerably beyond the rudimentary legs."[14] In the +embryos of all air-breathing vertebrates, certain glands called the +corpora Wolffiana, correspond with and act like the kidneys of mature +fishes.[15] Even at a later embryonic period, some striking resemblances +between man and the lower animals may be observed. Bischoff says that +the convolutions of the brain in a human foetus at the end of the +seventh month reach about the same stage of development as in a baboon +when adult.[16] The great toe, as Prof. Owen remarks,[17] "which forms +the fulcrum when standing or walking, is perhaps the most +characteristic peculiarity in the human structure;" but in an embryo, +about an inch in length, Prof. Wyman[18] found "that the great toe was +shorter than the others, and, instead of being parallel to them, +projected at an angle from the side of the foot, thus corresponding with +the permanent condition of this part in the quadrumana." I will conclude +with a quotation from Huxley,[19] who after asking, does man originate +in a different way from a dog, bird, frog or fish? says, "the reply is +not doubtful for a moment; without question, the mode of origin and the +early stages of the development of man are identical with those of the +animals immediately below him in the scale: without a doubt in these +respects, he is far nearer to apes, than the apes are to the dog." + + +_Rudiments._--This subject, though not intrinsically more important than +the two last, will for several reasons be here treated with more +fullness.[20] Not one of the higher animals can be named which does not +bear some part in a rudimentary condition; and man forms no exception to +the rule. Rudimentary organs must be distinguished from those that are +nascent; though in some cases the distinction is not easy. The former +are either absolutely useless, such as the mammae of male quadrupeds, or +the incisor teeth of ruminants which never cut through the gums; or they +are of such slight service to their present possessors, that we cannot +suppose that they were developed under the conditions which now exist. +Organs in this latter state are not strictly rudimentary, but they are +tending in this direction. Nascent organs, on the other hand, though not +fully developed, are of high service to their possessors, and are +capable of further development. Rudimentary organs are eminently +variable; and this is partly intelligible, as they are useless or nearly +useless, and consequently are no longer subjected to natural selection. +They often become wholly suppressed. When this occurs, they are +nevertheless liable to occasional reappearance through reversion; and +this is a circumstance well worthy of attention. + +Disuse at that period of life, when an organ is chiefly used, and this +is generally during maturity, together with inheritance at a +corresponding period of life, seem to have been the chief agents in +causing organs to become rudimentary. The term "disuse" does not relate +merely to the lessened action of muscles, but includes a diminished flow +of blood to a part or organ, from being subjected to fewer alternations +of pressure, or from becoming in any way less habitually active. +Rudiments, however, may occur in one sex of parts normally present in +the other sex; and such rudiments, as we shall hereafter see, have often +originated in a distinct manner. In some cases organs have been reduced +by means of natural selection, from having become injurious to the +species under changed habits of life. The process of reduction is +probably often aided through the two principles of compensation and +economy of growth; but the later stages of reduction, after disuse has +done all that can fairly be attributed to it, and when the saving to be +effected by the economy of growth would be very small,[21] are difficult +to understand. The final and complete suppression of a part, already +useless and much reduced in size, in which case neither compensation nor +economy can come into play, is perhaps intelligible by the aid of the +hypothesis of pangenesis, and apparently in no other way. But as the +whole subject of rudimentary organs has been fully discussed and +illustrated in my former works,[22] I need here say no more on this +head. + +Rudiments of various muscles have been observed in many parts of the +human body;[23] and not a few muscles, which are regularly present in +some of the lower animals can occasionally be detected in man in a +greatly reduced condition. Every one must have noticed the power which +many animals, especially horses, possess of moving or twitching their +skin; and this is effected by the panniculus carnosus. Remnants of this +muscle in an efficient state are found in various parts of our bodies; +for instance, on the forehead, by which the eyebrows are raised. The +_platysma myoides_, which is well developed on the neck, belongs to this +system, but cannot be voluntarily brought into action. Prof. Turner, of +Edinburgh, has occasionally detected, as he informs me, muscular +fasciculi in five different situations, namely in the axillae, near the +scapulae, &c., all of which must be referred to the system of the +panniculus. He has also shewn[24] that the _musculus sternalis_ or +_sternalis brutorum_, which is not an extension of the _rectus +abdominalis_, but is closely allied to the panniculus, occurred in the +proportion of about 3 per cent. in upwards of 600 bodies: he adds, that +this muscle affords "an excellent illustration of the statement that +occasional and rudimentary structures are especially liable to variation +in arrangement." + +Some few persons have the power of contracting the superficial muscles +on their scalps; and these muscles are in a variable and partially +rudimentary condition. M. A. de Candolle has communicated to me a +curious instance of the long-continued persistence or inheritance of +this power, as well as of its unusual development. He knows a family, in +which one member, the present head of a family, could, when a youth, +pitch several heavy books from his head by the movement of the scalp +alone; and he won wagers by performing this feat. His father, uncle, +grandfather, and all his three children possess the same power to the +same unusual degree. This family became divided eight generations ago +into two branches; so that the head of the above-mentioned branch is +cousin in the seventh degree to the head of the other branch. This +distant cousin resides in another part of France, and on being asked +whether he possessed the same faculty, immediately exhibited his power. +This case offers a good illustration how persistently an absolutely +useless faculty may be transmitted. + +The extrinsic muscles which serve to move the whole external ear, and +the intrinsic muscles which move the different parts, all of which +belong to the system of the panniculus, are in a rudimentary condition +in man; they are also variable in development, or at least in function. +I have seen one man who could draw his ears forwards, and another who +could draw them backwards;[25] and from what one of these persons told +me, it is probable that most of us by often touching our ears and thus +directing our attention towards them, could by repeated trials recover +some power of movement. The faculty of erecting the ears and of +directing them to different points of the compass, is no doubt of the +highest service to many animals, as they thus perceive the point of +danger; but I have never heard of a man who possessed the least power of +erecting his ears,--the one movement which might be of use to him. The +whole external shell of the ear may be considered a rudiment, together +with the various folds and prominences (helix and anti-helix, tragus and +anti-tragus, &c.) which in the lower animals strengthen and support the +ear when erect, without adding much to its weight. Some authors, +however, suppose that the cartilage of the shell serves to transmit +vibrations to the acoustic nerve; but Mr. Toynbee,[26] after collecting +all the known evidence on this head, concludes that the external shell +is of no distinct use. The ears of the chimpanzee and orang are +curiously like those of man, and I am assured by the keepers in the +Zoological Gardens that these animals never move or erect them; so that +they are in an equally rudimentary condition, as far as function is +concerned, as in man. Why these animals, as well as the progenitors of +man, should have lost the power of erecting their ears we cannot say. It +may be, though I am not quite satisfied with this view, that owing to +their arboreal habits and great strength they were but little exposed to +danger, and so during a lengthened period moved their ears but little, +and thus gradually lost the power of moving them. This would be a +parallel case with that of those large and heavy birds, which from +inhabiting oceanic islands have not been exposed to the attacks of +beasts of prey, and have consequently lost the power of using their +wings for flight. + +[Illustration: Fig. 2. Human Ear, modelled and drawn by Mr. Woolner. + +_a._ The projecting point.] + +The celebrated sculptor, Mr. Woolner, informs me of one little +peculiarity in the external ear, which he has often observed both in men +and women, and of which he perceived the full signification. His +attention was first called to the subject whilst at work on his figure +of Puck, to which he had given pointed ears. He was thus led to examine +the ears of various monkeys, and subsequently more carefully those of +man. The peculiarity consists in a little blunt point, projecting from +the inwardly folded margin, or helix. Mr. Woolner made an exact model of +one such case, and has sent me the accompanying drawing. (Fig. 2.) These +points not only project inwards, but often a little outwards, so that +they are visible when the head is viewed from directly in front or +behind. They are variable in size and somewhat in position, standing +either a little higher or lower; and they sometimes occur on one ear and +not on the other. Now the meaning of these projections is not, I think, +doubtful; but it may be thought that they offer too trifling a character +to be worth notice. This thought, however, is as false as it is natural. +Every character, however slight, must be the result of some definite +cause; and if it occurs in many individuals deserves consideration. The +helix obviously consists of the extreme margin of the ear folded +inwards; and this folding appears to be in some manner connected with +the whole external ear being permanently pressed backwards. In many +monkeys, which do not stand high in the order, as baboons and some +species of macacus,[27] the upper portion of the ear is slightly +pointed, and the margin is not at all folded inwards; but if the margin +were to be thus folded, a slight point would necessarily project inwards +and probably a little outwards. This could actually be observed in a +specimen of the _Ateles beelzebuth_ in the Zoological Gardens; and we +may safely conclude that it is a similar structure--a vestige of +formerly pointed ears--which occasionally reappears in man. + +The nictitating membrane, or third eyelid, with its accessory muscles +and other structures, is especially well developed in birds, and is of +much functional importance to them, as it can be rapidly drawn across +the whole eye-ball. It is found in some reptiles and amphibians, and in +certain fishes, as in sharks. It is fairly well developed in the two +lower divisions of the mammalian series, namely, in the monotremata and +marsupials, and in some few of the higher mammals, as in the walrus. But +in man, the quadrumana, and most other mammals, it exists, as is +admitted by all anatomists, as a mere rudiment, called the semilunar +fold.[28] + +The sense of smell is of the highest importance to the greater number of +mammals--to some, as the ruminants, in warning them of danger; to +others, as the carnivora, in finding their prey; to others, as the wild +boar, for both purposes combined. But the sense of smell is of extremely +slight service, if any, even to savages, in whom it is generally more +highly developed than in the civilised races. It does not warn them of +danger, nor guide them to their food; nor does it prevent the Esquimaux +from sleeping in the most fetid atmosphere, nor many savages from eating +half-putrid meat. Those who believe in the principle of gradual +evolution, will not readily admit that this sense in its present state +was originally acquired by man, as he now exists. No doubt he inherits +the power in _an_ enfeebled and so far rudimentary condition, from some +early progenitor, to whom it was highly serviceable and by whom it was +continually used. We can thus perhaps understand how it is, as Dr. +Maudsley has truly remarked,[29] that the sense of smell in man "is +singularly effective in recalling vividly the ideas and images of +forgotten scenes and places;" for we see in those animals, which have +this sense highly developed, such as dogs and horses, that old +recollections of persons and places are strongly associated with their +odour. + +Man differs conspicuously from all the other Primates in being almost +naked. But a few short straggling hairs are found over the greater part +of the body in the male sex, and fine down on that of the female sex, In +individuals belonging to the same race these hairs are highly variable, +not only in abundance, but likewise in position: thus the shoulders in +some Europeans are quite naked, whilst in others they bear thick tufts +of hair.[30] There can be little doubt that the hairs thus scattered +over the body are the rudiments of the uniform hairy coat of the lower +animals. This view is rendered all the more probable, as it is known +that fine, short, and pale-coloured hairs on the limbs and other parts +of the body occasionally become developed into "thickset, long, and +rather coarse dark hairs," when abnormally nourished near old-standing +inflamed surfaces.[31] + +I am informed by Mr. Paget that persons belonging to the same family +often have a few hairs in their eyebrows much longer than the others; so +that this slight peculiarity seems to be inherited. These hairs +apparently represent the vibrissae, which are used as organs of touch by +many of the lower animals. In a young chimpanzee I observed that a few +upright, rather long, hairs, projected above the eyes, where the true +eyebrows, if present, would have stood. + +The fine wool-like hair, or so-called lanugo, with which the human +foetus during the sixth month is thickly covered, offers a more +curious case. It is first developed, during the fifth month, on the +eyebrows and face, and especially round the mouth, where it is much +longer than that on the head. A moustache of this kind was observed by +Eschricht[32] on a female foetus; but this is not so surprising a +circumstance as it may at first appear, for the two sexes generally +resemble each other in all external characters during an early period of +growth. The direction and arrangement of the hairs on all parts of the +foetal body are the same as in the adult, but are subject to much +variability. The whole surface, including even the forehead and ears, is +thus thickly clothed; but it is a significant fact that the palms of the +hands and the soles of the feet are quite naked, like the inferior +surfaces of all four extremities in most of the lower animals. As this +can hardly be an accidental coincidence, we must consider the woolly +covering of the foetus to be the rudimental representative of the +first permanent coat of hair in those mammals which are born hairy. This +representation is much more complete, in accordance with the usual law +of embryological development, than that afforded by the straggling hairs +on the body of the adult. + +It appears as if the posterior molar or wisdom-teeth were tending to +become rudimentary in the more civilised races of man. These teeth are +rather smaller than the other molars, as is likewise the case with the +corresponding teeth in the chimpanzee and orang; and they have only two +separate fangs. They do not cut through the gums till about the +seventeenth year, and I am assured by dentists that they are much more +liable to decay, and are earlier lost, than the other teeth. It is also +remarkable that they are much more liable to vary both in structure and +in the period of their development than the other teeth.[33] In the +Melanian races, on the other hand, the wisdom-teeth are usually +furnished with three separate fangs, and are generally sound: they also +differ from the other molars in size less than in the Caucasian +races.[34] Prof. Schaaffhausen accounts for this difference between the +races by "the posterior dental portion of the jaw being always +shortened" in those that are civilised,[35] and this shortening may, I +presume, be safely attributed to civilised men habitually feeding on +soft, cooked food, and thus using their jaws less. I am informed by Mr. +Brace that it is becoming quite a common practice in the United States +to remove some of the molar teeth of children, as the jaw does not grow +large enough for the perfect development of the normal number. + +With respect to the alimentary canal I have met with an account of only +a single rudiment, namely the vermiform appendage of the caecum. The +caecum is a branch or diverticulum of the intestine, ending in a +cul-de-sac, and it is extremely long in many of the lower +vegetable-feeding mammals. In the marsupial koala it is actually more +than thrice as long as the whole body.[36] It is sometimes produced into +a long gradually-tapering point, and is sometimes constricted in parts. +It appears as if, in consequence of changed diet or habits, the caecum +had become much shortened in various animals, the vermiform appendage +being left as a rudiment of the shortened part. That this appendage is a +rudiment, we may infer from its small size, and from the evidence which +Prof. Canestrini[37] has collected of its variability in man. It is +occasionally quite absent, or again is largely developed. The passage is +sometimes completely closed for half or two-thirds of its length, with +the terminal part consisting of a flattened solid expansion. In the +orang this appendage is long and convoluted: in man it arises from the +end of the short caecum, and is commonly from four to five inches in +length, being only about the third of an inch in diameter. Not only is +it useless, but it is sometimes the cause of death, of which fact I have +lately heard two instances: this is due to small hard bodies, such as +seeds, entering the passage and causing inflammation.[38] + +In the Quadrumana and some other orders of mammals, especially in the +Carnivora, there is a passage near the lower end of the humerus, called +the supra-condyloid foramen, through which the great nerve of the fore +limb passes, and often the great artery. Now in the humerus of man, as +Dr. Struthers[39] and others have shewn, there is generally a trace of +this passage, and it is sometimes fairly well developed, being formed by +a depending hook-like process of bone, completed by a band of ligament. +When present the great nerve invariably passes through it, and this +clearly indicates that it is the homologue and rudiment of the +supra-condyloid foramen of the lower animals. Prof. Turner estimates, as +he informs me, that it occurs in about one per cent. of recent +skeletons; but during ancient times it appears to have been much more +common. Mr. Busk[40] has collected the following evidence on this head: +Prof. Broca "noticed the perforation in four and a half per cent. of the +arm-bones collected in the 'Cimetiere du Sud' at Paris; and in the +Grotto of Orrony, the contents of which are referred to the Bronze +period, as many as eight humeri out of thirty-two were perforated; but +this extraordinary proportion, he thinks, might be due to the cavern +having been a sort of 'family vault.' Again, M. Dupont found 30 per +cent. of perforated bones in the caves of the Valley of the Lesse, +belonging to the Reindeer period; whilst M. Leguay, in a sort of +_dolmen_ at Argenteuil, observed twenty-five per cent. to be perforated; +and M. Pruner-Bey found twenty-six per cent. in the same condition in +bones from Vaureal. Nor should it be left unnoticed that M. Pruner-Bey +states that this condition is common in Guanche skeletons." The fact +that ancient races, in this and several other cases, more frequently +present structures which resemble those of the lower animals than do the +modern races, is interesting. One chief cause seems to be that ancient +races stand somewhat nearer than modern races in the long line of +descent to their remote animal-like progenitors. + +The os coccyx in man, though functionless as a tail, plainly represents +this part in other vertebrate animals. At an early embryonic period it +is free, and, as we have seen, projects beyond the lower extremities. In +certain rare and anomalous cases it has been known, according to Isidore +Geoffroy St.-Hilaire and others,[41] to form a small external rudiment +of a tail. The os coccyx is short, usually including only four vertebrae: +and these are in a rudimental condition, for they consist, with the +exception of the basal one, of the centrum alone.[42] They are furnished +with some small muscles; one of which, as I am informed by Prof. Turner, +has been expressly described by Theile as a rudimentary repetition of +the extensor of the tail, which is so largely developed in many mammals. + +The spinal cord in man extends only as far downwards as the last dorsal +or first lumbar vertebra; but a thread-like structure (the _filum +terminale_) runs down the axis of the sacral part of the spinal canal, +and even along the back of the coccygeal bones. The upper part of this +filament, as Prof. Turner informs me, is undoubtedly homologous with the +spinal cord; but the lower part apparently consists merely of the _pia +mater_, or vascular investing membrane. Even in this case the os coccyx +may be said to possess a vestige of so important a structure as the +spinal cord, though no longer enclosed within a bony canal. The +following fact, for which I am also indebted to Prof. Turner, shews how +closely the os coccyx corresponds with the true tail in the lower +animals: Luschka has recently discovered at the extremity of the +coccygeal bones a very peculiar convoluted body, which is continuous +with the middle sacral artery; and this discovery led Krause and Meyer +to examine the tail of a monkey (Macacus) and of a cat, in both of which +they found, though not at the extremity, a similarly convoluted body. + +The reproductive system offers various rudimentary structures; but these +differ in one important respect from the foregoing cases. We are not +here concerned with a vestige of a part which does not belong to the +species in an efficient state; but with a part which is always present +and efficient in the one sex, being represented in the other by a mere +rudiment. Nevertheless, the occurrence of such rudiments is as difficult +to explain on the belief of the separate creation of each species, as in +the foregoing cases. Hereafter I shall have to recur to these rudiments, +and shall shew that their presence generally depends merely on +inheritance; namely, on parts acquired by one sex having been partially +transmitted to the other. Here I will only give some instances of such +rudiments. It is well known that in the males of all mammals, including +man, rudimentary mammae exist. These in several instances have become +well developed, and have yielded a copious supply of milk. Their +essential identity in the two sexes is likewise shewn by their +occasional sympathetic enlargement in both during an attack of the +measles. The _vesicula prostatica_, which has been observed in many +male mammals, is now universally acknowledged to be the homologue of the +female uterus, together with the connected passage. It is impossible to +read Leuckart's able description of this organ, and his reasoning, +without admitting the justness of his conclusion. This is especially +clear in the case of those mammals in which the true female uterus +bifurcates, for in the males of these the vesicula likewise +bifurcates.[43] Some additional rudimentary structures belonging to the +reproductive system might here have been adduced.[44] + + +The bearing of the three great classes of facts now given is +unmistakeable. But it would be superfluous here fully to recapitulate +the line of argument given in detail in my 'Origin of Species.' The +homological construction of the whole frame in the members of the same +class is intelligible, if we admit their descent from a common +progenitor, together with their subsequent adaptation to diversified +conditions. On any other view the similarity of pattern between the hand +of a man or monkey, the foot of a horse, the flipper of a seal, the wing +of a bat, &c., is utterly inexplicable. It is no scientific explanation +to assert that they have all been formed on the same ideal plan. With +respect to development, we can clearly understand, on the principle of +variations supervening at a rather late embryonic period, and being +inherited at a corresponding period, how it is that the embryos of +wonderfully different forms should still retain, more or less perfectly, +the structure of their common progenitor. No other explanation has ever +been given of the marvellous fact that the embryo of a man, dog, seal, +bat, reptile, &c., can at first hardly be distinguished from each other. +In order to understand the existence of rudimentary organs, we have only +to suppose that a former progenitor possessed the parts in question in a +perfect state, and that under changed habits of life they became greatly +reduced, either from simple disuse, or through the natural selection of +those individuals which were least encumbered with a superfluous part, +aided by the other means previously indicated. + +Thus we can understand how it has come to pass that man and all other +vertebrate animals have been constructed on the same general model, why +they pass through the same early stages of development, and why they +retain certain rudiments in common. Consequently we ought frankly to +admit their community of descent: to take any other view, is to admit +that our own structure and that of all the animals around us, is a mere +snare laid to entrap our judgment. This conclusion is greatly +strengthened, if we look to the members of the whole animal series, and +consider the evidence derived from their affinities or classification, +their geographical distribution and geological succession. It is only +our natural prejudice, and that arrogance which made our forefathers +declare that they were descended from demi-gods, which leads us to demur +to this conclusion. But the time will before long come when it will be +thought wonderful, that naturalists, who were well acquainted with the +comparative structure and development of man and other mammals, should +have believed that each was the work of a separate act of creation. + + + + +CHAPTER II. + +COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER ANIMALS. + + + The difference in mental power between the highest ape and the + lowest savage, immense--Certain instincts in common--The + emotions--Curiosity--Imitation--Attention--Memory--Imagination + --Reason--Progressive improvement--Tools and weapons used by + animals--Language--Self-consciousness--Sense of beauty--Belief + in God, spiritual agencies, superstitions. + + +We have seen in the last chapter that man bears in his bodily structure +clear traces of his descent from some lower form; but it may be urged +that, as man differs so greatly in his mental power from all other +animals, there must be some error in this conclusion. No doubt the +difference in this respect is enormous, even if we compare the mind of +one of the lowest savages, who has no words to express any number higher +than four, and who uses no abstract terms for the commonest objects or +affections,[45] with that of the most highly organised ape. The +difference would, no doubt, still remain immense, even if one of the +higher apes had been improved or civilised as much as a dog has been in +comparison with its parent-form, the wolf or jackal. The Fuegians rank +amongst the lowest barbarians; but I was continually struck with +surprise how closely the three natives on board H.M.S. "Beagle," who had +lived some years in England and could talk a little English, resembled +us in disposition and in most of our mental faculties. If no organic +being excepting man had possessed any mental power, or if his powers had +been of a wholly different nature from those of the lower animals, then +we should never have been able to convince ourselves that our high +faculties had been gradually developed. But it can be clearly shewn that +there is no fundamental difference of this kind. We must also admit that +there is a much wider interval in mental power between one of the lowest +fishes, as a lamprey or lancelet, and one of the higher apes, than +between an ape and man; yet this immense interval is filled up by +numberless gradations. + +Nor is the difference slight in moral disposition between a barbarian, +such as the man described by the old navigator Byron, who dashed his +child on the rocks for dropping a basket of sea-urchins, and a Howard or +Clarkson; and in intellect, between a savage who does not use any +abstract terms, and a Newton or Shakspeare. Differences of this kind +between the highest men of the highest races and the lowest savages, are +connected by the finest gradations. Therefore it is possible that they +might pass and be developed into each other. + +My object in this chapter is solely to shew that there is no fundamental +difference between man and the higher mammals in their mental faculties. +Each division of the subject might have been extended into a separate +essay, but must here be treated briefly. As no classification of the +mental powers has been universally accepted, I shall arrange my remarks +in the order most convenient for my purpose; and will select those facts +which have most struck me, with the hope that they may produce some +effect on the reader. + +With respect to animals very low in the scale, I shall have to give some +additional facts under Sexual Selection, shewing that their mental +powers are higher than might have been expected. The variability of the +faculties in the individuals of the same species is an important point +for us, and some few illustrations will here be given. But it would be +superfluous to enter into many details on this head, for I have found on +frequent enquiry, that it is the unanimous opinion of all those who have +long attended to animals of many kinds, including birds, that the +individuals differ greatly in every mental characteristic. In what +manner the mental powers were first developed in the lowest organisms, +is as hopeless an enquiry as how life first originated. These are +problems for the distant future, if they are ever to be solved by man. + +As man possesses the same senses with the lower animals, his fundamental +intuitions must be the same. Man has also some few instincts in common, +as that of self-preservation, sexual love, the love of the mother for +her new-born offspring, the power possessed by the latter of sucking, +and so forth. But man, perhaps, has somewhat fewer instincts than those +possessed by the animals which come next to him in the series. The orang +in the Eastern islands, and the chimpanzee in Africa, build platforms on +which they sleep; and, as both species follow the same habit, it might +be argued that this was due to instinct, but we cannot feel sure that it +is not the result of both animals having similar wants and possessing +similar powers of reasoning. These apes, as we may assume, avoid the +many poisonous fruits of the tropics, and man has no such knowledge; but +as our domestic animals, when taken to foreign lands and when first +turned out in the spring, often eat poisonous herbs, which they +afterwards avoid, we cannot feel sure that the apes do not learn from +their own experience or from that of their parents what fruits to +select. It is however certain, as we shall presently see, that apes +have an instinctive dread of serpents, and probably of other dangerous +animals. + +The fewness and the comparative simplicity of the instincts in the +higher animals are remarkable in contrast with those of the lower +animals. Cuvier maintained that instinct and intelligence stand in an +inverse ratio to each other; and some have thought that the intellectual +faculties of the higher animals have been gradually developed from their +instincts. But Pouchet, in an interesting essay,[46] has shewn that no +such inverse ratio really exists. Those insects which possess the most +wonderful instincts are certainly the most intelligent. In the +vertebrate series, the least intelligent members, namely fishes and +amphibians, do not possess complex instincts; and amongst mammals the +animal most remarkable for its instincts, namely the beaver, is highly +intelligent, as will be admitted by every one who has read Mr. Morgan's +excellent account of this animal.[47] + +Although the first dawnings of intelligence, according to Mr. Herbert +Spencer,[48] have been developed through the multiplication and +co-ordination of reflex actions, and although many of the simpler +instincts graduate into actions of this kind and can hardly be +distinguished from them, as in the case of young animals sucking, yet +the more complex instincts seem to have originated independently of +intelligence. I am, however, far from wishing to deny that instinctive +actions may lose their fixed and untaught character, and be replaced by +others performed by the aid of the free will. On the other hand, some +intelligent actions--as when, birds on oceanic islands first learn to +avoid man--after being performed during many generations, become +converted into instincts and are inherited. They may then be said to be +degraded in character, for they are no longer performed through reason +or from experience. But the greater number of the more complex instincts +appear to have been gained in a wholly different manner, through the +natural selection of variations of simpler instinctive actions. Such +variations appear to arise from the same unknown causes acting on the +cerebral organisation, which induce slight variations or individual +differences in other parts of the body; and these variations, owing to +our ignorance, are often said to arise spontaneously. We can, I think, +come to no other conclusion with respect to the origin of the more +complex instincts, when we reflect on the marvellous instincts of +sterile worker-ants and bees, which leave no offspring to inherit the +effects of experience and of modified habits. + +Although a high degree of intelligence is certainly compatible with the +existence of complex instincts, as we see in the insects just named and +in the beaver, it is not improbable that they may to a certain extent +interfere with each other's development. Little is known about the +functions of the brain, but we can perceive that as the intellectual +powers become highly developed, the various parts of the brain must be +connected by the most intricate channels of intercommunication; and as a +consequence each separate part would perhaps tend to become less well +fitted to answer in a definite and uniform, that is instinctive, manner +to particular sensations or associations. + +I have thought this digression worth giving, because we may easily +underrate the mental powers of the higher animals, and especially of +man, when we compare their actions founded on the memory of past events, +on foresight, reason, and imagination, with exactly similar actions +instinctively performed by the lower animals; in this latter case the +capacity of performing such actions having been gained, step by step, +through the variability of the mental organs and natural selection, +without any conscious intelligence on the part of the animal during each +successive generation. No doubt, as Mr. Wallace has argued,[49] much of +the intelligent work done by man is due to imitation and not to reason; +but there is this great difference between his actions and many of those +performed by the lower animals, namely, that man cannot, on his first +trial, make, for instance, a stone hatchet or a canoe, through his power +of imitation. He has to learn his work by practice; a beaver, on the +other hand, can make its dam or canal, and a bird its nest, as well, or +nearly as well, the first time it tries, as when old and experienced. + +To return to our immediate subject: the lower animals, like man, +manifestly feel pleasure and pain, happiness and misery. Happiness is +never better exhibited than by young animals, such as puppies, kittens, +lambs, &c., when playing together, like our own children. Even insects +play together, as has been described by that excellent observer, P. +Huber,[50] who saw ants chasing and pretending to bite each other, like +so many puppies. + +The fact that the lower animals are excited by the same emotions as +ourselves is so well established, that it will not be necessary to weary +the reader by many details. Terror acts in the same manner on them as on +us, causing the muscles to tremble, the heart to palpitate, the +sphincters to be relaxed, and the hair to stand on end. Suspicion, the +offspring of fear, is eminently characteristic of most wild animals. +Courage and timidity are extremely variable qualities in the +individuals of the same species, as is plainly seen in our dogs. Some +dogs and horses are ill-tempered and easily turn sulky; others are +good-tempered; and these qualities are certainly inherited. Every one +knows how liable animals are to furious rage, and how plainly they show +it. Many anecdotes, probably true, have been published on the +long-delayed and artful revenge of various animals. The accurate Rengger +and Brehm[51] state that the American and African monkeys which they +kept tame, certainly revenged themselves. The love of a dog for his +master is notorious; in the agony of death he has been known to caress +his master, and every one has heard of the dog suffering under +vivisection, who licked the hand of the operator; this man, unless he +had a heart of stone, must have felt remorse to the last hour of his +life. As Whewell[52] has remarked, "who that reads the touching +instances of maternal affection, related so often of the women of all +nations, and of the females of all animals, can doubt that the principle +of action is the same in the two cases?" + +We see maternal affection exhibited in the most trifling details; thus +Rengger observed an American monkey (a Cebus) carefully driving away the +flies which plagued her infant; and Duvaucel saw a Hylobates washing the +faces of her young ones in a stream. So intense is the grief of female +monkeys for the loss of their young, that it invariably caused the death +of certain kinds kept under confinement by Brehm in N. Africa. +Orphan-monkeys were always adopted and carefully guarded by the other +monkeys, both males and females. One female baboon had so capacious a +heart that she not only adopted young monkeys of other species, but +stole young dogs and cats, which she continually carried about. Her +kindness, however, did not go so far as to share her food with her +adopted offspring, at which Brehm was surprised, as his monkeys always +divided everything quite fairly with their own young ones. An adopted +kitten scratched the above-mentioned affectionate baboon, who certainly +had a fine intellect, for she was much astonished at being scratched, +and immediately examined the kitten's feet, and without more ado bit off +the claws. In the Zoological Gardens, I heard from the keeper that an +old baboon (_C. chacma_) had adopted a Rhesus monkey; but when a young +drill and mandrill were placed in the cage, she seemed to perceive that +these monkeys, though distinct species, were her nearer relatives, for +she at once rejected the Rhesus and adopted both of them. The young +Rhesus, as I saw, was greatly discontented at being thus rejected, and +it would, like a naughty child, annoy and attack the young drill and +mandrill whenever it could do so with safety; this conduct exciting +great indignation in the old baboon. Monkeys will also, according to +Brehm, defend their master when attacked by any one, as well as dogs to +whom they are attached, from the attacks of other dogs. But we here +trench on the subject of sympathy, to which I shall recur. Some of +Brehm's monkeys took much delight in teasing, in various ingenious ways, +a certain old dog whom they disliked, as well as other animals. + +Most of the more complex emotions are common to the higher animals and +ourselves. Every one has seen how jealous a dog is of his master's +affection, if lavished on any other creature; and I have observed the +same fact with monkeys. This shews that animals not only love, but have +the desire to be loved. Animals manifestly feel emulation. They love +approbation or praise; and a dog carrying a basket for his master +exhibits in a high degree self-complacency or pride. There can, I think, +be no doubt that a dog feels shame, as distinct from fear, and something +very like modesty when begging too often for food. A great dog scorns +the snarling of a little dog, and this may be called magnanimity. +Several observers have stated that monkeys certainly dislike being +laughed at; and they sometimes invent imaginary offences. In the +Zoological Gardens I saw a baboon who always got into a furious rage +when his keeper took out a letter or book and read it aloud to him; and +his rage was so violent that, as I witnessed on one occasion, he bit his +own leg till the blood flowed. + +We will now turn to the more intellectual emotions and faculties, which +are very important, as forming the basis for the development of the +higher mental powers. Animals manifestly enjoy excitement and suffer +from ennui, as may be seen with dogs, and, according to Rengger, with +monkeys. All animals feel Wonder, and many exhibit Curiosity. They +sometimes suffer from this latter quality, as when the hunter plays +antics and thus attracts them; I have witnessed this with deer, and so +it is with the wary chamois, and with some kinds of wild-ducks. Brehm +gives a curious account of the instinctive dread which his monkeys +exhibited towards snakes; but their curiosity was so great that they +could not desist from occasionally satiating their horror in a most +human fashion, by lifting up the lid of the box in which the snakes were +kept. I was so much surprised at his account, that I took a stuffed and +coiled-up snake into the monkey-house at the Zoological Gardens, and the +excitement thus caused was one of the most curious spectacles which I +ever beheld. Three species of Cercopithecus were the most alarmed; they +dashed about their cages and uttered sharp signal-cries of danger, which +were understood by the other monkeys. A few young monkeys and one old +Anubis baboon alone took no notice of the snake. I then placed the +stuffed specimen on the ground in one of the larger compartments. After +a time all the monkeys collected round it in a large circle, and staring +intently, presented a most ludicrous appearance. They became extremely +nervous; so that when a wooden ball, with which they were familiar as a +plaything, was accidently moved in the straw, under which it was partly +hidden, they all instantly started away. These monkeys behaved very +differently when a dead fish, a mouse, and some other new objects were +placed in their cages; for though at first frightened, they soon +approached, handled and examined them. I then placed a live snake in a +paper bag, with the mouth loosely closed, in one of the larger +compartments. One of the monkeys immediately approached, cautiously +opened the bag a little, peeped in, and instantly dashed away. Then I +witnessed what Brehm has described, for monkey after monkey, with head +raised high and turned on one side, could not resist taking momentary +peeps into the upright bag, at the dreadful object lying quiet at the +bottom. It would almost appear as if monkeys had some notion of +zoological affinities, for those kept by Brehm exhibited a strange, +though mistaken, instinctive dread of innocent lizards and frogs. An +orang, also, has been known to be much alarmed at the first sight of a +turtle.[53] + +The principle of _Imitation_ is strong in man, and especially in man in +a barbarous state. Desor[54] has remarked that no animal voluntarily +imitates an action performed by man, until in the ascending scale we +come to monkeys, which are well-known to be ridiculous mockers. Animals, +however, sometimes imitate each others' actions: thus two species of +wolves, which had been reared by dogs, learned to bark, as does +sometimes the jackal,[55] but whether this can be called voluntary +imitation is another question. From one account which I have read, there +is reason to believe that puppies nursed by cats sometimes learn to lick +their feet and thus to clean their faces: it is at least certain, as I +hear from a perfectly trustworthy friend, that some dogs behave in this +manner. Birds imitate the songs of their parents, and sometimes those of +other birds; and parrots are notorious imitators of any sound which they +often hear. + +Hardly any faculty is more important for the intellectual progress of +man than the power of _Attention_. Animals clearly manifest this power, +as when a cat watches by a hole and prepares to spring on its prey. Wild +animals sometimes become so absorbed when thus engaged, that they may be +easily approached. Mr. Bartlett has given me a curious proof how +variable this faculty is in monkeys. A man who trains monkeys to act +used to purchase common kinds from the Zoological Society at the price +of five pounds for each; but he offered to give double the price, if he +might keep three or four of them for a few days, in order to select one. +When asked how he could possibly so soon learn whether a particular +monkey would turn out a good actor, he answered that it all depended on +their power of attention. If when he was talking and explaining anything +to a monkey, its attention was easily distracted, as by a fly on the +wall or other trifling object, the case was hopeless. If he tried by +punishment to make an inattentive monkey act, it turned sulky. On the +other hand, a monkey which carefully attended to him could always be +trained. + +It is almost superfluous to state that animals have excellent _Memories_ +for persons and places. A baboon at the Cape of Good Hope, as I have +been informed by Sir Andrew Smith, recognised him with joy after an +absence of nine months. I had a dog who was savage and averse to all +strangers, and I purposely tried his memory after an absence of five +years and two days. I went near the stable where he lived, and shouted +to him in my old manner; he showed no joy, but instantly followed me out +walking and obeyed me, exactly as if I had parted with him only +half-an-hour before. A train of old associations, dormant during five +years, had thus been instantaneously awakened in his mind. Even ants, as +P. Huber[56] has clearly shewn, recognised their fellow-ants belonging +to the same community after a separation of four months. Animals can +certainly by some means judge of the intervals of time between recurrent +events. + +The _Imagination_ is one of the highest prerogatives of man. By this +faculty he unites, independently of the will, former images and ideas, +and thus creates brilliant and novel results. A poet, as Jean Paul +Richter remarks,[57] "who must reflect whether he shall make a +character say yes or no--to the devil with him; he is only a stupid +corpse." Dreaming gives us the best notion of this power; as Jean Paul +again says, "The dream is an involuntary art of poetry." The value of +the products of our imagination depends of course on the number, +accuracy, and clearness of our impressions; on our judgment and taste in +selecting or rejecting the involuntary combinations, and to a certain +extent on our power of voluntarily combining them. As dogs, cats, +horses, and probably all the higher animals, even birds, as is stated on +good authority,[58] have vivid dreams, and this is shewn by their +movements and voice, we must admit that they possess some power of +imagination. + +Of all the faculties of the human mind, it will, I presume, be admitted +that _Reason_ stands at the summit. Few persons any longer dispute that +animals possess some power of reasoning. Animals may constantly be seen +to pause, deliberate, and resolve. It is a significant fact, that the +more the habits of any particular animal are studied by a naturalist, +the more he attributes to reason and the less to unlearnt instincts.[59] +In future chapters we shall see that some animals extremely low in the +scale apparently display a certain amount of reason. No doubt it is +often difficult to distinguish between the power of reason and that of +instinct. Thus Dr. Hayes, in his work on 'The Open Polar Sea,' +repeatedly remarks that his dogs, instead of continuing to draw the +sledges in a compact body, diverged and separated when they came to thin +ice, so that their weight might be more evenly distributed. This was +often the first warning and notice which the travellers received that +the ice was becoming thin and dangerous. Now, did the dogs act thus from +the experience of each individual, or from the example of the older and +wiser dogs, or from an inherited habit, that is from an instinct? This +instinct might possibly have arisen since the time, long ago, when dogs +were first employed by the natives in drawing their sledges; or the +Arctic wolves, the parent-stock of the Esquimaux dog, may have acquired +this instinct, impelling them not to attack their prey in a close pack +when on thin ice. Questions of this kind are most difficult to answer. + +So many facts have been recorded in various works shewing that animals +possess some degree of reason, that I will here give only two or three +instances, authenticated by Rengger, and relating to American monkeys, +which stand low in their order. He states that when he first gave eggs +to his monkeys, they smashed them and thus lost much of their contents; +afterwards they gently hit one end against some hard body, and picked +off the bits of shell with their fingers. After cutting themselves only +once with any sharp tool, they would not touch it again, or would handle +it with the greatest care. Lumps of sugar were often given them wrapped +up in paper; and Rengger sometimes put a live wasp in the paper, so that +in hastily unfolding it they got stung; after this had once happened, +they always first held the packet to their ears to detect any movement +within. Any one who is not convinced by such facts as these, and by what +he may observe with his own dogs, that animals can reason, would not be +convinced by anything that I could add. Nevertheless I will give one +case with respect to dogs, as it rests on two distinct observers, and +can hardly depend on the modification of any instinct. + +Mr. Colquhoun[60] winged two wild-ducks, which fell on the opposite side +of a stream; his retriever tried to bring over both at once, but could +not succeed; she then, though never before known to ruffle a feather, +deliberately killed one, brought over the other, and returned for the +dead bird. Col. Hutchinson relates that two partridges were shot at +once, one being killed, the other wounded; the latter ran away, and was +caught by the retriever, who on her return came across the dead bird; +"she stopped, evidently greatly puzzled, and after one or two trials, +finding she could not take it up without permitting the escape of the +winged bird, she considered a moment, then deliberately murdered it by +giving it a severe crunch, and afterwards brought away both together. +This was the only known instance of her ever having wilfully injured any +game." Here we have reason, though not quite perfect, for the retriever +might have brought the wounded bird first and then returned for the dead +one, as in the case of the two wild-ducks. + +The muleteers in S. America say, "I will not give you the mule whose +step is easiest, but _la mas rational_,--the one that reasons best;" and +Humboldt[61] adds, "this popular expression, dictated by long +experience, combats the system of animated machines, better perhaps than +all the arguments of speculative philosophy." + + +It has, I think, now been shewn that man and the higher animals, +especially the Primates, have some few instincts in common. All have the +same senses, intuitions and sensations--similar passions, affections, +and emotions, even the more complex ones; they feel wonder and +curiosity; they possess the same faculties of imitation, attention, +memory, imagination, and reason, though in very different degrees. +Nevertheless many authors have insisted that man is separated through +his mental faculties by an impassable barrier from all the lower +animals. I formerly made a collection of above a score of such +aphorisms, but they are not worth giving, as their wide difference and +number prove the difficulty, if not the impossibility, of the attempt. +It has been asserted that man alone is capable of progressive +improvement; that he alone makes use of tools or fire, domesticates +other animals, possesses property, or employs language; that no other +animal is self-conscious, comprehends itself, has the power of +abstraction, or possesses general ideas; that man alone has a sense of +beauty, is liable to caprice, has the feeling of gratitude, mystery, +&c.; believes in God, or is endowed with a conscience. I will hazard a +few remarks on the more important and interesting of these points. + +Archbishop Sumner formerly maintained[62] that man alone is capable of +progressive improvement. With animals, looking first to the individual, +every one who has had any experience in setting traps knows that young +animals can be caught much more easily than old ones; and they can be +much more easily approached by an enemy. Even with respect to old +animals, it is impossible to catch many in the same place and in the +same kind of trap, or to destroy them by the same kind of poison; yet it +is improbable that all should have partaken of the poison, and +impossible that all should have been caught in the trap. They must learn +caution by seeing their brethren caught or poisoned. In North America, +where the fur-bearing animals have long been pursued, they exhibit, +according to the unanimous testimony of all observers, an almost +incredible amount of sagacity, caution, and cunning; but trapping has +been there so long carried on that inheritance may have come into play. + +If we look to successive generations, or to the race, there is no doubt +that birds and other animals gradually both acquire and lose caution in +relation to man or other enemies;[63] and this caution is certainly in +chief part an inherited habit or instinct, but in part the result of +individual experience. A good observer, Leroy,[64] states that in +districts where foxes are much hunted, the young when they first leave +their burrows are incontestably much more wary than the old ones in +districts where they are not much disturbed. + +Our domestic dogs are descended from wolves and jackals,[65] and though +they may not have gained in cunning, and may have lost in waryness and +suspicion, yet they have progressed in certain moral qualities, such as +in affection, trust-worthiness, temper, and probably in general +intelligence. The common rat has conquered and beaten several other +species throughout Europe, in parts of North America, New Zealand, and +recently in Formosa, as well as on the mainland of China. Mr. +Swinhoe,[66] who describes these latter cases, attributes the victory of +the common rat over the large _Mus coninga_ to its superior cunning; and +this latter quality may be attributed to the habitual exercise of all +its faculties in avoiding extirpation by man, as well as to nearly all +the less cunning or weak-minded rats having been successively destroyed +by him. To maintain, independently of any direct evidence, that no +animal during the course of ages has progressed in intellect or other +mental faculties, is to beg the question of the evolution of species. +Hereafter we shall see that, according to Lartet, existing mammals +belonging to several orders have larger brains than their ancient +tertiary prototypes. + +It has often been said that no animal uses any tool; but the chimpanzee +in a state of nature cracks a native fruit, somewhat like a walnut, with +a stone.[67] Rengger[68] easily taught an American monkey thus to break +open hard palm-nuts, and afterwards of its own accord it used stones to +open other kinds of nuts, as well as boxes. It thus also removed the +soft rind of fruit that had a disagreeable flavour. Another monkey was +taught to open the lid of a large box with a stick, and afterwards it +used the stick as a lever to move heavy bodies; and I have myself seen a +young orang put a stick into a crevice, slip his hand to the other end, +and use it in the proper manner as a lever. In the cases just mentioned +stones and sticks were employed as implements; but they are likewise +used as weapons. Brehm[69] states, on the authority of the well-known +traveller Schimper, that in Abyssinia when the baboons belonging to one +species (_C. gelada_) descend in troops from the mountains to plunder +the fields, they sometimes encounter troops of another species (_C. +hamadryas_), and then a fight ensues. The Geladas roll down great +stones, which the Hamadryas try to avoid, and then, both species, +making a great uproar, rush furiously against each other. Brehm, when +accompanying the Duke of Coburg-Gotha, aided in an attack with fire-arms +on a troop of baboons in the pass of Mensa in Abyssinia. The baboons in +return rolled so many stones down the mountain, some as large as a man's +head, that the attackers had to beat a hasty retreat; and the pass was +actually for a time closed against the caravan. It deserves notice that +these baboons thus acted in concert. Mr. Wallace[70] on three occasions +saw female orangs, accompanied by their young, "breaking off branches +and the great spiny fruit of the Durian tree, with every appearance of +rage; causing such a shower of missiles as effectually kept us from +approaching too near the tree." + +In the Zoological Gardens a monkey which had weak teeth used to break +open nuts with a stone; and I was assured by the keepers that this +animal, after using the stone, hid it in the straw, and would not let +any other monkey touch it. Here, then, we have the idea of property; but +this idea is common to every dog with a bone, and to most or all birds +with their nests. + +The Duke of Argyll[71] remarks, that the fashioning of an implement for +a special purpose is absolutely peculiar to man; and he considers that +this forms an immeasurable gulf between him and the brutes. It is no +doubt a very important distinction, but there appears to me much truth +in Sir J. Lubbock's suggestion,[72] that when primeval man first used +flint-stones for any purpose, he would have accidentally splintered +them, and would then have used the sharp fragments. From this step it +would be a small one to intentionally break the flints, and not a very +wide step to rudely fashion them. This latter advance, however, may have +taken long ages, if we may judge by the immense interval of time which +elapsed before the men of the neolithic period took to grinding and +polishing their stone tools. In breaking the flints, as Sir J. Lubbock +likewise remarks, sparks would have been emitted, and in grinding them +heat would have been evolved: "thus the two usual methods of obtaining +fire may have originated." The nature of fire would have been known in +the many volcanic regions where lava occasionally flows through forests. +The anthropomorphous apes, guided probably by instinct, build for +themselves temporary platforms; but as many instincts are largely +controlled by reason, the simpler ones, such as this of building a +platform, might readily pass into a voluntary and conscious act. The +orang is known to cover itself at night with the leaves of the Pandanus; +and Brehm states that one of his baboons used to protect itself from the +heat of the sun by throwing a straw-mat over its head. In these latter +habits, we probably see the first steps towards some of the simpler +arts; namely rude architecture and dress, as they arose amongst the +early progenitors of man. + + +_Language._--This faculty has justly been considered as one of the chief +distinctions between man and the lower animals. But man, as a highly +competent judge, Archbishop Whately remarks, "is not the only animal +that can make use of language to express what is passing in his mind, +and can understand, more or less, what is so expressed by another."[73] +In Paraguay the _Cebus Azarae_ when excited utters at least six distinct +sounds, which excite in other monkeys similar emotions.[74] The +movements of the features and gestures of monkeys are understood by us, +and they partly understand ours, as Rengger and others declare. It is a +more remarkable fact that the dog, since being domesticated, has learnt +to bark[75] in at least four or five distinct tones. Although barking is +a new art, no doubt the wild species, the parents of the dog, expressed +their feelings by cries of various kinds. With the domesticated dog we +have the bark of eagerness, as in the chase; that of anger; the yelping +or howling bark of despair, as when shut up; that of joy, as when +starting on a walk with his master; and the very distinct one of demand +or supplication, as when wishing for a door or window to be opened. + +Articulate language is, however, peculiar to man; but he uses in common +with the lower animals inarticulate cries to express his meaning, aided +by gestures and the movements of the muscles of the face.[76] This +especially holds good with the more simple and vivid feelings, which are +but little connected with our higher intelligence. Our cries of pain, +fear, surprise, anger, together with their appropriate actions, and the +murmur of a mother to her beloved child, are more expressive than any +words. It is not the mere power of articulation that distinguishes man +from other animals, for as every one knows, parrots can talk; but it is +his large power of connecting definite sounds with definite ideas; and +this obviously depends on the development of the mental faculties. + +As Horne Tooke, one of the founders of the noble science of philology, +observes, language is an art, like brewing or baking; but writing would +have been a much more appropriate simile. It certainly is not a true +instinct, as every language has to be learnt. It differs, however, +widely from all ordinary arts, for man has an instinctive tendency to +speak, as we see in the babble of our young children; whilst no child +has an instinctive tendency to brew, bake, or write. Moreover, no +philologist now supposes that any language has been deliberately +invented; each has been slowly and unconsciously developed by many +steps. The sounds uttered by birds offer in several respects the nearest +analogy to language, for all the members of the same species utter the +same instinctive cries expressive of their emotions; and all the kinds +that have the power of singing exert this power instinctively; but the +actual song, and even the call-notes, are learnt from their parents or +foster-parents. These sounds, as Daines Barrington[77] has proved, "are +no more innate than language is in man." The first attempts to sing "may +be compared to the imperfect endeavour in a child to babble." The young +males continue practising, or, as the bird-catchers say, recording, for +ten or eleven months. Their first essays show hardly a rudiment of the +future song; but as they grow older we can perceive what they are aiming +at; and at last they are said "to sing their song round." Nestlings +which have learnt the song of a distinct species, as with the +canary-birds educated in the Tyrol, teach and transmit their new song to +their offspring. The slight natural differences of song in the same +species inhabiting different districts may be appositely compared, as +Barrington remarks, "to provincial dialects;" and the songs of allied, +though distinct species may be compared with the languages of distinct +races of man. I have given the foregoing details to shew that an +instinctive tendency to acquire an art is not a peculiarity confined to +man. + +With respect to the origin of articulate language, after having read on +the one side the highly interesting works of Mr. Hensleigh Wedgwood, the +Rev. F. Farrar, and Prof. Schleicher,[78] and the celebrated lectures of +Prof. Max Mueller on the other side, I cannot doubt that language owes +its origin to the imitation and modification, aided by signs and +gestures, of various natural sounds, the voices of other animals, and +man's own instinctive cries. When we treat of sexual selection we shall +see that primeval man, or rather some early progenitor of man, probably +used his voice largely, as does one of the gibbon-apes at the present +day, in producing true musical cadences, that is in singing; we may +conclude from a widely-spread analogy that this power would have been +especially exerted during the courtship of the sexes, serving to express +various emotions, as love, jealousy, triumph, and serving as a challenge +to their rivals. The imitation by articulate sounds of musical cries +might have given rise to words expressive of various complex emotions. +As bearing on the subject of imitation, the strong tendency in our +nearest allies, the monkeys, in microcephalous idiots,[79] and in the +barbarous races of mankind, to imitate whatever they hear deserves +notice. As monkeys certainly understand much that is said to them by +man, and as in a state of nature they utter signal-cries of danger to +their fellows,[80] it does not appear altogether incredible, that some +unusually wise ape-like animal should have thought of imitating the +growl of a beast of prey, so as to indicate to his fellow monkeys the +nature of the expected danger. And this would have been a first step in +the formation of a language. + +As the voice was used more and more, the vocal organs would have been +strengthened and perfected through the principle of the inherited +effects of use; and this would have reacted on the power of speech. But +the relation between the continued use of language and the development +of the brain has no doubt been far more important. The mental powers in +some early progenitor of man must have been more highly developed than +in any existing ape, before even the most imperfect form of speech could +have come into use; but we may confidently believe that the continued +use and advancement of this power would have reacted on the mind by +enabling and encouraging it to carry on long trains of thought. A long +and complex train of thought can no more be carried on without the aid +of words, whether spoken or silent, than a long calculation without the +use of figures or algebra. It appears, also, that even ordinary trains +of thought almost require some form of language, for the dumb, deaf, and +blind girl, Laura Bridgman, was observed to use her fingers whilst +dreaming.[81] Nevertheless a long succession of vivid and connected +ideas, may pass through the mind without the aid of any form of +language, as we may infer from the prolonged dreams of dogs. We have, +also, seen that retriever-dogs are able to reason to a certain extent; +and this they manifestly do without the aid of language. The intimate +connection between the brain, as it is now developed in us, and the +faculty of speech, is well shewn by those curious cases of +brain-disease, in which speech is specially affected, as when the power +to remember substantives is lost, whilst other words can be correctly +used.[82] There is no more improbability in the effects of the continued +use of the vocal and mental organs being inherited, than in the case of +handwriting, which depends partly on the structure of the hand and +partly on the disposition of the mind; and handwriting is certainly +inherited.[83] + +Why the organs now used for speech should have been originally perfected +for this purpose, rather than any other organs, it is not difficult to +see. Ants have considerable powers of intercommunication by means of +their antennae, as shewn by Huber, who devotes a whole chapter to their +language. We might have used our fingers as efficient instruments, for a +person with practice can report to a deaf man every word of a speech +rapidly delivered at a public meeting; but the loss of our hands, whilst +thus employed, would have been a serious inconvenience. As all the +higher mammals possess vocal organs constructed on the same general +plan with ours, and which are used as a means of communication, it was +obviously probable, if the power of communication had to be improved, +that these same organs would have been still further developed; and this +has been effected by the aid of adjoining and well-adapted parts, namely +the tongue and lips.[84] The fact of the higher apes not using their +vocal organs for speech, no doubt depends on their intelligence not +having been sufficiently advanced. The possession by them of organs, +which with long-continued practice might have been used for speech, +although not thus used, is paralleled by the case of many birds which +possess organs fitted for singing, though they never sing. Thus, the +nightingale and crow have vocal organs similarly constructed, these +being used by the former for diversified song, and by the latter merely +for croaking.[85] + +The formation of different languages and of distinct species, and the +proofs that both have been developed through a gradual process, are +curiously the same.[86] But we can trace the origin of many words +further back than in the case of species, for we can perceive that they +have arisen from the imitation of various sounds, as in alliterative +poetry. We find in distinct languages striking homologies due to +community of descent, and analogies due to a similar process of +formation. The manner in which certain letters or sounds change when +others change is very like correlated growth. We have in both cases the +reduplication of parts, the effects of long-continued use, and so forth. +The frequent presence of rudiments, both in languages and in species, is +still more remarkable. The letter _m_ in the word _am_, means _I_; so +that in the expression _I am_, a superfluous and useless rudiment has +been retained. In the spelling also of words, letters often remain as +the rudiments of ancient forms of pronunciation. Languages, like organic +beings, can be classed in groups under groups; and they can be classed +either naturally according to descent, or artificially by other +characters. Dominant languages and dialects spread widely and lead to +the gradual extinction of other tongues. A language, like a species, +when once extinct, never, as Sir C. Lyell remarks, reappears. The same +language never has two birthplaces. Distinct languages may be crossed +or blended together.[87] We see variability in every tongue, and new +words are continually cropping up; but as there is a limit to the powers +of the memory, single words, like whole languages, gradually become +extinct. As Max Mueller[88] has well remarked:--"A struggle for life is +constantly going on amongst the words and grammatical forms in each +language. The better, the shorter, the easier forms are constantly +gaining the upper hand, and they owe their success to their own inherent +virtue." To these more important causes of the survival of certain +words, mere novelty may, I think, be added; for there is in the mind of +man a strong love for slight changes in all things. The survival or +preservation of certain favoured words in the struggle for existence is +natural selection. + +The perfectly regular and wonderfully complex construction of the +languages of many barbarous nations has often been advanced as a proof, +either of the divine origin of these languages, or of the high art and +former civilisation of their founders. Thus F. von Schlegel writes: "In +those languages which appear to be at the lowest grade of intellectual +culture, we frequently observe a very high and elaborate degree of art +in their grammatical structure. This is especially the case with the +Basque and the Lapponian, and many of the American languages."[89] But +it is assuredly an error to speak of any language as an art in the sense +of its having been elaborately and methodically formed. Philologists now +admit that conjugations, declensions, &c., originally existed as +distinct words, since joined together; and as such words express the +most obvious relations between objects and persons, it is not surprising +that they should have been used by the men of most races during the +earliest ages. With respect to perfection, the following illustration +will best shew how easily we may err: a Crinoid sometimes consists of no +less than 150,000 pieces of shell,[90] all arranged with perfect +symmetry in radiating lines; but a naturalist does not consider an +animal of this kind as more perfect than a bilateral one with +comparatively few parts, and with none of these alike, excepting on the +opposite sides of the body. He justly considers the differentiation and +specialisation of organs as the test of perfection. So with languages, +the most symmetrical and complex ought not to be ranked above irregular, +abbreviated, and bastardised languages, which have borrowed expressive +words and useful forms of construction from various conquering, or +conquered, or immigrant races. + +From these few and imperfect remarks I conclude that the extremely +complex and regular construction of many barbarous languages, is no +proof that they owe their origin to a special act of creation.[91] Nor, +as we have seen, does the faculty of articulate speech in itself offer +any insuperable objection to the belief that man has been developed from +some lower form. + + +_Self-consciousness, Individuality, Abstraction, General Ideas, &c._--It +would be useless to attempt discussing these high faculties, which, +according to several recent writers, make the sole and complete +distinction between man and the brutes, for hardly two authors agree in +their definitions. Such faculties could not have been fully developed in +man until his mental powers had advanced to a high standard, and this +implies the use of a perfect language. No one supposes that one of the +lower animals reflects whence he comes or whither he goes,--what is +death or what is life, and so forth. But can we feel sure that an old +dog with an excellent memory and some power of imagination, as shewn by +his dreams, never reflects on his past pleasures in the chase? and this +would be a form of self-consciousness. On the other hand, as Buechner[92] +has remarked, how little can the hard-worked wife of a degraded +Australian savage, who uses hardly any abstract words and cannot count +above four, exert her self-consciousness, or reflect on the nature of +her own existence. + +That animals retain their mental individuality is unquestionable. When +my voice awakened a train of old associations in the mind of the +above-mentioned dog, he must have retained his mental individuality, +although every atom of his brain had probably undergone change more than +once during the interval of five years. This dog might have brought +forward the argument lately advanced to crush all evolutionists, and +said, "I abide amid all mental moods and all material changes.... The +teaching that atoms leave their impressions as legacies to other atoms +falling into the places they have vacated is contradictory of the +utterance of consciousness, and is therefore false; but it is the +teaching necessitated by evolutionism, consequently the hypothesis is a +false one."[93] + + +_Sense of Beauty._--This sense has been declared to be peculiar to man. +But when we behold male birds elaborately displaying their plumes and +splendid colours before the females, whilst other birds not thus +decorated make no such display, it is impossible to doubt that the +females admire the beauty of their male partners. As women everywhere +deck themselves with these plumes, the beauty of such ornaments cannot +be disputed. The Bower-birds by tastefully ornamenting their +playing-passages with gaily-coloured objects, as do certain +humming-birds their nests, offer additional evidence that they possess a +sense of beauty. So with the song of birds, the sweet strains poured +forth by the males during the season of love are certainly admired by +the females, of which fact evidence will hereafter be given. If female +birds had been incapable of appreciating the beautiful colours, the +ornaments, and voices of their male partners, all the labour and +anxiety exhibited by them in displaying their charms before the females +would have been thrown away; and this it is impossible to admit. Why +certain bright colours and certain sounds should excite pleasure, when +in harmony, cannot, I presume, be explained any more than why certain +flavours and scents are agreeable; but assuredly the same colours and +the same sounds are admired by us and by many of the lower animals. + +The taste for the beautiful, at least as far as female beauty is +concerned, is not of a special nature in the human mind; for it differs +widely in the different races of man, as will hereafter be shewn, and is +not quite the same even in the different nations of the same race. +Judging from the hideous ornaments and the equally hideous music admired +by most savages, it might be urged that their aesthetic faculty was not +so highly developed as in certain animals, for instance, in birds. +Obviously no animal would be capable of admiring such scenes as the +heavens at night, a beautiful landscape, or refined music; but such high +tastes, depending as they do on culture and complex associations, are +not enjoyed by barbarians or by uneducated persons. + +Many of the faculties, which have been of inestimable service to man for +his progressive advancement, such as the powers of the imagination, +wonder, curiosity, an undefined sense of beauty, a tendency to +imitation, and the love of excitement or novelty, could not fail to have +led to the most capricious changes of customs and fashions. I have +alluded to this point, because a recent writer[94] has oddly fixed on +Caprice "as one of the most remarkable and typical differences between +savages and brutes." But not only can we perceive how it is that roan is +capricious, but the lower animals are, as we shall hereafter see, +capricious in their affections, aversions, and sense of beauty. There is +also good reason to suspect that they love novelty, for its own sake. + + +_Belief in God--Religion._--There is no evidence that man was +aboriginally endowed with the ennobling belief in the existence of an +Omnipotent God. On the contrary there is ample evidence, derived not +from hasty travellers, but from men who have long resided with savages, +that numerous races have existed and still exist, who have no idea of +one or more gods, and who have no words in their languages to express +such an idea.[95] The question is of course wholly distinct from that +higher one, whether there exists a Creator and Ruler of the universe; +and this has been answered in the affirmative by the highest intellects +that have ever lived. + +If, however, we include under the term "religion" the belief in unseen +or spiritual agencies, the case is wholly different; for this belief +seems to be almost universal with the less civilised races. Nor is it +difficult to comprehend how it arose. As soon as the important faculties +of the imagination, wonder, and curiosity, together with some power of +reasoning, had become partially developed, man would naturally have +craved to understand what was passing around him, and have vaguely +speculated on his own existence. As Mr. M'Lennan[96] has remarked, +"Some explanation of the phenomena of life, a man must feign for +himself; and to judge from the universality of it, the simplest +hypothesis, and the first to occur to men, seems to have been that +natural phenomena are ascribable to the presence in animals, plants, and +things, and in the forces of nature, of such spirits prompting to action +as men are conscious they themselves possess." It is probable, as Mr. +Tylor has clearly shewn, that dreams may have first given rise to the +notion of spirits; for savages do not readily distinguish between +subjective and objective impressions. When a savage dreams, the figures +which appear before him are believed to have come from a distance and to +stand over him; or "the soul of the dreamer goes out on its travels, and +comes home with a remembrance of what it has seen."[97] But until the +above-named faculties of imagination, curiosity, reason, &c., had been +fairly well developed in the mind of man, his dreams would not have led +him to believe in spirits, any more than in the case of a dog. + +The tendency in savages to imagine that natural objects and agencies +are animated by spiritual or living essences, is perhaps illustrated by +a little fact which I once noticed: my dog, a full-grown and very +sensible animal, was lying on the lawn during a hot and still day; but +at a little distance a slight breeze occasionally moved an open parasol, +which would have been wholly disregarded by the dog, had any one stood +near it. As it was, every time that the parasol slightly moved, the dog +growled fiercely and barked. He must, I think, have reasoned to himself +in a rapid and unconscious manner, that movement without any apparent +cause indicated the presence of some strange living agent, and no +stranger had a right to be on his territory. + +The belief in spiritual agencies would easily pass into the belief in +the existence of one or more gods. For savages would naturally attribute +to spirits the same passions, the same love of vengeance or simplest +form of justice, and the same affections which they themselves +experienced. The Fuegians appear to be in this respect in an +intermediate condition, for when the surgeon on board the "Beagle" shot +some young ducklings as specimens, York Minster declared in the most +solemn manner, "Oh! Mr. Bynoe, much rain, much snow, blow much;" and +this was evidently a retributive punishment for wasting human food. So +again he related how, when his brother killed a "wild man," storms long +raged, much rain and snow fell. Yet we could never discover that the +Fuegians believed in what we should call a God, or practised any +religious rites; and Jemmy Button, with justifiable pride, stoutly +maintained that there was no devil in his land. This latter assertion is +the more remarkable, as with savages the belief in bad spirits is far +more common than the belief in good spirits. + +The feeling of religious devotion is a highly complex one, consisting of +love, complete submission to an exalted and mysterious superior, a +strong sense of dependence,[98] fear, reverence, gratitude, hope for the +future, and perhaps other elements. No being could experience so complex +an emotion until advanced in his intellectual and moral faculties to at +least a moderately high level. Nevertheless we see some distant approach +to this state of mind, in the deep love of a dog for his master, +associated with complete submission, some fear, and perhaps other +feelings. The behaviour of a dog when returning to his master after an +absence, and, as I may add, of a monkey to his beloved keeper, is widely +different from that towards their fellows. In the latter case the +transports of joy appear to be somewhat less, and the sense of equality +is shewn in every action. Professor Braubach[99] goes so far as to +maintain that a dog looks on his master as on a god. + +The same high mental faculties which first led man to believe in unseen +spiritual agencies, then in fetishism, polytheism, and ultimately in +monotheism, would infallibly lead him, as long as his reasoning powers +remained poorly developed, to various strange superstitions and customs. +Many of these are terrible to think of--such as the sacrifice of human +beings to a blood-loving god; the trial of innocent persons by the +ordeal of poison or fire; witchcraft, &c.--yet it is well occasionally +to reflect on these superstitions, for they shew us what an infinite +debt of gratitude we owe to the improvement of our reason, to science, +and our accumulated knowledge.[100] As Sir J. Lubbock has well +observed, "it is not too much to say that the horrible dread of unknown +evil hangs like a thick cloud over savage life, and embitters every +pleasure." These miserable and indirect consequences of our highest +faculties may be compared with the incidental and occasional mistakes of +the instincts of the lower animals. + + + + +CHAPTER III. + +COMPARISON OF THE MENTAL POWERS OF MAN AND THE LOWER +ANIMALS--_continued_. + + + The moral sense--Fundamental proposition--The qualities of + social animals--Origin of sociability--Struggle between opposed + instincts--Man a social animal--The more enduring social + instincts conquer other less persistent instincts--The social + virtues alone regarded by savages--The self-regarding virtues + acquired at a later stage of development--The importance of the + judgment of the members of the same community on + conduct--Transmission of moral tendencies--Summary. + + +I fully subscribe to the judgment of those writers[101] who maintain +that of all the differences between man and the lower animals, the moral +sense or conscience is by far the most important. This sense, as +Mackintosh[102] remarks, "has a rightful supremacy over every other +principle of human action;" it is summed up in that short but imperious +word _ought_, so full of high significance. It is the most noble of all +the attributes of man, leading him without a moment's hesitation to risk +his life for that of a fellow-creature; or after due deliberation, +impelled simply by the deep feeling of right or duty, to sacrifice it in +some great cause. Immanuel Kant exclaims, "Duty! Wondrous thought, that +workest neither by fond insinuation, flattery, nor by any threat, but +merely by holding up thy naked law in the soul, and so extorting for +thyself always reverence, if not always obedience; before whom all +appetites are dumb, however secretly they rebel; whence thy +original?"[103] + +This great question has been discussed by many writers[104] of +consummate ability; and my sole excuse for touching on it is the +impossibility of here passing it over, and because, as far as I know, no +one has approached it exclusively from the side of natural history. The +investigation possesses, also, some independent interest, as an attempt +to see how far the study of the lower animals can throw light on one of +the highest psychical faculties of man. + +The following proposition seems to me in a high degree probable--namely, +that any animal whatever, endowed with well-marked social +instincts,[105] would inevitably acquire a moral sense or conscience, as +soon as its intellectual powers had become as well developed, or nearly +as well developed, as in man. For, _firstly_, the social instincts lead +an animal to take pleasure in the society of its fellows, to feel a +certain amount of sympathy with them, and to perform various services +for them. The services may be of a definite and evidently instinctive +nature; or there may be only a wish and readiness, as with most of the +higher social animals, to aid their fellows in certain general ways. But +these feelings and services are by no means extended to all the +individuals of the same species, only to those of the same association. +_Secondly_, as soon as the mental faculties had become highly developed, +images of all past actions and motives would be incessantly passing +through the brain of each individual; and that feeling of +dissatisfaction which invariably results, as we shall hereafter see, +from any unsatisfied instinct, would arise, as often as it was perceived +that the enduring and always present social instinct had yielded to some +other instinct, at the time stronger, but neither enduring in its +nature, nor leaving behind it a very vivid impression. It is clear that +many instinctive desires, such as that of hunger, are in their nature of +short duration; and after being satisfied are not readily or vividly +recalled. _Thirdly_, after the power of language had been acquired and +the wishes of the members of the same community could be distinctly +expressed, the common opinion how each member ought to act for the +public good, would naturally become to a large extent the guide to +action. But the social instincts would still give the impulse to act for +the good of the community, this impulse being strengthened, directed, +and sometimes even deflected by public opinion, the power of which +rests, as we shall presently see, on instinctive sympathy. _Lastly_, +habit in the individual would ultimately play a very important part in +guiding the conduct of each member; for the social instincts and +impulses, like all other instincts, would be greatly strengthened by +habit, as would obedience to the wishes and judgment of the community. +These several subordinate propositions must now be discussed; and some +of them at considerable length. + +It may be well first to premise that I do not wish to maintain that any +strictly social animal, if its intellectual faculties were to become as +active and as highly developed as in man, would acquire exactly the same +moral sense as ours. In the same manner as various animals have some +sense of beauty, though they admire widely different objects, so they +might have a sense of right and wrong, though led by it to follow widely +different lines of conduct. If, for instance, to take an extreme case, +men were reared under precisely the same conditions as hive-bees, there +can hardly be a doubt that our unmarried females would, like the +worker-bees, think it a sacred duty to kill their brothers, and mothers +would strive to kill their fertile daughters; and no one would think of +interfering. Nevertheless the bee, or any other social animal, would in +our supposed case gain, as it appears to me, some feeling of right and +wrong, or a conscience. For each individual would have an inward sense +of possessing certain stronger or more enduring instincts, and others +less strong or enduring; so that there would often be a struggle which +impulse should be followed; and satisfaction or dissatisfaction would be +felt, as past impressions were compared during their incessant passage +through the mind. In this case an inward monitor would tell the animal +that it would have been better to have followed the one impulse rather +than the other. The one course ought to have been followed: the one +would have been right and the other wrong; but to these terms I shall +have to recur. + + +_Sociability._--Animals of many kinds are social; we find even distinct +species living together, as with some American monkeys, and with the +united flocks of rooks, jackdaws, and starlings. Man shows the same +feeling in his strong love for the dog, which the dog returns with +interest. Every one must have noticed how miserable horses, dogs, sheep, +&c. are when separated from their companions; and what affection at +least the two former kinds show on their reunion. It is curious to +speculate on the feelings of a dog, who will rest peacefully for hours +in a room with his master or any of the family, without the least notice +being taken of him; but if left for a short time by himself, barks or +howls dismally. We will confine our attention to the higher social +animals, excluding insects, although these aid each other in many +important ways. The most common service which the higher animals perform +for each other, is the warning each other of danger by means of the +united senses of all. Every sportsman knows, as Dr. Jaeger remarks,[106] +how difficult it is to approach animals in a herd or troop. Wild horses +and cattle do not, I believe, make any danger-signal; but the attitude +of any one who first discovers an enemy, warns the others. Rabbits stamp +loudly on the ground with their hind-feet as a signal: sheep and chamois +do the same, but with their fore-feet, uttering likewise a whistle. Many +birds and some mammals post sentinels, which in the case of seals are +said[107] generally to be the females. The leader of a troop of monkeys +acts as the sentinel, and utters cries expressive both of danger and of +safety.[108] Social animals perform many little services for each +other: horses nibble, and cows lick each other, on any spot which +itches: monkeys search for each other's external parasites; and Brehm +states that after a troop of the _Cercopithecus griseo-viridis_ has +rushed through a thorny brake, each monkey stretches itself on a branch, +and another monkey sitting by "conscientiously" examines its fur and +extracts every thorn or burr. + +Animals also render more important services to each other: thus wolves +and some other beasts of prey hunt in packs, and aid each other in +attacking their victims. Pelicans fish in concert. The Hamadryas baboons +turn over stones to find insects, &c.; and when they come to a large +one, as many as can stand round, turn it over together and share the +booty. Social animals mutually defend each other. The males of some +ruminants come to the front when there is danger and defend the herd +with their horns. I shall also in a future chapter give cases of two +young wild bulls attacking an old one in concert, and of two stallions +together trying to drive away a third stallion from a troop of mares. +Brehm encountered in Abyssinia a great troop of baboons which were +crossing a valley: some had already ascended the opposite mountain, and +some were still in the valley: the latter were attacked by the dogs, but +the old males immediately hurried down from the rocks, and with mouths +widely opened roared so fearfully, that the dogs precipitately +retreated. They were again encouraged to the attack; but by this time +all the baboons had reascended the heights, excepting a young one, about +six months old, who, loudly calling for aid, climbed on a block of rock +and was surrounded. Now one of the largest males, a true hero, came down +again from the mountain, slowly went to the young one, coaxed him, and +triumphantly led him away--the dogs being too much astonished to make an +attack. I cannot resist giving another scene which was witnessed by this +same naturalist; an eagle seized a young Cercopithecus, which, by +clinging to a branch, was not at once carried off; it cried loudly for +assistance, upon which the other members of the troop with much uproar +rushed to the rescue, surrounded the eagle, and pulled out so many +feathers, that he no longer thought of his prey, but only how to escape. +This eagle, as Brehm remarks, assuredly would never again attack a +monkey in a troop. + +It is certain that associated animals have a feeling of love for each +other which is not felt by adult and non-social animals. How far in most +cases they actually sympathise with each other's pains and pleasures is +more doubtful, especially with respect to the latter. Mr. Buxton, +however, who had excellent means of observation,[109] states that his +macaws, which lived free in Norfolk, took "an extravagant interest" in a +pair with a nest, and whenever the female left it, she was surrounded by +a troop "screaming horrible acclamations in her honour." It is often +difficult to judge whether animals have any feeling for each other's +sufferings. Who can say what cows feel, when they surround and stare +intently on a dying or dead companion? That animals sometimes are far +from feeling any sympathy is too certain; for they will expel a wounded +animal from the herd, or gore or worry it to death. This is almost the +blackest fact in natural history, unless indeed the explanation which +has been suggested is true, that their instinct or reason leads them to +expel an injured companion, lest beasts of prey, including man, should +be tempted to follow the troop. In this case their conduct is not much +worse than that of the North American Indians who leave their feeble +comrades to perish on the plains, or the Feegeans, who, when their +parents get old or fall ill, bury them alive.[110] + +Many animals, however, certainly sympathise with each other's distress +or danger. This is the case even with birds; Capt. Stansbury[111] found +on a salt lake in Utah an old and completely blind pelican, which was +very fat, and must have been long and well fed by his companions. Mr. +Blyth, as he informs me, saw Indian crows feeding two or three of their +companions which were blind; and I have heard of an analogous case with +the domestic cock. We may, if we choose, call these actions instinctive; +but such cases are much too rare for the development of any special +instinct.[112] I have myself seen a dog, who never passed a great friend +of his, a cat which lay sick in a basket, without giving her a few licks +with his tongue, the surest sign of kind feeling in a dog. + +It must be called sympathy that leads a courageous dog to fly at any one +who strikes his master, as he certainly will. I saw a person pretending +to beat a lady who had a very timid little dog on her lap, and the trial +had never before been made. The little creature instantly jumped away, +but after the pretended beating was over, it was really pathetic to see +how perseveringly he tried to lick his mistress's face and comfort her. +Brehm[113] states that when a baboon in confinement was pursued to be +punished, the others tried to protect him. It must have been sympathy in +the cases above given which led the baboons and Cercopitheci to defend +their young comrades from the dogs and the eagle. I will give only one +other instance of sympathetic and heroic conduct in a little American +monkey. Several years ago a keeper at the Zoological Gardens, showed me +some deep and scarcely healed wounds on the nape of his neck, inflicted +on him whilst kneeling on the floor by a fierce baboon. The little +American monkey, who was a warm friend of this keeper, lived in the same +large compartment, and was dreadfully afraid of the great baboon. +Nevertheless, as soon as he saw his friend the keeper in peril, he +rushed to the rescue, and by screams and bites so distracted the baboon +that the man was able to escape, after running great risk, as the +surgeon who attended him thought, of his life. + +Besides love and sympathy, animals exhibit other qualities which in us +would be called moral; and I agree with Agassiz[114] that dogs possess +something very like a conscience. They certainly possess some power of +self-command, and this does not appear to be wholly the result of fear. +As Braubach[115] remarks, a dog will refrain from stealing food in the +absence of his master. Dogs have long been accepted as the very type of +fidelity and obedience. All animals living in a body which defend each +other or attack their enemies in concert, must be in some degree +faithful to each other; and those that follow a leader must be in some +degree obedient. When the baboons in Abyssinia[116] plunder a garden, +they silently follow their leader; and if an imprudent young animal +makes a noise, he receives a slap from the others to teach him silence +and obedience; but as soon as they are sure that there is no danger, all +show their joy by much clamour. + +With respect to the impulse which leads certain animals to associate +together, and to aid each other in many ways, we may infer that in most +cases they are impelled by the same sense of satisfaction or pleasure +which they experience in performing other instinctive actions; or by the +same sense of dissatisfaction, as in other cases of prevented +instinctive actions. We see this in innumerable instances, and it is +illustrated in a striking manner by the acquired instincts of our +domesticated animals; thus a young shepherd-dog delights in driving and +running round a flock of sheep, but not in worrying them; a young +foxhound delights in hunting a fox, whilst some other kinds of dogs as I +have witnessed, utterly disregard foxes. What a strong feeling of inward +satisfaction must impel a bird, so full of activity, to brood day after +day over her eggs. Migratory birds are miserable if prevented from +migrating, and perhaps they enjoy starting on their long flight. Some +few instincts are determined solely by painful feelings, as by fear, +which leads to self-preservation, or is specially directed against +certain enemies. No one, I presume, can analyse the sensations of +pleasure or pain. In many cases, however, it is probable that instincts +are persistently followed from the mere force of inheritance, without +the stimulus of either pleasure or pain. A young pointer, when it first +scents game, apparently cannot help pointing. A squirrel in a cage who +pats the nuts which it cannot eat, as if to bury them in the ground, can +hardly be thought to act thus either from pleasure or pain. Hence the +common assumption that men must be impelled to every action by +experiencing some pleasure or pain may be erroneous. Although a habit +may be blindly and implicitly followed, independently of any pleasure or +pain felt at the moment, yet if it be forcibly and abruptly checked, a +vague sense of dissatisfaction is generally experienced; and this is +especially true in regard to persons of feeble intellect. + +It has often been assumed that animals were in the first place rendered +social, and that they feel as a consequence uncomfortable when separated +from each other, and comfortable whilst together; but it is a more +probable view that these sensations were first developed, in order that +those animals which would profit by living in society, should be induced +to live together. In the same manner as the sense of hunger and the +pleasure of eating were, no doubt, first acquired in order to induce +animals to eat. The feeling of pleasure from society is probably an +extension of the parental or filial affections; and this extension may +be in chief part attributed to natural selection, but perhaps in part to +mere habit. For with those animals which were benefited by living in +close association, the individuals which took the greatest pleasure in +society would best escape various dangers; whilst those that cared least +for their comrades and lived solitary would perish in greater numbers. +With respect to the origin of the parental and filial affections, which +apparently lie at the basis of the social affections, it is hopeless to +speculate; but we may infer that they have been to a large extent +gained through natural selection. So it has almost certainly been with +the unusual and opposite feeling of hatred between the nearest +relations, as with the worker-bees which kill their brother-drones, and +with the queen-bees which kill their daughter-queens; the desire to +destroy, instead of loving, their nearest relations having been here of +service to the community. + +The all-important emotion of sympathy is distinct from that of love. A +mother may passionately love her sleeping and passive infant, but she +can then hardly be said to feel sympathy for it. The love of a man for +his dog is distinct from sympathy, and so is that of a dog for his +master. Adam Smith formerly argued, as has Mr. Bain recently, that the +basis of sympathy lies in our strong retentiveness of former states of +pain or pleasure. Hence, "the sight of another person enduring hunger, +cold, fatigue, revives in us some recollection of these states, which +are painful even in idea." We are thus impelled to relieve the +sufferings of another, in order that our own painful feelings may be at +the same time relieved. In like manner we are led to participate in the +pleasures of others.[117] But I cannot see how this view explains the +fact that sympathy is excited in an immeasurably stronger degree by a +beloved than by an indifferent person. The mere sight of suffering, +independently of love, would suffice to call up in us vivid +recollections and associations. Sympathy may at first have originated in +the manner above suggested; but it seems now to have become an instinct, +which is especially directed towards beloved objects, in the same manner +as fear with animals is especially directed against certain enemies. As +sympathy is thus directed, the mutual love of the members of the same +community will extend its limits. No doubt a tiger or lion feels +sympathy for the sufferings of its own young, but not for any other +animal. With strictly social animals the feeling will be more or less +extended to all the associated members, as we know to be the case. With +mankind selfishness, experience, and imitation probably add, as Mr. Bain +has shewn, to the power of sympathy; for we are led by the hope of +receiving good in return to perform acts of sympathetic kindness to +others; and there can be no doubt that the feeling of sympathy is much +strengthened by habit. In however complex a manner this feeling may have +originated, as it is one of high importance to all those animals which +aid and defend each other, it will have been increased, through natural +selection; for those communities, which included the greatest number of +the most sympathetic members, would flourish best and rear the greatest +number of offspring. + +In many cases it is impossible to decide whether certain social +instincts have been acquired through natural selection, or are the +indirect result of other instincts and faculties, such as sympathy, +reason, experience, and a tendency to imitation; or again, whether they +are simply the result of long-continued habit. So remarkable an instinct +as the placing sentinels to warn the community of danger, can hardly +have been the indirect result of any other faculty; it must therefore +have been directly acquired. On the other hand, the habit followed by +the males of some social animals, of defending the community and of +attacking their enemies or their prey in concert, may perhaps have +originated from mutual sympathy; but courage, and in most cases +strength, must have been previously acquired, probably through natural +selection. + +Of the various instincts and habits, some are much stronger than others, +that is, some either give more pleasure in their performance and more +distress in their prevention than others; or, which is probably quite as +important, they are more persistently followed through inheritance +without exciting any special feeling of pleasure or pain. We are +ourselves conscious that some habits are much more difficult to cure or +change than others. Hence a struggle may often be observed in animals +between different instincts, or between an instinct and some habitual +disposition; as when a dog rushes after a hare, is rebuked, pauses, +hesitates, pursues again or returns ashamed to his master; or as between +the love of a female dog for her young puppies and for her master, for +she may be seen to slink away to them, as if half ashamed of not +accompanying her master. But the most curious instance known to me of +one instinct conquering another, is the migratory instinct conquering +the maternal instinct. The former is wonderfully strong; a confined bird +will at the proper season beat her breast against the wires of her cage, +until it is bare and bloody. It causes young salmon to leap out of the +fresh water, where they could still continue to live, and thus +unintentionally to commit suicide. Every one knows how strong the +maternal instinct is, leading even timid birds to face great danger, +though with hesitation and in opposition to the instinct of +self-preservation. Nevertheless the migratory instinct is so powerful +that late in the autumn swallows and house-martins frequently desert +their tender young, leaving them to perish miserably in their +nests.[118] + +We can perceive that an instinctive impulse, if it be in any way more +beneficial to a species than some other or opposed instinct, would be +rendered the more potent of the two through natural selection; for the +individuals which had it most strongly developed would survive in larger +numbers. Whether this is the case with the migratory in comparison with +the maternal instinct, may well be doubted. The great persistence or +steady action of the former at certain seasons of the year during the +whole day, may give it for a time paramount force. + + +_Man a social animal._--Most persons admit that man is a social being. +We see this in his dislike of solitude, and in his wish for society +beyond that of his own family. Solitary confinement is one of the +severest punishments which can be inflicted. Some authors suppose that +man primevally lived in single families; but at the present day, though +single families, or only two or three together, roam the solitudes of +some savage lands, they are always, as far as I can discover, friendly +with other families inhabiting the same district. Such families +occasionally meet in council, and they unite for their common defence. +It is no argument against savage man being a social animal, that the +tribes inhabiting adjacent districts are almost always at war with each +other; for the social instincts never extend to all the individuals of +the same species. Judging from the analogy of the greater number of the +Quadrumana, it is probable that the early ape-like progenitors of man +were likewise social; but this is not of much importance for us. +Although man, as he now exists, has few special instincts, having lost +any which his early progenitors may have possessed, this is no reason +why he should not have retained from an extremely remote period some +degree of instinctive love and sympathy for his fellows. We are indeed +all conscious that we do possess such sympathetic feelings;[119] but our +consciousness does not tell us whether they are instinctive, having +originated long ago in the same manner as with the lower animals, or +whether they have been acquired by each of us during our early years. As +man is a social animal, it is also probable that he would inherit a +tendency to be faithful to his comrades, for this quality is common to +most social animals. He would in like manner possess some capacity for +self-command, and perhaps of obedience to the leader of the community. +He would from an inherited tendency still be willing to defend, in +concert with others, his fellow-men, and would be ready to aid them in +any way which did not too greatly interfere with his own welfare or his +own strong desires. + +The social animals which stand at the bottom of the scale are guided +almost exclusively, and those which stand higher in the scale are +largely guided, in the aid which they give to the members of the same +community, by special instincts; but they are likewise in part impelled +by mutual love and sympathy, assisted apparently by some amount of +reason. Although man, as just remarked, has no special instincts to tell +him how to aid his fellow-men, he still has the impulse, and with his +improved intellectual faculties would naturally be much guided in this +respect by reason and experience. Instinctive sympathy would, also, +cause him to value highly the approbation of his fellow-men; for, as Mr. +Bain has clearly shewn,[120] the love of praise and the strong feeling +of glory, and the still stronger horror of scorn and infamy, "are due to +the workings of sympathy." Consequently man would be greatly influenced +by the wishes, approbation, and blame of his fellow-men, as expressed by +their gestures and language. Thus the social instincts, which must have +been acquired by man in a very rude state, and probably even by his +early ape-like progenitors, still give the impulse to many of his best +actions; but his actions are largely determined by the expressed wishes +and judgment of his fellow-men, and unfortunately still oftener by his +own strong, selfish desires. But as the feelings of love and sympathy +and the power of self-command become strengthened by habit, and as the +power of reasoning becomes clearer so that man can appreciate the +justice of the judgments of his fellow-men, he will feel himself +impelled, independently of any pleasure or pain felt at the moment, to +certain lines of conduct. He may then say, I am the supreme judge of my +own conduct, and in the words of Kant, I will not in my own person +violate the dignity of humanity. + + +_The more enduring Social Instincts conquer the less Persistent +Instincts._--We have, however, not as yet considered the main point, on +which the whole question of the moral sense hinges. Why should a man +feel that he ought to obey one instinctive desire rather than another? +Why does he bitterly regret if he has yielded to the strong sense of +self-preservation, and has not risked his life to save that of a +fellow-creature; or why does he regret having stolen food from severe +hunger? + +It is evident in the first place, that with mankind the instinctive +impulses have different degrees of strength; a young and timid mother +urged by the maternal instinct will, without a moment's hesitation, run +the greatest danger for her infant, but not for a mere fellow-creature. +Many a man, or even boy, who never before risked his life for another, +but in whom courage and sympathy were well developed, has, disregarding +the instinct of self-preservation, instantaneously plunged into a +torrent to save a drowning fellow-creature. In this case man is impelled +by the same instinctive motive, which caused the heroic little American +monkey, formerly described, to attack the great and dreaded baboon, to +save his keeper. Such actions as the above appear to be the simple +result of the greater strength of the social or maternal instincts than +of any other instinct or motive; for they are performed too +instantaneously for reflection, or for the sensation of pleasure or +pain; though if prevented distress would be caused. + +I am aware that some persons maintain that actions performed +impulsively, as in the above cases, do not come under the dominion of +the moral sense, and cannot be called moral. They confine this term to +actions done deliberately, after a victory over opposing desires, or to +actions prompted by some lofty motive. But it appears scarcely possible +to draw any clear line of distinction of this kind; though the +distinction may be real. As far as exalted motives are concerned, many +instances have been recorded of barbarians, destitute of any feeling of +general benevolence towards mankind, and not guided by any religious +motive, who have deliberately as prisoners sacrificed their lives,[121] +rather than betray their comrades; and surely their conduct ought to be +considered as moral. As far as deliberation and the victory over +opposing motives are concerned, animals may be seen doubting between +opposed instincts, as in rescuing their offspring or comrades from +danger; yet their actions, though done for the good of others, are not +called moral. Moreover, an action repeatedly performed by us, will at +last be done without deliberation or hesitation, and can then hardly be +distinguished from an instinct; yet surely no one will pretend that an +action thus done ceases to be moral. On the contrary, we all feel that +an act cannot be considered as perfect, or as performed in the most +noble manner, unless it be done impulsively, without deliberation or +effort, in the same manner as by a man in whom the requisite qualities +are innate. He who is forced to overcome his fear or want of sympathy +before he acts, deserves, however, in one way higher credit than the man +whose innate disposition leads him to a good act without effort. As we +cannot distinguish between motives, we rank all actions of a certain +class as moral, when they are performed by a moral being. A moral being +is one who is capable of comparing his past and future actions or +motives, and of approving or disapproving of them. We have no reason to +suppose that any of the lower animals have this capacity; therefore +when a monkey faces danger to rescue its comrade, or takes charge of an +orphan-monkey, we do not call its conduct moral. But in the case of man, +who alone can with certainty be ranked as a moral being, actions of a +certain class are called moral, whether performed deliberately after a +struggle with opposing motives, or from the effects of slowly-gained +habit, or impulsively through instinct. + +But to return to our more immediate subject; although some instincts are +more powerful than others, thus leading to corresponding actions, yet it +cannot be maintained that the social instincts are ordinarily stronger +in man, or have become stronger through long-continued habit, than the +instincts, for instance, of self-preservation, hunger, lust, vengeance, +&c. Why then does man regret, even though he may endeavour to banish any +such regret, that he has followed the one natural impulse, rather than +the other; and why does he further feel that he ought to regret his +conduct? Man in this respect differs profoundly from the lower animals. +Nevertheless we can, I think, see with some degree of clearness the +reason of this difference. + +Man, from the activity of his mental faculties, cannot avoid reflection: +past impressions and images are incessantly passing through his mind +with distinctness. Now with those animals which live permanently in a +body, the social instincts are ever present and persistent. Such animals +are always ready to utter the danger-signal, to defend the community, +and to give aid to their fellows in accordance with their habits; they +feel at all times, without the stimulus of any special passion or +desire, some degree of love and sympathy for them; they are unhappy if +long separated from them, and always happy to be in their company. So it +is with ourselves. A man who possessed no trace of such feelings would +be an unnatural monster. On the other hand, the desire to satisfy +hunger, or any passion, such as vengeance, is in its nature temporary, +and can for a time be fully satisfied. Nor is it easy, perhaps hardly +possible, to call up with complete vividness the feeling, for instance, +of hunger; nor indeed, as has often been remarked, of any suffering. The +instinct of self-preservation is not felt except in the presence of +danger; and many a coward has thought himself brave until he has met his +enemy face to face. The wish for another man's property is perhaps as +persistent a desire as any that can be named; but even in this case the +satisfaction of actual possession is generally a weaker feeling than the +desire: many a thief, if not an habitual one, after success has wondered +why he stole some article. + +Thus, as man cannot prevent old impressions continually repassing +through his mind, he will be compelled to compare the weaker impressions +of, for instance, past hunger, or of vengeance satisfied or danger +avoided at the cost of other men, with the instinct of sympathy and +good-will to his fellows, which is still present and ever in some degree +active in his mind. He will then feel in his imagination that a stronger +instinct has yielded to one which now seems comparatively weak; and then +that sense of dissatisfaction will inevitably be felt with which man is +endowed, like every other animal, in order that his instincts may be +obeyed. The case before given, of the swallow, affords an illustration, +though of a reversed nature, of a temporary though for the time strongly +persistent instinct conquering another instinct which is usually +dominant over all others. At the proper season these birds seem all day +long to be impressed with the desire to migrate; their habits change; +they become restless, are noisy, and congregate in flocks. Whilst the +mother-bird is feeding or brooding over her nestlings, the maternal +instinct is probably stronger than the migratory; but the instinct which +is more persistent gains the victory, and at last, at a moment when her +young ones are not in sight, she takes flight and deserts them. When +arrived at the end of her long journey, and the migratory instinct +ceases to act, what an agony of remorse each bird would feel, if, from +being endowed with great mental activity, she could not prevent the +image continually passing before her mind of her young ones perishing in +the bleak north from cold and hunger. + +At the moment of action, man will no doubt be apt to follow the stronger +impulse; and though this may occasionally prompt him to the noblest +deeds, it will far more commonly lead him to gratify his own desires at +the expense of other men. But after their gratification, when past and +weaker impressions are contrasted with the ever-enduring social +instincts, retribution will surely come. Man will then feel dissatisfied +with himself, and will resolve with more or less force to act +differently for the future. This is conscience; for conscience looks +backwards and judges past actions, inducing that kind of +dissatisfaction, which if weak we call regret, and if severe remorse. + +These sensations are, no doubt, different from those experienced when +other instincts or desires are left unsatisfied; but every unsatisfied +instinct has its own proper prompting sensation, as we recognise with +hunger, thirst, &c. Man thus prompted, will through long habit acquire +such perfect self-command, that his desires and passions will at last +instantly yield to his social sympathies, and there will no longer be a +struggle between them. The still hungry, or the still revengeful man +will not think of stealing food, or of wreaking his vengeance. It is +possible, or, as we shall hereafter see, even probable, that the habit +of self-command may, like other habits, be inherited. Thus at last man +comes to feel, through acquired and perhaps inherited habit, that it is +best for him to obey his more persistent instincts. The imperious word +_ought_ seems merely to imply the consciousness of the existence of a +persistent instinct, either innate or partly acquired, serving him as a +guide, though liable to be disobeyed. We hardly use the word _ought_ in +a metaphorical sense, when we say hounds ought to hunt, pointers to +point, and retrievers to retrieve their game. If they fail thus to act, +they fail in their duty and act wrongly. + +If any desire or instinct, leading to an action opposed to the good of +others, still appears to a man, when recalled to mind, as strong as, or +stronger than, his social instinct, he will feel no keen regret at +having followed it; but he will be conscious that if his conduct were +known to his fellows, it would meet with their disapprobation; and few +are so destitute of sympathy as not to feel discomfort when this is +realised. If he has no such sympathy, and if his desires leading to bad +actions are at the time strong, and when recalled are not overmastered +by the persistent social instincts, then he is essentially a bad +man;[122] and the sole restraining motive left is the fear of +punishment, and the conviction that in the long run it would be best for +his own selfish interests to regard the good of others rather than his +own. + +It is obvious that every one may with an easy conscience gratify his own +desires, if they do not interfere with his social instincts, that is +with the good of others; but in order to be quite free from +self-reproach, or at least of anxiety, it is almost necessary for him to +avoid the disapprobation, whether reasonable or not, of his fellow men. +Nor must he break through the fixed habits of his life, especially if +these are supported by reason; for if he does, he will assuredly feel +dissatisfaction. He must likewise avoid the reprobation of the one God +or gods, in whom according to his knowledge or superstition he may +believe; but in this case the additional fear of divine punishment often +supervenes. + + +_The strictly Social Virtues at first alone regarded._--The above view +of the first origin and nature of the moral sense, which tells us what +we ought to do, and of the conscience which reproves us if we disobey +it, accords well with what we see of the early and undeveloped condition +of this faculty in mankind. The virtues which must be practised, at +least generally, by rude men, so that they may associate in a body, are +those which are still recognised as the most important. But they are +practised almost exclusively in relation to the men of the same tribe; +and their opposites are not regarded as crimes in relation to the men of +other tribes. No tribe could hold together if murder, robbery, +treachery, &c., were common; consequently such crimes within the limits +of the same tribe "are branded with everlasting infamy;"[123] but excite +no such sentiment beyond these limits. A North-American Indian is well +pleased with himself, and is honoured by others, when he scalps a man of +another tribe; and a Dyak cuts off the head of an unoffending person +and dries it as a trophy. The murder of infants has prevailed on the +largest scale throughout the world,[124] and has met with no reproach; +but infanticide, especially of females, has been thought to be good for +the tribe, or at least not injurious. Suicide during former times was +not generally considered as a crime,[125] but rather from the courage +displayed as an honourable act; and it is still largely practised by +some semi-civilised nations without reproach, for the loss to a nation +of a single individual is not felt: whatever the explanation may be, +suicide, as I hear from Sir J. Lubbock, is rarely practised by the +lowest barbarians. It has been recorded that an Indian Thug +conscientiously regretted that he had not strangled and robbed as many +travellers as did his father before him. In a rude state of civilisation +the robbery of strangers is, indeed, generally considered as honourable. + +The great sin of Slavery has been almost universal, and slaves have +often been treated in an infamous manner. As barbarians do not regard +the opinion of their women, wives are commonly treated like slaves. Most +savages are utterly indifferent to the sufferings of strangers, or even +delight in witnessing them. It is well known that the women and children +of the North-American Indians aided in torturing their enemies. Some +savages take a horrid pleasure in cruelty to animals,[126] and humanity +with them is an unknown virtue. Nevertheless, feelings of sympathy and +kindness are common, especially during sickness, between the members of +the same tribe, and are sometimes extended beyond the limits of the +tribe. Mungo Park's touching account of the kindness of the negro women +of the interior to him is well known. Many instances could be given of +the noble fidelity of savages towards each other, but not to strangers; +common experience justifies the maxim of the Spaniard, "Never, never +trust an Indian." There cannot be fidelity without truth; and this +fundamental virtue is not rare between the members of the same tribe: +thus Mungo Park heard the negro women teaching their young children to +love the truth. This, again, is one of the virtues which becomes so +deeply rooted in the mind that it is sometimes practised by savages even +at a high cost, towards strangers; but to lie to your enemy has rarely +been thought a sin, as the history of modern diplomacy too plainly +shews. As soon as a tribe has a recognised leader, disobedience becomes +a crime, and even abject submission is looked at as a sacred virtue. + +As during rude times no man can be useful or faithful to his tribe +without courage, this quality has universally been placed in the highest +rank; and although, in civilised countries, a good, yet timid, man may +be far more useful to the community than a brave one, we cannot help +instinctively honouring the latter above a coward, however benevolent. +Prudence, on the other hand, which does not concern the welfare of +others, though a very useful virtue, has never been highly esteemed. As +no man can practise the virtues necessary for the welfare of his tribe +without self-sacrifice, self-command, and the power of endurance, these +qualities have been at all times highly and most justly valued. The +American savage voluntarily submits without a groan to the most horrid +tortures to prove and strengthen his fortitude and courage; and we +cannot help admiring him, or even an Indian Fakir, who, from a foolish +religious motive, swings suspended by a hook buried in his flesh. + +The other self-regarding virtues, which do not obviously, though they +may really, affect the welfare of the tribe, have never been esteemed by +savages, though now highly appreciated by civilised nations. The +greatest intemperance with savages is no reproach. Their utter +licentiousness, not to mention unnatural crimes, is something +astounding.[127] As soon, however, as marriage, whether polygamous or +monogamous, becomes common, jealousy will lead to the inculcation of +female virtue; and this being honoured will tend to spread to the +unmarried females. How slowly it spreads to the male sex we see at the +present day. Chastity eminently requires self-command; therefore it has +been honoured from a very early period in the moral history of civilised +man. As a consequence of this, the senseless practice of celibacy has +been ranked from a remote period as a virtue.[128] The hatred of +indecency, which appears to us so natural as to be thought innate, and +which is so valuable an aid to chastity, is a modern virtue, +appertaining exclusively, as Sir G. Staunton remarks,[129] to civilised +life. This is shewn by the ancient religious rites of various nations, +by the drawings on the walls of Pompeii, and by the practices of many +savages. + +We have now seen that actions are regarded by savages, and were probably +so regarded by primeval man, as good or bad, solely as they affect in an +obvious manner the welfare of the tribe,--not that of the species, nor +that of man as an individual member of the tribe. This conclusion +agrees well with the belief that the so-called moral sense is +aboriginally derived from the social instincts, for both relate at first +exclusively to the community. The chief causes of the low morality of +savages, as judged by our standard, are, firstly, the confinement of +sympathy to the same tribe. Secondly, insufficient powers of reasoning, +so that the bearing of many virtues, especially of the self-regarding +virtues, on the general welfare of the tribe is not recognised. Savages, +for instance, fail to trace the multiplied evils consequent on a want of +temperance, chastity, &c. And, thirdly, weak power of self-command; for +this power has not been strengthened through long-continued, perhaps +inherited, habit, instruction and religion. + +I have entered into the above details on the immorality of savages,[130] +because some authors have recently taken a high view of their moral +nature, or have attributed most of their crimes to mistaken +benevolence.[131] These authors appear to rest their conclusion on +savages possessing, as they undoubtedly do possess, and often in a high +degree, those virtues which are serviceable, or even necessary, for the +existence of a tribal community. + + +_Concluding Remarks._--Philosophers of the derivative[132] school of +morals formerly assumed that the foundation of morality lay in a form of +Selfishness; but more recently in the "Greatest Happiness principle." +According to the view given above, the moral sense is fundamentally +identical with the social instincts; and in the case of the lower +animals it would be absurd to speak of these instincts as having been +developed from selfishness, or for the happiness of the community. They +have, however, certainly been developed for the general good of the +community. The term, general good, may be defined as the means by which +the greatest possible number of individuals can be reared in full vigour +and health, with all their faculties perfect, under the conditions to +which they are exposed. As the social instincts both of man and the +lower animals have no doubt been developed by the same steps, it would +be advisable, if found practicable, to use the same definition in both +cases, and to take as the test of morality, the general good or welfare +of the community, rather than the general happiness; but this definition +would perhaps require some limitation on account of political ethics. + +When a man risks his life to save that of a fellow-creature, it seems +more appropriate to say that he acts for the general good or welfare, +rather than for the general happiness of mankind. No doubt the welfare +and the happiness of the individual usually coincide; and a contented, +happy tribe will flourish better than one that is discontented and +unhappy. We have seen that at an early period in the history of man, the +expressed wishes of the community will have naturally influenced to a +large extent the conduct of each member; and as all wish for happiness, +the "greatest happiness principle" will have become a most important +secondary guide and object; the social instincts, including sympathy, +always serving as the primary impulse and guide. Thus the reproach of +laying the foundation of the most noble part of our nature in the base +principle of selfishness is removed; unless indeed the satisfaction +which every animal feels when it follows its proper instincts, and the +dissatisfaction felt when prevented, be called selfish. + +The expression of the wishes and judgment of the members of the same +community, at first by oral and afterwards by written language, serves, +as just remarked, as a most important secondary guide of conduct, in aid +of the social instincts, but sometimes in opposition to them. This +latter fact is well exemplified by the _Law of Honour_, that is the law +of the opinion of our equals, and not of all our countrymen. The breach +of this law, even when the breach is known to be strictly accordant with +true morality, has caused many a man more agony than a real crime. We +recognise the same influence in the burning sense of shame which most of +us have felt even after the interval of years, when calling to mind some +accidental breach of a trifling though fixed rule of etiquette. The +judgment of the community will generally be guided by some rude +experience of what is best in the long run for all the members; but this +judgment will not rarely err from ignorance and from weak powers of +reasoning. Hence the strangest customs and superstitions, in complete +opposition to the true welfare and happiness of mankind, have become +all-powerful throughout the world. We see this in the horror felt by a +Hindoo who breaks his caste, in the shame of a Mahometan woman who +exposes her face, and in innumerable other instances. It would be +difficult to distinguish between the remorse felt by a Hindoo who has +eaten unclean food, from that felt after committing a theft; but the +former would probably be the more severe. + +How so many absurd rules of conduct, as well as so many absurd religious +beliefs, have originated we do not know; nor how it is that they have +become, in all quarters of the world, so deeply impressed on the mind +of men; but it is worthy of remark that a belief constantly inculcated +during the early years of life, whilst the brain is impressible, appears +to acquire almost the nature of an instinct; and the very essence of an +instinct is that it is followed independently of reason. Neither can we +say why certain admirable virtues, such as the love of truth, are much +more highly appreciated by some savage tribes than by others;[133] nor, +again, why similar differences prevail even amongst civilised nations. +Knowing how firmly fixed many strange customs and superstitions have +become, we need feel no surprise that the self-regarding virtues should +now appear to us so natural, supported as they are by reason, as to be +thought innate, although they were not valued by man in his early +condition. + +Notwithstanding many sources of doubt, man can generally and readily +distinguish between the higher and lower moral rules. The higher are +founded on the social instincts, and relate to the welfare of others. +They are supported by the approbation of our fellow-men and by reason. +The lower rules, though some of them when implying self-sacrifice hardly +deserve to be called lower, relate chiefly to self, and owe their origin +to public opinion, when matured by experience and cultivated; for they +are not practised by rude tribes. + +As man advances in civilisation, and small tribes are united into larger +communities, the simplest reason would tell each individual that he +ought to extend his social instincts and sympathies to all the members +of the same nation, though personally unknown to him. This point being +once reached, there is only an artificial barrier to prevent his +sympathies extending to the men of all nations and races. If, indeed, +such men are separated from him by great differences in appearance or +habits, experience unfortunately shews us how long it is before we look +at them as our fellow-creatures. Sympathy beyond the confines of man, +that is humanity to the lower animals, seems to be one of the latest +moral acquisitions. It is apparently unfelt by savages, except towards +their pets. How little the old Romans knew of it is shewn by their +abhorrent gladiatorial exhibitions. The very idea of humanity, as far as +I could observe, was new to most of the Gauchos of the Pampas. This +virtue, one of the noblest with which man is endowed, seems to arise +incidentally from our sympathies becoming more tender and more widely +diffused, until they are extended to all sentient beings. As soon as +this virtue is honoured and practised by some few men, it spreads +through instruction and example to the young, and eventually through +public opinion. + +The highest stage in moral culture at which we can arrive, is when we +recognise that we ought to control our thoughts, and "not even in inmost +thought to think again the sins that made the past so pleasant to +us."[134] Whatever makes any bad action familiar to the mind, renders +its performance by so much the easier. As Marcus Aurelius long ago said, +"Such as are thy habitual thoughts, such also will be the character of +thy mind; for the soul is dyed by the thoughts."[135] + +Our great philosopher, Herbert Spencer, has recently explained his views +on the moral sense. He says,[136] "I believe that the experiences of +utility organised and consolidated through all past generations of the +human race, have been producing corresponding modifications, which, by +continued transmission and accumulation, have become in us certain +faculties of moral intuition--certain emotions responding to right and +wrong conduct, which have no apparent basis in the individual +experiences of utility." There is not the least inherent improbability, +as it seems to me, in virtuous tendencies being more or less strongly +inherited; for, not to mention the various dispositions and habits +transmitted by many of our domestic animals, I have heard of cases in +which a desire to steal and a tendency to lie appeared to run in +families of the upper ranks; and as stealing is so rare a crime in the +wealthy classes, we can hardly account by accidental coincidence for the +tendency occurring in two or three members of the same family. If bad +tendencies are transmitted, it is probable that good ones are likewise +transmitted. Excepting through the principle of the transmission of +moral tendencies, we cannot understand the differences believed to exist +in this respect between the various races of mankind. We have, however, +as yet, hardly sufficient evidence on this head. + +Even the partial transmission of virtuous tendencies would be an immense +assistance to the primary impulse derived directly from the social +instincts, and indirectly from the approbation of our fellow-men. +Admitting for the moment that virtuous tendencies are inherited, it +appears probable, at least in such cases as chastity, temperance, +humanity to animals, &c., that they become first impressed on the mental +organisation through habit, instruction, and example, continued during +several generations in the same family, and in a quite subordinate +degree, or not at all, by the individuals possessing such virtues, +having succeeded best in the struggle for life. My chief source of doubt +with respect to any such inheritance, is that senseless customs, +superstitions, and tastes, such as the horror of a Hindoo for unclean +food, ought on the same principle to be transmitted. Although this in +itself is perhaps not less probable than that animals should acquire +inherited tastes for certain kinds of food or fear of certain foes, I +have not met with any evidence in support of the transmission of +superstitious customs or senseless habits. + + +Finally, the social instincts which no doubt were acquired by man, as by +the lower animals, for the good of the community, will from the first +have given to him some wish to aid his fellows, and some feeling of +sympathy. Such impulses will have served him at a very early period as a +rude rule of right and wrong. But as man gradually advanced in +intellectual power and was enabled to trace the more remote consequences +of his actions; as he acquired sufficient knowledge to reject baneful +customs and superstitions; as he regarded more and more not only the +welfare but the happiness of his fellow-men; as from habit, following on +beneficial experience, instruction, and example, his sympathies became +more tender and widely diffused, so as to extend to the men of all +races, to the imbecile, the maimed, and other useless members of +society, and finally to the lower animals,--so would the standard of his +morality rise higher and higher. And it is admitted by moralists of the +derivative school and by some intuitionists, that the standard of +morality has risen since an early period in the history of man.[137] + +As a struggle may sometimes be seen going on between the various +instincts of the lower animals, it is not surprising that there should +be a struggle in man between his social instincts, with their derived +virtues, and his lower, though at the moment, stronger impulses or +desires. This, as Mr. Galton[138] has remarked, is all the less +surprising, as man has emerged from a state of barbarism within a +comparatively recent period. After having yielded to some temptation we +feel a sense of dissatisfaction, analogous to that felt from other +unsatisfied instincts, called in this case conscience; for we cannot +prevent past images and impressions continually passing through our +minds, and these in their weakened state we compare with the +ever-present social instincts, or with habits gained in early youth and +strengthened during our whole lives, perhaps inherited, so that they are +at last rendered almost as strong as instincts. Looking to future +generations, there is no cause to fear that the social instincts will +grow weaker, and we may expect that virtuous habits will grow stronger, +becoming perhaps fixed by inheritance. In this case the struggle between +our higher and lower impulses will be less severe, and virtue will be +triumphant. + + +_Summary of the two last Chapters._--There can be no doubt that the +difference between the mind of the lowest man and that of the highest +animal is immense. An anthropomorphous ape, if he could take a +dispassionate view of his own case, would admit that though he could +form an artful plan to plunder a garden--though he could use stones for +fighting or for breaking open nuts, yet that the thought of fashioning +a stone into a tool was quite beyond his scope. Still less, as he would +admit, could he follow out a train of metaphysical reasoning, or solve a +mathematical problem, or reflect on God, or admire a grand natural +scene. Some apes, however, would probably declare that they could and +did admire the beauty of the coloured skin and fur of their partners in +marriage. They would admit, that though they could make other apes +understand by cries some of their perceptions and simpler wants, the +notion of expressing definite ideas by definite sounds had never crossed +their minds. They might insist that they were ready to aid their +fellow-apes of the same troop in many ways, to risk their lives for +them, and to take charge of their orphans; but they would be forced to +acknowledge that disinterested love for all living creatures, the most +noble attribute of man, was quite beyond their comprehension. + +Nevertheless the difference in mind between man and the higher animals, +great as it is, is certainly one of degree and not of kind. We have seen +that the senses and intuitions, the various emotions and faculties, such +as love, memory, attention, curiosity, imitation, reason, &c., of which +man boasts, may be found in an incipient, or even sometimes in a +well-developed condition, in the lower animals. They are also capable of +some inherited improvement, as we see in the domestic dog compared with +the wolf or jackal. If it be maintained that certain powers, such as +self-consciousness, abstraction, &c., are peculiar to man, it may well +be that these are the incidental results of other highly-advanced +intellectual faculties; and these again are mainly the result of the +continued use of a highly developed language. At what age does the +new-born infant possess the power of abstraction, or become +self-conscious and reflect on its own existence? We cannot answer; nor +can we answer in regard to the ascending organic scale. The half-art and +half-instinct of language still bears the stamp of its gradual +evolution. The ennobling belief in God is not universal with man; and +the belief in active spiritual agencies naturally follows from his other +mental powers. The moral sense perhaps affords the best and highest +distinction between man and the lower animals; but I need not say +anything on this head, as I have so lately endeavoured to shew +that the social instincts,--the prime principle of man's moral +constitution[139]--with the aid of active intellectual powers and the +effects of habit, naturally lead to the golden rule, "As ye would that +men should do to you, do ye to them likewise;" and this lies at the +foundation of morality. + +In a future chapter I shall make some few remarks on the probable steps +and means by which the several mental and moral faculties of man have +been gradually evolved. That this at least is possible ought not to be +denied, when we daily see their development in every infant; and when we +may trace a perfect gradation from the mind of an utter idiot, lower +than that of the lowest animal, to the mind of a Newton. + + + + +CHAPTER IV. + +ON THE MANNER OF DEVELOPMENT OF MAN FROM SOME LOWER FORM. + + + Variability of body and mind in man--Inheritance--Causes of + variability--Laws of variation the same in man as in the lower + animals--Direct action of the conditions of life--Effects of + the increased use and disuse of parts--Arrested + development--Reversion--Correlated variation--Rate of + increase--Checks to increase--Natural selection--Man the most + dominant animal in the world--Importance of his corporeal + structure--The causes which have led to his becoming + erect--Consequent changes of structure--Decrease in size of the + canine teeth--Increased size and altered shape of the + skull--Nakedness--Absence of a tail--Defenceless condition of + man. + + +We have seen in the first chapter that the homological structure of man, +his embryological development and the rudiments which he still retains, +all declare in the plainest manner that he is descended from some lower +form. The possession of exalted mental powers is no insuperable +objection to this conclusion. In order that an ape-like creature should +have been transformed into man, it is necessary that this early form, as +well as many successive links, should all have varied in mind and body. +It is impossible to obtain direct evidence on this head; but if it can +be shewn that man now varies--that his variations are induced by the +same general causes, and obey the same general laws, as in the case of +the lower animals--there can be little doubt that the preceding +intermediate links varied in a like manner. The variations at each +successive stage of descent must, also, have been in some manner +accumulated and fixed. + +The facts and conclusions to be given in this chapter relate almost +exclusively to the probable means by which the transformation of man has +been effected, as far as his bodily structure is concerned. The +following chapter will be devoted to the development of his intellectual +and moral faculties. But the present discussion likewise bears on the +origin of the different races or species of mankind, whichever term may +be preferred. + +It is manifest that man is now subject to much variability. No two +individuals of the same race are quite alike. We may compare millions of +faces, and each will be distinct. There is an equally great amount of +diversity in the proportions and dimensions of the various parts of the +body; the length of the legs being one of the most variable points.[140] +Although in some quarters of the world an elongated skull, and in other +quarters a short skull prevails, yet there is great diversity of shape +even within the limits of the same race, as with the aborigines of +America and South Australia,--the latter a race "probably as pure and +homogeneous in blood, customs, and language as any in existence"--and +even with the inhabitants of so confined an area as the Sandwich +Islands.[141] An eminent dentist assures me that there is nearly as much +diversity in the teeth, as in the features. The chief arteries so +frequently run in abnormal courses, that it has been found useful for +surgical purposes to calculate from 12,000 corpses how often each +course prevails.[142] The muscles are eminently variable: thus those of +the foot were found by Prof. Turner[143] not to be strictly alike in any +two out of fifty bodies; and in some the deviations were considerable. +Prof. Turner adds that the power of performing the appropriate movements +must have been modified in accordance with the several deviations. Mr. +J. Wood has recorded[144] the occurrence of 295 muscular variations in +thirty-six subjects, and in another set of the same number no less than +558 variations, reckoning both sides of the body as one. In the last +set, not one body out of the thirty-six was "found totally wanting in +departures from the standard descriptions of the muscular system given +in anatomical text-books." A single body presented the extraordinary +number of twenty-five distinct abnormalities. The same muscle sometimes +varies in many ways: thus Prof. Macalister describes[145] no less than +twenty distinct variations in the _palmaris accessorius_. + +The famous old anatomist, Wolff,[146] insists that the internal viscera +are more variable than the external parts: _Nulla particula est quae non +aliter et aliter in aliis se habeat hominibus._ He has even written a +treatise on the choice of typical examples of the viscera for +representation. A discussion on the beau-ideal of the liver, lungs, +kidneys, &c., as of the human face divine, sounds strange in our ears. + +The variability or diversity of the mental faculties in men of the same +race, not to mention the greater differences between the men of +distinct races, is so notorious that not a word need here be said. So it +is with the lower animals, as has been illustrated by a few examples in +the last chapter. All who have had charge of menageries admit this fact, +and we see it plainly in our dogs and other domestic animals. Brehm +especially insists that each individual monkey of those which he kept +under confinement in Africa had its own peculiar disposition and temper: +he mentions one baboon remarkable for its high intelligence; and the +keepers in the Zoological Gardens pointed out to me a monkey, belonging +to the New World division, equally remarkable for intelligence. Rengger, +also, insists on the diversity in the various mental characters of the +monkeys of the same species which he kept in Paraguay; and this +diversity, as he adds, is partly innate, and partly the result of the +manner in which they have been treated or educated.[147] + +I have elsewhere[148] so fully discussed the subject of Inheritance that +I need here add hardly anything. A greater number of facts have been +collected with respect to the transmission of the most trifling, as well +as of the most important characters in man than in any of the lower +animals; though the facts are copious enough with respect to the latter. +So in regard to mental qualities, their transmission is manifest in our +dogs, horses, and other domestic animals. Besides special tastes and +habits, general intelligence, courage, bad and good temper, &c., are +certainly transmitted. With man we see similar facts in almost every +family; and we now know through the admirable labours of Mr. Galton[149] +that genius, which implies a wonderfully complex combination of high +faculties, tends to be inherited; and, on the other hand, it is too +certain that insanity and deteriorated mental powers likewise run in the +same families. + +With respect to the causes of variability we are in all cases very +ignorant; but we can see that in man as in the lower animals, they stand +in some relation with the conditions to which each species has been +exposed during several generations. Domesticated animals vary more than +those in a state of nature; and this is apparently due to the +diversified and changing nature of their conditions. The different races +of man resemble in this respect domesticated animals, and so do the +individuals of the same race when inhabiting a very wide area, like that +of America. We see the influence of diversified conditions in the more +civilised nations, the members of which belong to different grades of +rank and follow different occupations, presenting a greater range of +character than the members of barbarous nations. But the uniformity of +savages has often been exaggerated, and in some cases can hardly be said +to exist.[150] It is nevertheless an error to speak of man, even if we +look only to the conditions to which he has been subjected, as "far more +domesticated"[151] than any other animal. Some savage races, such as +the Australians, are not exposed to more diversified conditions than are +many species which have very wide ranges. In another and much more +important respect, man differs widely from any strictly domesticated +animal; for his breeding has not been controlled, either through +methodical or unconscious selection. No race or body of men has been so +completely subjugated by other men, that certain individuals have been +preserved and thus unconsciously selected, from being in some way more +useful to their masters. Nor have certain male and female individuals +been intentionally picked out and matched, except in the well-known case +of the Prussian grenadiers; and in this case man obeyed, as might have +been expected, the law of methodical selection; for it is asserted that +many tall men were reared in the villages inhabited by the grenadiers +with their tall wives. + +If we consider all the races of man, as forming a single species, his +range is enormous; but some separate races, as the Americans and +Polynesians, have very wide ranges. It is a well-known law that +widely-ranging species are much more variable than species with +restricted ranges; and the variability of man may with more truth be +compared with that of widely-ranging species, than with that of +domesticated animals. + +Not only does variability appear to be induced in man and the lower +animals by the same general causes, but in both the same characters are +affected in a closely analogous manner. This has been proved in such +full detail by Godron and Quatrefages, that I need here only refer to +their works.[152] Monstrosities, which graduate into slight variations, +are likewise so similar in man and the lower animals, that the same +classification and the same terms can be used for both, as may be seen +in Isidore Geoffroy St.-Hilaire's great work.[153] This is a necessary +consequence of the same laws of change prevailing throughout the animal +kingdom. In my work on the variation of domestic animals, I have +attempted to arrange in a rude fashion the laws of variation under the +following heads:--The direct and definite action of changed conditions, +as shewn by all or nearly all the individuals of the same species +varying in the same manner under the same circumstances. The effects of +the long-continued use or disuse of parts. The cohesion of homologous +parts. The variability of multiple parts. Compensation of growth; but of +this law I have found no good instances in the case of man. The effects +of the mechanical pressure of one part on another; as of the pelvis on +the cranium of the infant in the womb. Arrests of development, leading +to the diminution or suppression of parts. The reappearance of long-lost +characters through reversion. And lastly, correlated variation. All +these so-called laws apply equally to man and the lower animals; and +most of them even to plants. It would be superfluous here to discuss all +of them;[154] but several are so important for us, that they must be +treated at considerable length. + + +_The direct and definite action of changed conditions._--This is a most +perplexing subject. It cannot be denied that changed conditions produce +some effect, and occasionally a considerable effect, on organisms of all +kinds; and it seems at first probable that if sufficient time were +allowed this would be the invariable result. But I have failed to obtain +clear evidence in favour of this conclusion; and valid reasons may be +urged on the other side, at least as far as the innumerable structures +are concerned, which are adapted for special ends. There can, however, +be no doubt that changed conditions induce an almost indefinite amount +of fluctuating variability, by which the whole organisation is rendered +in some degree plastic. + +In the United States, above 1,000,000 soldiers, who served in the late +war, were measured, and the States in which they were born and reared +recorded.[155] From this astonishing number of observations it is proved +that local influences of some kind act directly on stature; and we +further learn that "the State where the physical growth has in great +measure taken place, and the State of birth, which indicates the +ancestry, seem to exert a marked influence on the stature." For instance +it is established, "that residence in the Western States, during the +years of growth, tends to produce increase of stature." On the other +hand, it is certain that with sailors, their manner of life delays +growth, as shewn "by the great difference between the statures of +soldiers and sailors at the ages of 17 and 18 years." Mr. B. A. Gould +endeavoured to ascertain the nature of the influences which thus act on +stature; but he arrived only at negative results, namely, that they did +not relate to climate, the elevation of the land, soil, nor even "in any +controlling degree" to the abundance or need of the comforts of life. +This latter conclusion is directly opposed to that arrived at by +Villerme from the statistics of the height of the conscripts in +different parts of France. When we compare the differences in stature +between the Polynesian chiefs and the lower orders within the same +islands, or between the inhabitants of the fertile volcanic and low +barren coral islands of the same ocean,[156] or again between the +Fuegians on the eastern and western shores of their country, where the +means of subsistence are very different, it is scarcely possible to +avoid the conclusion that better food and greater comfort do influence +stature. But the preceding statements shew how difficult it is to arrive +at any precise result. Dr. Beddoe has lately proved that, with the +inhabitants of Britain, residence in towns and certain occupations have +a deteriorating influence on height; and he infers that the result is to +a certain extent inherited, as is likewise the case in the United +States. Dr. Beddoe further believes that wherever a "race attains its +maximum of physical development, it rises highest in energy and moral +vigour."[157] + +Whether external conditions produce any other direct effect on man is +not known. It might have been expected that differences of climate would +have had a marked influence, as the lungs and kidneys are brought into +fuller activity under a low temperature, and the liver and skin under a +high one.[158] It was formerly thought that the colour of the skin and +the character of the hair were determined by light or heat; and +although it can hardly be denied that some effect is thus produced, +almost all observers now agree that the effect has been very small, even +after exposure during many ages. But this subject will be more properly +discussed when we treat of the different races of mankind. With our +domestic animals there are grounds for believing that cold and damp +directly affect the growth of the hair; but I have not met with any +evidence on this head in the case of man. + + +_Effects of the increased Use and Disuse of Parts._--It is well known +that use strengthens the muscles in the individual, and complete disuse, +or the destruction of the proper nerve, weakens them. When the eye is +destroyed the optic nerve often becomes atrophied. When an artery is +tied, the lateral channels increase not only in diameter, but in the +thickness and strength of their coats. When one kidney ceases acting +from disease, the other increases in size and does double work. Bones +increase not only in thickness, but in length, from carrying a greater +weight.[159] Different occupations habitually followed lead to changed +proportions in various parts of the body. Thus it was clearly +ascertained by the United States Commission[160] that the legs of the +sailors employed in the late war were longer by 0.217 of an inch than +those of the soldiers, though the sailors were on an average shorter +men; whilst their arms were shorter by 1.09 of an inch, and therefore +out of proportion shorter in relation to their lesser height. This +shortness of the arms is apparently due to their greater use, and is an +unexpected result; but sailors chiefly use their arms in pulling and not +in supporting weights. The girth of the neck and the depth of the instep +are greater, whilst the circumference of the chest, waist, and hips is +less in sailors than in soldiers. + +Whether the several foregoing modifications would become hereditary, if +the same habits of life were followed during many generations, is not +known, but is probable. Rengger[161] attributes the thin legs and thick +arms of the Payaguas Indians to successive generations having passed +nearly their whole lives in canoes, with their lower extremities +motionless. Other writers have come to a similar conclusion in other +analogous cases. According to Cranz,[162] who lived for a long time with +the Esquimaux, "the natives believe that ingenuity and dexterity in +seal-catching (their highest art and virtue) is hereditary; there is +really something in it, for the son of a celebrated seal-catcher will +distinguish himself though he lost his father in childhood." But in this +case it is mental aptitude, quite as much as bodily structure, which +appears to be inherited. It is asserted that the hands of English +labourers are at birth larger than those of the gentry.[163] From the +correlation which exists, at least in some cases,[164] between the +development of the extremities and of the jaws, it is possible that in +those classes which do not labour much with their hands and feet, the +jaws would be reduced in size from this cause. That they are generally +smaller in refined and civilised men than in hard-working men or +savages, is certain. But with savages, as Mr. Herbert Spencer[165] has +remarked, the greater use of the jaws in chewing coarse, uncooked food, +would act in a direct manner on the masticatory muscles and on the bones +to which they are attached. In infants long before birth, the skin on +the soles of the feet is thicker than on any other part of the +body;[166] and it can hardly be doubted that this is due to the +inherited effects of pressure during a long series of generations. + +It is familiar to every one that watchmakers and engravers are liable to +become short-sighted, whilst sailors and especially savages are +generally long-sighted. Short-sight and long-sight certainly tend to be +inherited.[167] The inferiority of Europeans, in comparison with +savages, in eyesight and in the other senses, is no doubt the +accumulated and transmitted effect of lessened use during many +generations; for Rengger[168] states that he has repeatedly observed +Europeans, who had been brought up and spent their whole lives with the +wild Indians, who nevertheless did not equal them in the sharpness of +their senses. The same naturalist observes that the cavities in the +skull for the reception of the several sense-organs are larger in the +American aborigines than in Europeans; and this no doubt indicates a +corresponding difference in the dimensions of the organs themselves. +Blumenbach has also remarked on the large size of the nasal cavities in +the skulls of the American aborigines, and connects this fact with their +remarkably acute power of smell. The Mongolians of the plains of +Northern Asia, according to Pallas, have wonderfully perfect senses; and +Prichard believes that the great breadth of their skulls across the +zygomas follows from their highly-developed sense-organs.[169] + +The Quechua Indians inhabit the lofty plateaux of Peru, and Alcide +d'Orbigny states[170] that from continually breathing a highly rarefied +atmosphere they have acquired chests and lungs of extraordinary +dimensions. The cells, also, of the lungs are larger and more numerous +than in Europeans. These observations have been doubted; but Mr. D. +Forbes carefully measured many Aymaras, an allied race, living at the +height of between ten and fifteen thousand feet; and he informs me[171] +that they differ conspicuously from the men of all other races seen by +him, in the circumference and length of their bodies. In his table of +measurements, the stature of each man is taken at 1000, and the other +measurements are reduced to this standard. It is here seen that the +extended arms of the Aymaras are shorter than those of Europeans, and +much shorter than those of Negroes. The legs are likewise shorter, and +they present this remarkable peculiarity, that in every Aymara measured +the femur is actually shorter than the tibia. On an average the length +of the femur to that of the tibia is as 211 to 252; whilst in two +Europeans measured at the same time, the femora to the tibiae were as +244 to 230; and in three Negroes as 258 to 241. The humerus is likewise +shorter relatively to the fore-arm. This shortening of that part of the +limb which is nearest to the body, appears to be, as suggested to me by +Mr. Forbes, a case of compensation in relation with the greatly +increased length of the trunk. The Aymaras present some other singular +points of structure, for instance, the very small projection of the +heel. + +These men are so thoroughly acclimatised to their cold and lofty abode, +that when formerly carried down by the Spaniards to the low Eastern +plains, and when now tempted down by high wages to the gold-washings, +they suffer a frightful rate of mortality. Nevertheless Mr. Forbes found +a few pure families which had survived during two generations; and he +observed that they still inherited their characteristic peculiarities. +But it was manifest, even without measurement, that these peculiarities +had all decreased; and on measurement their bodies were found not to be +so much elongated as those of the men on the high plateau; whilst their +femora had become somewhat lengthened, as had their tibiae but in a less +degree. The actual measurements may be seen by consulting Mr. Forbes' +memoir. From these valuable observations, there can, I think, be no +doubt that residence during many generations at a great elevation tends, +both directly and indirectly, to induce inherited modifications in the +proportions of the body.[172] + +Although man may not have been much modified during the latter stages of +his existence through the increased or decreased use of parts, the +facts now given shew that his liability in this respect has not been +lost; and we positively know that the same law holds good with the lower +animals. Consequently we may infer, that when at a remote epoch the +progenitors of man were in a transitional state, and were changing from +quadrupeds into bipeds, natural selection would probably have been +greatly aided by the inherited effects of the increased or diminished +use of the different parts of the body. + + +_Arrests of Development._--Arrested development differs from arrested +growth, as parts in the former state continue to grow whilst still +retaining their early condition. Various monstrosities come under this +head, and some are known to be occasionally inherited, as a +cleft-palate. It will suffice for our purpose to refer to the arrested +brain-development of microcephalous idiots, as described in Vogt's great +memoir.[173] Their skulls are smaller, and the convolutions of the brain +are less complex than in normal men. The frontal sinus, or the +projection over the eyebrows, is largely developed, and the jaws are +prognathous to an "_effrayant_" degree; so that these idiots somewhat +resemble the lower types of mankind. Their intelligence and most of +their mental faculties are extremely feeble. They cannot acquire the +power of speech, and are wholly incapable of prolonged attention, but +are much given to imitation. They are strong and remarkably active, +continually gamboling and jumping about, and making grimaces. They often +ascend stairs on all-fours; and are curiously fond of climbing up +furniture or trees. We are thus reminded of the delight shewn by almost +all boys in climbing trees; and this again reminds us how lambs and +kids, originally alpine animals, delight to frisk on any hillock, +however small. + + +_Reversion._--Many of the cases to be here given might have been +introduced under the last heading. Whenever a structure is arrested in +its development, but still continues growing until it closely resembles +a corresponding structure in some lower and adult member of the same +group, we may in one sense consider it as a case of reversion. The lower +members in a group give us some idea how the common progenitor of the +group was probably constructed; and it is hardly credible that a part +arrested at an early phase of embryonic development should be enabled to +continue growing so as ultimately to perform its proper function, unless +it had acquired this power of continued growth during some earlier state +of existence, when the present exceptional or arrested structure was +normal. The simple brain of a microcephalous idiot, in as far as it +resembles that of an ape, may in this sense be said to offer a case of +reversion. There are other cases which come more strictly under our +present heading of reversion. Certain structures, regularly occurring in +the lower members of the group to which man belongs, occasionally make +their appearance in him, though not found in the normal human embryo; +or, if present in the normal human embryo, they become developed in an +abnormal manner, though this manner of development is proper to the +lower members of the same group. These remarks will be rendered clearer +by the following illustrations. + +In various mammals the uterus graduates from a double organ with two +distinct orifices and two passages, as in the marsupials, into a single +organ, showing no signs of doubleness except a slight internal fold, as +in the higher apes and man. The rodents exhibit a perfect series of +gradations between these two extreme states. In all mammals the uterus +is developed from two simple primitive tubes, the inferior portions of +which form the cornua; and it is in the words of Dr. Farre "by the +coalescence of the two cornua at their lower extremities that the body +of the uterus is formed in man; while in those animals in which no +middle portion or body exists, the cornua remain ununited. As the +development of the uterus proceeds, the two cornua become gradually +shorter, until at length they are lost, or, as it were, absorbed into +the body of the uterus." The angles of the uterus are still produced +into cornua, even so high in the scale as in the lower apes, and their +allies the lemurs. + +Now in women anomalous cases are not very infrequent, in which the +mature uterus is furnished with cornua, or is partially divided into two +organs; and such cases, according to Owen, repeat "the grade of +concentrative development," attained by certain rodents. Here perhaps we +have an instance of a simple arrest of embryonic development, with +subsequent growth and perfect functional development, for either side of +the partially double uterus is capable of performing the proper office +of gestation. In other and rarer cases, two distinct uterine cavities +are formed, each having its proper orifice and passage.[174] No such +stage is passed through during the ordinary development of the embryo, +and it is difficult to believe, though perhaps not impossible, that the +two simple, minute, primitive tubes could know how (if such an +expression may be used) to grow into two distinct uteri, each with a +well-constructed orifice and passage, and each furnished with numerous +muscles, nerves, glands and vessels, if they had not formerly passed +through a similar course of development, as in the case of existing +marsupials. No one will pretend that so perfect a structure as the +abnormal double uterus in woman could be the result of mere chance. But +the principle of reversion, by which long-lost dormant structures are +called back into existence, might serve as the guide for the full +development of the organ, even after the lapse of an enormous interval +of time. + +Professor Canestrini,[175] after discussing the foregoing and various +analogous cases, arrives at the same conclusion as that just given. He +adduces, as another instance, the malar bone, which, in some of the +Quadrumana and other mammals, normally consists of two portions. This is +its condition in the two-months-old human foetus; and thus it +sometimes remains, through arrested development, in man when adult, more +especially in the lower prognathous races. Hence Canestrini concludes +that some ancient progenitor of man must have possessed this bone +normally divided into two portions, which subsequently became fused +together. In man the frontal bone consists of a single piece, but in the +embryo and in children, and in almost all the lower mammals, it consists +of two pieces separated by a distinct suture. This suture occasionally +persists, more or less distinctly, in man after maturity, and more +frequently in ancient than in recent crania, especially as Canestrini +has observed in those exhumed from the Drift and belonging to the +brachycephalic type. Here again he comes to the same conclusion as in +the analogous case of the malar bones. In this and other instances +presently to be given, the cause of ancient races approaching the lower +animals in certain characters more frequently than do the modern races, +appears to be that the latter stand at a somewhat greater distance in +the long line of descent from their early semi-human progenitors. + +Various other anomalies in man, more or less analogous with the +foregoing, have been advanced by different authors[176] as cases of +reversion; but these seem not a little doubtful, for we have to descend +extremely low in the mammalian series before we find such structures +normally present.[177] + +In man the canine teeth are perfectly efficient instruments for +mastication. But their true canine character, as Owen[178] remarks, "is +indicated by the conical form of the crown, which terminates in an +obtuse point, is convex outward and flat or sub-concave within, at the +base of which surface there is a feeble prominence. The conical form is +best expressed in the Melanian races, especially the Australian. The +canine is more deeply implanted, and by a stronger fang than the +incisors." Nevertheless this tooth no longer serves man as a special +weapon for tearing his enemies or prey; it may, therefore, as far as its +proper function is concerned, be considered as rudimentary. In every +large collection of human skulls some may be found, as Haeckel[179] +observes, with the canine teeth projecting considerably beyond the +others in the same manner, but in a less degree, as in the +anthropomorphous apes. In these cases, open spaces between the teeth in +the one jaw are left for the reception of the canines belonging to the +opposite jaw. An interspace of this kind in a Kaffir skull, figured by +Wagner, is surprisingly wide.[180] Considering how few ancient skulls +have been examined in comparison with recent skulls, it is an +interesting fact that in at least three cases the canines project +largely; and in the Naulette jaw they are spoken of as enormous.[181] + +The males alone of the anthropomorphous apes have their canines fully +developed; but in the female gorilla, and in a less degree in the female +orang, these teeth project considerably beyond the others; therefore the +fact that women sometimes have, as I have been assured, considerably +projecting canines, is no serious objection to the belief that their +occasional great development in man is a case of reversion to an +ape-like progenitor. He who rejects with scorn the belief that the shape +of his own canines, and their occasional great development in other men, +are due to our early progenitors having been provided with these +formidable weapons, will probably reveal by sneering the line of his +descent. For though he no longer intends, nor has the power, to use +these teeth as weapons, he will unconsciously retract his "snarling +muscles" (thus named by Sir C. Bell)[182] so as to expose them ready for +action, like a dog prepared to fight. + +Many muscles are occasionally developed in man, which are proper to the +Quadrumana or other mammals. Professor Vlacovich[183] examined forty +male subjects, and found a muscle, called by him the ischiopubic, in +nineteen of them; in three others there was a ligament which represented +this muscle; and in the remaining eighteen no trace of it. Out of thirty +female subjects this muscle was developed on both sides in only two, but +in three others the rudimentary ligament was present. This muscle, +therefore, appears to be much more common in the male than in the female +sex; and on the principle of the descent of man from some lower form, +its presence can be understood; for it has been detected in several of +the lower animals, and in all of these it serves exclusively to aid the +male in the act of reproduction. + +Mr. J. Wood, in his valuable series of papers,[184] has minutely +described a vast number of muscular variations in man, which resemble +normal structures in the lower animals. Looking only to the muscles +which closely resemble those regularly present in our nearest allies, +the Quadrumana, they are too numerous to be here even specified. In a +single male subject, having a strong bodily frame and well-formed skull, +no less than seven muscular variations were observed, all of which +plainly represented muscles proper to various kinds of apes. This man, +for instance, had on both sides of his neck a true and powerful +"_levator claviculae_," such as is found in all kinds of apes, and which +is said to occur in about one out of sixty human subjects.[185] Again, +this man had "a special abductor of the metatarsal bone of the fifth +digit, such as Professor Huxley and Mr. Flower have shewn to exist +uniformly in the higher and lower apes." The hands and arms of man are +eminently characteristic structures, but their muscles are extremely +liable to vary, so as to resemble the corresponding muscles in the lower +animals.[186] Such resemblances are either complete and perfect or +imperfect, yet in this latter case manifestly of a transitional nature. +Certain variations are more common in man, and others in woman, without +our being able to assign any reason. Mr. Wood, after describing numerous +cases, makes the following pregnant remark: "Notable departures from the +ordinary type of the muscular structures run in grooves or directions, +which must be taken to indicate some unknown factor, of much importance +to a comprehensive knowledge of general and scientific anatomy."[187] + +That this unknown factor is reversion to a former state of existence may +be admitted as in the highest degree probable. It is quite incredible +that a man should through mere accident abnormally resemble, in no less +than seven of his muscles, certain apes, if there had been no genetic +connection between them. On the other hand, if man is descended from +some ape-like creature, no valid reason can be assigned why certain +muscles should not suddenly reappear after an interval of many thousand +generations, in the same manner as with horses, asses, and mules, +dark-coloured stripes suddenly reappear on the legs and shoulders, after +an interval of hundreds, or more probably thousands, of generations. + +These various cases of reversion are so closely related to those of +rudimentary organs given in the first chapter, that many of them might +have been indifferently introduced in either chapter. Thus a human +uterus furnished with cornua may be said to represent in a rudimentary +condition the same organ in its normal state in certain mammals. Some +parts which are rudimental in man, as the os coccyx in both sexes and +the mammae in the male sex, are always present; whilst others, such as +the supra-condyloid foramen, only occasionally appear, and therefore +might have been introduced under the head of reversion. These several +reversionary, as well as the strictly rudimentary, structures reveal the +descent of man from some lower form in an unmistakeable manner. + + +_Correlated Variation._--In man, as in the lower animals, many +structures are so intimately related, that when one part varies so does +another, without our being able, in most cases, to assign any reason. We +cannot say whether the one part governs the other, or whether both are +governed by some earlier developed part. Various monstrosities, as I. +Geoffroy repeatedly insists, are thus intimately connected. Homologous +structures are particularly liable to change together, as we see on the +opposite sides of the body, and in the upper and lower extremities. +Meckel long ago remarked that when the muscles of the arm depart from +their proper type, they almost always imitate those of the leg; and so +conversely with the muscles of the legs. The organs of sight and +hearing, the teeth and hair, the colour of the skin and hair, colour and +constitution, are more or less correlated.[188] Professor Schaaffhausen +first drew attention to the relation apparently existing between a +muscular frame and strongly-pronounced supra-orbital ridges, which are +so characteristic of the lower races of man. + +Besides the variations which can be grouped with more or less +probability under the foregoing heads, there is a large class of +variations which may be provisionally called spontaneous, for they +appear, owing to our ignorance, to arise without any exciting cause. It +can, however, be shewn that such variations, whether consisting of +slight individual differences, or of strongly-marked and abrupt +deviations of structure, depend much more on the constitution of the +organism than on the nature of the conditions to which it has been +subjected.[189] + + +_Rate of Increase._--Civilised populations have been known under +favourable conditions, as in the United States, to double their number +in twenty-five years; and according to a calculation by Euler, this +might occur in a little over twelve years.[190] At the former rate the +present population of the United States, namely, thirty millions, would +in 657 years cover the whole terraqueous globe so thickly, that four men +would have to stand on each square yard of surface. The primary or +fundamental check to the continued increase of man is the difficulty of +gaining subsistence and of living in comfort. We may infer that this is +the case from what we see, for instance, in the United States, where +subsistence is easy and there is plenty of room. If such means were +suddenly doubled in Great Britain, our number would be quickly doubled. +With civilised nations the above primary check acts chiefly by +restraining marriages. The greater death-rate of infants in the poorest +classes is also very important; as well as the greater mortality at all +ages, and from various diseases, of the inhabitants of crowded and +miserable houses. The effects of severe epidemics and wars are soon +counterbalanced, and more than counterbalanced, in nations placed under +favourable conditions. Emigration also comes in aid as a temporary +check, but not to any great extent with the extremely poor classes. + +There is reason to suspect, as Malthus has remarked, that the +reproductive power is actually less in barbarous than in civilised +races. We know nothing positively on this head, for with savages no +census has been taken; but from the concurrent testimony of +missionaries, and of others who have long resided with such people, it +appears that their families are usually small, and large ones rare. This +may be partly accounted for, as it is believed, by the women suckling +their infants for a prolonged period; but it is highly probable that +savages, who often suffer much hardship, and who do not obtain so much +nutritious food as civilised men, would be actually less prolific. I +have shewn in a former work,[191] that all our domesticated quadrupeds +and birds, and all our cultivated plants, are more fertile than the +corresponding species in a state of nature. It is no valid objection to +this conclusion that animals suddenly supplied with an excess of food, +or when rendered very fat, and that most plants when suddenly removed +from very poor to very rich soil, are rendered more or less sterile. We +might, therefore, expect that civilised men, who in one sense are highly +domesticated, would be more prolific than wild men. It is also probable +that the increased fertility of civilised nations would become, as with +our domestic animals, an inherited character: it is at least known that +with mankind a tendency to produce twins runs in families.[192] + +Notwithstanding that savages appear to be less prolific than civilised +people, they would no doubt rapidly increase if their numbers were not +by some means rigidly kept down. The Santali, or hill-tribes of India, +have recently afforded a good illustration of this fact; for they have +increased, as shewn by Mr. Hunter,[193] at an extraordinary rate since +vaccination has been introduced, other pestilences mitigated, and war +sternly repressed. This increase, however, would not have been possible +had not these rude people spread into the adjoining districts and worked +for hire. Savages almost always marry; yet there is some prudential +restraint, for they do not commonly marry at the earliest possible age. +The young men are often required to show that they can support a wife, +and they generally have first to earn the price with which to purchase +her from her parents. With savages the difficulty of obtaining +subsistence occasionally limits their number in a much more direct +manner than with civilised people, for all tribes periodically suffer +from severe famines. At such times savages are forced to devour much bad +food, and their health can hardly fail to be injured. Many accounts have +been published of their protruding stomachs and emaciated limbs after +and during famines. They are then, also, compelled to wander much about, +and their infants, as I was assured in Australia, perish in large +numbers. As famines are periodical, depending chiefly on extreme +seasons, all tribes must fluctuate in number. They cannot steadily and +regularly increase, as there is no artificial increase in the supply of +food. Savages when hardly pressed encroach on each other's territories, +and war is the result; but they are indeed almost always at war with +their neighbours. They are liable to many accidents on land and water in +their search for food; and in some countries they must suffer much from +the larger beasts of prey. Even in India, districts have been +depopulated by the ravages of tigers. + +Malthus has discussed these several checks, but he does not lay stress +enough on what is probably the most important of all, namely +infanticide, especially of female infants, and the habit of procuring +abortion. These practices now prevail in many quarters of the world, and +infanticide seems formerly to have prevailed, as Mr. M'Lennan[194] has +shewn, on a still more extensive scale. These practices appear to have +originated in savages recognising the difficulty, or rather the +impossibility of supporting all the infants that are born. +Licentiousness may also be added to the foregoing checks; but this does +not follow from failing means of subsistence; though there is reason to +believe that in some cases (as in Japan) it has been intentionally +encouraged as a means of keeping down the population. + +If we look back to an extremely remote epoch, before man had arrived at +the dignity of manhood, he would have been guided more by instinct and +less by reason than are savages at the present time. Our early +semi-human progenitors would not have practised infanticide, for the +instincts of the lower animals are never so perverted as to lead them +regularly to destroy their own offspring. There would have been no +prudential restraint from marriage, and the sexes would have freely +united at an early age. Hence the progenitors of man would have tended +to increase rapidly, but checks of some kind, either periodical or +constant, must have kept down their numbers, even more severely than +with existing savages. What the precise nature of these checks may have +been, we cannot say, any more than with most other animals. We know that +horses and cattle, which are not highly prolific animals, when first +turned loose in South America, increased at an enormous rate. The +slowest breeder of all known animals, namely the elephant, would in a +few thousand years stock the whole world. The increase of every species +of monkey must be checked by some means; but not, as Brehm remarks, by +the attacks of beasts of prey. No one will assume that the actual power +of reproduction in the wild horses and cattle of America, was at first +in any sensible degree increased; or that, as each district became fully +stocked, this same power was diminished. No doubt in this case and in +all others, many checks concur, and different checks under different +circumstances; periodical dearths, depending on unfavourable seasons, +being probably the most important of all. So it will have been with the +early progenitors of man. + + +_Natural Selection._--We have now seen that man is variable in body and +mind; and that the variations are induced, either directly or +indirectly, by the same general causes, and obey the same general laws, +as with the lower animals. Man has spread widely over the face of the +earth, and must have been exposed, during his incessant migrations,[195] +to the most diversified conditions. The inhabitants of Tierra del +Fuego, the Cape of Good Hope, and Tasmania in the one hemisphere, and of +the Arctic regions in the other, must have passed through many climates +and changed their habits many times, before they reached their present +homes.[196] The early progenitors of man must also have tended, like all +other animals, to have increased beyond their means of subsistence; they +must therefore occasionally have been exposed to a struggle for +existence, and consequently to the rigid law of natural selection. +Beneficial variations of all kinds will thus, either occasionally or +habitually, have been preserved, and injurious ones eliminated. I do not +refer to strongly-marked deviations of structure, which occur only at +long intervals of time, but to mere individual differences. We know, for +instance, that the muscles of our hands and feet, which determine our +powers of movement, are liable, like those of the lower animals,[197] to +incessant variability. If then the ape-like progenitors of man which +inhabited any district, especially one undergoing some change in its +conditions, were divided into two equal bodies, the one half which +included all the individuals best adapted by their powers of movement +for gaining subsistence or for defending themselves, would on an average +survive in greater number and procreate more offspring than the other +and less well endowed half. + +Man in the rudest state in which he now exists is the most dominant +animal that has ever appeared on the earth. He has spread more widely +than any other highly organised form; and all others have yielded +before him. He manifestly owes this immense superiority to his +intellectual faculties, his social habits, which lead him to aid and +defend his fellows, and to his corporeal structure. The supreme +importance of these characters has been proved by the final arbitrament +of the battle for life. Through his powers of intellect, articulate +language has been evolved; and on this his wonderful advancement has +mainly depended. He has invented and is able to use various weapons, +tools, traps, &c., with which he defends himself, kills or catches prey, +and otherwise obtains food. He has made rafts or canoes on which to fish +or cross over to neighbouring fertile islands. He has discovered the art +of making fire, by which hard and stringy roots can be rendered +digestible, and poisonous roots or herbs innocuous. This last discovery, +probably the greatest, excepting language, ever made by man, dates from +before the dawn of history. These several inventions, by which man in +the rudest state has become so preeminent, are the direct result of the +development of his powers of observation, memory, curiosity, +imagination, and reason. I cannot, therefore, understand how it is that +Mr. Wallace[198] maintains, that "natural selection could only have +endowed the savage with a brain a little superior to that of an ape." + +Although the intellectual powers and social habits of man are of +paramount importance to him, we must not underrate the importance of his +bodily structure, to which subject the remainder of this chapter will be +devoted. The development of the intellectual and social or moral +faculties will be discussed in the following chapter. + +Even to hammer with precision is no easy matter, as every one who has +tried to learn carpentry will admit. To throw a stone with as true an +aim as can a Fuegian in defending himself, or in killing birds, requires +the most consummate perfection in the correlated action of the muscles +of the hand, arm, and shoulder, not to mention a fine sense of touch. In +throwing a stone or spear, and in many other actions, a man must stand +firmly on his feet; and this again demands the perfect coadaptation of +numerous muscles. To chip a flint into the rudest tool, or to form a +barbed spear or hook from a bone, demands the use of a perfect hand; +for, as a most capable judge, Mr. Schoolcraft,[199] remarks, the shaping +fragments of stone into knives, lances, or arrow-heads, shews +"extraordinary ability and long practice." We have evidence of this in +primeval men having practised a division of labour; each man did not +manufacture his own flint tools or rude pottery; but certain individuals +appear to have devoted themselves to such work, no doubt receiving in +exchange the produce of the chase. Archaeologists are convinced that an +enormous interval of time elapsed before our ancestors thought of +grinding chipped flints into smooth tools. A man-like animal who +possessed a hand and arm sufficiently perfect to throw a stone with +precision or to form a flint into a rude tool, could, it can hardly be +doubted, with sufficient practice make almost anything, as far as +mechanical skill alone is concerned, which a civilised man can make. The +structure of the hand in this respect may be compared with that of the +vocal organs, which in the apes are used for uttering various +signal-cries, or, as in one species, musical cadences; but in man +closely similar vocal organs have become adapted through the inherited +effects of use for the utterance of articulate language. + +Turning now to the nearest allies of man, and therefore to the best +representatives of our early progenitors, we find that the hands in the +Quadrumana are constructed on the same general pattern as in us, but are +far less perfectly adapted for diversified uses. Their hands do not +serve so well as the feet of a dog for locomotion; as may be seen in +those monkeys which walk on the outer margins of the palms, or on the +backs of their bent fingers, as in the chimpanzee and orang.[200] Their +hands, however, are admirably adapted for climbing trees. Monkeys seize +thin branches or ropes, with the thumb on one side and the fingers and +palm on the other side, in the same manner as we do. They can thus also +carry rather large objects, such as the neck of a bottle, to their +mouths. Baboons turn over stones and scratch up roots with their hands. +They seize nuts, insects, or other small objects with the thumb in +opposition to the fingers, and no doubt they thus extract eggs and the +young from the nests of birds. American monkeys beat the wild oranges on +the branches until the rind is cracked, and then tear it off with the +fingers of the two hands. Other monkeys open mussel-shells with the two +thumbs. With their fingers they pull out thorns and burrs, and hunt for +each other's parasites. In a state of nature they break open hard fruits +with the aid of stones. They roll down stones or throw them at their +enemies; nevertheless, they perform these various actions clumsily, and +they are quite unable, as I have myself seen, to throw a stone with +precision. + +It seems to me far from true that because "objects are grasped clumsily" +by monkeys, "a much less specialised organ of prehension" would have +served them[201] as well as their present hands. On the contrary, I see +no reason to doubt that a more perfectly constructed hand would have +been an advantage to them, provided, and it is important to note this, +that their hands had not thus been rendered less well adapted for +climbing trees. We may suspect that a perfect hand would have been +disadvantageous for climbing; as the most arboreal monkeys in the world, +namely Ateles in America and Hylobates in Asia, either have their thumbs +much reduced in size and even rudimentary, or their fingers +partially coherent, so that their hands are converted into mere +grasping-hooks.[202] + +As soon as some ancient member in the great series of the Primates came, +owing to a change in its manner of procuring subsistence, or to a change +in the conditions of its native country, to live somewhat less on trees +and more on the ground, its manner of progression would have been +modified; and in this case it would have had to become either more +strictly quadrupedal or bipedal. Baboons frequent hilly and rocky +districts, and only from necessity climb up high trees;[203] and they +have acquired almost the gait of a dog. Man alone has become a biped; +and we can, I think, partly see how he has come to assume his erect +attitude, which forms one of the most conspicuous differences between +him and his nearest allies. Man could not have attained his present +dominant position in the world without the use of his hands which are so +admirably adapted to act in obedience to his will. As Sir C. Bell[204] +insists "the hand supplies all instruments, and by its correspondence +with the intellect gives him universal dominion." But the hands and arms +could hardly have become perfect enough to have manufactured weapons, or +to have hurled stones and spears with a true aim, as long as they were +habitually used for locomotion and for supporting the whole weight of +the body, or as long as they were especially well adapted, as previously +remarked, for climbing trees. Such rough treatment would also have +blunted the sense of touch, on which their delicate use largely depends. +From these causes alone it would have been an advantage to man to have +become a biped; but for many actions it is almost necessary that both +arms and the whole upper part of the body should be free; and he must +for this end stand firmly on his feet. To gain this great advantage, the +feet have been rendered flat, and the great toe peculiarly modified, +though this has entailed the loss of the power of prehension. It accords +with the principle of the division of physiological labour, which +prevails throughout the animal kingdom, that as the hands became +perfected for prehension, the feet should have become perfected for +support and locomotion. With some savages, however, the foot has not +altogether lost its prehensile power, as shewn by their manner of +climbing trees and of using them in other ways.[205] + +If it be an advantage to man to have his hands and arms free and to +stand firmly on his feet, of which there can be no doubt from his +preeminent success in the battle of life, then I can see no reason why +it should not have been advantageous to the progenitors of man to have +become more and more erect or bipedal. They would thus have been better +able to have defended themselves with stones or clubs, or to have +attacked their prey, or otherwise obtained food. The best constructed +individuals would in the long run have succeeded best, and have survived +in larger numbers. If the gorilla and a few allied forms had become +extinct, it might have been argued with great force and apparent truth, +that an animal could not have been gradually converted from a quadruped +into a biped; as all the individuals in an intermediate condition would +have been miserably ill-fitted for progression. But we know (and this is +well worthy of reflection) that several kinds of apes are now actually +in this intermediate condition; and no one doubts that they are on the +whole well adapted for their conditions of life. Thus the gorilla runs +with a sidelong shambling gait, but more commonly progresses by resting +on its bent hands. The long-armed apes occasionally use their arms like +crutches, swinging their bodies forward between them, and some kinds of +Hylobates, without having been taught, can walk or run upright with +tolerable quickness; yet they move awkwardly, and much less securely +than man. We see, in short, with existing monkeys various gradations +between a form of progression strictly like that of a quadruped and that +of a biped or man. + +As the progenitors of man became more and more erect, with their hands +and arms more and more modified for prehension and other purposes, with +their feet and legs at the same time modified for firm support and +progression, endless other changes of structure would have been +necessary. The pelvis would have had to be made broader, the spine +peculiarly curved and the head fixed in an altered position, and all +these changes have been attained by man. Prof. Schaaffhausen[206] +maintains that "the powerful mastoid processes of the human skull are +the result of his erect position;" and these processes are absent in the +orang, chimpanzee, &c., and are smaller in the gorilla than in man. +Various other structures might here have been specified, which appear +connected with man's erect position. It is very difficult to decide how +far all these correlated modifications are the result of natural +selection, and how far of the inherited effects of the increased use of +certain parts, or of the action of one part on another. No doubt these +means of change act and react on each other: thus when certain muscles, +and the crests of bone to which they are attached, become enlarged by +habitual use, this shews that certain fictions are habitually performed +and must be serviceable. Hence the individuals which performed them +best, would tend to survive in greater numbers. + +The free use of the arms and hands, partly the cause and partly the +result of man's erect position, appears to have led in an indirect +manner to other modifications of structure. The early male progenitors +of man were, as previously stated, probably furnished with great canine +teeth; but as they gradually acquired the habit of using stones, clubs, +or other weapons, for fighting with their enemies, they would have used +their jaws and teeth less and less. In this case, the jaws, together +with the teeth, would have become reduced in size, as we may feel sure +from innumerable analogous cases. In a future chapter we shall meet with +a closely-parallel case, in the reduction or complete disappearance of +the canine teeth in male ruminants, apparently in relation with the +development of their horns; and in horses, in relation with their habit +of fighting with their incisor teeth and hoofs. + +In the adult male anthropomorphous apes, as Ruetimeyer,[207] and others +have insisted, it is precisely the effect which the jaw-muscles by their +great development have produced on the skull, that causes it to differ +so greatly in many respects from that of man, and has given to it "a +truly frightful physiognomy." Therefore as the jaws and teeth in the +progenitors of man gradually become reduced in size, the adult skull +would have presented nearly the same characters which it offers in the +young of the anthropomorphous apes, and would thus have come to resemble +more nearly that of existing man. A great reduction of the canine teeth +in the males would almost certainly, as we shall hereafter see, have +affected through inheritance the teeth of the females. + +As the various mental faculties were gradually developed, the brain +would almost certainly have become larger. No one, I presume, doubts +that the large size of the brain in man, relatively to his body, in +comparison with that of the gorilla or orang, is closely connected with +his higher mental powers. We meet with closely analogous facts with +insects, in which the cerebral ganglia are of extraordinary dimensions +in ants; these ganglia in all the Hymenoptera being many times larger +than in the less intelligent orders, such as beetles.[208] On the other +hand, no one supposes that the intellect of any two animals or of any +two men can be accurately gauged by the cubic contents of their skulls. +It is certain that there may be extraordinary mental activity with an +extremely small absolute mass of nervous matter: thus the wonderfully +diversified instincts, mental powers, and affections of ants are +generally known, yet their cerebral ganglia are not so large as the +quarter of a small pin's head. Under this latter point of view, the +brain of an ant is one of the most marvellous atoms of matter in the +world, perhaps more marvellous than the brain of man. + +The belief that there exists in man some close relation between the size +of the brain and the development of the intellectual faculties is +supported by the comparison of the skulls of savage and civilised races, +of ancient and modern people, and by the analogy of the whole +vertebrate series. Dr. J. Barnard Davis has proved[209] by many careful +measurements, that the mean internal capacity of the skull in Europeans +is 92.3 cubic inches; in Americans 87.5; in Asiatics 87.1; and in +Australians only 81.9 inches. Professor Broca[210] found that skulls +from graves in Paris of the nineteenth century, were larger than those +from vaults of the twelfth century, in the proportion of 1484 to 1426; +and Prichard is persuaded that the present inhabitants of Britain have +"much more capacious brain-cases" than the ancient inhabitants. +Nevertheless it must be admitted that some skulls of very high +antiquity, such as the famous one of Neanderthal, are well developed and +capacious. With respect to the lower animals, M. E. Lartet,[211] by +comparing the crania of tertiary and recent mammals, belonging to the +same groups, has come to the remarkable conclusion that the brain is +generally larger and the convolutions more complex in the more recent +form. On the other hand I have shewn[212] that the brains of domestic +rabbits are considerably reduced in bulk, in comparison with those of +the wild rabbit or hare; and this may be attributed to their having been +closely confined during many generations, so that they have exerted but +little their intellect, instincts, senses, and voluntary movements. + +The gradually increasing weight of the brain and skull in man must have +influenced the development of the supporting spinal column, more +especially whilst he was becoming erect. As this change of position was +being brought about, the internal pressure of the brain, will, also, +have influenced the form of the skull; for many facts shew how easily +the skull is thus affected. Ethnologists believe that it is modified by +the kind of cradle in which infants sleep. Habitual spasms of the +muscles and a cicatrix from a severe burn have permanently modified the +facial bones. In young persons whose heads from disease have become +fixed either sideways or backwards, one of the eyes has changed its +position, and the bones of the skull have been modified; and this +apparently results from the brain pressing in a new direction.[213] I +have shewn that with long-eared rabbits, even so trifling a cause as the +lopping forward of one ear drags forward on that side almost every bone +of the skull; so that the bones on the opposite sides no longer strictly +correspond. Lastly, if any animal were to increase or diminish much in +general size, without any change in its mental powers; or if the mental +powers were to be much increased or diminished without any great change +in the size of the body; the shape of the skull would almost certainly +be altered. I infer this from my observations on domestic rabbits, some +kinds of which have become very much larger than the wild animal, whilst +others have retained nearly the same size, but in both cases the brain +has been much reduced relatively to the size of the body. Now I was at +first much surprised by finding that in all these rabbits the skull had +become elongated or dolichocephalic; for instance, of two skulls of +nearly equal breadth, the one from a wild rabbit and the other from a +large domestic kind, the former was only 3.15 and the latter 4.3 inches +in length.[214] One of the most marked distinctions in different races +of man is that the skull in some is elongated, and in others rounded; +and here the explanation suggested by the case of the rabbits may +partially hold good; for Welcker finds that short "men incline more to +brachycephaly, and tall men to dolichocephaly;"[215] and tall men may be +compared with the larger and longer-bodied rabbits, all of which have +elongated skulls, or are dolichocephalic. + +From these several facts we can to a certain extent understand the means +through which the great size and more or less rounded form of the skull +has been acquired by man; and these are characters eminently distinctive +of him in comparison with the lower animals. + +Another most conspicuous difference between man and the lower animals is +the nakedness of his skin. Whales and dolphins (Cetacea), dugongs +(Sirenia) and the hippopotamus are naked; and this may be advantageous +to them for gliding through the water; nor would it be injurious to them +from the loss of warmth, as the species which inhabit the colder regions +are protected by a thick layer of blubber, serving the same purpose as +the fur of seals and otters. Elephants and rhinoceroses are almost +hairless; and as certain extinct species which formerly lived under an +arctic climate were covered with long wool or hair, it would almost +appear as if the existing species of both genera had lost their hairy +covering from exposure to heat. This appears the more probable, as the +elephants in India which live on elevated and cool districts are more +hairy[216] than those on the lowlands. May we then infer that man became +divested of hair from having aboriginally inhabited some tropical land? +The fact of the hair being chiefly retained in the male sex on the chest +and face, and in both sexes at the junction of all four limbs with the +trunk, favours this inference, assuming that the hair was lost before +man became erect; for the parts which now retain most hair would then +have been most protected from the heat of the sun. The crown of the +head, however, offers a curious exception, for at all times it must have +been one of the most exposed parts, yet it is thickly clothed with hair. +In this respect man agrees with the great majority of quadrupeds, which +generally have their upper and exposed surfaces more thickly clothed +than the lower surface. Nevertheless, the fact that the other members of +the order of Primates, to which man belongs, although inhabiting various +hot regions, are well clothed with hair, generally thickest on the upper +surface,[217] is strongly opposed to the supposition that man became +naked through the action of the sun. I am inclined to believe, as we +shall see under sexual selection, that man, or rather primarily woman, +became divested of hair for ornamental purposes; and according to this +belief it is not surprising that man should differ so greatly in +hairiness from all his lower brethren, for characters gained through +sexual selection often differ in closely-related forms to an +extraordinary degree. + +According to a popular impression, the absence of a tail is eminently +distinctive of man; but as those apes which come nearest to man are +destitute of this organ, its disappearance does not especially concern +us. Nevertheless it may be well to own that no explanation, as far as I +am aware, has ever been given of the loss of the tail by certain apes +and man. Its loss, however, is not surprising, for it sometimes differs +remarkably in length in species of the same genera: thus in some species +of Macacus the tail is longer than the whole body, consisting of +twenty-four vertebrae; in others it consists of a scarcely visible stump, +containing only three or four vertebrae. In some kinds of baboons there +are twenty-five, whilst in the mandrill there are ten very small stunted +caudal vertebrae, or, according to Cuvier,[218] sometimes only five. This +great diversity in the structure and length of the tail in animals +belonging to the same genera, and following nearly the same habits of +life, renders it probable that the tail is not of much importance to +them; and if so, we might have expected that it would sometimes have +become more or less rudimentary, in accordance with what we incessantly +see with other structures. The tail almost always tapers towards the end +whether it be long or short; and this, I presume, results from the +atrophy, through disuse, of the terminal muscles together with their +arteries and nerves, leading to the atrophy of the terminal bones. With +respect to the os coccyx, which in man and the higher apes manifestly +consists of the few basal and tapering segments of an ordinary tail, I +have heard it asked how could these have become completely embedded +within the body; but there is no difficulty in this respect, for in many +monkeys the basal segments of the true tail are thus embedded. For +instance, Mr. Murie informs me that in the skeleton of a not full-grown +_Macacus inornatus_, he counted nine or ten caudal vertebrae, which +altogether were only 1.8 inch in length. Of these the three basal ones +appeared to have been embedded; the remainder forming the free part of +the tail, which was only one inch in length, and half an inch in +diameter. Here, then, the three embedded caudal vertebrae plainly +correspond with the four coalesced vertebrae of the human os coccyx. + + +I have now endeavoured to shew that some of the most distinctive +characters of man have in all probability been acquired, either +directly, or more commonly indirectly, through natural selection. We +should bear in mind that modifications in structure or constitution, +which are of no service to an organism in adapting it to its habits of +life, to the food which it consumes, or passively to the surrounding +conditions, cannot have been thus acquired. We must not, however, be too +confident in deciding what modifications are of service to each being: +we should remember how little we know about the use of many parts, or +what changes in the blood or tissues may serve to fit an organism for a +new climate or some new kind of food. Nor must we forget the principle +of correlation, by which, as Isidore Geoffroy has shewn in the case of +man, many fit-range deviations of structure are tied together. +Independently of correlation, a change in one part often leads through +the increased or decreased use of other parts, to other changes of a +quite unexpected nature. It is also well to reflect on such facts, as +the wonderful growth of galls on plants caused by the poison of an +insect, and on the remarkable changes of colour in the plumage of +parrots when fed on certain fishes, or inoculated with the poison of +toads;[219] for we can thus see that the fluids of the system, if +altered for some special purpose, might induce other strange changes. We +should especially bear in mind that modifications acquired and +continually used during past ages for some useful purpose would probably +become firmly fixed and might be long inherited. + +Thus a very large yet undefined extension may safely be given to the +direct and indirect results of natural selection; but I now admit, after +reading the essay by Naegeli on plants, and the remarks by various +authors with respect to animals, more especially those recently made by +Professor Broca, that in the earlier editions of my 'Origin of Species' +I probably attributed too much to the action of natural selection or the +survival of the fittest. I have altered the fifth edition of the Origin +so as to confine my remarks to adaptive changes of structure. I had not +formerly sufficiently considered the existence of many structures which +appear to be, as far as we can judge, neither beneficial nor injurious; +and this I believe to be one of the greatest oversights as yet detected +in my work. I may be permitted to say as some excuse, that I had two +distinct objects in view, firstly, to shew that species had not been +separately created, and secondly, that natural selection had been the +chief agent of change, though largely aided by the inherited effects of +habit, and slightly by the direct action of the surrounding conditions. +Nevertheless I was not able to annul the influence of my former belief, +then widely prevalent, that each species had been purposely created; and +this led to my tacitly assuming that every detail of structure, +excepting rudiments, was of some special, though unrecognised, service. +Any one with this assumption in his mind would naturally extend the +action of natural selection, either during past or present times, too +far. Some of those who admit the principle of evolution, but reject +natural selection, seem to forget, when criticising my book, that I had +the above two objects in view; hence if I have erred in giving to +natural selection great power, which I am far from admitting, or in +having exaggerated its power, which is in itself probable, I have at +least, as I hope, done good service in aiding to overthrow the dogma of +separate creations. + +That all organic beings, including man, present many modifications of +structure which are of no service to them at present, nor have been +formerly, is, as I can now see, probable. We know not what produces the +numberless slight differences between the individuals of each species, +for reversion only carries the problem a few steps backwards; but each +peculiarity must have had its own efficient cause. If these causes, +whatever they may be, were to act more uniformly and energetically +during a lengthened period (and no reason can be assigned why this +should not sometimes occur), the result would probably be not mere +slight individual differences, but well-marked, constant modifications. +Modifications which are in no way beneficial cannot have been kept +uniform through natural selection, though any which were injurious would +have been thus eliminated. Uniformity of character would, however, +naturally follow from, the assumed uniformity of the exciting causes, +and likewise from the free intercrossing of many individuals. The same +organism might acquire in this manner during successive periods +successive modifications, and these would be transmitted in a nearly +uniform state as long as the exciting causes remained the same and there +was free intercrossing. With respect to the exciting causes we can only +say, as when speaking of so-called spontaneous variations, that they +relate much more closely to the constitution of the varying organism, +than to the nature of the conditions to which it has been subjected. + + +_Conclusion._--In this chapter we have seen that as man at the present +day is liable, like every other animal, to multiform individual +differences or slight variations, so no doubt were the early progenitors +of man; the variations being then as now induced by the same general +causes, and governed by the same general and complex laws. As all +animals tend to multiply beyond their means of subsistence, so it must +have been with the progenitors of man; and this will inevitably have led +to a struggle for existence and to natural selection. This latter +process will have been greatly aided by the inherited effects of the +increased use of parts; these two processes incessantly reacting on each +other. It appears, also, as we shall hereafter see, that various +unimportant characters have been acquired by man through sexual +selection. An unexplained residuum of change, perhaps a large one, must +be left to the assumed uniform action of those unknown agencies, which +occasionally induce strongly-marked and abrupt deviations of structure +in our domestic productions. + +Judging from the habits of savages and of the greater number of the +Quadrumana, primeval men, and even the ape-like progenitors of man, +probably lived in society. With strictly social animals, natural +selection sometimes acts indirectly on the individual, through the +preservation of variations which are beneficial only to the community. A +community including a large number of well-endowed individuals increases +in number and is victorious over other and less well-endowed +communities; although each separate member may gain no advantage over +the other members of the same community. With associated insects many +remarkable structures, which are of little or no service to the +individual or its own offspring, such as the pollen-collecting +apparatus, or the sting of the worker-bee, or the great jaws of +soldier-ants, have been thus acquired. With the higher social animals, I +am not aware that any structure has been modified solely for the good of +the community, though some are of secondary service to it. For instance, +the horns of ruminants and the great canine teeth of baboons appear to +have been acquired by the males as weapons for sexual strife, but they +are used in defence of the herd or troop. In regard to certain mental +faculties the case, as we shall see in the following chapter, is wholly +different; for these faculties have been chiefly, or even exclusively, +gained for the benefit of the community; the individuals composing the +community being at the same time indirectly benefited. + +It has often been objected to such views as the foregoing, that man is +one of the most helpless and defenceless creatures in the world; and +that during his early and less well-developed condition he would have +been still more helpless. The Duke of Argyll, for instance, insists[220] +that "the human frame has diverged from the structure of brutes, in the +direction of greater physical helplessness and weakness. That is to say, +it is a divergence which of all others it is most impossible to ascribe +to mere natural selection." He adduces the naked and unprotected state +of the body, the absence of great teeth or claws for defence, the little +strength of man, his small speed in running, and his slight power of +smell, by which to discover food or to avoid danger. To these +deficiencies there might have been added the still more serious loss of +the power of quickly climbing trees, so as to escape from enemies. +Seeing that the unclothed Fuegians can exist under their wretched +climate, the loss of hair would not have been a great injury to primeval +man, if he inhabited a warm country. When we compare defenceless man +with the apes, many of which are provided with formidable canine teeth, +we must remember that these in their fully-developed condition are +possessed by the males alone, being chiefly used by them for fighting +with their rivals; yet the females which are not thus provided, are able +to survive. + +In regard to bodily size or strength, we do not know whether man is +descended from some comparatively small species, like the chimpanzee, or +from one as powerful as the gorilla; and, therefore, we cannot say +whether man has become larger and stronger, or smaller and weaker, in +comparison with his progenitors. We should, however, bear in mind that +an animal possessing great size, strength, and ferocity, and which, like +the gorilla, could defend itself from all enemies, would probably, +though not necessarily, have failed to become social; and this would +most effectually have checked the acquirement by man of his higher +mental qualities, such as sympathy and the love of his fellow-creatures. +Hence it might have been an immense advantage to man to have sprung +from some comparatively weak creature. + +The slight corporeal strength of man, his little speed, his want of +natural weapons, &c., are more than counterbalanced, firstly by his +intellectual powers, through which he has, whilst still remaining in a +barbarous state, formed for himself weapons, tools, &c., and secondly by +his social qualities which lead him to give aid to his fellow-men and to +receive it in return. No country in the world abounds in a greater +degree with dangerous beasts than Southern Africa; no country presents +more fearful physical hardships than the Arctic regions; yet one of the +puniest races, namely, the Bushmen, maintain themselves in Southern +Africa, as do the dwarfed Esquimaux in the Arctic regions. The early +progenitors of man were, no doubt, inferior in intellect, and probably +in social disposition, to the lowest existing savages; but it is quite +conceivable that they might have existed, or even flourished, if, whilst +they gradually lost their brute-like powers, such as climbing trees, +&c., they at the same time advanced in intellect. But granting that the +progenitors of man were far more helpless and defenceless than any +existing savages, if they had inhabited some warm continent or large +island, such as Australia or New Guinea, or Borneo (the latter island +being now tenanted by the orang), they would not have been exposed to +any special danger. In an area as large as one of these islands, the +competition between tribe and tribe would have been sufficient, under +favourable conditions, to have raised man, through the survival of the +fittest, combined with the inherited effects of habit, to his present +high position in the organic scale. + + + + +CHAPTER V. + +ON THE DEVELOPMENT OF THE INTELLECTUAL AND MORAL FACULTIES DURING +PRIMEVAL AND CIVILISED TIMES. + + + The advancement of the intellectual powers through natural + selection--Importance of imitation--Social and moral + faculties--Their development within the limits of the same + tribe--Natural selection as affecting civilised + nations--Evidence that civilised nations were once barbarous. + + +The subjects to be discussed in this chapter are of the highest +interest, but are treated by me in a most imperfect and fragmentary +manner. Mr. Wallace, in an admirable paper before referred to,[221] +argues that man after he had partially acquired those intellectual and +moral faculties which distinguish him from the lower animals, would have +been but little liable to have had his bodily structure modified through +natural selection or any other means. For man is enabled through his +mental faculties "to keep with an unchanged body in harmony with the +changing universe." He has great power of adapting his habits to new +conditions of life. He invents weapons, tools and various stratagems, by +which he procures food and defends himself. When he migrates into a +colder climate he uses clothes, builds sheds, and makes fires; and, by +the aid of fire, cooks food otherwise indigestible. He aids his +fellow-men in many ways, and anticipates future events. Even at a remote +period he practised some subdivision of labour. + +The lower animals, on the other hand, must have their bodily structure +modified in order to survive under greatly changed conditions. They must +be rendered stronger, or acquire more effective teeth or claws, in order +to defend themselves from new enemies; or they must be reduced in size +so as to escape detection and danger. When they migrate into a colder +climate they must become clothed with thicker fur, or have their +constitutions altered. If they fail to be thus modified, they will cease +to exist. + +The case, however, is widely different, as Mr. Wallace has with justice +insisted, in relation to the intellectual and moral faculties of man. +These faculties are variable; and we have every reason to believe that +the variations tend to be inherited. Therefore, if they were formerly of +high importance to primeval man and to his ape-like progenitors, they +would have been perfected or advanced through natural selection. Of the +high importance of the intellectual faculties there can be no doubt, for +man mainly owes to them his preeminent position in the world. We can +see that, in the rudest state of society, the individuals who were the +most sagacious, who invented and used the best weapons or traps, and who +were best able to defend themselves, would rear the greatest number of +offspring. The tribes which included the largest number of men thus +endowed would increase in number and supplant other tribes. Numbers +depend primarily on the means of subsistence, and this, partly on the +physical nature of the country, but in a much higher degree on the arts +which are there practised. As a tribe increases and is victorious, it is +often still further increased by the absorption of other tribes.[222] +The stature and strength of the men of a tribe are likewise of some +importance for its success, and these depend in part on the nature and +amount of the food which can be obtained. In Europe the men of the +Bronze period were supplanted by a more powerful and, judging from their +sword-handles, larger-handed race;[223] but their success was probably +due in a much higher degree to their superiority in the arts. + +All that we know about savages, or may infer from their traditions and +from old monuments, the history of which is quite forgotten by the +present inhabitants, shew that from the remotest times successful tribes +have supplanted other tribes. Relics of extinct or forgotten tribes have +been discovered throughout the civilised regions of the earth, on the +wild plains of America, and on the isolated islands in the Pacific +Ocean. At the present day civilised nations are everywhere supplanting +barbarous nations, excepting where the climate opposes a deadly barrier; +and they succeed mainly, though not exclusively, through their arts, +which are the products of the intellect. It is, therefore, highly +probable that with mankind the intellectual faculties have been +gradually perfected through natural selection; and this conclusion is +sufficient for our purpose. Undoubtedly it would have been very +interesting to have traced the development of each separate faculty from +the state in which it exists in the lower animals to that in which it +exists in man; but neither my ability nor knowledge permit the attempt. + +It deserves notice that as soon as the progenitors of man became social +(and this probably occurred at a very early period), the advancement of +the intellectual faculties will have been aided and modified in an +important manner, of which we see only traces in the lower animals, +namely, through the principle of imitation, together with reason and +experience. Apes are much given to imitation, as are the lowest savages; +and the simple fact previously referred to, that after a time no animal +can be caught in the same place by the same sort of trap, shews that +animals learn by experience, and imitate each others' caution. Now, if +some one man in a tribe, more sagacious than the others, invented a new +snare or weapon, or other means of attack or defence, the plainest +self-interest, without the assistance of much reasoning power, would +prompt the other members to imitate him; and all would thus profit. The +habitual practice of each new art must likewise in some slight degree +strengthen the intellect. If the new invention were an important one, +the tribe would increase in number, spread, and supplant other tribes. +In a tribe thus rendered more numerous there would always be a rather +better chance of the birth of other superior and inventive members. If +such men left children to inherit their mental superiority, the chance +of the birth of still more ingenious members would be somewhat better, +and in a very small tribe decidedly better. Even if they left no +children, the tribe would still include their blood-relations; and it +has been ascertained by agriculturists[224] that by preserving and +breeding from the family of an animal, which when slaughtered was found +to be valuable, the desired character has been obtained. + + +Turning now to the social and moral faculties. In order that primeval +men, or the ape-like progenitors of man, should have become social, they +must have acquired the same instinctive feelings which impel other +animals to live in a body; and they no doubt exhibited the same general +disposition. They would have felt uneasy when separated from their +comrades, for whom they would have felt some degree of love; they would +have warned each other of danger, and have given mutual aid in attack or +defence. All this implies some degree of sympathy, fidelity, and +courage. Such social qualities, the paramount importance of which to the +lower animals is disputed by no one, were no doubt acquired by the +progenitors of man in a similar manner, namely, through natural +selection, aided by inherited habit. When two tribes of primeval man, +living in the same country, came into competition, if the one tribe +included (other circumstances being equal) a greater number of +courageous, sympathetic, and faithful members, who were always ready to +warn each other of danger, to aid and defend each other, this tribe +would without doubt succeed best and conquer the other. Let it be borne +in mind how all-important, in the never-ceasing wars of savages, +fidelity and courage must be. The advantage which disciplined soldiers +have over undisciplined hordes follows chiefly from the confidence which +each man feels in his comrades. Obedience, as Mr. Bagehot has well +shewn,[225] is of the highest value, for any form of government is +better than none. Selfish and contentious people will not cohere, and +without coherence nothing can be effected. A tribe possessing the above +qualities in a high degree would spread and be victorious over other +tribes; but in the course of time it would, judging from all past +history, be in its turn overcome by some other and still more highly +endowed tribe. Thus the social and moral qualities would tend slowly to +advance and be diffused throughout the world. + +But it may be asked, how within the limits of the same tribe did a large +number of members first become endowed with these social and moral +qualities, and how was the standard of excellence raised? It is +extremely doubtful whether the offspring of the more sympathetic and +benevolent parents, or of those which were the most faithful to their +comrades, would be reared in greater number than the children of selfish +and treacherous parents of the same tribe. He who was ready to sacrifice +his life, as many a savage has been, rather than betray his comrades, +would often leave no offspring to inherit his noble nature. The bravest +men, who were always willing to come to the front in war, and who freely +risked their lives for others, would on an average perish in larger +number than other men. Therefore it seems scarcely possible (bearing in +mind that we are not here speaking of one tribe being victorious over +another) that the number of men gifted with such virtues, or that the +standard of their excellence, could be increased through natural +selection, that is, by the survival of the fittest. + +Although the circumstances which lead to an increase in the number of +men thus endowed within the same tribe are too complex to be clearly +followed out, we can trace some of the probable steps. In the first +place, as the reasoning powers and foresight of the members became +improved, each man would soon learn from experience that if he aided his +fellow-men, he would commonly receive aid in return. From this low +motive he might acquire the habit of aiding his fellows; and the habit +of performing benevolent actions certainly strengthens the feeling of +sympathy, which gives the first impulse to benevolent actions. Habits, +moreover, followed during many generations probably tend to be +inherited. + +But there is another and much more powerful stimulus to the development +of the social virtues, namely, the praise and the blame of our +fellow-men. The love of approbation and the dread of infamy, as well as +the bestowal of praise or blame, are primarily due, as we have seen in +the third chapter, to the instinct of sympathy; and this instinct no +doubt was originally acquired, like all the other social instincts, +through natural selection. At how early a period the progenitors of man, +in the course of their development, became capable of feeling and being +impelled by the praise or blame of their fellow-creatures, we cannot, of +course, say. But it appears that even dogs appreciate encouragement, +praise, and blame. The rudest savages feel the sentiment of glory, as +they clearly show by preserving the trophies of their prowess, by their +habit of excessive boasting, and even by the extreme care which they +take of their personal appearance and decorations; for unless they +regarded the opinion of their comrades, such habits would be senseless. + +They certainly feel shame at the breach of some of their lesser rules; +but how far they experience remorse is doubtful. I was at first +surprised that I could not recollect any recorded instances of this +feeling in savages; and Sir J. Lubbock[226] states that he knows of +none. But if we banish from our minds all cases given in novels and +plays and in death-bed confessions made to priests, I doubt whether many +of us have actually witnessed remorse; though we may have often seen +shame and contrition for smaller offences. Remorse is a deeply hidden +feeling. It is incredible that a savage, who will sacrifice his life +rather than betray his tribe, or one who will deliver himself up as a +prisoner rather than break his parole,[227] would not feel remorse in +his inmost soul, though he might conceal it, if he had failed in a duty +which he held sacred. + +We may therefore conclude that primeval man, at a very remote period, +would have been influenced by the praise and blame of his fellows. It is +obvious, that the members of the same tribe would approve of conduct +which appeared to them to be for the general good, and would reprobate +that which appeared evil. To do good unto others--to do unto others as +ye would they should do unto you,--is the foundation-stone of morality. +It is, therefore, hardly possible to exaggerate the importance during +rude times of the love of praise and the dread of blame. A man who was +not impelled by any deep, instinctive feeling, to sacrifice his life for +the good of others, yet was roused to such actions by a sense of glory, +would by his example excite the same wish for glory in other men, and +would strengthen by exercise the noble feeling of admiration. He might +thus do far more good to his tribe than by begetting offspring with a +tendency to inherit his own high character. + +With increased experience and reason, man perceives the more remote +consequences of his actions, and the self-regarding virtues, such as +temperance, chastity, &c., which during early times are, as we have +before seen, utterly disregarded, come to be highly esteemed or even +held sacred. I need not, however, repeat what I have said on this head +in the third chapter. Ultimately a highly complex sentiment, having its +first origin in the social instincts, largely guided by the approbation +of our fellow-men, ruled by reason, self-interest, and in later times by +deep religious feelings, confirmed by instruction and habit, all +combined, constitute our moral sense or conscience. + +It must not be forgotten that although a high standard of morality gives +but a slight or no advantage to each individual man and his children +over the other men of the same tribe, yet that an advancement in the +standard of morality and an increase in the number of well-endowed men +will certainly give an immense advantage to one tribe over another. +There can be no doubt that a tribe including many members who, from +possessing in a high degree the spirit of patriotism, fidelity, +obedience, courage, and sympathy, were always ready to give aid to each +other and to sacrifice themselves for the common good, would be +victorious over most other tribes; and this would be natural selection. +At all times throughout the world tribes have supplanted other tribes; +and as morality is one element in their success, the standard of +morality and the number of well-endowed men will thus everywhere tend to +rise and increase. + +It is, however, very difficult to form any judgment why one particular +tribe and not another has been successful and has risen in the scale of +civilisation. Many savages are in the same condition as when first +discovered several centuries ago. As Mr. Bagehot has remarked, we are +apt to look at progress as the normal rule in human society; but history +refutes this. The ancients did not even entertain the idea; nor do the +oriental nations at the present day. According to another high +authority, Mr. Maine,[228] "the greatest part of mankind has never +shewn a particle of desire that its civil institutions should be +improved." Progress seems to depend on many concurrent favourable +conditions, far too complex to be followed out. But it has often been +remarked, that a cool climate from leading to industry and the various +arts has been highly favourable, or even indispensable for this end. The +Esquimaux, pressed by hard necessity, have succeeded in many ingenious +inventions, but their climate has been too severe for continued +progress. Nomadic habits, whether over wide plains, or through the dense +forests of the tropics, or along the shores of the sea, have in every +case been highly detrimental. Whilst observing the barbarous inhabitants +of Tierra del Fuego, it struck me that the possession of some property, +a fixed abode, and the union of many families under a chief, were the +indispensable requisites for civilisation. Such habits almost +necessitate the cultivation of the ground; and the first steps in +cultivation would probably result, as I have elsewhere shewn,[229] from +some such accident as the seeds of a fruit-tree falling on a heap of +refuse and producing an unusually fine variety. The problem, however, of +the first advance of savages towards civilisation is at present much too +difficult to be solved. + + +_Natural Selection as affecting Civilised Nations._--In the last and +present chapters I have considered the advancement of man from a former +semi-human condition to his present state as a barbarian. But some +remarks on the agency of natural selection on civilised nations may be +here worth adding. This subject has been ably discussed by Mr. W. R. +Greg,[230] and previously by Mr. Wallace and Mr. Galton.[231] Most of +my remarks are taken from these three authors. With savages, the weak in +body or mind are soon eliminated; and those that survive commonly +exhibit a vigorous state of health. We civilised men, on the other hand, +do our utmost to check the process of elimination; we build asylums for +the imbecile, the maimed, and the sick; we institute poor-laws; and our +medical men exert their utmost skill to save the life of every one to +the last moment. There is reason to believe that vaccination has +preserved thousands, who from a weak constitution would formerly have +succumbed to small-pox. Thus the weak members of civilised societies +propagate their kind. No one who has attended to the breeding of +domestic animals will doubt that this must be highly injurious to the +race of man. It is surprising how soon a want of care, or care wrongly +directed, leads to the degeneration of a domestic race; but excepting in +the case of man himself, hardly any one is so ignorant as to allow his +worst animals to breed. + +The aid which we feel impelled to give to the helpless is mainly an +incidental result of the instinct of sympathy, which was originally +acquired as part of the social instincts, but subsequently rendered, in +the manner previously indicated, more tender and more widely diffused. +Nor could we check our sympathy, if so urged by hard reason, without +deterioration in the noblest part of our nature. The surgeon may harden +himself whilst performing an operation, for he knows that he is acting +for the good of his patient; but if we were intentionally to neglect the +weak and helpless, it could only be for a contingent benefit, with a +certain and great present evil. Hence we must bear without complaining +the undoubtedly bad effects of the weak surviving and propagating their +kind; but there appears to be at least one check in steady action, +namely the weaker and inferior members of society not marrying so freely +as the sound; and this check might be indefinitely increased, though +this is more to be hoped for than expected, by the weak in body or mind +refraining from marriage. + +In all civilised countries man accumulates property and bequeaths it to +his children. So that the children in the same country do not by any +means start fair in the race for success. But this is far from an +unmixed evil; for without the accumulation of capital the arts could not +progress; and it is chiefly through their power that the civilised races +have extended, and are now everywhere extending, their range, so as to +take the place of the lower races. Nor does the moderate accumulation of +wealth interfere with the process of selection. When a poor man becomes +rich, his children enter trades or professions in which there is +struggle enough, so that the able in body and mind succeed best. The +presence of a body of well-instructed men, who have not to labour for +their daily bread, is important to a degree which cannot be +over-estimated; as all high intellectual work is carried on by them, and +on such work material progress of all kinds mainly depends, not to +mention other and higher advantages. No doubt wealth when very great +tends to convert men into useless drones, but their number is never +large; and some degree of elimination here occurs, as we daily see rich +men, who happen to be fools or profligate, squandering away all their +wealth. + +Primogeniture with entailed estates is a more direct evil, though it may +formerly have been a great advantage by the creation of a dominant +class, and any government is better than anarchy. The eldest sons, +though they may be weak in body or mind, generally marry, whilst the +younger sons, however superior in these respects, do not so generally +marry. Nor can worthless eldest sons with entailed estates squander +their wealth. But here, as elsewhere, the relations of civilised life +are so complex that some compensatory checks intervene. The men who are +rich through primogeniture are able to select generation after +generation the more beautiful and charming women; and these must +generally be healthy in body and active in mind. The evil consequences, +such as they may be, of the continued preservation of the same line of +descent, without any selection, are checked by men of rank always +wishing to increase their wealth and power; and this they effect by +marrying heiresses. But the daughters of parents who have produced +single children, are themselves, as Mr. Galton has shewn,[232] apt to be +sterile; and thus noble families are continually cut off in the direct +line, and their wealth flows into some side channel; but unfortunately +this channel is not determined by superiority of any kind. + +Although civilisation thus checks in many ways the action of natural +selection, it apparently favours, by means of improved food and the +freedom from occasional hardships, the better development of the body. +This may be inferred from civilised men having been found, wherever +compared, to be physically stronger than savages. They appear also to +have equal powers of endurance, as has been proved in many adventurous +expeditions. Even the great luxury of the rich can be but little +detrimental; for the expectation of life of our aristocracy, at all ages +and of both sexes, is very little inferior to that of healthy English +lives in the lower classes.[233] + +We will now look to the intellectual faculties alone. If in each grade +of society the members were divided into two equal bodies, the one +including the intellectually superior and the other the inferior, there +can be little doubt that the former would succeed best in all +occupations and rear a greater number of children. Even in the lowest +walks of life, skill and ability must be of some advantage, though in +many occupations, owing to the great division of labour, a very small +one. Hence in civilised nations there will be some tendency to an +increase both in the number and in the standard of the intellectually +able. But I do not wish to assert that this tendency may not be more +than counterbalanced in other ways, as by the multiplication of the +reckless and improvident; but even to such as these, ability must be +some advantage. + +It has often been objected to views like the foregoing, that the most +eminent men who have ever lived have left no offspring to inherit their +great intellect. Mr. Galton says,[234] "I regret I am unable to solve +the simple question whether, and how far, men and women who are +prodigies of genius are infertile. I have, however, shewn that men of +eminence are by no means so." + +Great lawgivers, the founders of beneficent religions, great +philosophers and discoverers in science, aid the progress of mankind in +a far higher degree by their works than by leaving a numerous progeny. +In the case of corporeal structures, it is the selection of the slightly +better-endowed and the elimination of the slightly less well-endowed +individuals, and not the preservation of strongly-marked and rare +anomalies, that leads to the advancement of a species.[235] So it will +be with the intellectual faculties, namely from the somewhat more able +men in each grade of society succeeding rather better than the less +able, and consequently increasing in number, if not otherwise prevented. +When in any nation the standard of intellect and the number of +intellectual men have increased, we may expect from the law of the +deviation from an average, as shewn by Mr. Galton, that prodigies of +genius will appear somewhat more frequently than before. + +In regard to the moral qualities, some elimination of the worst +dispositions is always in progress even in the most civilised nations. +Malefactors are executed, or imprisoned for long periods, so that they +cannot freely transmit their bad qualities. Melancholic and insane +persons are confined, or commit suicide. Violent and quarrelsome men +often come to a bloody end. Restless men who will not follow any steady +occupation--and this relic of barbarism is a great check to +civilisation[236]--emigrate to newly-settled countries, where they prove +useful pioneers. Intemperance is so highly destructive, that the +expectation of life of the intemperate, at the age, for instance, of +thirty, is only 13.8 years; whilst for the rural labourers of England at +the same age it is 40.59 years.[237] Profligate women bear few children, +and profligate men rarely marry; both suffer from disease. In the +breeding of domestic animals, the elimination of those individuals, +though few in number, which are in any marked manner inferior, is by no +means an unimportant element towards success. This especially holds good +with injurious characters which tend to reappear through reversion, such +as blackness in sheep; and with mankind some of the worst dispositions, +which occasionally without any assignable cause make their appearance in +families, may perhaps be reversions to a savage state, from which we are +not removed by very many generations. This view seems indeed recognised +in the common expression that such men are the black sheep of the +family. + +With civilised nations, as far as an advanced standard of morality, and +an increased number of fairly well-endowed men are concerned, natural +selection apparently effects but little; though the fundamental social +instincts were originally thus gained. But I have already said enough, +whilst treating of the lower races, on the causes which lead to the +advance of morality, namely, the approbation of our fellow-men--the +strengthening of our sympathies by habit--example and imitation-- +reason--experience and even self-interest--instruction during +youth, and religious feelings. + +A most important obstacle in civilised countries to an increase in the +number of men of a superior class has been strongly urged by Mr. Greg +and Mr. Galton,[238] namely, the fact that the very poor and reckless, +who are often degraded by vice, almost invariably marry early, whilst +the careful and frugal, who are generally otherwise virtuous, marry late +in life, so that they may be able to support themselves and their +children in comfort. Those who marry early produce within a given period +not only a greater number of generations, but, as shewn by Dr. +Duncan,[239] they produce many more children. The children, moreover, +that are born by mothers during the prime of life are heavier and +larger, and therefore probably more vigorous, than those born at other +periods. Thus the reckless, degraded, and often vicious members of +society, tend to increase at a quicker rate than the provident and +generally virtuous members. Or as Mr. Greg puts the case: "The careless, +squalid, unaspiring Irishman multiplies like rabbits: the frugal, +foreseeing, self-respecting, ambitious Scot, stern in his morality, +spiritual in his faith, sagacious and disciplined in his intelligence, +passes his best years in struggle and in celibacy, marries late, and +leaves few behind him. Given a land originally peopled by a thousand +Saxons and a thousand Celts--and in a dozen generations five-sixths of +the population would be Celts, but five-sixths of the property, of the +power, of the intellect, would belong to the one-sixth of Saxons that +remained. In the eternal 'struggle for existence,' it would be the +inferior and _less_ favoured race that had prevailed--and prevailed by +virtue not of its good qualities but of its faults." + +There are, however, some checks to this downward tendency. We have seen +that the intemperate suffer from a high rate of mortality, and the +extremely profligate leave few offspring. The poorest classes crowd into +towns, and it has been proved by Dr. Stark from the statistics of ten +years in Scotland,[240] that at all ages the death-rate is higher in +towns than in rural districts, "and during the first five years of life +the town death-rate is almost exactly double that of the rural +districts." As these returns include both the rich and the poor, no +doubt more than double the number of births would be requisite to keep +up the number of the very poor inhabitants in the towns, relatively to +those in the country. With women, marriage at too early an age is highly +injurious; for it has been found in France that, "twice as many wives +under twenty die in the year, as died out of the same number of the +unmarried." The mortality, also, of husbands under twenty is +"excessively high,"[241] but what the cause of this may be seems +doubtful. Lastly, if the men who prudently delay marrying until they can +bring up their families in comfort, were to select, as they often do, +women in the prime of life, the rate of increase in the better class +would be only slightly lessened. + +It was established from an enormous body of statistics, taken during +1853, that the unmarried men throughout France, between the ages of +twenty and eighty, die in a much larger proportion than the married: for +instance, out of every 1000 unmarried men, between the ages of twenty +and thirty, 11.3 annually died, whilst of the married only 6.5 +died.[242] A similar law was proved to hold good, during the years 1863 +and 1864, with the entire population above the age of twenty in +Scotland: for instance, out of every 1000 unmarried men, between the +ages of twenty and thirty, 14.97 annually died, whilst of the married +only 7.24 died, that is less than half.[243] Dr. Stark remarks on this, +"Bachelorhood is more destructive to life than the most unwholesome +trades, or than residence in an unwholesome house or district where +there has never been the most distant attempt at sanitary improvement." +He considers that the lessened mortality is the direct result of +"marriage, and the more regular domestic habits which attend that +state." He admits, however, that the intemperate, profligate, and +criminal classes, whose duration of life is low, do not commonly marry; +and it must likewise be admitted that men with a weak constitution, +ill health, or any great infirmity in body or mind, will often not wish +to marry, or will be rejected. Dr. Stark seems to have come to the +conclusion that marriage in itself is a main cause of prolonged life, +from finding that aged married men still have a considerable advantage +in this respect over the unmarried of the same advanced age; but every +one must have known instances of men, who with weak health during youth +did not marry, and yet have survived to old age, though remaining weak +and therefore always with a lessened chance of life. There is another +remarkable circumstance which seems to support Dr. Stark's conclusion, +namely, that widows and widowers in France suffer in comparison with the +married a very heavy rate of mortality; but Dr. Farr attributes this to +the poverty and evil habits consequent on the disruption of the family, +and to grief. On the whole we may conclude with Dr. Farr that the lesser +mortality of married than of unmarried men, which seems to be a general +law, "is mainly due to the constant elimination of imperfect types, and +to the skilful selection of the finest individuals out of each +successive generation;" the selection relating only to the marriage +state, and acting on all corporeal, intellectual, and moral qualities. +We may, therefore, infer that sound and good men who out of prudence +remain for a time unmarried do not suffer a high rate of mortality. + +If the various checks specified in the two last paragraphs, and perhaps +others as yet unknown, do not prevent the reckless, the vicious and +otherwise inferior members of society from increasing at a quicker rate +than the better class of men, the nation will retrograde, as has +occurred too often in the history of the world. We must remember that +progress is no invariable rule. It is most difficult to say why one +civilised nation rises, becomes more powerful, and spreads more widely, +than another; or why the same nation progresses more at one time than at +another. We can only say that it depends on an increase in the actual +number of the population, on the number of the men endowed with high +intellectual and moral faculties, as well as on their standard of +excellence. Corporeal structure, except so far as vigour of body leads +to vigour of mind, appears to have little influence. + +It has been urged by several writers that as high intellectual powers +are advantageous to a nation, the old Greeks, who stood some grades +higher in intellect than any race that has ever existed,[244] ought to +have risen, if the power of natural selection were real, still higher +in the scale, increased in number, and stocked the whole of Europe. Here +we have the tacit assumption, so often made with respect to corporeal +structures, that there is some innate tendency towards continued +development in mind and body. But development of all kinds depends on +many concurrent favourable circumstances. Natural selection acts only in +a tentative manner. Individuals and races may have acquired certain +indisputable advantages, and yet have perished from failing in other +characters. The Greeks may have retrograded from a want of coherence +between the many small states, from the small size of their whole +country, from the practice of slavery, or from extreme sensuality; for +they did not succumb until "they were enervated and corrupt to the very +core."[245] The western nations of Europe, who now so immeasurably +surpass their former savage progenitors and stand at the summit of +civilisation, owe little or none of their superiority to direct +inheritance from the old Greeks; though they owe much to the written +works of this wonderful people. + +Who can positively say why the Spanish nation, so dominant at one time, +has been distanced in the race. The awakening of the nations of Europe +from the dark ages is a still more perplexing problem. At this early +period, as Mr. Galton[246] has remarked, almost all the men of a gentle +nature, those given to meditation or culture of the mind, had no refuge +except in the bosom of the Church which demanded celibacy; and this +could hardly fail to have had a deteriorating influence on each +successive generation. During this same period the Holy Inquisition +selected with extreme care the freest and boldest men in order to burn +or imprison them. In Spain alone some of the best men--those who doubted +and questioned, and without doubting there can be no progress--were +eliminated during three centuries at the rate of a thousand a year. The +evil which the Catholic Church has thus effected, though no doubt +counterbalanced to a certain, perhaps large extent in other ways, is +incalculable; nevertheless, Europe has progressed at an unparalleled +rate. + +The remarkable success of the English as colonists over other European +nations, which is well illustrated by comparing the progress of the +Canadians of English and French extraction, has been ascribed to their +"daring and persistent energy;" but who can say how the English gained +their energy. There is apparently much truth in the belief that the +wonderful progress of the United States, as well as the character of the +people, are the results of natural selection; the more energetic, +restless, and courageous men from all parts of Europe having emigrated +during the last ten or twelve generations to that great country, and +having there succeeded best.[247] Looking to the distant future, I do +not think that the Rev. Mr. Zincke takes an exaggerated view when he +says:[248] "All other series of events--as that which resulted in the +culture of mind in Greece, and that which resulted in the empire of +Rome--only appear to have purpose and value when viewed in connection +with, or rather as subsidiary to ... the great stream of Anglo-Saxon +emigration to the west." + +Obscure as is the problem of the advance of civilisation, we can at +least see that a nation which produced during a lengthened period the +greatest number of highly intellectual, energetic, brave, patriotic, and +benevolent men, would generally prevail over less favoured nations. + +Natural selection follows from the struggle for existence; and this from +a rapid rate of increase. It is impossible not bitterly to regret, but +whether wisely is another question, the rate at which man tends to +increase; for this leads in barbarous tribes to infanticide and many +other evils, and in civilised nations to abject poverty, celibacy, and +to the late marriages of the prudent. But as man suffers from the same +physical evils with the lower animals, he has no right to expect an +immunity from the evils consequent on the struggle for existence. Had he +not been subjected to natural selection, assuredly he would never have +attained to the rank of manhood. When we see in many parts of the world +enormous areas of the most fertile land peopled by a few wandering +savages, but which are capable of supporting numerous happy homes, it +might be argued that the struggle for existence had not been +sufficiently severe to force man upwards to his highest standard. +Judging from all that we know of man and the lower animals, there has +always been sufficient variability in the intellectual and moral +faculties, for their steady advancement through natural selection. No +doubt such advancement demands many favourable concurrent circumstances; +but it may well be doubted whether the most favourable would have +sufficed, had not the rate of increase been rapid, and the consequent +struggle for existence severe to an extreme degree. + + +_On the evidence that all civilised nations were once barbarous._--As we +have had to consider the steps by which some semi-human creature has +been gradually raised to the rank of man in his most perfect state, the +present subject cannot be quite passed over. But it has been treated in +so full and admirable a manner by Sir J. Lubbock,[249] Mr. Tylor, Mr. +M'Lennan, and others, that I need here give only the briefest summary of +their results. The arguments recently advanced by the Duke of +Argyll[250] and formerly by Archbishop Whately, in favour of the belief +that man came into the world as a civilised being and that all savages +have since undergone degradation, seem to me weak in comparison with +those advanced on the other side. Many nations, no doubt, have fallen +away in civilisation, and some may have lapsed into utter barbarism, +though on this latter head I have not met with any evidence. The +Fuegians were probably compelled by other conquering hordes to settle in +their inhospitable country, and they may have become in consequence +somewhat more degraded; but it would be difficult to prove that they +have fallen much below the Botocudos who inhabit the finest parts of +Brazil. + +The evidence that all civilised nations are the descendants of +barbarians, consists, on the one side, of clear traces of their former +low condition in still-existing customs, beliefs, language, &c.; and on +the other side, of proofs that savages are independently able to raise +themselves a few steps in the scale of civilisation, and have actually +thus risen. The evidence on the first head is extremely curious, but +cannot be here given: I refer to such cases as that, for instance, of +the art of enumeration, which, as Mr. Tylor clearly shews by the words +still used in some places, originated in counting the fingers, first of +one hand and then of the other, and lastly of the toes. We have traces +of this in our own decimal system, and in the Roman numerals, which +after reaching to the number V., change into VI., &c., when the other +hand no doubt was used. So again, "when we speak of three-score and ten, +we are counting by the vigesimal system, each score thus ideally made, +standing for 20--for 'one man' as a Mexican or Carib would put it."[251] +According to a large and increasing school of philologists, every +language bears the marks of its slow and gradual evolution. So it is +with the art of writing, as letters are rudiments of pictorial +representations. It is hardly possible to read Mr. M'Lennan's work[252] +and not admit that almost all civilised nations still retain some traces +of such rude habits as the forcible capture of wives. What ancient +nation, as the same author asks, can be named that was originally +monogamous? The primitive idea of justice, as shewn by the law of battle +and other customs of which traces still remain, was likewise most rude. +Many existing superstitions are the remnants of former false religious +beliefs. The highest form of religion--the grand idea of God hating sin +and loving righteousness--was unknown during primeval times. + +Turning to the other kind of evidence: Sir J. Lubbock has shewn that +some savages have recently improved a little in some of their simpler +arts. From the extremely curious account which he gives of the weapons, +tools, and arts, used or practised by savages in various parts of the +world, it cannot be doubted that these have nearly all been independent +discoveries, excepting perhaps the art of making fire.[253] The +Australian boomerang is a good instance of one such independent +discovery. The Tahitians when first visited had advanced in many +respects beyond the inhabitants of most of the other Polynesian islands. +There are no just grounds for the belief that the high culture of the +native Peruvians and Mexicans was derived from any foreign source;[254] +many native plants were there cultivated, and a few native animals +domesticated. We should bear in mind that a wandering crew from some +semi-civilised land, if washed to the shores of America, would not, +judging from the small influence of most missionaries, have produced any +marked effect on the natives, unless they had already become somewhat +advanced. Looking to a very remote period in the history of the world, +we find, to use Sir J. Lubbock's well-known terms, a paleolithic and +neolithic period; and no one will pretend that the art of grinding rough +flint tools was a borrowed one. In all parts of Europe, as far east as +Greece, in Palestine, India, Japan, New Zealand, and Africa, including +Egypt, flint tools have been discovered in abundance; and of their use +the existing inhabitants retain no tradition. There is also indirect +evidence of their former use by the Chinese and ancient Jews. Hence +there can hardly be a doubt that the inhabitants of these many +countries, which include nearly the whole civilised world, were once in +a barbarous condition. To believe that man was aboriginally civilised +and then suffered utter degradation in so many regions, is to take a +pitiably low view of human nature. It is apparently a truer and more +cheerful view that progress has been much more general than +retrogression; that man has risen, though by slow and interrupted steps, +from a lowly condition to the highest standard as yet attained by him in +knowledge, morals, and religion. + + + + +CHAPTER VI. + +ON THE AFFINITIES AND GENEALOGY OF MAN. + + + Position of man in the animal series--The natural system + genealogical--Adaptive characters of slight value--Various + small points of resemblance between man and the + Quadrumana--Rank of man in the natural system--Birthplace and + antiquity of man--Absence of fossil connecting-links--Lower + stages in the genealogy of man, as inferred, firstly from his + affinities and secondly from his structure--Early androgynous + condition of the Vertebrata--Conclusion. + + +Even if it be granted that the difference between man and his nearest +allies is as great in corporeal structure as some naturalists maintain, +and although we must grant that the difference between them is immense +in mental power, yet the facts given in the previous chapters declare, +as it appears to me, in the plainest manner, that man is descended from +some lower form, notwithstanding that connecting-links have not hitherto +been discovered. + +Man is liable to numerous, slight, and diversified variations, which are +induced by the same general causes, are governed and transmitted in +accordance with the same general laws, as in the lower animals. Man +tends to multiply at so rapid a rate that his offspring are necessarily +exposed to a struggle for existence, and consequently to natural +selection. He has given rise to many races, some of which are so +different that they have often been ranked by naturalists as distinct +species. His body is constructed on the same homological plan as that of +other mammals, independently of the uses to which the several parts may +be put. He passes through the same phases of embryological development. +He retains many rudimentary and useless structures, which no doubt were +once serviceable. Characters occasionally make their reappearance in +him, which we have every reason to believe were possessed by his early +progenitors. If the origin of man had been wholly different from that of +all other animals, these various appearances would be mere empty +deceptions; but such an admission is incredible. These appearances, on +the other hand, are intelligible, at least to a large extent, if man is +the co-descendant with other mammals of some unknown and lower form. + +Some naturalists, from being deeply impressed with the mental and +spiritual powers of man, have divided the whole organic world into three +kingdoms, the Human, the Animal, and the Vegetable, thus giving to man a +separate kingdom.[255] Spiritual powers cannot be compared or classed by +the naturalist; but he may endeavour to shew, as I have done, that the +mental faculties of man and the lower animals do not differ in kind, +although immensely in degree. A difference in degree, however great, +does not justify us in placing man in a distinct kingdom, as will +perhaps be best illustrated by comparing the mental powers of two +insects, namely, a coccus or scale-insect and an ant, which undoubtedly +belong to the same class. The difference is here greater, though of a +somewhat different kind, than that between man and the highest mammal. +The female coccus, whilst young, attaches itself by its proboscis to a +plant; sucks the sap but never moves again; is fertilised and lays eggs; +and this is its whole history. On the other hand, to describe the habits +and mental powers of a female ant, would require, as Pierre Huber has +shewn, a large volume; I may, however, briefly specify a few points. +Ants communicate information to each other, and several unite for the +same work, or games of play. They recognise their fellow-ants after +months of absence. They build great edifices, keep them clean, close the +doors in the evening, and post sentries. They make roads, and even +tunnels under rivers. They collect food for the community, and when an +object, too large for entrance, is brought to the nest, they enlarge the +door, and afterwards build it up again.[256] They go out to battle in +regular bands, and freely sacrifice their lives for the common weal. +They emigrate in accordance with a preconcerted plan. They capture +slaves. They keep Aphides as milch-cows. They move the eggs of their +aphides, as well as their own eggs and cocoons, into warm parts of the +nest, in order that they may be quickly hatched; and endless similar +facts could be given. On the whole, the difference in mental power +between an ant and a coccus is immense; yet no one has ever dreamed of +placing them in distinct classes, much less in distinct kingdoms. No +doubt this interval is bridged over by the intermediate mental powers of +many other insects; and this is not the case with man and the higher +apes. But we have every reason to believe that breaks in the series are +simply the result of many forms having become extinct. + +Professor Owen, relying chiefly on the structure of the brain, has +divided the mammalian series into four sub-classes. One of these he +devotes to man; in another he places both the marsupials and the +monotremata; so that he makes man as distinct from all other mammals as +are these two latter groups conjoined. This view has not been accepted, +as far as I am aware, by any naturalist capable of forming an +independent judgment, and therefore need not here be further considered. + +We can understand why a classification founded on any single character +or organ--even an organ so wonderfully complex and important as the +brain--or on the high development of the mental faculties, is almost +sure to prove unsatisfactory. This principle has indeed been tried with +hymenopterous insects; but when thus classed by their habits or +instincts, the arrangement proved thoroughly artificial.[257] +Classifications may, of course, be based on any character whatever, as +on size, colour, or the element inhabited; but naturalists have long +felt a profound conviction that there is a natural system. This system, +it is now generally admitted, must be, as far as possible, genealogical +in arrangement,--that is, the co-descendants of the same form must be +kept together in one group, separate from the co-descendants of any +other form; but if the parent-forms are related, so will be their +descendants, and the two groups together will form a larger group. The +amount of difference between the several groups--that is the amount of +modification which each has undergone--will be expressed by such terms +as genera, families, orders, and classes. As we have no record of the +lines of descent, these lines can be discovered only by observing the +degrees of resemblance between the beings which are to be classed. For +this object numerous points of resemblance are of much more importance +than the amount of similarity or dissimilarity in a few points. If two +languages were found to resemble each other in a multitude of words and +points of construction, they would be universally recognised as having +sprung from a common source, notwithstanding that they differed greatly +in some few words or points of construction. But with organic beings the +points of resemblance must not consist of adaptations to similar habits +of life: two animals may, for instance, have had their whole frames +modified for living in the water, and yet they will not be brought any +nearer to each other in the natural system. Hence we can see how it is +that resemblances in unimportant structures, in useless and rudimentary +organs, and in parts not as yet fully developed or functionally active, +are by far the most serviceable for classification; for they can hardly +be due to adaptations within a late period; and thus they reveal the old +lines of descent or of true affinity. + +We can further see why a great amount of modification in some one +character ought not to lead us to separate widely any two organisms. A +part which already differs much from the same part in other allied forms +has already, according to the theory of evolution, varied much; +consequently it would (as long as the organism remained exposed to the +same exciting conditions) be liable to further variations of the same +kind; and these, if beneficial, would be preserved, and thus continually +augmented. In many cases the continued development of a part, for +instance, of the beak of a bird, or of the teeth of a mammal, would not +be advantageous to the species for gaining its food, or for any other +object; but with man we can see no definite limit, as far as advantage +is concerned, to the continued development of the brain and mental +faculties. Therefore in determining the position of man in the natural +or genealogical system, the extreme development of his brain ought not +to outweigh a multitude of resemblances in other less important or +quite unimportant points. + +The greater number of naturalists who have taken into consideration the +whole structure of man, including his mental faculties, have followed +Blumenbach and Cuvier, and have placed man in a separate Order, under +the title of the Bimana, and therefore on an equality with the Orders of +the Quadrumana, Carnivora, &c. Recently many of our best naturalists +have recurred to the view first propounded by Linnaeus, so remarkable for +his sagacity, and have placed man in the same Order with the Quadrumana, +under the title of the Primates. The justice of this conclusion will be +admitted if, in the first place, we bear in mind the remarks just made +on the comparatively small importance for classification of the great +development of the brain in man; bearing, also, in mind that the +strongly-marked differences between the skulls of man and the Quadrumana +(lately insisted upon by Bischoff, Aeby, and others) apparently follow +from their differently developed brains. In the second place, we must +remember that nearly all the other and more important differences +between man and the Quadrumana are manifestly adaptive in their nature, +and relate chiefly to the erect position of man; such as the structure +of his hand, foot, and pelvis, the curvature of his spine, and the +position of his head. The family of seals offers a good illustration of +the small importance of adaptive characters for classification. These +animals differ from all other Carnivora in the form of their bodies and +in the structure of their limbs, far more than does man from the higher +apes; yet in every system, from that of Cuvier to the most recent one by +Mr. Flower,[258] seals are ranked as a mere family in the Order of the +Carnivora. If man had not been his own classifier, he would never have +thought of founding a separate order for his own reception. + +It would be beyond my limits, and quite beyond my knowledge, even to +name the innumerable points of structure in which man agrees with the +other Primates. Our great anatomist and philosopher, Prof. Huxley, has +fully discussed this subject,[259] and has come to the conclusion that +man in all parts of his organisation differs less from the higher apes, +than these do from the lower members of the same group. Consequently +there "is no justification for placing man in a distinct order." + +In an early part of this volume I brought forward various facts, shewing +how closely man agrees in constitution with the higher mammals; and this +agreement, no doubt, depends on our close similarity in minute structure +and chemical composition. I gave, as instances, our liability to the +same diseases, and to the attacks of allied parasites; our tastes in +common for the same stimulants, and the similar effects thus produced, +as well as by various drugs; and other such facts. + +As small unimportant points of resemblance between man and the higher +apes are not commonly noticed in systematic works, and as, when +numerous, they clearly reveal our relationship, I will specify a few +such points. The relative position of the features are manifestly the +same in man and the Quadrumana; and the various emotions are displayed +by nearly similar movements of the muscles and skin, chiefly above the +eyebrows and round the mouth. Some few expressions are, indeed, almost +the same, as in the weeping of certain kinds of monkeys, and in the +laughing noise made by others, during which the corners of the mouth are +drawn backwards, and the lower eyelids wrinkled. The external ears are +curiously alike. In man the nose is much more prominent than in most +monkeys; but we may trace the commencement of an aquiline curvature in +the nose of the Hoolock Gibbon; and this in the _Semnopithecus nasica_ +is carried to a ridiculous extreme. + +The faces of many monkeys are ornamented with beards, whiskers, or +moustaches. The hair on the head grows to a great length in some species +of Semnopithecus;[260] and in the Bonnet monkey (_Macacus radiatus_) it +radiates from a point on the crown, with a parting down the middle, as +in man. It is commonly said that the forehead gives to man his noble and +intellectual appearance; but the thick hair on the head of the Bonnet +monkey terminates abruptly downwards, and is succeeded by such short and +fine hair, or down, that at a little distance the forehead, with the +exception of the eyebrows, appears quite naked. It has been erroneously +asserted that eyebrows are not present in any monkey. In the species +just named the degree of nakedness of the forehead differs in different +individuals; and Eschricht states[261] that in our children the limit +between the hairy scalp and the naked forehead is sometimes not well +defined; so that here we seem to have a trifling case of reversion to a +progenitor, in whom the forehead had not as yet become quite naked. + +It is well known that the hair on our arms tends to converge from above +and below to a point at the elbow. This curious arrangement, so unlike +that in most of the lower mammals, is common to the gorilla, chimpanzee, +orang, some species of Hylobates, and even to some few American monkeys. +But in _Hylobates agilis_ the hair on the fore-arm is directed +downwards or towards the wrist in the ordinary manner; and in _H. lar_ +it is nearly erect, with only a very slight forward inclination; so that +in this latter species it is in a transitional state. It can hardly be +doubted that with most mammals the thickness of the hair and its +direction on the back is adapted to throw off the rain; even the +transverse hairs on the fore-legs of a dog may serve for this end when +he is coiled up asleep. Mr. Wallace remarks that the convergence of the +hair towards the elbow on the arms of the orang (whose habits he has so +carefully studied) serves to throw off the rain, when, as is the custom +of this animal, the arms are bent, with the hands clasped round a branch +or over its own head. We should, however, bear in mind that the attitude +of an animal may perhaps be in part determined by the direction of the +hair; and not the direction of the hair by the attitude. If the above +explanation is correct in the case of the orang, the hair on our +fore-arms offers a curious record of our former state; for no one +supposes that it is now of any use in throwing off the rain, nor in our +present erect condition is it properly directed for this purpose. + +It would, however, be rash to trust too much to the principle of +adaptation in regard to the direction of the hair in man or his early +progenitors; for it is impossible to study the figures given by +Eschricht of the arrangement of the hair on the human foetus (this +being the same as in the adult) and not agree with this excellent +observer that other and more complex causes have intervened. The points +of convergence seem to stand in some relation to those points in the +embryo which are last closed in during development. There appears, also, +to exist some relation between the arrangement of the hair on the +limbs, and the course of the medullary arteries.[262] + +It must not be supposed that the resemblances between man and certain +apes in the above and many other points--such as in having a naked +forehead, long tresses on the head, &c.--are all necessarily the result +of unbroken inheritance from a common progenitor thus characterised, or +of subsequent reversion. Many of these resemblances are more probably +due to analogous variation, which follows, as I have elsewhere attempted +to shew,[263] from co-descended organisms having a similar constitution +and having been acted on by similar causes inducing variability. With +respect to the similar direction of the hair on the fore-arms of man and +certain monkeys, as this character is common to almost all the +anthropomorphous apes, it may probably be attributed to inheritance; but +not certainly so, as some very distinct American monkeys are thus +characterised. The same remark is applicable to the tailless condition +of man; for the tail is absent in all the anthropomorphous apes. +Nevertheless this character cannot with certainty be attributed to +inheritance, as the tail, though not absent, is rudimentary in several +other Old World and in some New World species, and is quite absent in +several species belonging to the allied group of Lemurs. + +Although, as we have now seen, man has no just right to form a separate +Order for his own reception, he may perhaps claim a distinct Sub-order +or Family. Prof. Huxley, in his last work,[264] divides the Primates +into three Sub-orders; namely, the Anthropidae with man alone, the +Simiadae including monkeys of all kinds, and the Lemuridae with the +diversified genera of lemurs. As far as differences in certain important +points of structure are concerned, man may no doubt rightly claim the +rank of a Sub-order; and this rank is too low, if we look chiefly to his +mental faculties. Nevertheless, under a genealogical point of view it +appears that this rank is too high, and that man ought to form merely a +Family, or possibly even only a Sub-family. If we imagine three lines of +descent proceeding from a common source, it is quite conceivable that +two of them might after the lapse of ages be so slightly changed as +still to remain as species of the same genus; whilst the third line +might become so greatly modified as to deserve to rank as a distinct +Sub-family, Family, or even Order. But in this case it is almost certain +that the third line would still retain through inheritance numerous +small points of resemblance with the other two lines. Here then would +occur the difficulty, at present insoluble, how much weight we ought to +assign in our classifications to strongly-marked differences in some few +points,--that is to the amount of modification undergone; and how much +to close resemblance in numerous unimportant points, as indicating the +lines of descent or genealogy. The former alternative is the most +obvious, and perhaps the safest, though the latter appears the most +correct as giving a truly natural classification. + +To form a judgment on this head, with reference to man we must glance at +the classification of the Simiadae. This family is divided by almost all +naturalists into the Catarhine group, or Old World monkeys, all of which +are characterised (as their name expresses) by the peculiar structure of +their nostrils and by having four premolars in each jaw; and into the +Platyrhine group or New World monkeys (including two very distinct +sub-groups), all of which are characterised by differently-constructed +nostrils and by having six premolars in each jaw. Some other small +differences might be mentioned. Now man unquestionably belongs in his +dentition, in the structure of his nostrils, and some other respects, to +the Catarhine or Old World division; nor does he resemble the +Platyrhines more closely than the Catarhines in any characters, +excepting in a few of not much importance and apparently of an adaptive +nature. Therefore it would be against all probability to suppose that +some ancient New World species had varied, and had thus produced a +man-like creature with all the distinctive characters proper to the Old +World division; losing at the same time all its own distinctive +characters. There can consequently hardly be a doubt that man is an +offshoot from the Old World Simian stem; and that under a genealogical +point of view, he must be classed with the Catarhine division.[265] + +The anthropomorphous apes, namely the gorilla, chimpanzee, orang, and +hylobates, are separated as a distinct sub-group from the other Old +World monkeys by most naturalists. I am aware that Gratiolet, relying on +the structure of the brain, does not admit the existence of this +sub-group, and no doubt it is a broken one; thus the orang, as Mr. St. +G. Mivart remarks,[266] "is one of the most peculiar and aberrant forms +to be found in the Order." The remaining, non-anthropomorphous, Old +World monkeys, are again divided by some naturalists into two or three +smaller sub-groups; the genus Semnopithecus, with its peculiar +sacculated stomach, being the type of one such sub-group. But it appears +from M. Gaudry's wonderful discoveries in Attica, that during the +Miocene period a form existed there, which connected Semnopithecus and +Macacus; and this probably illustrates the manner in which the other and +higher groups were once blended together. + +If the anthropomorphous apes be admitted to form a natural sub-group, +then as man agrees with them, not only in all those characters which he +possesses in common with the whole Catarhine group, but in other +peculiar characters, such as the absence of a tail and of callosities +and in general appearance, we may infer that some ancient member of the +anthropomorphous sub-group gave birth to man. It is not probable that a +member of one of the other lower sub-groups should, through the law of +analogous variation, have given rise to a man-like creature, resembling +the higher anthropomorphous apes in so many respects. No doubt man, in +comparison with most of his allies, has undergone an extraordinary +amount of modification, chiefly in consequence of his greatly developed +brain and erect position; nevertheless we should bear in mind that he +"is but one of several exceptional forms of Primates."[267] + +Every naturalist, who believes in the principle of evolution, will +grant that the two main divisions of the Simiadae, namely the Catarhine +and Platyrhine monkeys, with their sub-groups, have all proceeded from +some one extremely ancient progenitor. The early descendants of this +progenitor, before they had diverged to any considerable extent from +each other, would still have formed a single natural group; but some of +the species or incipient genera would have already begun to indicate by +their diverging characters the future distinctive marks of the Catarhine +and Platyrhine divisions. Hence the members of this supposed ancient +group would not have been so uniform in their dentition or in the +structure of their nostrils, as are the existing Catarhine monkeys in +one way and the Platyrhines in another way, but would have resembled in +this respect the allied Lemuridae which differ greatly from each other in +the form of their muzzles,[268] and to an extraordinary degree in their +dentition. + +The Catarhine and Platyrhine monkeys agree in a multitude of characters, +as is shewn by their unquestionably belonging to one and the same Order. +The many characters which they possess in common can hardly have been +independently acquired by so many distinct species; so that these +characters must have been inherited. But an ancient form which possessed +many characters common to the Catarhine and Platyrhine monkeys, and +others in an intermediate condition, and some few perhaps distinct from +those now present in either group, would undoubtedly have been ranked, +if seen by a naturalist, as an ape or monkey. And as man under a +genealogical point of view belongs to the Catarhine or Old World stock, +we must conclude, however much the conclusion may revolt our pride, +that our early progenitors would have been properly thus +designated.[269] But we must not fall into the error of supposing that +the early progenitor of the whole Simian stock, including man, was +identical with, or even closely resembled, any existing ape or monkey. + + +_On the Birthplace and Antiquity of Man._--We are naturally led to +enquire where was the birthplace of man at that stage of descent when +our progenitors diverged from the Catarhine stock. The fact that they +belonged to this stock clearly shews that they inhabited the Old World; +but not Australia nor any oceanic island, as we may infer from the laws +of geographical distribution. In each great region of the world the +living mammals are closely related to the extinct species of the same +region. It is therefore probable that Africa was formerly inhabited by +extinct apes closely allied to the gorilla and chimpanzee; and as these +two species are now man's nearest allies, it is somewhat more probable +that our early progenitors lived on the African continent than +elsewhere. But it is useless to speculate on this subject, for an ape +nearly as large as a man, namely the Dryopithecus of Lartet, which was +closely allied to the anthropomorphous Hylobates, existed in Europe +during the Upper Miocene period; and since so remote a period the earth +has certainly undergone many great revolutions, and there has been ample +time for migration on the largest scale. + +At the period and place, whenever and wherever it may have been, when +man first lost his hairy covering, he probably inhabited a hot country; +and this would have been favourable for a frugiferous diet, on which, +judging from analogy, he subsisted. We are far from knowing how long ago +it was when man first diverged from the Catarhine stock; but this may +have occurred at an epoch as remote as the Eocene period; for the higher +apes had diverged from the lower apes as early as the Upper Miocene +period, as shewn by the existence of the Dryopithecus. We are also quite +ignorant at how rapid a rate organisms, whether high or low in the +scale, may under favourable circumstances be modified: we know, however, +that some have retained the same form during an enormous lapse of time. +From what we see going on under domestication, we learn that within the +same period some of the co-descendants of the same species may be not at +all changed, some a little, and some greatly changed. Thus it may have +been with man, who has undergone a great amount of modification in +certain characters in comparison with the higher apes. + +The great break in the organic chain between man and his nearest allies, +which cannot be bridged over by any extinct or living species, has often +been advanced as a grave objection to the belief that man is descended +from some lower form; but this objection will not appear of much weight +to those who, convinced by general reasons, believe in the general +principle of evolution. Breaks incessantly occur in all parts of the +series, some being wide, sharp and defined, others less so in various +degrees; as between the orang and its nearest allies--between the +Tarsius and the other Lemuridae--between the elephant and in a more +striking manner between the Ornithorhynchus or Echidna, and other +mammals. But all these breaks depend merely on the number of related +forms which have become extinct. At some future period, not very distant +as measured by centuries, the civilised races of man will almost +certainly exterminate and replace throughout the world the savage races. +At the same time the anthropomorphous apes, as Professor Schaaffhausen +has remarked,[270] will no doubt be exterminated. The break will then be +rendered wider, for it will intervene between man in a more civilised +state, as we may hope, than the Caucasian, and some ape as low as a +baboon, instead of as at present between the negro or Australian and the +gorilla. + +With respect to the absence of fossil remains, serving to connect man +with his ape-like progenitors, no one will lay much stress on this fact, +who will read Sir C. Lyell's discussion,[271] in which he shews that in +all the vertebrate classes the discovery of fossil remains has been an +extremely slow and fortuitous process. Nor should it be forgotten that +those regions which are the most likely to afford remains connecting man +with some extinct ape-like creature, have not as yet been searched by +geologists. + + +_Lower Stages in the Genealogy of Man._--We have seen that man appears +to have diverged from the Catarhine or Old World division of the +Simiadae, after these had diverged from the New World division. We will +now endeavour to follow the more remote traces of his genealogy, +trusting in the first place to the mutual affinities between the various +classes and orders, with some slight aid from the periods, as far as +ascertained, of their successive appearance on the earth. The Lemuridae +stand below and close to the Simiadae, constituting a very distinct +family of the Primates, or, according to Haeckel, a distinct Order. This +group is diversified and broken to an extraordinary degree, and includes +many aberrant forms. It has, therefore, probably suffered much +extinction. Most of the remnants survive on islands, namely in +Madagascar and in the islands of the Malayan archipelago, where they +have not been exposed to such severe competition as they would have been +on well-stocked continents. This group likewise presents many +gradations, leading, as Huxley remarks,[272] "insensibly from the crown +and summit of the animal creation down to creatures from which there is +but a step, as it seems, to the lowest, smallest, and least intelligent +of the placental mammalia." From these various considerations it is +probable that the Simiadae were originally developed from the progenitors +of the existing Lemuridae; and these in their turn from forms standing +very low in the mammalian series. + +The Marsupials stand in many important characters below the placental +mammals. They appeared at an earlier geological period, and their range +was formerly much more extensive than what it now is. Hence the +Placentata are generally supposed to have been derived from the +Implacentata or Marsupials; not, however, from forms closely like the +existing Marsupials, but from their early progenitors. The Monotremata +are plainly allied to the Marsupials; forming a third and still lower +division in the great mammalian series. They are represented at the +present day solely by the Ornithorhynchus and Echidna; and these two +forms may be safely considered as relics of a much larger group which +have been preserved in Australia through some favourable concurrence of +circumstances. The Monotremata are eminently interesting, as in several +important points of structure they lead towards the class of reptiles. + +In attempting to trace the genealogy of the Mammalia, and therefore of +man, lower down in the series, we become involved in greater and greater +obscurity. He who wishes to see what ingenuity and knowledge can effect, +may consult Prof. Haeckel's works.[273] I will content myself with a few +general remarks. Every evolutionist will admit that the five great +vertebrate classes, namely, mammals, birds, reptiles, amphibians, and +fishes, are all descended from some one prototype; for they have much in +common, especially during their embryonic state. As the class of fishes +is the most lowly organised and appeared before the others, we may +conclude that all the members of the vertebrate kingdom are derived from +some fish-like animal, less highly organised than any as yet found in +the lowest known formations. The belief that animals so distinct as a +monkey or elephant and a humming-bird, a snake, frog, and fish, &c., +could all have sprung from the same parents, will appear monstrous to +those who have not attended to the recent progress of natural history. +For this belief implies the former existence of links closely binding +together all these forms, now so utterly unlike. + +Nevertheless it is certain that groups of animals have existed, or do +now exist, which serve to connect more or less closely the several great +vertebrate classes. We have seen that the Ornithorhynchus graduates +towards reptiles; and Prof. Huxley has made the remarkable discovery, +confirmed by Mr. Cope and others, that the old Dinosaurians are +intermediate in many important respects between certain reptiles and +certain birds--the latter consisting of the ostrich-tribe (itself +evidently a widely-diffused remnant of a larger group) and of the +Archeopteryx, that strange Secondary bird having a long tail like that +of the lizard. Again, according to Prof. Owen,[274] the +Ichthyosaurians--great sea-lizards furnished with paddles--present many +affinities with fishes, or rather, according to Huxley, with amphibians. +This latter class (including in its highest division frogs and toads) is +plainly allied to the Ganoid fishes. These latter fishes swarmed during +the earlier geological periods, and were constructed on what is called a +highly generalised type, that is they presented diversified affinities +with other groups of organisms. The amphibians and fishes are also so +closely united by the Lepidosiren, that naturalists long disputed in +which of these two classes it ought to be placed. The Lepidosiren and +some few Ganoid fishes have been preserved from utter extinction by +inhabiting our rivers, which are harbours of refuge, bearing the same +relation to the great waters of the ocean that islands bear to +continents. + +Lastly, one single member of the immense and diversified class of +fishes, namely the lancelet or amphioxus, is so different from all other +fishes, that Haeckel maintains that it ought to form a distinct class in +the vertebrate kingdom. This fish is remarkable for its negative +characters; it can hardly be said to possess a brain, vertebral column, +or heart, &c.; so that it was classed by the older naturalists amongst +the worms. Many years ago Prof. Goodsir perceived that the lancelet +presented some affinities with the Ascidians, which are invertebrate, +hermaphrodite, marine creatures permanently attached to a support. They +hardly appear like animals, and consist of a simple, tough, leathery +sack, with two small projecting orifices. They belong to the Molluscoida +of Huxley--a lower division of the great kingdom of the Mollusca; but +they have recently been placed by some naturalists amongst the Vermes or +worms. Their larvae somewhat resemble tadpoles in shape,[275] and have +the power of swimming freely about. Some observations lately made by M. +Kowalevsky,[276] since confirmed by Prof. Kuppfer, will form a discovery +of extraordinary interest, if still further extended, as I hear from M. +Kowalevsky in Naples he has now effected. The discovery is that the +larvae of Ascidians are related to the Vertebrata, in their manner of +development, in the relative position of the nervous system, and in +possessing a structure closely like the _chorda dorsalis_ of vertebrate +animals. It thus appears, if we may rely on embryology, which has always +proved the safest guide in classification, that we have at last gained a +clue to the source whence the Vertebrata have been derived. We should +thus be justified in believing that at an extremely remote period a +group of animals existed, resembling in many respects the larvae of our +present Ascidians, which diverged into two great branches--the one +retrograding in development and producing the present class of +Ascidians, the other rising to the crown and summit of the animal +kingdom by giving birth to the Vertebrata. + + +We have thus far endeavoured rudely to trace the genealogy of the +Vertebrata by the aid of their mutual affinities. We will now look to +man as he exists; and we shall, I think, be able partially to restore +during successive periods, but not in due order of time, the structure +of our early progenitors. This can be effected by means of the rudiments +which man still retains, by the characters which occasionally make their +appearance in him through reversion, and by the aid of the principles of +morphology and embryology. The various facts, to which I shall here +allude, have been given in the previous chapters. The early progenitors +of man were no doubt once covered with hair, both sexes having beards; +their ears were pointed and capable of movement; and their bodies were +provided with a tail, having the proper muscles. Their limbs and bodies +were also acted on by many muscles which now only occasionally reappear, +but are normally present in the Quadrumana. The great artery and nerve +of the humerus ran through a supra-condyloid foramen. At this or some +earlier period, the intestine gave forth a much larger diverticulum or +caecum than that now existing. The foot, judging from the condition of +the great toe in the foetus, was then prehensile; and our progenitors, +no doubt, were arboreal in their habits, frequenting some warm, +forest-clad land. The males were provided with great canine teeth, +which served them as formidable weapons. + +At a much earlier period the uterus was double; the excreta were voided +through a cloaca; and the eye was protected by a third eyelid or +nictitating membrane. At a still earlier period the progenitors of man +must have been aquatic in their habits; for morphology plainly tells us +that our lungs consist of a modified swim-bladder, which once served as +a float. The clefts on the neck in the embryo of man show where the +branchiae once existed. At about this period the true kidneys were +replaced by the corpora Wolffiana. The heart existed as a simple +pulsating vessel; and the chorda dorsalis took the place of a vertebral +column. These early predecessors of man, thus seen in the dim recesses +of time, must have been as lowly organised as the lancelet or amphioxus, +or even still more lowly organised. + +There is one other point deserving a fuller notice. It has long been +known that in the vertebrate kingdom one sex bears rudiments of various +accessory parts, appertaining to the reproductive system, which properly +belong to the opposite sex; and it has now been ascertained that at a +very early embryonic period both sexes possess true male and female +glands. Hence some extremely remote progenitor of the whole vertebrate +kingdom appears to have been hermaphrodite or androgynous.[277] But here +we encounter a singular difficulty. In the mammalian class the males +possess in their vesiculae prostaticae rudiments of a uterus with the +adjacent passage; they bear also rudiments of mammae, and some male +marsupials have rudiments of a marsupial sack.[278] Other analogous +facts could be added. Are we, then, to suppose that some extremely +ancient mammal possessed organs proper to both sexes, that is, continued +androgynous after it had acquired the chief distinctions of its proper +class, and therefore after it had diverged from the lower classes of the +vertebrate kingdom? This seems improbable in the highest degree; for had +this been the case, we might have expected that some few members of the +two lower classes, namely fishes[279] and amphibians, would still have +remained androgynous. We must, on the contrary, believe that when the +five vertebrate classes diverged from their common progenitor the sexes +had already become separated. To account, however, for male mammals +possessing rudiments of the accessory female organs, and for female +mammals possessing rudiments of the masculine organs, we need not +suppose that their early progenitors were still androgynous after they +had assumed their chief mammalian characters. It is quite possible that +as the one sex gradually acquired the accessory organs proper to it, +some of the successive steps or modifications were transmitted to the +opposite sex. When we treat of sexual selection, we shall meet with +innumerable instances of this form of transmission,--as in the case of +the spurs, plumes, and brilliant colours, acquired by male birds for +battle or ornament, and transferred to the females in an imperfect or +rudimentary condition. + +The possession by male mammals of functionally imperfect mammary organs +is, in some respects, especially curious. The Monotremata have the +proper milk-secreting glands with orifices, but no nipples; and as these +animals stand at the very base of the mammalian series, it is probable +that the progenitors of the class possessed, in like manner, the +milk-secreting glands, but no nipples. This conclusion is supported by +what is known of their manner of development; for Professor Turner +informs me, on the authority of Koelliker and Lauger, that in the embryo +the mammary glands can be distinctly traced before the nipples are in +the least visible; and it should be borne in mind that the development +of successive parts in the individual generally seems to represent and +accord with the development of successive beings in the same line of +descent. The Marsupials differ from the Monotremata by possessing +nipples; so that these organs were probably first acquired by the +Marsupials after they had diverged from, and risen above, the +Monotremata, and were then transmitted to the placental mammals. No one +will suppose that after the Marsupials had approximately acquired their +present structure, and therefore at a rather late period in the +development of the mammalian series, any of its members still remained +androgynous. We seem, therefore, compelled to recur to the foregoing +view, and to conclude that the nipples were first developed in the +females of some very early marsupial form, and were then, in accordance +with a common law of inheritance, transferred in a functionally +imperfect condition to the males. + +Nevertheless a suspicion has sometimes crossed my mind that long after +the progenitors of the whole mammalian class had ceased to be +androgynous, both sexes might have yielded milk and thus nourished their +young; and in the case of the Marsupials, that both sexes might have +carried their young in marsupial sacks. This will not appear utterly +incredible, if we reflect that the males of syngnathous fishes receive +the eggs of the females in their abdominal pouches, hatch them, and +afterwards, as some believe, nourish the young;[280]--that certain other +male fishes hatch the eggs within their mouths or branchial +cavities;--that certain male toads take the chaplets of eggs from the +females and wind them round their own thighs, keeping them there until +the tadpoles are born;--that certain male birds undertake the whole duty +of incubation, and that male pigeons, as well as the females, feed their +nestlings with a secretion from their crops. But the above suspicion +first occurred to me from the mammary glands in male mammals being +developed so much more perfectly than the rudiments of those other +accessory reproductive parts, which are found in the one sex though +proper to the other. The mammary glands and nipples, as they exist in +male mammals, can indeed hardly be called rudimentary; they are simply +not fully developed and not functionally active. They are +sympathetically affected under the influence of certain diseases, like +the same organs in the female. At birth they often secrete a few drops +of milk; and they have been known occasionally in man and other mammals +to become well developed, and to yield a fair supply of milk. Now if we +suppose that during a former prolonged period male mammals aided the +females in nursing their offspring, and that afterwards from some cause, +as from a smaller number of young being produced, the males ceased +giving this aid, disuse of the organs during maturity would lead to +their becoming inactive; and from two well-known principles of +inheritance this state of inactivity would probably be transmitted to +the males at the corresponding age of maturity. But at all earlier ages +these organs would be left unaffected, so that they would be equally +well developed in the young of both sexes. + + +_Conclusion._--The best definition of advancement or progress in the +organic scale ever given, is that by Von Baer; and this rests on the +amount of differentiation and specialisation of the several parts of the +same being, when arrived, as I should be inclined to add, at maturity. +Now as organisms have become slowly adapted by means of natural +selection for diversified lines of life, their parts will have become, +from the advantage gained by the division of physiological labour, more +and more differentiated and specialised for various functions. The same +part appears often to have been modified first for one purpose, and then +long afterwards for some other and quite distinct purpose; and thus all +the parts are rendered more and more complex. But each organism will +still retain the general type of structure of the progenitor from which +it was aboriginally derived. In accordance with this view it seems, if +we turn to geological evidence, that organisation on the whole has +advanced throughout the world by slow and interrupted steps. In the +great kingdom of the Vertebrata it has culminated in man. It must not, +however, be supposed that groups of organic beings are always supplanted +and disappear as soon as they have given birth to other and more perfect +groups. The latter, though victorious over their predecessors, may not +have become better adapted for all places in the economy of nature. Some +old forms appear to have survived from inhabiting protected sites, where +they have not been exposed to very severe competition; and these often +aid us in constructing our genealogies, by giving us a fair idea of +former and lost populations. But we must not fall into the error of +looking at the existing members of any lowly-organised group as perfect +representatives of their ancient predecessors. + +The most ancient progenitors in the kingdom of the Vertebrata, at which +we are able to obtain an obscure glance, apparently consisted of a group +of marine animals,[281] resembling the larvae of existing Ascidians. +These animals probably gave rise to a group of fishes, as lowly +organised as the lancelet; and from these the Ganoids, and other fishes +like the Lepidosiren, must have been developed. From such fish a very +small advance would carry us on to the amphibians. We have seen that +birds and reptiles were once intimately connected together; and the +Monotremata now, in a slight degree, connect mammals with reptiles. But +no one can at present say by what line of descent the three higher and +related classes, namely, mammals, birds, and reptiles, were derived from +either of the two lower vertebrate classes, namely amphibians and +fishes. In the class of mammals the steps are not difficult to conceive +which led from the ancient Monotremata to the ancient Marsupials; and +from these to the early progenitors of the placental mammals. We may +thus ascend to the Lemuridae; and the interval is not wide from these to +the Simiadae. The Simiadae then branched off into two great stems, the New +World and Old World monkeys; and from the latter, at a remote period, +Man, the wonder and glory of the Universe, proceeded. + +Thus we have given to man a pedigree of prodigious length, but not, it +may be said, of noble quality. The world, it has often been remarked, +appears as if it had long been preparing for the advent of man; and +this, in one sense is strictly true, for he owes his birth to a long +line of progenitors. If any single link in this chain had never existed, +man would not have been exactly what he now is. Unless we wilfully close +our eyes, we may, with our present knowledge, approximately recognise +our parentage; nor need we feel ashamed of it. The most humble organism +is something much higher than the inorganic dust under our feet; and no +one with an unbiassed mind can study any living creature, however +humble, without being struck with enthusiasm at its marvellous structure +and properties. + + + + +CHAPTER VII. + +ON THE RACES OF MAN. + + + The nature and value of specific characters--Application to the + races of man--Arguments in favour of, and opposed to, ranking + the so-called races of man as distinct + species--Sub-species--Monogenists and polygenists--Convergence + of character--Numerous points of resemblance in body and mind + between the most distinct races of man--The state of man when + he first spread over the earth--Each race not descended from a + single pair--The extinction of races--The formation of + races--The effects of crossing--Slight influence of the direct + action of the conditions of life--Slight or no influence of + natural selection--Sexual selection. + + +It is not my intention here to describe the several so-called races of +men; but to inquire what is the value of the differences between them +under a classificatory point of view, and how they have originated. In +determining whether two or more allied forms ought to be ranked as +species or varieties, naturalists are practically guided by the +following considerations; namely, the amount of difference between them, +and whether such differences relate to few or many points of structure, +and whether they are of physiological importance; but more especially +whether they are constant. Constancy of character is what is chiefly +valued and sought for by naturalists. Whenever it can be shewn, or +rendered probable, that the forms in question have remained distinct for +a long period, this becomes an argument of much weight in favour of +treating them as species. Even a slight degree of sterility between any +two forms when first crossed, or in their offspring, is generally +considered as a decisive test of their specific distinctness; and their +continued persistence without blending within the same area, is usually +accepted as sufficient evidence, either of some degree of mutual +sterility, or in the case of animals of some repugnance to mutual +pairing. + +Independently of blending from intercrossing, the complete absence, in a +well-investigated region, of varieties linking together any two +closely-allied forms, is probably the most important of all the +criterions of their specific distinctness; and this is a somewhat +different consideration from mere constancy of character, for two forms +may be highly variable and yet not yield intermediate varieties. +Geographical distribution is often unconsciously and sometimes +consciously brought into play; so that forms living in two widely +separated areas, in which most of the other inhabitants are specifically +distinct, are themselves usually looked at as distinct; but in truth +this affords no aid in distinguishing geographical races from so-called +good or true species. + +Now let us apply these generally-admitted principles to the races of +man, viewing him in the same spirit as a naturalist would any other +animal. In regard to the amount of difference between the races, we must +make some allowance for our nice powers of discrimination gained by the +long habit of observing ourselves. In India, as Elphinstone +remarks,[282] although a newly-arrived European cannot at first +distinguish the various native races, yet they soon appear to him +extremely dissimilar; and the Hindoo cannot at first perceive any +difference between the several European nations. Even the most distinct +races of man, with the exception of certain negro tribes, are much more +like each other in form than would at first be supposed. This is well +shewn by the French photographs in the Collection Anthropologique du +Museum of the men belonging to various races, the greater number of +which, as many persons to whom I have shown them have remarked, might +pass for Europeans. Nevertheless, these men if seen alive would +undoubtedly appear very distinct, so that we are clearly much influenced +in our judgment by the mere colour of the skin and hair, by slight +differences in the features, and by expression. + +There is, however, no doubt that the various races, when carefully +compared and measured, differ much from each other,--as in the texture +of the hair, the relative proportions of all parts of the body,[283] the +capacity of the lungs, the form and capacity of the skull, and even in +the convolutions of the brain.[284] But it would be an endless task to +specify the numerous points of structural difference. The races differ +also in constitution, in acclimatisation, and in liability to certain +diseases. Their mental characteristics are likewise very distinct; +chiefly as it would appear in their emotional, but partly in their +intellectual, faculties. Every one who has had the opportunity of +comparison, must have been struck with the contrast between the +taciturn, even morose, aborigines of S. America and the light-hearted, +talkative negroes. There is a nearly similar contrast between the Malays +and the Papuans,[285] who live under the same physical conditions, and +are separated from each other only by a narrow space of sea. + +We will first consider the arguments which may be advanced in favour of +classing the races of man as distinct species, and then those on the +other side. If a naturalist, who had never before seen such beings, were +to compare a Negro, Hottentot, Australian, or Mongolian, he would at +once perceive that they differed in a multitude of characters, some of +slight and some of considerable importance. On inquiry he would find +that they were adapted to live under widely different climates, and that +they differed somewhat in bodily constitution and mental disposition. If +he were then told that hundreds of similar specimens could be brought +from the same countries, he would assuredly declare that they were as +good species as many to which he had been in the habit of affixing +specific names. This conclusion would be greatly strengthened as soon as +he had ascertained that these forms had all retained the same character +for many centuries; and that negroes, apparently identical with existing +negroes, had lived at least 4000 years ago.[286] He would also hear from +an excellent observer, Dr. Lund,[287] that the human skulls found in +the caves of Brazil, entombed with many extinct mammals, belonged to the +same type as that now prevailing throughout the American Continent. + +Our naturalist would then perhaps turn to geographical distribution, and +he would probably declare that forms differing not only in appearance, +but fitted for the hottest and dampest or driest countries, as well as +for the arctic regions, must be distinct species. He might appeal to the +fact that no one species in the group next to man, namely the +Quadrumana, can resist a low temperature or any considerable change of +climate; and that those species which come nearest to man have never +been reared to maturity, even under the temperate climate of Europe. He +would be deeply impressed with the fact, first noticed by Agassiz,[288] +that the different races of man are distributed over the world in the +same zoological provinces, as those inhabited by undoubtedly distinct +species and genera of mammals. This is manifestly the case with the +Australian, Mongolian, and Negro races of man; in a less well-marked +manner with the Hottentots; but plainly with the Papuans and Malays, who +are separated, as Mr. Wallace has shewn, by nearly the same line which +divides the great Malayan and Australian zoological provinces. The +aborigines of America range throughout the Continent; and this at first +appears opposed to the above rule, for most of the productions of the +Southern and Northern halves differ widely; yet some few living forms, +as the opossum, range from the one into the other, as did formerly some +of the gigantic Edentata. The Esquimaux, like other Arctic animals, +extend round the whole polar regions. It should be observed that the +mammalian forms which inhabit the several zoological provinces, do not +differ from each other in the same degree; so that it can hardly be +considered as an anomaly that the Negro differs more, and the American +much less, from the other races of man than do the mammals of the same +continents from those of the other provinces. Man, it may be added, does +not appear to have aboriginally inhabited any oceanic island; and in +this respect he resembles the other members of his class. + +In determining whether the varieties of the same kind of domestic animal +should be ranked as specifically distinct, that is, whether any of them +are descended from distinct wild species, every naturalist would lay +much stress on the fact, if established, of their external parasites +being specifically distinct. All the more stress would be laid on this +fact, as it would be an exceptional one, for I am informed by Mr. Denny +that the most different kinds of dogs, fowls, and pigeons, in England, +are infested by the same species of Pediculi or lice. Now Mr. A. Murray +has carefully examined the Pediculi collected in different countries +from the different races of man;[289] and he finds that they differ, not +only in colour, but in the structure of their claws and limbs. In every +case in which numerous specimens were obtained the differences were +constant. The surgeon of a whaling ship in the Pacific assured me that +when the Pediculi, with which some Sandwich Islanders on board swarmed, +strayed on to the bodies of the English sailors, they died in the course +of three or four days. These Pediculi were darker coloured and appeared +different from those proper to the natives of Chiloe in South America, +of which he gave me specimens. These, again, appeared larger and much +softer than European lice. Mr. Murray procured four kinds from Africa, +namely from the Negroes of the Eastern and Western coasts, from the +Hottentots and Caffres; two kinds from the natives of Australia; two +from North, and two from South America. In these latter cases it may be +presumed that the Pediculi came from natives inhabiting different +districts. With insects slight structural differences, if constant, are +generally esteemed of specific value: and the fact of the races of man +being infested by parasites, which appear to be specifically distinct, +might fairly be urged as an argument that the races themselves ought to +be classed as distinct species. + +Our supposed naturalist having proceeded thus far in his investigation, +would next inquire whether the races of men, when crossed, were in any +degree sterile. He might consult the work[290] of a cautious and +philosophical observer, Professor Broca; and in this he would find good, +evidence that some races were quite fertile together; but evidence of an +opposite nature in regard to other races. Thus it has been asserted that +the native women of Australia and Tasmania rarely produce children to +European men; the evidence, however, on this head has now been shewn to +be almost valueless. The half-castes are killed by the pure blacks; and +an account has lately been published of eleven half-caste youths +murdered and burnt at the same time, whose remains were found by the +police.[291] Again, it has often been said that when mulattoes +intermarry they produce few children; on the other hand, Dr. Bachman of +Charlestown[292] positively asserts that he has known mulatto families +which have intermarried for several generations, and have continued on +an average as fertile as either pure whites or pure blacks. Inquiries +formerly made by Sir C. Lyell on this subject led him, as he informs me, +to the same conclusion. In the United States the census for the year +1854 included, according to Dr. Bachman, 405,751 mulattoes; and this +number, considering all the circumstances of the case, seems small; but +it may partly be accounted for by the degraded and anomalous position of +the class, and by the profligacy of the women. A certain amount of +absorption of mulattoes into negroes must always be in progress; and +this would lead to an apparent diminution of the former. The inferior +vitality of mulattoes is spoken of in a trustworthy work[293] as a +well-known phenomenon; but this is a different consideration from their +lessened fertility; and can hardly be advanced as a proof of the +specific distinctness of the parent races. No doubt both animal and +vegetable hybrids, when produced from extremely distinct species, are +liable to premature death; but the parents of mulattoes cannot be put +under the category of extremely distinct species. The common Mule, so +notorious for long life and vigour, and yet so sterile, shews how little +necessary connection there is in hybrids between lessened fertility and +vitality: other analogous cases could be added. + +Even if it should hereafter be proved that all the races of men were +perfectly fertile together, he who was inclined from other reasons to +rank them as distinct species, might with justice argue that fertility +and sterility are not safe criterions of specific distinctness. We know +that these qualities are easily affected by changed conditions of life +or by close inter-breeding, and that they are governed by highly complex +laws, for instance that of the unequal fertility of reciprocal crosses +between the same two species. With forms which must be ranked as +undoubted species, a perfect series exists from those which are +absolutely sterile when crossed, to those which are almost or quite +fertile. The degrees of sterility do not coincide strictly with the +degrees of difference in external structure or habits of life. Man in +many respects may be compared with those animals which have long been +domesticated, and a large body of evidence can be advanced in favour of +the Pallasian doctrine[294] that domestication tends to eliminate the +sterility which is so general a result of the crossing of species in a +state of nature. From these several considerations, it may be justly +urged that the perfect fertility of the intercrossed races of man, if +established, would not absolutely preclude us from ranking them as +distinct species. + +Independently of fertility, the character of the offspring from a cross +has sometimes been thought to afford evidence whether the parent-forms +ought to be ranked as species or varieties; but after carefully studying +the evidence, I have come to the conclusion that no general rules of +this kind can be trusted. Thus with mankind the offspring of distinct +races resemble in all respects the offspring of true species and of +varieties. This is shewn, for instance, by the manner in which the +characters of both parents are blended, and by one form absorbing +another through repeated crosses. In this latter case the progeny both +of crossed species and varieties retain for a long period a tendency to +revert to their ancestors, especially to that one which is prepotent in +transmission. When any character has suddenly appeared in a race or +species as the result of a single act of variation, as is general with +monstrosities,[295] and this race is crossed with another not thus +characterised, the characters in question do not commonly appear in a +blended condition in the young, but are transmitted to them either +perfectly developed or not at all. As with the crossed races of man +cases of this kind rarely or never occur, this may be used as an +argument against the view suggested by some ethnologists, namely that +certain characters, for instance the blackness of the negro, first +appeared as a sudden variation or sport. Had this occurred, it is +probable that mulattoes would often have been born, either completely +black or completely white. + +We have now seen that a naturalist might feel himself fully justified in +ranking the races of man as distinct species; for he has found that they +are distinguished by many differences in structure and constitution, +some being of importance. These differences have, also, remained nearly +constant for very long periods of time. He will have been in some degree +influenced by the enormous range of man, which is a great anomaly in the +class of mammals, if mankind be viewed as a single species. He will have +been struck with the distribution of the several so-called races, in +accordance with that of other undoubtedly distinct species of mammals. +Finally he might urge that the mutual fertility of all the races has not +as yet been fully proved; and even if proved would not be an absolute +proof of their specific identity. + + +On the other side of the question, if our supposed naturalist were to +enquire whether the forms of man kept distinct like ordinary species, +when mingled together in large numbers in the same country, he would +immediately discover that this was by no means the case. In Brazil he +would behold an immense mongrel population of Negroes and Portuguese; in +Chiloe and other parts of South America, he would behold the whole +population consisting of Indians and Spaniards blended in various +degrees.[296] In many parts of the same continent he would meet with the +most complex crosses between Negroes, Indians, and Europeans; and such +triple crosses afford the severest test, judging from the vegetable +kingdom, of the mutual fertility of the parent-forms. In one island of +the Pacific he would find a small population of mingled Polynesian and +English blood; and in the Viti Archipelago a population of Polynesians +and Negritos crossed in all degrees. Many analogous cases could be +added, for instance, in South Africa. Hence the races of man are not +sufficiently distinct to co-exist without fusion; and this it is, which +in all ordinary cases affords the usual test of specific distinctness. + +Our naturalist would likewise be much disturbed as soon as he perceived +that the distinctive characters of every race of man were highly +variable. This strikes every one when he first beholds the negro-slaves +in Brazil, who have been imported from all parts of Africa. The same +remark holds good with the Polynesians, and with many other races. It +may be doubted whether any character can be named which is distinctive +of a race and is constant. Savages, even within the limits of the same +tribe, are not nearly so uniform in character, as has often been said. +Hottentot women offer certain peculiarities, more strongly marked than +those occurring in any other race, but these are known not to be of +constant occurrence. In the several American tribes, colour and +hairyness differ considerably; as does colour to a certain degree, and +the shape of the features greatly, in the Negroes of Africa. The shape +of the skull varies much in some races;[297] and so it is with every +other character. Now all naturalists have learnt by dearly-bought +experience, how rash it is to attempt to define species by the aid of +inconstant characters. + +But the most weighty of all the arguments against treating the races of +man as distinct species, is that they graduate into each other, +independently in many cases, as far as we can judge, of their having +intercrossed. Man has been studied more carefully than any other organic +being, and yet there is the greatest possible diversity amongst capable +judges whether he should be classed as a single species or race, or as +two (Virey), as three (Jacquinot), as four (Kant), five (Blumenbach), +six (Buffon), seven (Hunter), eight (Agassiz), eleven (Pickering), +fifteen (Bory St. Vincent), sixteen (Desmoulins), twenty-two (Morton), +sixty (Crawfurd), or as sixty-three, according to Burke.[298] This +diversity of judgment does not prove that the races ought not to be +ranked as species, but it shews that they graduate into each other, and +that it is hardly possible to discover clear distinctive characters +between them. + +Every naturalist who has had the misfortune to undertake the +description of a group of highly varying organisms, has encountered +cases (I speak after experience) precisely like that of man; and if of a +cautious disposition, he will end by uniting all the forms which +graduate into each other as a single species; for he will say to himself +that he has no right to give names to objects which he cannot define. +Cases of this kind occur in the Order which includes man, namely in +certain genera of monkeys; whilst in other genera, as in Cercopithecus, +most of the species can be determined with certainty. In the American +genus Cebus, the various forms are ranked by some naturalists as +species, by others as mere geographical races. Now if numerous specimens +of Cebus were collected from all parts of South America, and those forms +which at present appear to be specifically distinct, were found to +graduate into each other by close steps, they would be ranked by most +naturalists as mere varieties or races; and thus the greater number of +naturalists have acted with respect to the races of man. Nevertheless it +must be confessed that there are forms, at least in the vegetable +kingdom,[299] which we cannot avoid naming as species, but which are +connected together, independently of intercrossing, by numberless +gradations. + +Some naturalists have lately employed the term "sub-species" to +designate forms which possess many of the characteristics of true +species, but which hardly deserve so high a rank. Now if we reflect on +the weighty arguments, above given, for raising the races of man to the +dignity of species, and the insuperable difficulties on the other side +in defining them, the term "sub-species" might here be used with much +propriety. But from long habit the term "race" will perhaps always be +employed. The choice of terms is only so far important as it is highly +desirable to use, as far as that may be possible, the same terms for the +same degrees of difference. Unfortunately this is rarely possible; for +within the same family the larger genera generally include +closely-allied forms, which can be distinguished only with much +difficulty, whilst the smaller genera include forms that are perfectly +distinct; yet all must equally be ranked as species. So again the +species within the same large genus by no means resemble each other to +the same degree: on the contrary, in most cases some of them can be +arranged in little groups round other species, like satellites round +planets.[300] + + +The question whether mankind consists of one or several species has of +late years been much agitated by anthropologists, who are divided into +two schools of monogenists and polygenists. Those who do not admit the +principle of evolution, must look at species either as separate +creations or as in some manner distinct entities; and they must decide +what forms to rank as species by the analogy of other organic beings +which are commonly thus received. But it is a hopeless endeavour to +decide this point on sound grounds, until some definition of the term +"species" is generally accepted; and the definition must not include an +element which cannot possibly be ascertained, such as an act of +creation. We might as well attempt without any definition to decide +whether a certain number of houses should be called a village, or town, +or city. We have a practical illustration of the difficulty in the +never-ending doubts whether many closely-allied mammals, birds, +insects, and plants, which represent each other in North America and +Europe, should be ranked species or geographical races; and so it is +with the productions of many islands situated at some little distance +from the nearest continent. + +Those naturalists, on the other hand, who admit the principle of +evolution, and this is now admitted by the greater number of rising men, +will feel no doubt that all the races of man are descended from a single +primitive stock; whether or not they think fit to designate them as +distinct species, for the sake of expressing their amount of +difference.[301] With our domestic animals the question whether the +various races have arisen from one or more species is different. +Although all such races, as well as all the natural species within the +same genus, have undoubtedly sprung from the same primitive stock, yet +it is a fit subject for discussion, whether, for instance, all the +domestic races of the dog have acquired their present differences since +some one species was first domesticated and bred by man; or whether they +owe some of their characters to inheritance from distinct species, which +had already been modified in a state of nature. With mankind no such +question can arise, for he cannot be said to have been domesticated at +any particular period. + +When the races of man diverged at an extremely remote epoch from their +common progenitor, they will have differed but little from each other, +and been few in number; consequently they will then, as far as their +distinguishing characters are concerned, have had less claim to rank as +distinct species, than the existing so-called races. Nevertheless such +early races would perhaps have been ranked by some naturalists as +distinct species, so arbitrary is the term, if their differences, +although extremely slight, had been more constant than at present, and +had not graduated into each other. + +It is, however, possible, though far from probable, that the early +progenitors of man might at first have diverged much in character, until +they became more unlike each other than are any existing races; but that +subsequently, as suggested by Vogt,[302] they converged in character. +When man selects for the same object the offspring of two distinct +species, he sometimes induces, as far as general appearance is +concerned, a considerable amount of convergence. This is the case, as +shewn by Von Nathusius,[303] with the improved breeds of pigs, which are +descended from two distinct species; and in a less well-marked manner +with the improved breeds of cattle. A great anatomist, Gratiolet, +maintains that the anthropomorphous apes do not form a natural +sub-group; but that the orang is a highly developed gibbon or +semnopithecus; the chimpanzee a highly developed macacus; and the +gorilla a highly developed mandrill. If this conclusion, which rests +almost exclusively on brain-characters, be admitted, we should have a +case of convergence at least in external characters, for the +anthropomorphous apes are certainly more like each other in many points +than they are to other apes. All analogical resemblances, as of a whale +to a fish, may indeed be said to be cases of convergence; but this term +has never been applied to superficial and adaptive resemblances. It +would be extremely rash in most cases to attribute to convergence close +similarity in many points of structure in beings which had once been +widely different. The form of a crystal is determined solely by the +molecular forces, and it is not surprising that dissimilar substances +should sometimes assume the same form; but with organic beings we should +bear in mind that the form of each depends on an infinitude of complex +relations, namely on the variations which have arisen, these being due +to causes far too intricate to be followed out,--on the nature of the +variations which have been preserved, and this depends on the +surrounding physical conditions, and in a still higher degree on the +surrounding organisms with which each has come into competition,--and +lastly, on inheritance (in itself a fluctuating element) from +innumerable progenitors, all of which have had their forms determined +through equally complex relations. It appears utterly incredible that +two organisms, if differing in a marked manner, should ever afterwards +converge so closely as to lead to a near approach to identity throughout +their whole organisation. In the case of the convergent pigs above +referred to, evidence of their descent from two primitive stocks is +still plainly retained, according to Von Nathusius, in certain bones of +their skulls. If the races of man were descended, as supposed by some +naturalists, from two or more distinct species, which had differed as +much, or nearly as much, from each other, as the orang differs from the +gorilla, it can hardly be doubted that marked differences in the +structure of certain bones would still have been discoverable in man as +he now exists. + + +Although the existing races of man differ in many respects, as in +colour, hair, shape of skull, proportions of the body, &c., yet if their +whole organisation be taken into consideration they are found to +resemble each other closely in a multitude of points. Many of these +points are of so unimportant or of so singular a nature, that it is +extremely improbable that they should have been independently acquired +by aboriginally distinct species or races. The same remark holds good +with equal or greater force with respect to the numerous points of +mental similarity between the most distinct races of man. The American +aborigines, Negroes and Europeans differ as much from each other in mind +as any three races that can be named; yet I was incessantly struck, +whilst living with the Fuegians on board the "Beagle," with the many +little traits of character, shewing how similar their minds were to +ours; and so it was with a full-blooded negro with whom I happened once +to be intimate. + +He who will carefully read Mr. Tylor's and Sir J. Lubbock's interesting +works[304] can hardly fail to be deeply impressed with the close +similarity between the men of all races in tastes, dispositions and +habits. This is shewn by the pleasure which they all take in dancing, +rude music, acting, painting, tattooing, and otherwise decorating +themselves,--in their mutual comprehension of gesture-language--and, as +I shall be able to shew in a future essay, by the same expression in +their features, and by the same inarticulate cries, when they are +excited by various emotions. This similarity, or rather identity, is +striking, when contrasted with the different expressions which may be +observed in distinct species of monkeys. There is good evidence that the +art of shooting with bows and arrows has not been handed down from any +common progenitor of mankind, yet the stone arrow-heads, brought from +the most distant parts of the world and manufactured at the most remote +periods, are, as Nilsson has shewn,[305] almost identical; and this fact +can only be accounted for by the various races having similar inventive +or mental powers. The same observation has been made by +archaeologists[306] with respect to certain widely-prevalent ornaments, +such as zigzags, &c.; and with respect to various simple beliefs and +customs, such as the burying of the dead under megalithic structures. I +remember observing in South America,[307] that there, as in so many +other parts of the world, man has generally chosen the summits of lofty +hills, on which to throw up piles of stones, either for the sake of +recording some remarkable event, or for burying his dead. + +Now when naturalists observe a close agreement in numerous small details +of habits, tastes and dispositions between two or more domestic races, +or between nearly-allied natural forms, they use this fact as an +argument that all are descended from a common progenitor who was thus +endowed; and consequently that all should be classed under the same +species. The same argument may be applied with much force to the races +of man. + +As it is improbable that the numerous and unimportant points of +resemblance between the several races of man in bodily structure and +mental faculties (I do not here refer to similar customs) should all +have been independently acquired, they must have been inherited from +progenitors who were thus characterised. We thus gain some insight into +the early state of man, before he had spread step by step over the face +of the earth. The spreading of man to regions widely separated by the +sea, no doubt, preceded any considerable amount of divergence of +character in the several races; for otherwise we should sometimes meet +with the same race in distinct continents; and this is never the case. +Sir J. Lubbock, after comparing the arts now practised by savages in all +parts of the world, specifies those which man could not have known, when +he first wandered from his original birthplace; for if once learnt they +would never have been forgotten.[308] He thus shews that "the spear, +which is but a development of the knife-point, and the club, which is +but a long hammer, are the only things left." He admits, however, that +the art of making fire probably had already been discovered, for it is +common to all the races now existing, and was known to the ancient +cave-inhabitants of Europe. Perhaps the art of making rude canoes or +rafts was likewise known; but as man existed at a remote epoch, when the +land in many places stood at a very different level, he would have been +able, without the aid of canoes, to have spread widely. Sir J. Lubbock +further remarks how improbable it is that our earliest ancestors could +have "counted as high as ten, considering that so many races now in +existence cannot get beyond four." Nevertheless, at this early period, +the intellectual and social faculties of man could hardly have been +inferior in any extreme degree to those now possessed by the lowest +savages; otherwise primeval man could not have been so eminently +successful in the struggle for life, as proved by his early and wide +diffusion. + +From the fundamental differences between certain languages, some +philologists have inferred that when man first became widely diffused he +was not a speaking animal; but it may be suspected that languages, far +less perfect than any now spoken, aided by gestures, might have been +used, and yet have left no traces on subsequent and more +highly-developed tongues. Without the use of some language, however +imperfect, it appears doubtful whether man's intellect could have risen +to the standard implied by his dominant position at an early period. + +Whether primeval man, when he possessed very few arts of the rudest +kind, and when his power of language was extremely imperfect, would have +deserved to be called man, must depend on the definition which we +employ. In a series of forms graduating insensibly from some ape-like +creature to man as he now exists, it would be impossible to fix on any +definite point when the term "man" ought to be used. But this is a +matter of very little importance. So again it is almost a matter of +indifference whether the so-called races of man are thus designated, or +are ranked as species or sub-species; but the latter term appears the +most appropriate. Finally, we may conclude that when the principles of +evolution are generally accepted, as they surely will be before long, +the dispute between the monogenists and the polygenists will die a +silent and unobserved death. + + +One other question ought not to be passed over without notice, namely, +whether, as is sometimes assumed, each sub-species or race of man has +sprung from a single pair of progenitors. With our domestic animals a +new race can readily be formed from a single pair possessing some new +character, or even from a single individual thus characterised, by +carefully matching the varying offspring; but most of our races have +been formed, not intentionally from a selected pair, but unconsciously +by the preservation of many individuals which have varied, however +slightly, in some useful or desired manner. If in one country stronger +and heavier horses, and in another country lighter and fleeter horses, +were habitually preferred, we may feel sure that two distinct sub-breeds +would, in the course of time, be produced, without any particular pairs +or individuals having been separated and bred from in either country. +Many races have been thus formed, and their manner of formation is +closely analogous with that of natural species. We know, also, that the +horses which have been brought to the Falkland Islands have become, +during successive generations, smaller and weaker, whilst those which +have run wild on the Pampas have acquired larger and coarser heads; and +such changes are manifestly due, not to any one pair, but to all the +individuals having been subjected to the same conditions, aided, +perhaps, by the principle of reversion. The new sub-breeds in none of +these cases are descended from any single pair, but from many +individuals which have varied in different degrees, but in the same +general manner; and we may conclude that the races of man have been +similarly produced, the modifications being either the direct result of +exposure to different conditions, or the indirect result of some form of +selection. But to this latter subject we shall presently return. + + +_On the Extinction of the Races of Man._--The partial and complete +extinction of many races and sub-races of man are historically known +events. Humboldt saw in South America a parrot which was the sole living +creature that could speak the language of a lost tribe. Ancient +monuments and stone implements found in all parts of the world, of which +no tradition is preserved by the present inhabitants, indicate much +extinction. Some small and broken tribes, remnants of former races, +still survive in isolated and generally mountainous districts. In Europe +the ancient races were all, according to Schaaffhausen,[309] "lower in +the scale than the rudest living savages;" they must therefore have +differed, to a certain extent, from any existing race. The remains +described by Professor Broca[310] from Les Eyzies, though they +unfortunately appear to have belonged to a single family, indicate a +race with a most singular combination of low or simious and high +characteristics, and is "entirely different from any other race, ancient +or modern, that we have ever heard of." It differed, therefore, from the +quaternary race of the caverns of Belgium. + +Unfavourable physical conditions appear to have had but little effect in +the extinction of races.[311] Man has long lived in the extreme regions +of the North, with no wood wherewith to make his canoes or other +implements, and with blubber alone for burning and giving him warmth, +but more especially for melting the snow. In the Southern extremity of +America the Fuegians survive without the protection of clothes, or of +any building worthy to be called a hovel. In South Africa the aborigines +wander over the most arid plains, where dangerous beasts abound. Man can +withstand the deadly influence of the Terai at the foot of the Himalaya, +and the pestilential shores of tropical Africa. + +Extinction follows chiefly from the competition of tribe with tribe, +and race with race. Various checks are always in action, as specified in +a former chapter, which serve to keep down the numbers of each savage +tribe,--such as periodical famines, the wandering of the parents and the +consequent deaths of infants, prolonged suckling, the stealing of women, +wars, accidents, sickness, licentiousness, especially infanticide, and, +perhaps, lessened fertility from less nutritious food, and many +hardships. If from any cause any one of these checks is lessened, even +in a slight degree, the tribe thus favoured will tend to increase; and +when one of two adjoining tribes becomes more numerous and powerful than +the other, the contest is soon settled by war, slaughter, cannibalism, +slavery, and absorption. Even when a weaker tribe is not thus abruptly +swept away, if it once begins to decrease, it generally goes on +decreasing until it is extinct.[312] + +When civilised nations come into contact with barbarians the struggle is +short, except where a deadly climate gives its aid to the native race. +Of the causes which lead to the victory of civilised nations, some are +plain and some very obscure. We can see that the cultivation of the land +will be fatal in many ways to savages, for they cannot, or will not, +change their habits. New diseases and vices are highly destructive; and +it appears that in every nation a new disease causes much death, until +those who are most susceptible to its destructive influence are +gradually weeded out;[313] and so it may be with the evil effects from +spirituous liquors, as well as with the unconquerably strong taste for +them shewn by so many savages. It further appears, mysterious as is the +fact, that the first meeting of distinct and separated people generates +disease.[314] Mr. Sproat, who in Vancouver Island closely attended to +the subject of extinction, believes that changed habits of life, which +always follow from the advent of Europeans, induces much ill-health. He +lays, also, great stress on so trifling a cause as that the natives +become "bewildered and dull by the new life around them; they lose the +motives for exertion, and get no new ones in their place."[315] + +The grade of civilisation seems a most important element in the success +of nations which come in competition. A few centuries ago Europe feared +the inroads of Eastern barbarians; now, any such fear would be +ridiculous. It is a more curious fact, that savages did not formerly +waste away, as Mr. Bagehot has remarked, before the classical nations, +as they now do before modern civilised nations; had they done so, the +old moralists would have mused over the event; but there is no lament in +any writer of that period over the perishing barbarians.[316] + +Although the gradual decrease and final extinction of the races of man +is an obscure problem, we can see that it depends on many causes, +differing in different places and at different times. It is the same +difficult problem as that presented by the extinction of one of the +higher animals--of the fossil horse, for instance, which disappeared +from South America, soon afterwards to be replaced, within the same +districts, by countless troops of the Spanish horse. The New Zealander +seems conscious of this parallelism, for he compares his future fate +with that of the native rat almost exterminated by the European rat. The +difficulty, though great to our imagination, and really great if we wish +to ascertain the precise causes, ought not to be so to our reason, as +long as we keep steadily in mind that the increase of each species and +each race is constantly hindered by various checks; so that if any new +check, or cause of destruction, even a slight one, be superadded, the +race will surely decrease in number; and as it has everywhere been +observed that savages are much opposed to any change of habits, by which +means injurious checks could be counterbalanced, decreasing numbers will +sooner or later lead to extinction; the end, in most cases, being +promptly determined by the inroads of increasing and conquering tribes. + + +_On the Formation of the Races of Man._--It may be premised that when we +find the same race, though broken up into distinct tribes, ranging over +a great area, as over America, we may attribute their general +resemblance to descent from a common stock. In some cases the crossing +of races already, distinct has led to the formation of new races. The +singular fact that Europeans and Hindoos, who belong to the same Aryan +stock and speak a language fundamentally the same, differ widely in +appearance, whilst Europeans differ but little from Jews, who belong to +the Semitic stock and speak quite another language, has been accounted +for by Broca[317] through the Aryan branches having been largely crossed +during their wide diffusion by various indigenous tribes. When two races +in close contact cross, the first result is a heterogeneous mixture: +thus Mr. Hunter, in describing the Santali or hill-tribes of India, says +that hundreds of imperceptible gradations may be traced "from the black, +squat tribes of the mountains to the tall olive-coloured Brahman, with +his intellectual brow, calm eyes, and high but narrow head;" so that it +is necessary in courts of justice to ask the witnesses whether they are +Santalis or Hindoos.[318] Whether a heterogeneous people, such as the +inhabitants of some of the Polynesian islands, formed by the crossing of +two distinct races, with few or no pure members left, would ever become +homogeneous, is not known from direct evidence. But as with our +domesticated animals, a crossed breed can certainly, in the course of a +few generations, be fixed and made uniform by careful selection,[319] we +may infer that the free and prolonged intercrossing during many +generations of a heterogeneous mixture would supply the place of +selection, and overcome any tendency to reversion, so that a crossed +race would ultimately become homogeneous, though it might not partake in +an equal degree of the characters of the two parent-races. + +Of all the differences between the races of man, the colour of the skin +is the most conspicuous and one of the best marked. Differences of this +kind, it was formerly thought, could be accounted for by long exposure +under different climates; but Pallas first shewed that this view is not +tenable, and he has been followed by almost all anthropologists.[320] +The view has been rejected chiefly because the distribution of the +variously coloured races, most of whom must have long inhabited their +present homes, does not coincide with corresponding differences of +climate. Weight must also be given to such cases as that of the Dutch +families, who, as we hear on excellent authority,[321] have not +undergone the least change of colour, after residing for three centuries +in South Africa. The uniform appearance in various parts of the world of +gypsies and Jews, though the uniformity of the latter has been somewhat +exaggerated,[322] is likewise an argument on the same side. A very damp +or a very dry atmosphere has been supposed to be more influential in +modifying the colour of the skin than mere heat; but as D'Orbigny in +South America, and Livingstone in Africa, arrived at diametrically +opposite conclusions with respect to dampness and dryness, any +conclusion on this head must be considered as very doubtful.[323] + +Various facts, which I have elsewhere given, prove that the colour of +the skin and hair is sometimes correlated in a surprising manner with a +complete immunity from the action of certain vegetable poisons and from +the attacks of certain parasites. Hence it occurred to me, that negroes +and other dark races might have acquired their dark tints by the darker +individuals escaping during a long series of generations from the deadly +influence of the miasmas of their native countries. + +I afterwards found that the same idea had long ago occurred to Dr. +Wells.[324] That negroes, and even mulattoes, are almost completely +exempt from the yellow-fever, which is so destructive in tropical +America, has long been known.[325] They likewise escape to a large +extent the fatal intermittent fevers that prevail along, at least, 2600 +miles of the shores of Africa, and which annually cause one-fifth of the +white settlers to die, and another fifth to return home invalided.[326] +This immunity in the negro seems to be partly inherent, depending on +some unknown peculiarity of constitution, and partly the result of +acclimatisation. Pouchet[327] states that the negro regiments, borrowed +from the Viceroy of Egypt for the Mexican war, which had been recruited +near the Soudan, escaped the yellow-fever almost equally well with the +negroes originally brought from various parts of Africa, and accustomed +to the climate of the West Indies. That acclimatisation plays a part is +shewn by the many cases in which negroes, after having resided for some +time in a colder climate, have become to a certain extent liable to +tropical fevers.[328] The nature of the climate under which the white +races have long resided, likewise has some influence on them; for during +the fearful epidemic of yellow-fever in Demerara during 1837, Dr. Blair +found that the death-rate of the immigrants was proportional to the +latitude of the country whence they had come. With the negro the +immunity, as far as it is the result of acclimatisation, implies +exposure during a prodigious length of time; for the aborigines of +tropical America, who have resided there from time immemorial, are not +exempt from yellow-fever; and the Rev. B. Tristram states, that there +are districts in Northern Africa which the native inhabitants are +compelled annually to leave, though the negroes can remain with safety. + +That the immunity of the negro is in any degree correlated with the +colour of his skin is a mere conjecture: it may be correlated with some +difference in his blood, nervous system, or other tissues. Nevertheless, +from the facts above alluded to, and from some connection apparently +existing between complexion and a tendency to consumption, the +conjecture seemed to me not improbable. Consequently I endeavoured, with +but little success,[329] to ascertain how far it held good. The late +Dr. Daniell, who had long lived on the West Coast of Africa, told me +that he did not believe in any such relation. He was himself unusually +fair, and had withstood the climate in a wonderful manner. When he first +arrived as a boy on the coast, an old and experienced negro chief +predicted from his appearance that this would prove the case. Dr. +Nicholson, of Antigua, after having attended to this subject, wrote to +me that he did not think that dark-coloured Europeans escaped the +yellow-fever better than those that were light-coloured. Mr. J. M. +Harris altogether denies[330] that Europeans with dark hair withstand a +hot climate better than other men; on the contrary, experience has +taught him in making a selection of men for service on the coast of +Africa, to choose those with red hair. As far, therefore, as these +slight indications serve, there seems no foundation for the hypothesis, +which has been accepted by several writers, that the colour of the black +races may have resulted from darker and darker individuals having +survived in greater numbers, during their exposure to the +fever-generating miasmas of their native countries. + +Although with our present knowledge we cannot account for the +strongly-marked differences in colour between the races of man, either +through correlation with constitutional peculiarities, or through the +direct action of climate; yet we must not quite ignore the latter +agency, for there is good reason to believe that some inherited effect +is thus produced.[331] + +We have seen in our third chapter that the conditions of life, such as +abundant food and general comfort, affect in a direct manner the +development of the bodily frame, the effects being transmitted. Through +the combined influences of climate and changed habits of life, European +settlers, in the United States undergo, as is generally admitted, a +slight but extraordinarily rapid change of appearance. There is, also, a +considerable body of evidence shewing that in the Southern States the +house-slaves of the third generation present a markedly different +appearance from the field-slaves.[332] + +If, however, we look to the races of man, as distributed over the world, +we must infer that their characteristic differences cannot be accounted +for by the direct action of different conditions of life, even after +exposure to them for an enormous period of time. The Esquimaux live +exclusively on animal food; they are clothed in thick fur, and are +exposed to intense cold and to prolonged darkness; yet they do not +differ in any extreme degree from the inhabitants of Southern China, who +live entirely on vegetable food and are exposed almost naked to a hot, +glaring climate. The unclothed Fuegians live on the marine productions +of their inhospitable shores; the Botocudos of Brazil wander about the +hot forests of the interior and live chiefly on vegetable productions; +yet these tribes resemble each other so closely that the Fuegians on +board the "Beagle" were mistaken by some Brazilians for Botocudos. The +Botocudos again, as well as the other inhabitants of tropical America, +are wholly different from the Negroes who inhabit the opposite shores of +the Atlantic, are exposed to a nearly similar climate, and follow nearly +the same habits of life. + +Nor can the differences between the races of man be accounted for, +except to a quite insignificant degree, by the inherited effects of the +increased or decreased use of parts. Men who habitually live in canoes, +may have their legs somewhat stunted; those who inhabit lofty regions +have their chests enlarged; and those who constantly use certain +sense-organs have the cavities in which they are lodged somewhat +increased in size, and their features consequently a little modified. +With civilised nations, the reduced size of the jaws from lessened use, +the habitual play of different muscles serving to express different +emotions, and the increased size of the brain from greater intellectual +activity, have together produced a considerable effect on their general +appearance in comparison with savages.[333] It is also possible that +increased bodily stature, with no corresponding increase in the size of +the brain, may have given to some races (judging from the previously +adduced cases of the rabbits) an elongated skull of the dolichocephalic +type. + +Lastly, the little-understood principle of correlation will almost +certainly have come into action, as in the case of great muscular +development and strongly projecting supra-orbital ridges. It is not +improbable that the texture of the hair, which differs much in the +different races, may stand in some kind of correlation with the +structure of the skin; for the colour of the hair and skin are certainly +correlated, as is its colour and texture with the Mandans.[334] The +colour of the skin and the odour emitted by it are likewise in some +manner connected. With the breeds of sheep the number of hairs within a +given space and the number of the excretory pores stand in some relation +to each other.[335] If we may judge from the analogy of our domesticated +animals, many modifications of structure in man probably come under this +principle of correlated growth. + + +We have now seen that the characteristic differences between the races +of man cannot be accounted for in a satisfactory manner by the direct +action of the conditions of life, nor by the effects of the continued +use of parts, nor through the principle of correlation. We are therefore +led to inquire whether slight individual differences, to which man is +eminently liable, may not have been preserved and augmented during a +long series of generations through natural selection. But here we are at +once met by the objection that beneficial variations alone can be thus +preserved; and as far as we are enabled to judge (although always liable +to error on this head) not one of the external differences between the +races of man are of any direct or special service to him. The +intellectual and moral or social faculties must of course be excepted +from this remark; but differences in these faculties can have had little +or no influence on external characters. The variability of all the +characteristic differences between the races, before referred to, +likewise indicates that these differences cannot be of much importance; +for, had they been important, they would long ago have been either fixed +and preserved, or eliminated. In this respect man resembles those forms, +called by naturalists protean or polymorphic, which have remained +extremely variable, owing, as it seems, to their variations being of an +indifferent nature, and consequently to their having escaped the action +of natural selection. + +We have thus far been baffled in all our attempts to account for the +differences between the races of man; but there remains one important +agency, namely Sexual Selection, which appears to have acted as +powerfully on man, as on many other animals. I do not intend to assert +that sexual selection will account for all the differences between the +races. An unexplained residuum is left, about which we can in our +ignorance only say, that as individuals are continually born with, for +instance, heads a little rounder or narrower, and with noses a little +longer or shorter, such slight differences might become fixed and +uniform, if the unknown agencies which induced them were to act in a +more constant manner, aided by long-continued intercrossing. Such +modifications come under the provisional class, alluded to in our fourth +chapter, which for the want of a better term have been called +spontaneous variations. Nor do I pretend that the effects of sexual +selection can be indicated with scientific precision; but it can be +shewn that it would be an inexplicable fact if man had not been modified +by this agency, which has acted so powerfully on innumerable animals, +both high and low in the scale. It can further be shewn that the +differences between the races of man, as in colour, hairyness, form of +features, &c., are of the nature which it might have been expected would +have been acted on by sexual selection. But in order to treat this +subject in a fitting manner, I have found it necessary to pass the whole +animal kingdom in review; I have therefore devoted to it the Second Part +of this work. At the close I shall return to man, and, after attempting +to shew how far he has been modified through sexual selection, will give +a brief summary of the chapters in this First Part. + + + + +PART II.--SEXUAL SELECTION. + + +CHAPTER VIII. + +PRINCIPLES OF SEXUAL SELECTION. + + + Secondary sexual characters--Sexual selection--Manner of + action--Excess of males--Polygamy--The male alone generally + modified through sexual selection--Eagerness of the + male--Variability of the male--Choice exerted by the + female--Sexual compared with natural selection--Inheritance, at + corresponding periods of life, at corresponding seasons of the + year, and as limited by sex--Relations between the several + forms of inheritance--Causes why one sex and the young are not + modified through sexual selection--Supplement on the + proportional numbers of the two sexes throughout the animal + kingdom--On the limitation of the numbers of the two sexes + through natural selection. + + +With animals which have their sexes separated, the males necessarily +differ from the females in their organs of reproduction; and these +afford the primary sexual characters. But the sexes often differ in what +Hunter has called secondary sexual characters, which are not directly +connected with the act of reproduction; for instance, in the male +possessing certain organs of sense or locomotion, of which the female is +quite destitute, or in having them more highly-developed, in order that +he may readily find or reach her; or again, in the male having special +organs of prehension so as to hold her securely. These latter organs of +infinitely diversified kinds graduate into, and in some cases can hardly +be distinguished from, those which are commonly ranked as primary, such +as the complex appendages at the apex of the abdomen in male insects. +Unless indeed we confine the term "primary" to the reproductive glands, +it is scarcely possible to decide, as far as the organs of prehension +are concerned, which ought to be called primary and which secondary. + +The female often differs from the male in having organs for the +nourishment or protection of her young, as the mammary glands of +mammals, and the abdominal sacks of the marsupials. The male, also, in +some few cases differs from the female in possessing analogous organs, +as the receptacles for the ova possessed by the males of certain fishes, +and those temporarily developed in certain male frogs. Female bees have +a special apparatus for collecting and carrying pollen, and their +ovipositor is modified into a sting for the defence of their larvae and +the community. In the females of many insects the ovipositor is modified +in the most complex manner for the safe placing of the eggs. Numerous +similar cases could be given, but they do not here concern us. There +are, however, other sexual differences quite disconnected with the +primary organs with which we are more especially concerned--such as the +greater size, strength, and pugnacity of the male, his weapons of +offence or means of defence against rivals, his gaudy colouring and +various ornaments, his power of song, and other such characters. + +Besides the foregoing primary and secondary sexual differences, the male +and female sometimes differ in structures connected with different +habits of life, and not at all, or only indirectly, related to the +reproductive functions. Thus the females of certain flies (Culicidae and +Tabanidae) are blood-suckers, whilst the males live on flowers and have +their mouths destitute of mandibles.[336] The males alone of certain +moths and of some crustaceans (_e.g._ Tanais) have imperfect, closed +mouths, and cannot feed. The Complemental males of certain cirripedes +live like epiphytic plants either on the female or hermaphrodite form, +and are destitute of a mouth and prehensile limbs. In these cases it is +the male which has been modified and has lost certain important organs, +which the other members of the same group possess. In other cases it is +the female which has lost such parts; for instance, the female glow-worm +is destitute of wings, as are many female moths, some of which never +leave their cocoons. Many female parasitic crustaceans have lost their +natatory legs. In some weevil-beetles (Curculionidae) there is a great +difference between the male and female in the length of the rostrum or +snout;[337] but the meaning of this and of many analogous differences, +is not at all understood. Differences of structure between the two sexes +in relation to different habits of life are generally confined to the +lower animals; but with some few birds the beak of the male differs from +that of the female. No doubt in most, but apparently not in all these +cases, the differences are indirectly connected with the propagation of +the species: thus a female which has to nourish a multitude of ova will +require more food than the male, and consequently will require special +means for procuring it. A male animal which lived for a very short time +might without detriment lose through disuse its organs for procuring +food; but he would retain his locomotive organs in a perfect state, so +that he might reach the female. The female, on the other hand, might +safely lose her organs for flying, swimming, or walking, if she +gradually acquired habits which rendered such powers useless. + +We are, however, here concerned only with that kind of selection, which +I have called sexual selection. This depends on the advantage which +certain individuals have over other individuals of the same sex and +species, in exclusive relation to reproduction. When the two sexes +differ in structure in relation to different habits of life, as in the +cases above mentioned, they have no doubt been modified through natural +selection, accompanied by inheritance limited to one and the same sex. +So again the primary sexual organs, and those for nourishing or +protecting the young, come under this same head; for those individuals +which generated or nourished their offspring best, would leave, _caeteris +paribus_, the greatest number to inherit their superiority; whilst those +which generated or nourished their offspring badly, would leave but few +to inherit their weaker powers. As the male has to search for the +female, he requires for this purpose organs of sense and locomotion, but +if these organs are necessary for the other purposes of life, as is +generally the case, they will have been developed through natural +selection. When the male has found the female he sometimes absolutely +requires prehensile organs to hold her; thus Dr. Wallace informs me that +the males of certain moths cannot unite with the females if their tarsi +or feet are broken. The males of many oceanic crustaceans have their +legs and antennae modified in an extraordinary manner for the prehension +of the female; hence we may suspect that owing to these animals being +washed about by the waves of the open sea, they absolutely require these +organs in order to propagate their kind, and if so their development +will have been the result of ordinary or natural selection. + +When the two sexes follow exactly the same habits of life, and the male +has more highly developed sense or locomotive organs than the female, it +may be that these in their perfected state are indispensable to the male +for finding the female; but in the vast majority of cases, they serve +only to give one male an advantage over another, for the less +well-endowed males, if time were allowed them, would succeed in pairing +with the females; and they would in all other respects, judging from the +structure of the female, be equally well adapted for their ordinary +habits of life. In such cases sexual selection must have come into +action, for the males have acquired their present structure, not from +being better fitted to survive in the struggle for existence, but from +having gained an advantage over other males, and from having transmitted +this advantage to their male offspring alone. It was the importance of +this distinction which led me to designate this form of selection as +sexual selection. So again, if the chief service rendered to the male by +his prehensile organs is to prevent the escape of the female before the +arrival of other males, or when assaulted by them, these organs will +have been perfected through sexual selection, that is by the advantage +acquired by certain males over their rivals. But in most cases it is +scarcely possible to distinguish between the effects of natural and +sexual selection. Whole chapters could easily be filled with details on +the differences between the sexes in their sensory, locomotive, and +prehensile organs. As, however, these structures are not more +interesting than others adapted for the ordinary purposes of life, I +shall almost pass them over, giving only a few instances under each +class. + +There are many other structures and instincts which must have been +developed through sexual selection--such as the weapons of offence and +the means of defence possessed by the males for fighting with and +driving away their rivals--their courage and pugnacity--their ornaments +of many kinds--their organs for producing vocal or instrumental +music--and their glands for emitting odours; most of these latter +structures serving only to allure or excite the female. That these +characters are the result of sexual and not of ordinary selection is +clear, as unarmed, unornamented, or unattractive males would succeed +equally well in the battle for life and in leaving a numerous progeny, +if better endowed males were not present. We may infer that this would +be the case, for the females, which are unarmed and unornamented, are +able to survive and procreate their kind. Secondary sexual characters of +the kind just referred to, will be fully discussed in the following +chapters, as they are in many respects interesting, but more especially +as they depend on the will, choice, and rivalry of the individuals of +either sex. When we behold two males fighting for the possession of the +female, or several male birds displaying their gorgeous plumage, and +performing the strangest antics before an assembled body of females, we +cannot doubt that, though led by instinct, they know what they are +about, and consciously exert their mental and bodily powers. + +In the same manner as man can improve the breed of his game-cocks by the +selection of those birds which are victorious in the cockpit, so it +appears that the strongest and most vigorous males, or those provided +with the best weapons, have prevailed under nature, and have led to the +improvement of the natural breed or species. Through repeated deadly +contests, a slight degree of variability, if it led to some advantage, +however slight, would suffice for the work of sexual selection; and it +is certain that secondary sexual characters are eminently variable. In +the same manner as man can give beauty, according to his standard of +taste, to his male poultry--can give to the Sebright bantam a new and +elegant plumage, an erect and peculiar carriage--so it appears that in a +state of nature female birds, by having long selected the more +attractive males, have added to their beauty. No doubt this implies +powers of discrimination and taste on the part of the female which will +at first appear extremely improbable; but I hope hereafter to shew that +this is not the case. + +From our ignorance on several points, the precise manner in which sexual +selection acts is to a certain extent uncertain. Nevertheless if those +naturalists who already believe in the mutability of species, will read +the following chapters, they will, I think, agree with me that sexual +selection has played an important part in the history of the organic +world. It is certain that with almost all animals there is a struggle +between the males for the possession of the female. This fact is so +notorious that it would be superfluous to give instances. Hence the +females, supposing that their mental capacity sufficed for the exertion +of a choice, could select one out of several males. But in numerous +cases it appears as if it had been specially arranged that there should +be a struggle between many males. Thus with migratory birds, the males +generally arrive before the females at their place of breeding, so that +many males are ready to contend for each female. The bird-catchers +assert that this is invariably the case with the nightingale and +blackcap, as I am informed by Mr. Jenner Weir, who confirms the +statement with respect to the latter species. + +Mr. Swaysland of Brighton, who has been in the habit, during the last +forty years, of catching our migratory birds on their first arrival, +writes to me that he has never known the females of any species to +arrive before their males. During one spring he shot thirty-nine males +of Ray's wagtail (_Budytes Raii_) before he saw a single female. Mr. +Gould has ascertained by dissection, as he informs me, that male snipes +arrive in this country before the females. In the case of fish, at the +period when the salmon ascend our rivers, the males in large numbers are +ready to breed before the females. So it apparently is with frogs and +toads. Throughout the great class of insects the males almost always +emerge from the pupal state before the other sex, so that they generally +swarm for a time before any females can be seen.[338] The cause of this +difference between the males and females in their periods of arrival and +maturity is sufficiently obvious. Those males which annually first +migrated into any country, or which in the spring were first ready to +breed, or were the most eager, would leave the largest number of +offspring; and these would tend to inherit similar instincts and +constitutions. On the whole there can be no doubt that with almost all +animals, in which the sexes are separate, there is a constantly +recurrent struggle between the males for the possession of the females. + +Our difficulty in regard to sexual selection lies in understanding how +it is that the males which conquer other males, or those which prove the +most attractive to the females, leave a greater number of offspring to +inherit their superiority than the beaten and less attractive males. +Unless this result followed, the characters which gave to certain males +an advantage over others, could not be perfected and augmented through +sexual selection. When the sexes exist in exactly equal numbers, the +worst-endowed males will ultimately find females (excepting where +polygamy prevails), and leave as many offspring, equally well fitted for +their general habits of life, as the best-endowed males. From various +facts and considerations, I formerly inferred that with most animals, in +which secondary sexual characters were well developed, the males +considerably exceeded the females in number; and this does hold good in +some few cases. If the males were to the females as two to one, or as +three to two, or even in a somewhat lower ratio, the whole affair would +be simple; for the better-armed or more attractive males would leave the +largest number of offspring. But after investigating, as far as +possible, the numerical proportions of the sexes, I do not believe that +any great inequality in number commonly exists. In most cases sexual +selection appears to have been effective in the following manner. + +Let us take any species, a bird for instance, and divide the females +inhabiting a district into two equal bodies: the one consisting of the +more vigorous and better-nourished individuals, and the other of the +less vigorous and healthy. The former, there can be little doubt, would +be ready to breed in the spring before the others; and this is the +opinion of Mr. Jenner Weir, who has during many years carefully attended +to the habits of birds. There can also be no doubt that the most +vigorous, healthy, and best-nourished females would on an average +succeed in rearing the largest number of offspring. The males, as we +have seen, are generally ready to breed before the females; of the males +the strongest, and with some species the best armed, drive away the +weaker males; and the former would then unite with the more vigorous and +best-nourished females, as these are the first to breed. Such vigorous +pairs would surely rear a larger number of offspring than the retarded +females, which would be compelled, supposing the sexes to be numerically +equal, to unite with the conquered and less powerful males; and this is +all that is wanted to add, in the course of successive generations, to +the size, strength and courage of the males, or to improve their +weapons. + +But in a multitude of cases the males which conquer other males, do not +obtain possession of the females, independently of choice on the part of +the latter. The courtship of animals is by no means so simple and short +an affair as might be thought. The females are most excited by, or +prefer pairing with, the more ornamented males, or those which are the +best songsters, or play the best antics; but it is obviously probable, +as has been actually observed in some cases, that they would at the same +time prefer the more vigorous and lively males.[339] Thus the more +vigorous females, which are the first to breed, will have the choice of +many males; and though they may not always select the strongest or best +armed, they will select those which are vigorous and well armed, and in +other respects the most attractive. Such early pairs would have the same +advantage in rearing offspring on the female side as above explained, +and nearly the same advantage on the male side. And this apparently has +sufficed during a long course of generations to add not only to the +strength and fighting-powers of the males, but likewise to their +various ornaments or other attractions. + +In the converse and much rarer case of the males selecting particular +females, it is plain that those which were the most vigorous and had +conquered others, would have the freest choice; and it is almost certain +that they would select vigorous as well as attractive females. Such +pairs would have an advantage in rearing offspring, more especially if +the male had the power to defend the female during the pairing-season, +as occurs with some of the higher animals, or aided in providing for the +young. The same principles would apply if both sexes mutually preferred +and selected certain individuals of the opposite sex; supposing that +they selected not only the more attractive, but likewise the more +vigorous individuals. + + +_Numerical Proportion of the Two Sexes._--I have remarked that sexual +selection would be a simple affair if the males considerably exceeded in +number the females. Hence I was led to investigate, as far as I could, +the proportions between the two sexes of as many animals as possible; +but the materials are scanty. I will here give only a brief abstract of +the results, retaining the details for a supplementary discussion, so as +not to interfere with the course of my argument. Domesticated animals +alone afford the opportunity of ascertaining the proportional numbers at +birth; but no records have been specially kept for this purpose. By +indirect means, however, I have collected a considerable body of +statistical data, from which it appears that with most of our domestic +animals the sexes are nearly equal at birth. Thus with race-horses, +25,560 births have been recorded during twenty-one years, and the male +births have been to the female births as 99.7 to 100. With greyhounds +the inequality is greater than with any other animal, for during twelve +years, out of 6878 births, the male births have been as 110.1 to 100 +female births. It is, however, in some degree doubtful whether it is +safe to infer that the same proportional numbers would hold good under +natural conditions as under domestication; for slight and unknown +differences in the conditions affect to a certain extent the proportion +of the sexes. Thus with mankind, the male births in England are as +104.5, in Russia as 108.9, and with the Jews of Livornia as 120 _to_ 100 +females. The proportion is also mysteriously affected by the +circumstance of the births being legitimate or illegitimate. + +For our present purpose we are concerned with the proportion of the +sexes, not at birth, but at maturity, and this adds another element of +doubt; for it is a well ascertained fact that with man a considerably +larger proportion of males than of females die before or during birth, +and during the first few years of infancy. So it almost certainly is +with male lambs, and so it may be with the males of other animals. The +males of some animals kill each other by fighting; or they drive each +other about until they become greatly emaciated. They must, also, whilst +wandering about in eager search for the females, be often exposed to +various dangers. With many kinds of fish the males are much smaller than +the females, and they are believed often to be devoured by the latter or +by other fishes. With some birds the females appear to die in larger +proportion than the males: they are also liable to be destroyed on their +nests, or whilst in charge of their young. With insects the female larvae +are often larger than those of the males, and would consequently be more +likely to be devoured: in some cases the mature females are less active +and less rapid in their movements than the males, and would not be so +well able to escape from danger. Hence, with animals in a state of +nature, in order to judge of the proportions of the sexes at maturity, +we must rely on mere estimation; and this, except perhaps when the +inequality is strongly marked, is but little trustworthy. Nevertheless, +as far as a judgment can be formed, we may conclude from the facts given +in the supplement, that the males of some few mammals, of many birds, of +some fish and insects, considerably exceed in number the females. + +The proportion between the sexes fluctuates slightly during successive +years: thus with race-horses, for every 100 females born, the males +varied from 107.1 in one year to 92.6 in another year, and with +greyhounds from 116.3 to 95.3. But had larger numbers been tabulated +throughout a more extensive area than England, these fluctuations would +probably have disappeared; and such as they are, they would hardly +suffice to lead under a state of nature to the effective action of +sexual selection. Nevertheless with some few wild animals, the +proportions seem, as shewn in the supplement, to fluctuate either during +different seasons or in different localities in a sufficient degree to +lead to such action. For it should be observed that any advantage gained +during certain years or in certain localities by those males which were +able to conquer other males, or were the most attractive to the females, +would probably be transmitted to the offspring and would not +subsequently be eliminated. During the succeeding seasons, when from the +equality of the sexes every male was everywhere able to procure a +female, the stronger or more attractive males previously produced would +still have at least as good a chance of leaving offspring as the less +strong or less attractive. + + +_Polygamy._--The practice of polygamy leads to the same results as +would follow from an actual inequality in the number of the sexes; for +if each male secures two or more females, many males will not be able to +pair; and the latter assuredly will be the weaker or less attractive +individuals. Many mammals and some few birds are polygamous, but with +animals belonging to the lower classes I have found no evidence of this +habit. The intellectual powers of such animals are, perhaps, not +sufficient to lead them to collect and guard a harem of females. That +some relation exists between polygamy and the development of secondary +sexual characters, appears nearly certain; and this supports the view +that a numerical preponderance of males would be eminently favourable to +the action of sexual selection. Nevertheless many animals, especially +birds, which are strictly monogamous, display strongly-marked secondary +sexual characters; whilst some few animals, which are polygamous, are +not thus characterised. + +We will first briefly run through the class of mammals, and then turn to +birds. The gorilla seems to be a polygamist, and the male differs +considerably from the female; so it is with some baboons which live in +herds containing twice as many adult females as males. In South America +the _Mycetes caraya_ presents well-marked sexual differences in colour, +beard, and vocal organs, and the male generally lives with two or three +wives: the male of the _Cebus capucinus_ differs somewhat from the +female, and appears to be polygamous.[340] Little is known on this head +with respect to most other monkeys, but some species are strictly +monogamous. The ruminants are eminently polygamous, and they more +frequently present sexual differences than almost any other group of +mammals, especially in their weapons, but likewise in other characters. +Most deer, cattle, and sheep are polygamous; as are most antelopes, +though some of the latter are monogamous. Sir Andrew Smith, in speaking +of the antelopes of South Africa, says that in herds of about a dozen +there was rarely more than one mature male. The Asiatic _Antilope saiga_ +appears to be the most inordinate polygamist in the world; for +Pallas[341] states that the male drives away all rivals, and collects a +herd of about a hundred, consisting of females and kids: the female is +hornless and has softer hair, but does not otherwise differ much from +the male. The horse is polygamous, but, except in his greater size and +in the proportions of his body, differs but little from the mare. The +wild boar, in his great tusks and some other characters, presents +well-marked sexual characters; in Europe and in India he leads a +solitary life, except during the breeding-season; but at this season he +consorts in India with several females, as Sir W. Elliot, who has had +large experience in observing this animal, believes: whether this holds +good in Europe is doubtful, but is supported by some statements. The +adult male Indian elephant, like the boar, passes much of his time in +solitude; but when associating with others, "it is rare to find," as Dr. +Campbell states, "more than one male with a whole herd of females." The +larger males expel or kill the smaller and weaker ones. The male differs +from the female by his immense tusks and greater size, strength, and +endurance; so great is the difference in these latter respects, that +the males when caught are valued at twenty per cent. above the +females.[342] With other pachydermatous animals the sexes differ very +little or not at all, and they are not, as far as known, polygamists. +Hardly a single species amongst the Cheiroptera and Edentata, or in the +great Orders of the Rodents and Insectivora, presents well-developed +secondary sexual differences; and I can find no account of any species +being polygamous, excepting, perhaps, the common rat, the males of +which, as some rat-catchers affirm, live with several females. + +The lion in South Africa, as I hear from Sir Andrew Smith, sometimes +lives with a single female, but generally with more than one, and, in +one case, was found with as many as five females, so that he is +polygamous. He is, as far as I can discover, the sole polygamist in the +whole group of the terrestrial Carnivora, and he alone presents +well-marked sexual characters. If, however, we turn to the marine +Carnivora, the case is widely different; for many species of seals +offer, as we shall hereafter see, extraordinary sexual differences, and +they are eminently polygamous. Thus the male sea-elephant of the +Southern Ocean, always possesses, according to Peron, several females, +and the sea-lion of Forster is said to be surrounded by from twenty to +thirty females. In the North, the male sea-bear of Steller is +accompanied by even a greater number of females. + +With respect to birds, many species, the sexes of which differ greatly +from each other, are certainly monogamous. In Great Britain we see +well-marked sexual differences in, for instance, the wild-duck which +pairs with a single female, with the common blackbird, and with the +bullfinch which is said to pair for life. So it is, as I am informed by +Mr. Wallace, with the Chatterers or Cotingidae of South America, and +numerous other birds. In several groups I have not been able to discover +whether the species are polygamous or monogamous. Lesson says that birds +of paradise, so remarkable for their sexual differences, are polygamous, +but Mr. Wallace doubts whether he had sufficient evidence. Mr. Salvin +informs me that he has been led to believe that humming-birds are +polygamous. The male widow-bird; remarkable for his caudal plumes, +certainly seems to be a polygamist.[343] I have been assured by Mr. +Jenner Weir and by others, that three starlings not rarely frequent the +same nest; but whether this is a case of polygamy or polyandry has not +been ascertained. + +The Gallinaceae present almost as strongly marked sexual differences as +birds of paradise or humming-birds, and many of the species are, as is +well known, polygamous; others being strictly monogamous. What a +contrast is presented between the sexes by the polygamous peacock or +pheasant, and the monogamous guinea-fowl or partridge! Many similar +cases could be given, as in the grouse tribe, in which the males of the +polygamous capercailzie and black-cock differ greatly from the females; +whilst the sexes of the monogamous red grouse and ptarmigan differ very +little. Amongst the Cursores, no great number of species offer +strongly-marked sexual differences, except the bustards, and the great +bustard (_Otis tarda_), is said to be polygamous. With the Grallatores, +extremely few species differ sexually, but the ruff (_Machetes pugnax_) +affords a strong exception, and this species is believed by Montagu to +be a polygamist. Hence it appears that with birds there often exists a +close relation between polygamy and the development of strongly-marked +sexual differences. On asking Mr. Bartlett, at the Zoological Gardens, +who has had such large experience with birds, whether the male tragopan +(one of the Gallinaceae) was polygamous, I was struck by his answering, +"I do not know, but should think so from his splendid colours." + +It deserves notice that the instinct of pairing with a single female is +easily lost under domestication. The wild-duck is strictly monogamous, +the domestic-duck highly polygamous. The Rev. W. D. Fox informs me that +with some half-tamed wild-ducks, kept on a large pond in his +neighbourhood, so many mallards were shot by the gamekeeper that only +one was left for every seven or eight females; yet unusually large +broods were reared. The guinea-fowl is strictly monogamous; but Mr. Fox +finds that his birds succeed best when he keeps one cock to two or three +hens.[344] Canary-birds pair in a state of nature, but the breeders in +England successfully put one male to four or five females; nevertheless +the first female, as Mr. Fox has been assured, is alone treated as the +wife, she and her young ones being fed by him; the others are treated as +concubines. I have noticed these cases, as it renders it in some degree +probable that monogamous species, in a state of nature, might readily +become either temporarily or permanently polygamous. + +With respect to reptiles and fishes, too little is known of their habits +to enable us to speak of their marriage arrangements. The stickle-back +Gasterosteus, however, is said to be a polygamist;[345] and the male +during the breeding-season differs conspicuously from the female. + +To sum up on the means through which, as far as we can judge, sexual +selection has led to the development of secondary sexual characters. It +has been shewn that the largest number of vigorous offspring will be +reared from the pairing of the strongest and best-armed males, which +have conquered other males, with the most vigorous and best-nourished +females, which are the first to breed in the spring. Such females, if +they select the more attractive, and at the same time vigorous, males, +will rear a larger number of offspring than the retarded females, which +must pair with the less vigorous and less attractive males. So it will +be if the more vigorous males select the more attractive and at the same +time healthy and vigorous females; and this will especially hold good if +the male defends the female, and aids in providing food for the young. +The advantage thus gained by the more vigorous pairs in rearing a larger +number of offspring has apparently sufficed to render sexual selection +efficient. But a large preponderance in number of the males over the +females would be still more efficient; whether the preponderance was +only occasional and local, or permanent; whether it occurred at birth, +or subsequently from the greater destruction of the females; or whether +it indirectly followed from the practice of polygamy. + + +_The Male generally more modified than the Female._--Throughout the +animal kingdom, when the sexes differ from each other in external +appearance, it is the male which, with rare exceptions, has been chiefly +modified; for the female still remains more like the young of her own +species, and more like the other members of the same group. The cause of +this seems to lie in the males of almost all animals having stronger +passions than the females. Hence it is the males that fight together and +sedulously display their charms before the females; and those which are +victorious transmit their superiority to their male offspring. Why the +males do not transmit their characters to both sexes will hereafter be +considered. That the males of all mammals eagerly pursue the females is +notorious to every one. So it is with birds; but many male birds do not +so much pursue the female, as display their plumage, perform strange +antics, and pour forth their song, in her presence. With the few fish +which have been observed, the male seems much more eager than the +female; and so it is with alligators, and apparently with Batrachians. +Throughout the enormous class of insects, as Kirby remarks,[346] "the +law is, that the male shall seek the female." With spiders and +crustaceans, as I hear from two great authorities, Mr. Blackwall and Mr. +C. Spence Bate, the males are more active and more erratic in their +habits than the females. With insects and crustaceans, when the organs +of sense or locomotion are present in the one sex and absent in the +other, or when, as is frequently the case, they are more highly +developed in the one than the other, it is almost invariably the male, +as far as I can discover, which retains such organs, or has them most +developed; and this shews that the male is the more active member in the +courtship of the sexes.[347] + +The female, on the other hand, with the rarest exception, is less eager +than the male. As the illustrious Hunter[348] long ago observed, she +generally "requires to be courted;" she is coy, and may often be seen +endeavouring for a long time to escape from the male. Every one who has +attended to the habits of animals will be able to call to mind instances +of this kind. Judging from various facts, hereafter to be given, and +from the results which may fairly be attributed to sexual selection, the +female, though comparatively passive, generally exerts some choice and +accepts one male in preference to others. Or she may accept, as +appearances would sometimes lead us to believe, not the male which is +the most attractive to her, but the one which is the least distasteful. +The exertion of some choice on the part of the female seems almost as +general a law as the eagerness of the male. + +We are naturally led to enquire why the male in so many and such widely +distinct classes has been rendered more eager than the female, so that +he searches for her and plays the more active part in courtship. It +would be no advantage and some loss of power if both sexes were mutually +to search for each other; but why should the male almost always be the +seeker? With plants, the ovules after fertilisation have to be nourished +for a time; hence the pollen is necessarily brought to the female +organs--being placed on the stigma, through the agency of insects or of +the wind, or by the spontaneous movements of the stamens; and with the +Algae, &c., by the locomotive power of the antherozooids. With +lowly-organised animals permanently affixed to the same spot and having +their sexes separate, the male element is invariably brought to the +female; and we can see the reason; for the ova, even if detached before +being fertilised and not requiring subsequent nourishment or protection, +would be, from their larger relative size, less easily transported than +the male element. Hence plants[349] and many of the lower animals are, +in this respect, analogous. In the case of animals not affixed to the +same spot, but enclosed within a shell with no power of protruding any +part of their bodies, and in the case of animals having little power of +locomotion, the males must trust the fertilising element to the risk of +at least a short transit through the waters of the sea. It would, +therefore, be a great advantage to such animals, as their organisation +became perfected, if the males when ready to emit the fertilising +element, were to acquire the habit of approaching the female as closely +as possible. The males of various lowly-organised animals having thus +aboriginally acquired the habit of approaching and seeking the females, +the same habit would naturally be transmitted to their more highly +developed male descendants; and in order that they should become +efficient seekers, they would have to be endowed with strong passions. +The acquirement of such passions would naturally follow from the more +eager males leaving a larger number of offspring than the less eager. + +The great eagerness of the male has thus indirectly led to the much +more frequent development of secondary sexual characters in the male +than in the female. But the development of such characters will have +been much aided, if the conclusion at which I arrived after studying +domesticated animals, can be trusted, namely, that the male is more +liable to vary than the female. I am aware how difficult it is to verify +a conclusion of this kind. Some slight evidence, however, can be gained +by comparing the two sexes in mankind, as man has been more carefully +observed than any other animal. During the Novara Expedition[350] a vast +number of measurements of various parts of the body in different races +were made, and the men were found in almost every case to present a +greater range of variation than the women; but I shall have to recur to +this subject in a future chapter. Mr. J. Wood,[351] who has carefully +attended to the variation of the muscles in man, puts in italics the +conclusion that "the greatest number of abnormalities in each subject is +found in the males." He had previously remarked that "altogether in 102 +subjects the varieties of redundancy were found to be half as many again +as in females, contrasting widely with the greater frequency of +deficiency in females before described." Professor Macalister like wise +remarks[352] that variations in the muscles "are probably more common in +males than females." Certain muscles which are not normally present in +mankind are also more frequently developed in the male than in the +female sex, although exceptions to this rule are said to occur. Dr. +Burt Wilder[353] has tabulated the cases of 152 individuals with +supernumerary digits, of which 86 were males, and 39, or less than half, +females; the remaining 27 being of unknown sex. It should not, however, +be overlooked that women would more frequently endeavour to conceal a +deformity of this kind than men. Whether the large proportional number +of deaths of the male offspring of man and apparently of sheep, compared +with the female offspring, before, during, and shortly after birth (see +supplement), has any relation to a stronger tendency in the organs of +the male to vary and thus to become abnormal in structure or function, I +will not pretend to conjecture. + +In various classes of animals a few exceptional cases occur, in which +the female instead of the male has acquired well pronounced secondary +sexual characters, such as brighter colours, greater size, strength, or +pugnacity. With birds, as we shall hereafter see, there has sometimes +been a complete transposition of the ordinary characters proper to each +sex; the females having become the more eager in courtship, the males +remaining comparatively passive, but apparently selecting, as we may +infer from the results, the more attractive females. Certain female +birds have thus been rendered more highly coloured or otherwise +ornamented, as well as more powerful and pugnacious than the males, +these characters being transmitted to the female offspring alone. + +It may be suggested that in some cases a double process of selection has +been carried on; the males having selected the more attractive females, +and the latter the more attractive males. This process however, though +it might lead to the modification of both sexes, would not make the one +sex different from the other, unless indeed their taste for the +beautiful differed; but this is a supposition too improbable in the case +of any animal, excepting man, to be worth considering. There are, +however, many animals, in which the sexes resemble each other, both +being furnished with the same ornaments, which analogy would lead us to +attribute to the agency of sexual selection. In such cases it may be +suggested with more plausibility, that there has been a double or mutual +process of sexual selection; the more vigorous and precocious females +having selected the more attractive and vigorous males, the latter +having rejected all except the more attractive females. But from what we +know of the habits of animals, this view is hardly probable, the male +being generally eager to pair with any female. It is more probable that +the ornaments common to both sexes were acquired by one sex, generally +the male, and then transmitted to the offspring of both sexes. If, +indeed, during a lengthened period the males of any species were greatly +to exceed the females in number, and then during another lengthened +period under different conditions the reverse were to occur, a double, +but not simultaneous, process of sexual selection might easily be +carried on, by which the two sexes might be rendered widely different. + +We shall hereafter see that many animals exist, of which neither sex is +brilliantly coloured or provided with special ornaments, and yet the +members of both sexes or of one alone have probably been modified +through sexual selection. The absence of bright tints or other ornaments +may be the result of variations of the right kind never having occurred, +or of the animals themselves preferring simple colours, such as plain +black or white. Obscure colours have often been acquired through natural +selection for the sake of protection, and the acquirement through +sexual selection of conspicuous colours, may have been checked from the +danger thus incurred. But in other cases the males have probably +struggled together during long ages, through brute force, or by the +display of their charms, or by both means combined, and yet no effect +will have been produced unless a larger number of offspring were left by +the more successful males to inherit their superiority, than by the less +successful males; and this, as previously shewn, depends on various +complex contingencies. + +Sexual selection acts in a less rigorous manner than natural selection. +The latter produces its effects by the life or death at all ages of the +more or less successful individuals. Death, indeed, not rarely ensues +from the conflicts of rival males. But generally the less successful +male merely fails to obtain a female, or obtains later in the season a +retarded and less vigorous female, or, if polygamous, obtains fewer +females; so that they leave fewer, or less vigorous, or no offspring. In +regard to structures acquired through ordinary or natural selection, +there is in most cases, as long as the conditions of life remain the +same, a limit to the amount of advantageous modification in relation to +certain special ends; but in regard to structures adapted to make one +male victorious over another, either in fighting or in charming the +female, there is no definite limit to the amount of advantageous +modification; so that as long as the proper variations arise the work of +sexual selection will go on. This circumstance may partly account for +the frequent and extraordinary amount of variability presented by +secondary sexual characters. Nevertheless, natural selection will +determine that characters of this kind shall not be acquired by the +victorious males, which would be injurious to them in any high degree, +either by expending too much of their vital powers, or by exposing them +to any great danger. The development, however, of certain structures--of +the horns, for instance, in certain stags--has been carried to a +wonderful extreme; and in some instances to an extreme which, as far as +the general conditions of life are concerned, must be slightly injurious +to the male. From this fact we learn that the advantages which favoured +males have derived from conquering other males in battle or courtship, +and thus leaving a numerous progeny, have been in the long run greater +than those derived from rather more perfect adaptation to the external +conditions of life. We shall further see, and this could never have been +anticipated, that the power to charm the female has been in some few +instances more important than the power to conquer other males in +battle. + + +LAWS OF INHERITANCE. + +In order to understand how sexual selection has acted, and in the course +of ages has produced conspicuous results with many animals of many +classes, it is necessary to bear in mind the laws of inheritance, as far +as they are known. Two distinct elements are included under the term +"inheritance," namely the transmission and the development of +characters; but as these generally go together, the distinction is often +overlooked. We see this distinction in those characters which are +transmitted through the early years of life, but are developed only at +maturity or during old age. We see the same distinction more clearly +with secondary sexual characters, for these are transmitted through both +sexes, though developed in one alone. That they are present in both +sexes, is manifest when two species, having strongly-marked sexual +characters, are crossed, for each transmits the characters proper to +its own male and female sex to the hybrid offspring of both sexes. The +same fact is likewise manifest, when characters proper to the male are +occasionally developed in the female when she grows old or becomes +diseased; and so conversely with the male. Again, characters +occasionally appear, as if transferred from the male to the female, as +when, in certain breeds of the fowl, spurs regularly appear in the young +and healthy females; but in truth they are simply developed in the +female; for in every breed each detail in the structure of the spur is +transmitted through the female to her male offspring. In all cases of +reversion, characters are transmitted through two, three, or many +generations, and are then under certain unknown favourable conditions +developed. This important distinction between transmission and +development will be easiest kept in mind by the aid of the hypothesis of +pangenesis, whether or not it be accepted as true. According to this +hypothesis, every unit or cell of the body throws off gemmules or +undeveloped atoms, which are transmitted to the offspring of both sexes, +and are multiplied by self-division. They may remain undeveloped during +the early years of life or during successive generations; their +development into units or cells, like those from which they were +derived, depending on their affinity for, and union with, other units or +cells previously developed in the due order of growth. + + +_Inheritance at Corresponding Periods of Life._--This tendency is well +established. If a new character appears in an animal whilst young, +whether it endures throughout life or lasts only for a time, it will +reappear, as a general rule, at the same age and in the same manner in +the offspring. If, on the other hand, a new character appears at +maturity, or even during old age, it tends to reappear in the offspring +at the same advanced age. When deviations from this rule occur, the +transmitted characters much oftener appear before than after the +corresponding age. As I have discussed this subject at sufficient length +in another work,[354] I will here merely give two or three instances, +for the sake of recalling the subject to the reader's mind. In several +breeds of the Fowl, the chickens whilst covered with down, in their +first true plumage, and in their adult plumage, differ greatly from each +other, as well as from their common parent-form, the _Gallus bankiva_; +and these characters are faithfully transmitted by each breed to their +offspring at the corresponding period of life. For instance, the +chickens of spangled Hamburghs, whilst covered with down, have a few +dark spots on the head and rump, but are not longitudinally striped, as +in many other breeds; in their first true plumage, "they are beautifully +pencilled," that is each feather is transversely marked by numerous dark +bars; but in their second plumage the feathers all become spangled or +tipped with a dark round spot.[355] Hence in this breed variations have +occurred and have been transmitted at three distinct periods of life. +The Pigeon offers a more remarkable case, because the aboriginal +parent-species does not undergo with advancing age any change of +plumage, excepting that at maturity the breast becomes more iridescent; +yet there are breeds which do not acquire their characteristic colours +until they have moulted two, three, or four times; and these +modifications of plumage are regularly transmitted. + + +_Inheritance at Corresponding Seasons of the Year._--With animals in a +state of nature innumerable instances occur of characters periodically +appearing at different seasons. We see this with the horns of the stag, +and with the fur of arctic animals which becomes thick and white during +the winter. Numerous birds acquire bright colours and other decorations +during the breeding-season alone. I can throw but little light on this +form of inheritance from facts observed under domestication. Pallas +states,[356] that in Siberia domestic cattle and horses periodically +become lighter-coloured during the winter; and I have observed a similar +marked change of colour in certain ponies in England. Although I do not +know that this tendency to assume a differently coloured coat during +different seasons of the year is transmitted, yet it probably is so, as +all shades of colour are strongly inherited by the horse. Nor is this +form of inheritance, as limited by season, more remarkable than +inheritance as limited by age or sex. + + +_Inheritance as Limited by Sex._--The equal transmission of characters +to both, sexes is the commonest form of inheritance, at least with those +animals which do not present strongly-marked sexual differences, and +indeed with many of these. But characters are not rarely transferred +exclusively to that sex, in which they first appeared. Ample evidence on +this head has been advanced in my work on Variation under +Domestication; but a few instances may here be given. There are breeds +of the sheep and goat, in which the horns of the male differ greatly in +shape from those of the female; and these differences, acquired under +domestication, are regularly transmitted to the same sex. With +tortoise-shell cats the females alone, as a general rule, are thus +coloured, the males being rusty-red. With most breeds of the fowl, the +characters proper to each sex are transmitted to the same sex alone. So +general is this form of transmission that it is an anomaly when we see +in certain breeds variations transmitted equally to both sexes. There +are also certain sub-breeds of the fowl in which the males can hardly be +distinguished from each other, whilst the females differ considerably in +colour. With the pigeon the sexes of the parent-species do not differ in +any external character; nevertheless in certain domesticated breeds the +male is differently coloured from the female.[357] The wattle in the +English Carrier pigeon and the crop in the Pouter are more highly +developed in the male than in the female; and although these characters +have been gained through long-continued selection by man, the difference +between the two sexes is wholly due to the form of inheritance which has +prevailed; for it has arisen, not from, but rather in opposition to, the +wishes of the breeder. + +Most of our domestic races have been formed by the accumulation of many +slight variations; and as some of the successive steps have been +transmitted to one sex alone, and some to both sexes, we find in the +different breeds of the same species all gradations between great sexual +dissimilarity and complete similarity. Instances have already been +given with the breeds of the fowl and pigeon; and under nature analogous +cases are of frequent occurrence. With animals under domestication, but +whether under nature I will not venture to say, one sex may lose +characters proper to it, and may thus come to resemble to a certain +extent the opposite sex; for instance, the males of some breeds of the +fowl have lost their masculine plumes and hackles. On the other hand the +differences between the sexes may be increased under domestication, as +with merino sheep, in which the ewes have lost their horns. Again, +characters proper to one sex may suddenly appear in the other sex; as +with those sub-breeds of the fowl in which the hens whilst young acquire +spurs; or, as in certain Polish sub-breeds, in which the females, as +there is reason to believe, originally acquired a crest, and +subsequently transferred it to the males. All these cases are +intelligible on the hypothesis of pangenesis; for they depend on the +gemmules of certain units of the body, although present in both sexes, +becoming through the influence of domestication dormant in the one sex; +or if naturally dormant, becoming developed. + +There is one difficult question which it will be convenient to defer to +a future chapter; namely, whether a character at first developed in both +sexes, can be rendered through selection limited in its development to +one sex alone. If, for instance, a breeder observed that some of his +pigeons (in which species characters are usually transferred in an equal +degree to both sexes) varied into pale blue; could he by long-continued +selection make a breed, in which the males alone should be of this tint, +whilst the females remained unchanged? I will here only say, that this, +though perhaps not impossible, would be extremely difficult; for the +natural result of breeding from the pale-blue males would be to change +his whole stock, including both sexes, into this tint. If, however, +variations of the desired tint appeared, which were from the first +limited in their development to the male sex, there would not be the +least difficulty in making a breed characterised by the two sexes being +of a different colour, as indeed has been effected with a Belgian breed, +in which the males alone are streaked with black. In a similar manner, +if any variation appeared in a female pigeon, which was from the first +sexually limited in its development, it would be easy to make a breed +with the females alone thus characterised; but if the variation was not +thus originally limited, the process would be extremely difficult, +perhaps impossible. + + +_On the Relation between the period of Development of a Character and +its transmission to one sex or to both sexes._--Why certain characters +should be inherited by both sexes, and other characters by one sex +alone, namely by that sex in which the character first appeared, is in +most cases quite unknown. We cannot even conjecture why with certain +sub-breeds of the pigeon, black striae, though transmitted through the +female, should be developed in the male alone, whilst every other +character is equally transferred to both sexes. Why, again, with cats, +the tortoise-shell colour should, with rare exceptions, be developed in +the female alone. The very same characters, such as deficient or +supernumerary digits, colour-blindness, &c., may with mankind be +inherited by the males alone of one family, and in another family by the +females alone, though in both cases transmitted through the opposite as +well as the same sex.[358] Although we are thus ignorant, two rules +often hold good, namely that variations which, first appear in either +sex at a late period of life, tend to be developed in the same sex +alone; whilst variations which first appear early in life in either sex +tend to be developed in both sexes. I am, however, far from supposing +that this is the sole determining cause. As I have not elsewhere +discussed this subject, and as it has an important bearing on sexual +selection, I must here enter into lengthy and somewhat intricate +details. + +It is in itself probable that any character appearing at an early age +would tend to be inherited equally by both sexes, for the sexes do not +differ much in constitution, before the power of reproduction is gained. +On the other hand, after this power has been gained and the sexes have +come to differ in constitution, the gemmules (if I may again use the +language of pangenesis) which are cast off from each varying part in the +one sex would be much more likely to possess the proper affinities for +uniting with the tissues of the same sex, and thus becoming developed, +than with those of the opposite sex. + +I was first led to infer that a relation of this kind exists, from the +fact that whenever and in whatever manner the adult male has come to +differ from the adult female, he differs in the same manner from the +young of both sexes. The generality of this fact is quite remarkable: it +holds good with almost all mammals, birds, amphibians, and fishes; also +with many crustaceans, spiders and some few insects, namely certain +orthoptera and libellulae. In all these cases the variations, through the +accumulation of which the male acquired his proper masculine characters, +must have occurred at a somewhat late period of life; otherwise the +young males would have been similarly characterised; and conformably +with our rule, they are transmitted to and developed in the adult males +alone. When, on the other hand, the adult male closely resembles the +young of both sexes (these, with rare exceptions, being alike), he +generally resembles the adult female; and in most of these cases the +variations through which the young and old acquired their present +characters, probably occurred in conformity with our rule during youth. +But there is here room for doubt, as characters are sometimes +transferred to the offspring at an earlier age than that at which they +first appeared in the parents, so that the parents may have varied when +adult, and have transferred their characters to their offspring whilst +young. There are, moreover, many animals, in which the two sexes closely +resemble each other, and yet both differ from their young; and here the +characters of the adults must have been acquired late in life; +nevertheless, these characters in apparent contradiction to our rule, +are transferred to both sexes. We must not, however, overlook the +possibility or even probability of successive variations of the same +nature sometimes occurring, under exposure to similar conditions, +simultaneously in both sexes at a rather late period of life; and in +this case the variations would be transferred to the offspring of both +sexes at a corresponding late age; and there would be no real +contradiction to our rule of the variations which occur late in life +being transferred exclusively to the sex in which they first appeared. +This latter rule seems to hold true more generally than the second rule, +namely, that variations which occur in either sex early in life tend to +be transferred to both sexes. As it was obviously impossible even to +estimate in how large a number of cases throughout the animal kingdom +these two propositions hold good, it occurred to me to investigate some +striking or crucial instances, and to rely on the result. + +An excellent case for investigation is afforded by the Deer Family. In +all the species, excepting one, the horns are developed in the male +alone, though certainly transmitted through the female, and capable of +occasional abnormal development in her. In the reindeer, on the other +hand, the female is provided with horns; so that in this species, the +horns ought, according to our rule, to appear early in life, long before +the two sexes had arrived at maturity and had come to differ much in +constitution. In all the other species of deer the horns ought to appear +later in life, leading to their development in that sex alone, in which +they first appeared in the progenitor of the whole Family. Now in seven +species, belonging to distinct sections of the family and inhabiting +different regions, in which the stags alone bear horns, I find that the +horns first appear at periods varying from nine months after birth in +the roebuck to ten or twelve or even more months in the stags of the six +other larger species.[359] But with the reindeer the case is widely +different, for as I hear from Prof. Nilsson, who kindly made special +enquiries for me in Lapland, the horns appear in the young animals +within four or five weeks after birth, and at the same time in both +sexes. So that here we have a structure, developed at a most unusually +early age in one species of the family, and common to both sexes in this +one species. + +In several kinds of antelopes the males alone are provided with horns, +whilst in the greater number both sexes have horns. With respect to the +period of development, Mr. Blyth informs me that there lived at one time +in the Zoological Gardens a young koodoo (_Ant. strepsiceros_), in which +species the males alone are horned, and the young of a closely-allied +species, viz. the eland (_Ant. oreas_), in which both sexes are horned. +Now in strict conformity with our rule, in the young male koodoo, +although arrived at the age of ten months, the horns were remarkably +small considering the size ultimately attained by them: whilst in the +young male eland, although only three months old, the horns were already +very much larger than in the koodoo. It is also worth notice that in the +prong-horned antelope,[360] in which species the horns, though present +in both sexes, are almost rudimentary in the female, they do not appear +until about five or six months after birth. With sheep, goats, and +cattle, in which the horns are well developed in both sexes, though not +quite equal in size, they can be felt, or even seen, at birth or soon +afterwards.[361] Our rule, however, fails in regard to some breeds of +sheep, for instance merinos, in which the rams alone are horned; for I +cannot find on enquiry,[362] that the horns are developed later in life +in this breed than in ordinary sheep in which both sexes are horned. But +with domesticated sheep the presence or absence of horns is not a firmly +fixed character; a certain proportion of the merino ewes bearing small +horns, and some of the rams being hornless; whilst with ordinary sheep +hornless ewes are occasionally produced. + +In most of the species of the splendid family of the Pheasants, the +males differ conspicuously from the females, and they acquire their +ornaments at a rather late period of life. The eared pheasant +(_Crossoptilon auritum_), however, offers a remarkable exception, for +both sexes possess the fine caudal plumes, the large ear-tufts and the +crimson velvet about the head; and I find on enquiry in the Zoological +Gardens that all these characters, in accordance with our rule, appear +very early in life. The adult male can, however, be distinguished from +the adult female by one character, namely by the presence of spurs; and +conformably with our rule, these do not begin to be developed, as I am +assured by Mr. Bartlett, before the age of six months, and even at this +age, can hardly be distinguished in the two sexes.[363] The male and +female Peacock differ conspicuously from each other in almost every +part of their plumage, except in the elegant head-crest, which is common +to both sexes; and this is developed very early in life, long before the +other ornaments which are confined to the male. The wild-duck offers an +analogous case, for the beautiful green speculum on the wings is common +to both sexes, though duller and somewhat smaller in the female, and it +is developed early in life, whilst the curled tail-feathers and other +ornaments peculiar to the male are developed later.[364] Between such +extreme cases of close sexual resemblance and wide dissimilarity, as +those of the Crossoptilon and peacock, many intermediate ones could be +given, in which the characters follow in their order of development our +two rules. + +As most insects emerge from their pupal state in a mature condition, it +is doubtful whether the period of development determines the +transference of their characters to one or both sexes. But we do not +know that the coloured scales, for instance, in two species of +butterflies, in one of which the sexes differ in colour, whilst in the +other they are alike, are developed at the same relative age in the +cocoon. Nor do we know whether all the scales are simultaneously +developed on the wings of the same species of butterfly, in which +certain coloured marks are confined to one sex, whilst other marks are +common to both sexes. A difference of this kind in the period of +development is not so improbable as it may at first appear; for with the +Orthoptera, which assume their adult state, not by a single +metamorphosis, but by a succession of moults, the young males of some +species at first resemble the females, and acquire their distinctive +masculine characters only during a later moult. Strictly analogous cases +occur during the successive moults of certain male crustaceans. + +We have as yet only considered the transference of characters, +relatively to their period of development, with species in a natural +state; we will now turn to domesticated animals; first touching on +monstrosities and diseases. The presence of supernumerary digits, and +the absence of certain phalanges, must be determined at an early +embryonic period--the tendency to profuse bleeding is at least +congenital, as is probably colour-blindness--yet these peculiarities, +and other similar ones, are often limited in their transmission to one +sex; so that the rule that characters which are developed at an early +period tend to be transmitted to both sexes, here wholly fails. But this +rule, as before remarked, does not appear to be nearly so generally true +as the converse proposition, namely, that characters which appear late +in life in one sex are transmitted exclusively to the same sex. From the +fact of the above abnormal peculiarities becoming attached to one sex, +long before the sexual functions are active, we may infer that there +must be a difference of some kind between the sexes at an extremely +early age. With respect to sexually-limited diseases, we know too little +of the period at which they originate, to draw any fair conclusion. +Gout, however, seems to fall under our rule; for it is generally caused +by intemperance after early youth, and is transmitted from the father to +his sons in a much more marked manner than to his daughters. + +In the various domestic breeds of sheep, goats, and cattle, the males +differ from their respective females in the shape or development of +their horns, forehead, mane, dewlap, tail, and hump on the shoulders; +and these peculiarities, in accordance with our rule, are not fully +developed until rather late in life. With dogs, the sexes do not differ, +except that in certain breeds, especially in the Scotch deer-hound, the +male is much larger and heavier than the female; and as we shall see in +a future chapter, the male goes on increasing in size to an unusually +late period of life, which will account, according to our rule, for his +increased size being transmitted to his male offspring alone. On the +other hand, the tortoise-shell colour of the hair, which is confined to +female cats, is quite distinct at birth, and this case violates our +rule. There is a breed of pigeons in which the males alone are streaked +with black, and the streaks can be detected even in the nestlings; but +they become more conspicuous at each successive moult, so that this case +partly opposes and partly supports the rule. With the English Carrier +and Pouter pigeon the full development of the wattle and the crop occurs +rather late in life, and these characters, conformably with our rule, +are transmitted in full perfection to the males alone. The following +cases perhaps come within the class previously alluded to, in which the +two sexes have varied in the same manner at a rather late period of +life, and have consequently transferred their new characters to both +sexes at a corresponding late period; and if so, such cases are not +opposed to our rule. Thus there are sub-breeds of the pigeon, described +by Neumeister,[365] both sexes of which change colour after moulting +twice or thrice, as does likewise the Almond Tumbler; nevertheless these +changes, though occurring rather late in life, are common to both sexes. +One variety of the Canary-bird, namely the London Prize, offers a nearly +analogous case. + +With the breeds of the Fowl the inheritance of various characters by one +sex or by both sexes, seems generally determined by the period at which +such characters are developed. Thus in all the many breeds in which the +adult male differs greatly in colour from the female and from the adult +male parent-species, he differs from the young male, so that the newly +acquired characters must have appeared at a rather late period of life. +On the other hand with most of the breeds in which the two sexes +resemble each other, the young are coloured in nearly the same manner as +their parents, and this renders it probable that their colours first +appeared early in life. We have instances of this fact in all black and +white breeds, in which the young and old of both sexes are alike; nor +can it be maintained that there is something peculiar in a black or +white plumage, leading to its transference to both sexes; for the males +alone of many natural species are either black or white, the females +being very differently coloured. With the so-called Cuckoo sub-breeds of +the fowl, in which the feathers are transversely pencilled with dark +stripes, both sexes and the chickens are coloured in nearly the same +manner. The laced plumage of the Sebright bantam is the same in both +sexes, and in the chickens the feathers are tipped with black, which +makes a near approach to lacing. Spangled Hamburghs, however, offer a +partial exception, for the two sexes, though not quite alike, resemble +each other more closely than do the sexes of the aboriginal +parent-species, yet they acquire their characteristic plumage late in +life, for the chickens are distinctly pencilled. Turning to other +characters besides colour: the males alone of the wild parent-species +and of most domestic breeds possess a fairly well developed comb, but in +the young of the Spanish fowl it is largely developed at a very early +age, and apparently in consequence of this it is of unusual size in the +adult females. In the Game breeds pugnacity is developed at a +wonderfully early age, of which curious proofs could be given; and this +character is transmitted to both sexes, so that the hens, from their +extreme pugnacity, are now generally exhibited in separate pens. With +the Polish breeds the bony protuberance of the skull which supports the +crest is partially developed even before the chickens are hatched, and +the crest itself soon begins to grow, though at first feebly;[366] and +in this breed a great bony protuberance and an immense crest +characterise the adults of both sexes. + +Finally, from what we have now seen of the relation which exists in many +natural species and domesticated races, between the period of the +development of their characters and the manner of their +transmission--for example the striking fact of the early growth of the +horns in the reindeer, in which both sexes have horns, in comparison +with their much later growth in the other species in which the male +alone bears horns--we may conclude that one cause, though not the sole +cause, of characters being exclusively inherited by one sex, is their +development at a late age. And secondly, that one, though apparently a +less efficient, cause of characters being inherited by both sexes is +their development at an early age, whilst the sexes differ but little in +constitution. It appears, however, that some difference must exist +between the sexes even during an early embryonic period, for characters +developed at this age not rarely become attached to one sex. + + +_Summary and concluding remarks._--From the foregoing discussion on the +various laws of inheritance, we learn that characters often or even +generally tend to become developed in the same sex, at the same age, and +periodically at the same season of the year, in which they first +appeared in the parents. But these laws, from unknown causes, are very +liable to change. Hence the successive steps in the modification of a +species might readily be transmitted in different ways; some of the +steps being transmitted to one sex, and some to both; some to the +offspring at one age, and some at all ages. Not only are the laws of +inheritance extremely complex, but so are the causes which induce and +govern variability. The variations thus caused are preserved and +accumulated by sexual selection, which is in itself an extremely complex +affair, depending, as it does, on ardour in love, courage, and the +rivalry of the males, and on the powers of perception, taste, and will +of the female. Sexual selection will also be dominated by natural +selection for the general welfare of the species. Hence the manner in +which the individuals of either sex or of both sexes are affected +through sexual selection cannot fail to be complex in the highest +degree. + +When variations occur late in life in one sex, and are transmitted to +the same sex at the same age, the other sex and the young are +necessarily left unmodified. When they occur late in life, but are +transmitted to both sexes at the same age, the young alone are left +unmodified. Variations, however, may occur at any period of life in one +sex or in both, and be transmitted to both sexes at all ages, and then +all the individuals of the species will be similarly modified. In the +following chapters it will be seen that all these cases frequently occur +under nature. + +Sexual selection can never act on any animal whilst young, before the +age for reproduction has arrived. From the great eagerness of the male +it has generally acted on this sex and not on the females. The males +have thus become provided with weapons for fighting with their rivals, +or with organs for discovering and securely holding the female, or for +exciting and charming her. When the sexes differ in these respects, it +is also, as we have seen, an extremely general law that the adult male +differs more or less from the young male; and we may conclude from this +fact that the successive variations, by which the adult male became +modified, cannot have occurred much before the age for reproduction. How +then are we to account for this general and remarkable coincidence +between the period of variability and that of sexual selection,-- +principles which are quite independent of each other? I +think we can see the cause: it is not that the males have never varied +at an early age, but that such variations have commonly been lost, +whilst those occurring at a later age have been preserved. + +All animals produce more offspring than can survive to maturity; and we +have every reason to believe that death falls heavily on the weak and +inexperienced young. If then a certain proportion of the offspring were +to vary at birth or soon afterwards, in some manner which at this age +was of no service to them, the chance of the preservation of such +variations would be small. We have good evidence under domestication how +soon variations of all kinds are lost, if not selected. But variations +which occurred at or near maturity, and which were of immediate service +to either sex, would probably be preserved; as would similar variations +occurring at an earlier period in any individuals which happened to +survive. As this principle has an important bearing on sexual selection, +it may be advisable to give an imaginary illustration. We will take a +pair of animals, neither very fertile nor the reverse, and assume that +after arriving at maturity they live on an average for five years, +producing each year five young. They would thus produce 25 offspring; +and it would not, I think, be an unfair estimate to assume that 18 or 20 +out of the 25 would perish before maturity, whilst still young and +inexperienced; the remaining seven or five sufficing to keep up the +stock of mature individuals. If so, we can see that variations which +occurred during youth, for instance in brightness, and which were not of +the least service to the young, would run a good chance of being utterly +lost. Whilst similar variations, which occurring at or near maturity in +the comparatively few individuals surviving to this age, and which +immediately gave an advantage to certain males, by rendering them more +attractive to the females, would be likely to be preserved. No doubt +some of the variations in brightness which occurred at an earlier age +would by chance be preserved, and eventually give to the male the same +advantage as those which appeared later; and this will account for the +young males commonly partaking to a certain extent (as may be observed +with many birds) of the bright colours of their adult male parents. If +only a few of the successive variations in brightness were to occur at a +late age, the adult male would be only a little brighter than the young +male; and such cases are common. + +In this illustration I have assumed that the young varied in a manner +which was of no service to them; but many characters proper to the adult +male would be actually injurious to the young,--as bright colours from +making them conspicuous, or horns of large size from expending much +vital force. Such variations in the young would promptly be eliminated +through natural selection. With the adult and experienced males, on the +other hand, the advantage thus derived in their rivalry with other males +would often more than counterbalance exposure to some degree of danger. +Thus we can understand how it is that variations which must originally +have appeared rather late in life have alone or in chief part been +preserved for the development of secondary sexual characters; and the +remarkable coincidence between the periods of variability and of sexual +selection is intelligible. + +As variations which give to the male an advantage in lighting with other +males, or in finding, securing, or charming the female, would be of no +use to the female, they will not have been preserved in this sex either +during youth or maturity. Consequently such variations would be +extremely liable to be lost; and the female, as far as these characters +are concerned, would be left unmodified, excepting in so far as she may +have received them by transference from the male. No doubt if the female +varied and transferred serviceable characters to her male offspring, +these would be favoured through sexual selection; and then both sexes +would thus far be modified in the same manner. But I shall hereafter +have to recur to these more intricate contingencies. + +In the following chapters, I shall treat of the secondary sexual +characters in animals of all classes, and shall endeavour in each case +to apply the principles explained in the present chapter. The lowest +classes will detain us for a very short time, but the higher animals, +especially birds, must be treated at considerable length. It should be +borne in mind that for reasons already assigned, I intend to give only a +few illustrative instances of the innumerable structures by the aid of +which the male finds the female, or, when found, holds her. On the other +hand, all structures and instincts by which the male conquers other +males, and by which he allures or excites the female, will be fully +discussed, as these are in many ways the most interesting. + + +_Supplement on the proportional numbers of the two sexes in animals +belonging to various classes._ + + +As no one, as far as I can discover, has paid attention to the relative +numbers of the two sexes throughout the animal kingdom, I will here give +such materials as I have been able to collect, although they are +extremely imperfect. They consist in only a few instances of actual +enumeration, and the numbers are not very large. As the proportions are +known with certainty on a large scale in the case of man alone, I will +first give them, as a standard of comparison. + + +_Man._--In England during ten years (from 1857 to 1866) 707,120 children +on an annual average have been born alive, in the proportion of 104.5 +males to 100 females. But in 1857 the male births throughout England +were as 105.2, and in 1865 as 104.0 to 100. Looking to separate +districts, in Buckinghamshire (where on an average 5000 children are +annually born) the _mean_ proportion of male to female births, during +the whole period of the above ten years, was as 102.8 to 100; whilst in +N. Wales (where the average annual births are 12,873) it was as high as +106.2 to 100. Taking a still smaller district, viz., Rutlandshire (where +the annual births average only 739), in 1864 the male births were as +114.6, and in 1862 as 97.0 to 100; but even in this small district the +average of the 7385 births during the whole ten years was as 104.5 to +100; that is in the same ratio as throughout England.[367] The +proportions are sometimes slightly disturbed by unknown causes; thus +Prof. Faye states "that in some districts of Norway there has been +during a decennial period a steady deficiency of boys, whilst in others +the opposite condition has existed." In France during forty-four years +the male to the female births have been as 106.2 to 100; but during this +period it has occurred five times in one department, and six times in +another, that the female births have exceeded the males. In Russia the +average proportion is as high as 108.9 to 100.[368] It is a singular +fact that with Jews the proportion of male births is decidedly larger +than with Christians: thus in Prussia the proportion is as 113, in +Breslau as 114, and in Livonia as 120 to 100; the Christian births in +these countries being the same as usual, for instance, in Livonia as 104 +to 100.[369] It is a still more singular fact that in different nations, +under different conditions and climates, in Naples, Prussia, Westphalia, +France and England, the excess of male over female births is less when +they are illegitimate than when legitimate.[370] + +In various parts of Europe, according to Prof. Faye and other authors, +"a still greater preponderance of males would be met with, if death +struck both sexes in equal proportion in the womb and during birth. But +the fact is, that for every 100 still-born females, we have in several +countries from 134.6 to 144.9 still-born males." Moreover during the +first four or five years of life more male children die than females; +"for example in England, during the first year, 126 boys die +for every 100 girls,--a proportion which in France is still more +unfavourable."[371] As a consequence of this excess in the death-rate of +male children, and of the exposure of men when adult to various dangers, +and of their tendency to emigrate, the females in all old-settled +countries, where statistical records have been kept,[372] are found to +preponderate considerably over the males. + +It has often been supposed that the relative ages of the parents +determine the sex of the offspring; and Prof. Leuckart[373] has advanced +what he considers sufficient evidence, with respect to man and certain +domesticated animals, to shew that this is one important factor in the +result. So again the period of impregnation has been thought to be the +efficient cause; but recent observations discountenance this belief. +Again, with mankind polygamy has been supposed to lead to the birth of a +greater proportion of female infants; but Dr. J. Campbell[374] carefully +attended to this subject in the harems of Siam, and he concludes that +the proportion of male to female births is the same as from monogamous +unions. Hardly any animal has been rendered so highly polygamous as our +English race-horses, and we shall immediately see that their male and +female offspring are almost exactly equal in number. + + + _Horses._--Mr. Tegetmeier has been so kind as to tabulate for + me from the 'Racing Calendar' the births of race-horses during + a period of twenty-one years, viz. from 1846 to 1867; 1849 + being omitted, as no returns were that year published. The + total births have been 25,560,[375] consisting of 12,763 males + and 12,797 females, or in the proportion of 99.7 males to 100 + females. As these numbers are tolerably large, and as they are + drawn from all parts of England, during several years, we may + with much confidence conclude that with the domestic horse, or + at least with the race-horse, the two sexes are produced in + almost equal numbers. The fluctuations in the proportions + during successive years are closely like those which occur with + mankind, when a small and thinly-populated area is considered: + thus in 1856 the male horses were as 107.1, and in 1867 as only + 92.6 to 100 females. In the tabulated returns the proportions + vary in cycles, for the males exceeded the females during six + successive years; and the females exceeded the males during two + periods each of four years: this, however, may be accidental; + at least I can detect nothing of the kind with man in the + decennial table in the Registrar's Report for 1866. I may add + that certain, mares, and this holds good with certain cows and + with women, tend to produce more of one sex than of the other; + Mr. Wright of Yeldersley House, informs me that one of his Arab + mares, though put seven times to different horses, produced + seven fillies. + + + _Dogs._--During a period of twelve years, from 1857 to 1868, + the births of a large number of greyhounds, throughout England, + have been sent to the 'Field' newspaper; and I am again + indebted to Mr. Tegetmeier for carefully tabulating the + results. The recorded, births have been 6878, consisting of + 3605 males and 3273 females, that is, in the proportion of + 110.1 males to 100 females. The greatest fluctuations occurred + in 1864, when the proportion was as 95.3 males, and in 1867, as + 116.3 males to 100 females. The above average proportion of + 110.1 to 100 is probably nearly correct in the case of the + greyhound, but whether it would hold with other domesticated + breeds is in some degree doubtful. Mr. Cupples has enquired + from several great breeders of dogs, and finds that all without + exception believe that females are produced in excess; he + suggests that this belief may have arisen from females being + less valued and the consequent disappointment producing a + stronger impression on the mind. + + + _Sheep._--The sexes of sheep are not ascertained by + agriculturists until several months after birth, at the period + when the males are castrated; so that the following returns do + not give the proportions at birth. Moreover, I find that + several great breeders in Scotland, who annually raise some + thousand sheep, are firmly convinced that a larger proportion + of males than of females die during the first one or two years; + therefore the proportion of males would be somewhat greater at + birth than at the age of castration. This is a remarkable + coincidence with what occurs, as we have seen, with mankind, + and both cases probably depend on some common cause. I have + received returns from four gentlemen in England who have bred + lowland sheep, chiefly Leicesters, during the last ten or + sixteen years; they amount altogether to 8965 births, + consisting of 4407 males and 4558 females; that is in the + proportion of 96.7 males to 100 females. With respect to + Cheviot and black-faced sheep bred in Scotland, I have received + returns from six breeders, two of them on a large scale, + chiefly for the years 1867-1869, but some of the returns + extending back to 1862. The total number recorded amounts to + 50,685, consisting of 25,071 males and 25,614 females, or in + the proportion of 97.9 males to 100 females. If we take the + English and Scotch returns together, the total number amounts + to 59,650, consisting of 29,478 males and 30,172 females, or + as 97.7 to 100. So that with sheep at the age of castration the + females are certainly in excess of the males; but whether this + would hold good at birth is doubtful, owing to the greater + liability in the males to early death.[376] + + Of _Cattle_ I have received returns from nine gentlemen of 982 + births, too few to be trusted; these consisted of 477 + bull-calves and 505 cow-calves; _i.e._ in the proportion of + 94.4 males to 100 females. The Rev. W. D. Fox informs me that + in 1867 out of 34 calves born on a farm in Derbyshire only one + was a bull. Mr. Harrison Weir writes to me that he has enquired + from several breeders of _Pigs_, and most of them estimate the + male to the female births as about 7 to 6. This same gentleman + has bred _Rabbits_ for many years, and has noticed that a far + greater number of bucks are produced than does. + + Of mammalia in a state of nature I have been able to learn very + little. In regard to the common rat, I have received + conflicting statements. Mr. R. Elliot of Laighwood, informs me + that a rat-catcher assured him that he had always found the + males in great excess, even with the young in the nest. In + consequence of this, Mr. Elliot himself subsequently examined + some hundred old ones, and found the statement true. Mr. F. + Buckland has bred a large number of white rats, and he also + believes that the males greatly exceed the females. In regard + to Moles, it is said that "the males are much more numerous + than the females;"[377] and as the catching of these animals is + a special occupation, the statement may perhaps be trusted. Sir + A. Smith, in describing an antelope of S. Africa[378] (_Kobus + ellipsiprymnus_), remarks, that in the herds of this and other + species, the males are few in number compared with the females: + the natives believe that they are born in this proportion; + others believe that the younger males are expelled from the + herds, and Sir A. Smith says, that though he has himself never + seen herds consisting of young males alone, others affirm that + this does occur. It appears probable that the young males when + expelled from the herd, would be likely to fell a prey to the + many beasts of prey of the country. + + + BIRDS. + + With respect to the _Fowl_, I have received only one account, + namely, that out of 1001 chickens of a highly-bred stock of + Cochins, reared during eight years by Mr. Stretch, 487 proved + males and 514 females: _i.e._ as 94.7 to 100. In regard to + domestic pigeons there is good evidence that the males are + produced in excess, or that their lives are longer; for these + birds invariably pair, and single males, as Mr. Tegetmeier + informs me, can always be purchased cheaper than females. + Usually the two birds reared from the two eggs laid in the same + nest consist of a male and female; but Mr. Harrison Weir, who + has been so large a breeder, says that he has often bred two + cocks from the same nest, and seldom two hens; moreover the hen + is generally the weaker of the two, and more liable to perish. + + With respect to birds in a state of nature, Mr. Gould and + others[379] are convinced that the males are generally the more + numerous; and as the young males of many species resemble the + females, the latter would naturally appear to be the most + numerous. Large numbers of pheasants are reared by Mr. Baker of + Leadenhall from eggs laid by wild birds, and he informs Mr. + Jenner Weir that four or five males to one female are generally + produced. An experienced observer remarks[380] that in + Scandinavia the broods of the capercailzie and black-cock + contain more males than females; and that with the Dal-ripa (a + kind of ptarmigan) more males than females attend the _leks_ or + places of courtship; but this latter circumstance is accounted + for by some observers by a greater number of hen birds being + killed by vermin. From various facts given by White of + Selbourne,[381] it seems clear that the males of the partridge + must be in considerable excess in the south of England; and I + have been assured that this is the case in Scotland. Mr. Weir + on enquiring from the dealers who receive at certain seasons + large numbers of ruffs (_Machetes pugnax_) was told that the + males are much the most numerous. This same naturalist has also + enquired for me from the bird-catchers, who annually catch an + astonishing number of various small species alive for the + London market, and he was unhesitatingly answered by an old and + trustworthy man, that with the chaffinch the males are in large + excess; he thought as high as 2 males to 1 female, or at least + as high as 5 to 3.[382] The males of the blackbird, he likewise + maintained, were by far the most numerous, whether caught by + traps or by netting at night. These statements may apparently + be trusted, because the same man said that the sexes are about + equal with the lark, the twite (_Linaria montana_), and + goldfinch. On the other hand he is certain that with the common + linnet, the females preponderate greatly, but unequally during + different years; during some years he has found the females to + the males as four to one. It should, however, be borne in mind, + that the chief season for catching birds does not begin till + September, so that with some species partial migrations may + have begun, and the flocks at this period often consist of hens + alone. Mr. Salvin paid particular attention to the sexes of the + humming-birds in Central America, and he is convinced that with + most of the species the males are in excess; thus one year he + procured 204 specimens belonging to ten species, and these + consisted of 166 males and of 38 females. With two other + species the females were in excess: but the proportions + apparently vary either during different seasons or in different + localities; for on one occasion the males of _Campylopterus + hemileucurus_ were to the females as five to two, and on + another occasion[383] in exactly the reversed ratio. As bearing + on this latter point, I may add, that Mr. Powys found in Corfu + and Epirus the sexes of the chaffinch keeping apart, and "the + females by far the most numerous;" whilst in Palestine Mr. + Tristram found "the male flocks appearing greatly to exceed the + female in number."[384] So again with the _Quiscalus major_, + Mr. G. Taylor[385] says, that in Florida there were "very few + females in proportion to the males," whilst in Honduras the + proportion was the other way, the species there having the + character of a polygamist. + + + FISH. + + With Fish the proportional numbers of the sexes can be + ascertained only by catching them in the adult or nearly adult + state; and there are many difficulties in arriving at any just + conclusion.[386] Infertile females might readily be mistaken + for males, as Dr. Guenther has remarked to me in regard to + trout. With some species the males are believed to die soon + after fertilising the ova. With many species the males are of + much smaller size than the females, so that a large number of + males would escape from the same net by which the females were + caught. M. Carbonnier,[387] who has especially attended to the + natural history of the pike (_Esox lucius_) states that many + males, owing to their small size, are devoured by the larger + females; and he believes that the males of almost all fish are + exposed from the same cause to greater danger than the females. + Nevertheless in the few cases in which the proportional numbers + have been actually observed, the males appear to be largely in + excess. Thus Mr. R. Buist, the superintendent of the + Stormontfield experiments, says that in 1865, out of 70 salmon + first landed for the purpose of obtaining the ova, upwards of + 60 were males. In 1867 he again "calls attention to the vast + disproportion of the males to the females. We had at the outset + at least ten males to one female." Afterwards sufficient + females for obtaining ova were procured. He adds, "from the + great proportion of the males, they are constantly fighting and + tearing each other on the spawning-beds."[388] This + disproportion, no doubt, can be accounted for in part, but + whether wholly is very doubtful, by the males ascending the + rivers before the females. Mr. F. Buckland remarks in regard to + trout, that "it is a curious fact that the males preponderate + very largely in number over the females. It _invariably_ + happens that when the first rush of fish is made to the net, + there will be at least seven or eight males to one female found + captive. I cannot quite account for this; either the males are + more numerous than the females, or the latter seek safety by + concealment rather than flight." He then adds, that by + carefully searching the banks, sufficient females for obtaining + ova can be found.[389] Mr. H. Lee informs me that out of 212 + trout, taken for this purpose in Lord Portsmouth's park, 150 + were males and 62 females. + + With the Cyprinidae the males likewise seem to be in excess; but + several members of this Family, viz., the carp, tench, bream + and minnow, appear regularly to follow the practice, rare in + the animal kingdom, of polyandry; for the female whilst + spawning is always attended by two males, one on each side, and + in the case of the bream by three or four males. This fact is + so well known, that it is always recommended to stock a pond + with two male tenches to one female, or at least with three + males to two females. With the minnow, an excellent observer + states, that on the spawning-beds the males are ten times as + numerous as the females; when a female comes amongst the males, + "she is immediately pressed closely by a male on each side; and + when they have been in that situation for a time, are + superseded by other two males."[390] + + + INSECTS. + + In this class, the Lepidoptera alone afford the means of + judging of the proportional numbers of the sexes; for they have + been collected with special care by many good observers, and + have been largely bred from the egg or caterpillar state. I had + hoped that some breeders of silk-moths might have kept an exact + record, but after writing to France and Italy, and consulting + various treatises, I cannot find that this has ever been done. + The general opinion appears to be that the sexes are nearly + equal, but in Italy as I hear from Professor Canestrini, many + breeders are convinced that the females are produced in excess. + The same naturalist, however, informs me, that in the two + yearly broods of the Ailanthus silk-moth (_Bombyx cynthia_), + the males greatly preponderate in the first, whilst in the + second the two sexes are nearly equal, or the females rather in + excess. + + In regard to Butterflies in a state of nature, several + observers have been much struck by the apparently enormous + preponderance of the males.[391] Thus Mr. Bates,[392] in + speaking of the species, no less than about a hundred in + number, which inhabit the Upper Amazons, says that the males + are much more numerous than the females, even in the proportion + of a hundred to one. In North America, Edwards, who had great + experience, estimates in the genus Papilio the males to the + females as four to one; and Mr. Walsh, who informed me of this + statement, says that with _P. turnus_ this is certainly the + case. In South Africa, Mr. R. Trimen found the males in excess + in 19 species;[393] and in one of these, which swarms in open + places, he estimated the number of males as fifty to one + female. With another species, in which the males are numerous + in certain localities, he collected during seven years only + five females. In the island of Bourbon, M. Maillard states that + the males of one species of Papilio are twenty times as + numerous as the females.[394] Mr. Trimen informs me that as far + as he has himself seen, or heard from others, it is rare for + the females of any butterfly to exceed in number the males; but + this is perhaps the case with three South African species. Mr. + Wallace[395] states that the females of _Ornithoptera + croesus_, in the Malay archipelago, are more common and more + easily caught than the males; but this is a rare butterfly. I + may here add, that in Hyperythra, a genus of moths, Guenee + says, that from four to five females are sent in collections + from India for one male. + + When this subject of the proportional numbers of the sexes of + insects was brought before the Entomological Society,[396] it + was generally admitted that the males of most Lepidoptera, in + the adult or imago state, are caught in greater numbers than + the females; but this fact was attributed by various observers + to the more retiring habits of the females, and to the males + emerging earlier from the cocoon. This latter circumstance is + well known to occur with most Lepidoptera, as well as with + other insects. So that, as M. Personnat remarks, the males of + the domesticated _Bombyx yamamai_, are lost at the beginning of + the season, and the females at the end, from the want of + mates.[397] I cannot however persuade myself that these causes + suffice to explain the great excess of males in the cases, + above given, of butterflies which are extremely common in their + native countries. Mr. Stainton, who has paid such close + attention during many years to the smaller moths, informs me + that when he collected them in the imago state, he thought that + the males were ten times as numerous as the females, but that + since he has reared them on a large scale from the caterpillar + state, he is convinced that the females are the most numerous. + Several entomologists concur in this view. Mr. Doubleday, + however, and some others, take an opposite view, and are + convinced that they have reared from the egg and caterpillar + states a larger proportion of males than of females. + + Besides the more active habits of the males, their earlier + emergence from the cocoon, and their frequenting in some cases + more open stations, other causes may be assigned for an + apparent or real difference in the proportional numbers of the + sexes of Lepidoptera, when captured in the imago state, and + when reared from the egg or caterpillar state. It is believed + by many breeders in Italy, as I hear from Professor Canestrini, + that the female caterpillar of the silk-moth suffers more from + the recent disease than the male; and Dr. Staudinger informs me + that in rearing Lepidoptera more females die in the cocoon than + males. With many species the female caterpillar is larger than + the male, and a collector would naturally choose the finest + specimens, and thus unintentionally collect a larger number of + females. Three collectors have told me that this was their + practice; but Dr. Wallace is sure that most collectors take all + the specimens which they can find of the rarer kinds, which + alone are worth the trouble of rearing. Birds when surrounded + by caterpillars would probably devour the largest; and + Professor Canestrini informs me that in Italy some breeders + believe, though on insufficient evidence, that in the first + brood of the Ailanthus silk-moth, the wasps destroy a larger + number of the female than of the male caterpillars. Dr. Wallace + further remarks that female caterpillars, from being larger + than the males, require more time for their development and + consume more food and moisture; and thus they would be exposed + during a longer time to danger from ichneumons, birds, &c., and + in times of scarcity would perish in greater numbers. Hence it + appears quite possible that, in a state of nature, fewer female + Lepidoptera may reach maturity than males; and for our special + object we are concerned with the numbers at maturity, when the + sexes are ready to propagate their kind. + + The manner in which the males of certain moths congregate in + extraordinary numbers round a single female, apparently + indicates a great excess of males, though this fact may perhaps + be accounted for by the earlier emergence of the males from + their cocoons. Mr. Stainton informs me that from twelve to + twenty males may often be seen congregated round a female + _Elachista rufocinerea_. It is well known that if a virgin + _Lasiocampa quercus_ or _Saturnia carpini_ be exposed in a + cage, vast numbers of males collect round her, and if confined + in a room will even come down the chimney to her. Mr. + Doubleday believes that he has seen from fifty to a hundred + males of both these species attracted in the course of a single + day by a female under confinement. Mr. Trimen exposed in the + Isle of Wight a box in which a female of the Lasiocampa had + been confined on the previous day, and five males soon + endeavoured to gain admittance. M. Verreaux, in Australia, + having placed the female of a small Bombyx in a box in his + pocket, was followed by a crowd of males, so that about 200 + entered the house with him.[398] + + Mr. Doubleday has called my attention to Dr. Staudinger's[399] + list of Lepidoptera, which gives the prices of the males and + females of 300 species or well-marked varieties of + (Rhopalocera) butterflies. The prices for both sexes of the + very common species are of course the same; but with 114 of the + rarer species they differ; the males being in all cases, + excepting one, the cheapest. On an average of the prices of the + 113 species, the price of the male to that of the female is as + 100 to 149; and this apparently indicates that inversely the + males exceed the females in number in the same proportion. + About 2000 species or varieties of moths (Heterocera) are + catalogued, those with wingless females being here excluded on + account of the difference in habits of the two sexes: of these + 2000 species, 141 differ in price according to sex, the males + of 130 being cheaper, and the males of only 11 being dearer + than the females. The average price of the males of the 130 + species, to that of the females, is as 100 to 143. With respect + to the butterflies in this priced list, Mr. Doubleday thinks + (and no man in England has had more experience), that there is + nothing in the habits of the species which can account for the + difference in the prices of the two sexes, and that it can be + accounted for only by an excess in the numbers of the males. + But I am bound to add that Dr. Staudinger himself, as he + informs me, is of a different opinion. He thinks that the less + active habits of the females and the earlier emergence of the + males will account for his collectors securing a larger number + of males than of females, and consequently for the lower prices + of the former With respect to specimens reared from the + caterpillar-state, Dr. Staudinger believes, as previously + stated, that a greater number of females than of males die + under confinement in the cocoons. He adds that with certain + species one sex seems to preponderate over the other during + certain years. + + Of direct observations on the sexes of Lepidoptera, reared + either from eggs or caterpillars, I have received only the few + following cases:-- + + -------------------------------------------------------------------------------------------------------------------------------------- + Males. | Females. + The Rev. J. Hellins[400] of Exeter reared, during | + 1868, imagos of 73 species, which consisted of 153 | 137 + | + Mr. Albert Jones of Eltham reared, during 1868, | + imagos of 9 species, which, consisted of 159 | 126 + | + During 1869 he reared imagos from 4 species, | + consisting of 114 | 112 + | + Mr. Buckler of Emsworth, Hants, during 1869, | + reared imagos from 74 species, consisting of 180 | 169 + | + Dr. Wallace of Colchester reared from one brood | + of Bombyx cynthia 52 | 48 + | + Dr. Wallace raised, from cocoons of Bombyx | + Pernyi sent from China, during 1869 224 | 123 + | + Dr. Wallace raised, during 1868 and 1869, | + from two lots of cocoons of Bombyx yamamai 52 | 46 + --------------+------------ + Total 934 | 761 + ------------------------------------------------------------------------------------------------------------------+------------------ + + So that in these eight lots of cocoons and eggs, males were + produced in excess. Taken together the proportion of males is + as 122.7 to 100 females. But the numbers are hardly large + enough to be trustworthy. + + On the whole, from the above various sources of evidence, all + pointing to the same direction, I infer that with most species + of Lepidoptera, the males in the imago state generally exceed + the females in number, whatever the proportions may be at their + first emergence from the egg. + + With reference to the other Orders of insects, I have been able + to collect very little reliable information. With the + stag-beetle (_Lucanus cervus_) "the males appear to be much + more numerous than the females;" but when, as Cornelius + remarked during 1867, an unusual number of these beetles + appeared in one part of Germany, the females appeared to exceed + the males as six so one. With one of the Elateridae, the males + are said to be much more numerous than the females, and "two or + three are often found united with one female;"[401] so that here + polyandry seems to prevail. With Siagonium (Staphylinidae), in + which the males are furnished with horns, "the females are far + more numerous than the opposite sex." Mr. Janson stated at the + Entomological Society that the females of the bark-feeding + _Tomicus villosus_ are so common as to be a plague, whilst the + males are so rare as to be hardly known. In other Orders, from + unknown causes, but apparently in some instances owing to + parthenogenesis, the males of certain species have never been + discovered or are excessively rare, as with several of the + Cynipidae.[402] In all the gall-making Cynipidae known to Mr. + Walsh, the females are four or five times as numerous as the + males; and so it is, as he informs me, with the gall-making + Cecidomyiiae (Diptera). With some common species of Saw-flies + (Tenthredinae) Mr. F. Smith has reared hundreds of specimens + from larvae of all sizes, but has never reared a single male: on + the other hand Curtis says,[403] that with certain species + (Athalia), bred by him, the males to the females were as six to + one; whilst exactly the reverse occurred with the mature + insects of the same species caught in the fields. With the + Neuroptera, Mr. Walsh states that in many, but by no means in + all, the species of the Odonatous groups (Ephemerina), there is + a great overplus of males: in the genus Hetaerina, also, the + males are generally at least four times as numerous as the + females. In certain species in the genus Gomphus the males are + equally numerous, whilst in two other species, the females are + twice or thrice as numerous as the males. In some European + species of Psocus thousands of females may be collected without + a single male, whilst with other species of the same genus both + sexes are common.[404] In England, Mr. MacLachlan has captured + hundreds of the female _Apatania muliebris_, but has never seen + the male; and of _Boreus hyemalis_ only four or five males have + been here seen.[405] With most of these species (excepting, as + I have heard, with the Tenthredinae) there is no reason to + suppose that the females are subject to parthenogenesis; and + thus we see how ignorant we are on the causes of the apparent + discrepancy in the proportional numbers of the two sexes. + + In the other Classes of the Articulata I have been able to + collect still less information. With Spiders, Mr. Blackwall, + who has carefully attended to this class during many years, + writes to me that the males from their more erratic habits are + more commonly seen, and therefore appear to be the more + numerous. This is actually the case with a few species; but he + mentions several species in six genera, in which the females + appear to be much more numerous than the males.[406] The small + size of the males in comparison with the females, which is + sometimes carried to an extreme degree, and their widely + different appearance, may account in some instances for their + rarity in collections.[407] + + Some of the lower Crustaceans are able to propagate their kind + asexually, and this will account for the extreme rarity of the + males. With some other forms (as with Tanais and Cypris) there + is reason to believe, as Fritz Mueller informs me, that the male + is much shorter-lived than the female, which, supposing the two + sexes to be at first equal in number, would explain the + scarcity of the males. On the other hand this same naturalist + has invariably taken, on the shores of Brazil, far more males + than females of the Diastylidae and of Cypridina; thus with a + species in the latter genus, 63 specimens caught the same day, + included 57 males; but he suggests that this preponderance may + be due to some unknown difference in the habits of the two + sexes. With one of the higher Brazilian crabs, namely a + Gelasimus, Fritz Mueller found the males to be more numerous + than the females. The reverse seems to be the case, according + to the large experience of Mr. C. Spence Bate, with six common + British crabs, the names of which he has given me. + + +_On the Power of Natural Selection to regulate the proportional Numbers +of the Sexes, and General Fertility._--In some peculiar cases, an excess +in the number of one sex over the other might be a great advantage to a +species, as with the sterile females of social insects, or with those +animals in which more than one male is requisite to fertilise the +female, as with certain cirripedes and perhaps certain fishes. An +inequality between the sexes in these cases might have been acquired +through natural selection, but from their rarity they need not here be +further considered. In all ordinary cases an inequality would be no +advantage or disadvantage to certain individuals more than to others; +and therefore it could hardly have resulted from natural selection. We +must attribute the inequality to the direct action of those unknown +conditions, which with mankind lead to the males being born in a +somewhat larger excess in certain countries than in others, or which +cause the proportion between the sexes to differ slightly in legitimate +and illegitimate births. + +Let us now take the case of a species producing from the unknown causes +just alluded to, an excess of one sex--we will say of males--these being +superfluous and useless, or nearly useless. Could the sexes be equalised +through natural selection? We may feel sure, from all characters being +variable, that certain pairs would produce a somewhat less excess of +males over females than other pairs. The former, supposing the actual +number of the offspring to remain constant, would necessarily produce +more females, and would therefore be more productive. On the doctrine of +chances a greater number of the offspring of the more productive pairs +would survive; and these would inherit a tendency to procreate fewer +males and more females. Thus a tendency towards the equalisation of the +sexes would be brought about. But our supposed species would by this +process be rendered, as just remarked, more productive; and this would +in many cases be far from an advantage; for whenever the limit to the +numbers which exist, depends, not on destruction by enemies, but on the +amount of food, increased fertility will lead to severer competition and +to most of the survivors being badly fed. In this case, if the sexes +were equalised by an increase in the number of the females, a +simultaneous decrease in the total number of the offspring would be +beneficial, or even necessary, for the existence of the species; and +this, I believe, could be effected through natural selection in the +manner hereafter to be described. The same train of reasoning is +applicable in the above, as well as in the following case, if we assume +that females instead of males are produced in excess, for such females +from not uniting with males would be superfluous and useless. So it +would be with polygamous species, if we assume the excess of females to +be inordinately great. + +An excess of either sex, we will again say of the males, could, however, +apparently be eliminated through natural selection in another and +indirect manner, namely by an actual diminution of the males, without +any increase of the females, and consequently without any increase in +the productiveness of the species. From the variability of all +characters, we may feel assured that some pairs, inhabiting any +locality, would produce a rather smaller excess of superfluous males, +but an equal number of productive females. When the offspring from the +more and the less male-productive parents were all mingled together, +none would have any direct advantage over the others; but those that +produced few superfluous males would have one great indirect advantage, +namely that their ova or embryos would probably be larger and finer, or +their young better nurtured in the womb and afterwards. We see this +principle illustrated with plants; as those which bear a vast number of +seed produce small ones; whilst those which bear comparatively few +seeds, often produce large ones well-stocked with nutriment for the use +of the seedlings.[408] Hence the offspring of the parents which had +wasted least force in producing superfluous males would be the most +likely to survive, and would inherit the same tendency not to produce +superfluous males, whilst retaining their full fertility in the +production of females. So it would be with the converse case of the +female sex. Any slight excess, however, of either sex could hardly be +checked in so indirect a manner. Nor indeed has a considerable +inequality between the sexes been always prevented, as we have seen in +some of the cases given in the previous discussion. In these cases the +unknown causes which determine the sex of the embryo, and which under +certain conditions lead to the production of one sex in excess over the +other, have not been mastered by the survival of those varieties which +were subjected to the least waste of organised matter and force by the +production of superfluous individuals of either sex. Nevertheless we may +conclude that natural selection will always tend, though sometimes +inefficiently, to equalise the relative numbers of the two sexes. + +Having said this much on the equalisation of the sexes, it may be well +to add a few remarks on the regulation through natural selection of the +ordinary fertility of species. Mr. Herbert Spencer has shewn in an able +discussion[409] that with all organisms a ratio exists between what he +calls individuation and genesis; whence it follows that beings which +consume much matter or force in their growth, complicated structure or +activity, or which produce ova and embryos of large size, or which +expend much energy in nurturing their young, cannot be so productive as +beings of an opposite nature. Mr. Spencer further shews that minor +differences in fertility will be regulated through natural selection. +Thus the fertility of each species will tend to increase, from the more +fertile pairs producing a larger number of offspring, and these from +their mere number will have the best chance of surviving, and will +transmit their tendency to greater fertility. The only check to a +continued augmentation of fertility in each organism seems to be either +the expenditure of more power and the greater risks run by the parents +that produce a more numerous progeny, or the contingency of very +numerous eggs and young being produced of smaller size, or less +vigorous, or subsequently not so well nurtured. To strike a balance in +any case between the disadvantages which follow from the production of a +numerous progeny, and the advantages (such as the escape of at least +some individuals from various dangers) is quite beyond our power of +judgment. + +When an organism has once been rendered extremely fertile, how its +fertility can be reduced through natural selection is not so clear as +how this capacity was first acquired. Yet it is obvious that if +individuals of a species, from a decrease of their natural enemies, were +habitually reared in larger numbers than could be supported, all the +members would suffer. Nevertheless the offspring from the less fertile +parents would have no direct advantage over the offspring from the more +fertile parents, when all were mingled together in the same district. +All the individuals would mutually tend to starve each other. The +offspring indeed of the less fertile parents would lie under one great +disadvantage, for from the simple fact of being produced in smaller +numbers, they would be the most liable to extermination. Indirectly, +however, they would partake of one great advantage; for under the +supposed condition of severe competition, when all were pressed for +food, it is extremely probable that those individuals which from some +variation in their constitution produced fewer eggs or young, would +produce them of greater size or vigour; and the adults reared from such +eggs or young would manifestly have the best chance of surviving, and +would inherit a tendency towards lessened fertility. The parents, +moreover, which had to nourish or provide for fewer offspring would +themselves be exposed to a less severe strain in the struggle for +existence, and would have a better chance of surviving. By these steps, +and by no others as far as I can see, natural selection under the above +conditions of severe competition for food, would lead to the formation +of a new race less fertile, but better adapted for survival, than the +parent-race. + + + + +CHAPTER IX. + +SECONDARY SEXUAL CHARACTERS IN THE LOWER CLASSES OF THE ANIMAL KINGDOM. + + + These characters absent in the lowest classes--Brilliant + colours--Mollusca--Annelids--Crustacea, secondary sexual + characters strongly developed; dimorphism; colour; characters + not acquired before maturity--Spiders, sexual colours of; + stridulation by the males--Myriapoda. + + +In the lowest classes the two sexes are not rarely united in the same +individual, and therefore secondary sexual characters cannot be +developed. In many cases in which the two sexes are separate, both are +permanently attached to some support, and the one cannot search or +struggle for the other. Moreover it is almost certain that these animals +have too imperfect senses and much too low mental powers to feel mutual +rivalry, or to appreciate each other's beauty or other attractions. + +Hence in these classes, such as the Protozoa, Coelenterata, +Echinodermata, Scolecida, true secondary sexual characters do not occur; +and this fact agrees with the belief that such characters in the higher +classes have been acquired through sexual selection, which depends on +the will, desires, and choice of either sex. Nevertheless some few +apparent exceptions occur; thus, as I hear from Dr. Baird, the males of +certain Entozoa, or internal parasitic worms, differ slightly in colour +from the females; but we have no reason to suppose that such differences +have been augmented through sexual selection. + +Many of the lower animals, whether hermaphrodites or with the sexes +separate, are ornamented with the most brilliant tints, or are shaded +and striped in an elegant manner. This is the case with many corals and +sea-anemonies (Actiniae), with some jelly-fish (Medusae, Porpita, &c.), +with some Planariae, Ascidians, numerous Star-fishes, Echini, &c.; but we +may conclude from the reasons already indicated, namely the union of the +two sexes in some of these animals, the permanently affixed condition of +others, and the low mental powers of all, that such colours do not serve +as a sexual attraction, and have not been acquired through sexual +selection. With the higher animals the case is very different; for with +them when one sex is much more brilliantly or conspicuously coloured +than the other, and there is no difference in the habits of the two +sexes which will account for this difference, we have reason to believe +in the influence of sexual selection; and this belief is strongly +confirmed when the more ornamented individuals, which are almost always +the males, display their attractions before the other sex. We may also +extend this conclusion to both sexes, when coloured alike, if their +colours are plainly analogous to those of one sex alone in certain other +species of the same group. + +How, then, are we to account for the beautiful or even gorgeous colours +of many animals in the lowest classes? It appears very doubtful whether +such colours usually serve as a protection; but we are extremely liable +to err in regard to characters of all kinds in relation to protection, +as will be admitted by every one who has read Mr. Wallace's excellent +essay on this subject. It would not, for instance, at first occur to any +one that the perfect transparency of the Medusae, or jelly-fishes, was of +the highest service to them as a protection; but when we are reminded +by Haeckel that not only the medusae but many floating mollusca, +crustaceans, and even small oceanic fishes partake of this same +glass-like structure, we can hardly doubt that they thus escape the +notice of pelagic birds and other enemies. + +Notwithstanding our ignorance how far colour in many cases serves as a +protection, the most probable view in regard to the splendid tints of +many of the lowest animals seems to be that their colours are the direct +result either of the chemical nature or the minute structure of their +tissues, independently of any benefit thus derived. Hardly any colour is +finer than that of arterial blood; but there is no reason to suppose +that the colour of the blood is in itself any advantage; and though it +adds to the beauty of the maiden's cheek, no one will pretend that it +has been acquired for this purpose. So again with many animals, +especially the lower ones, the bile is richly coloured; thus the extreme +beauty of the Eolidae (naked sea-slugs) is chiefly due, as I am informed +by Mr. Hancock, to the biliary glands seen through the translucent +integuments; this beauty being probably of no service to these animals. +The tints of the decaying leaves in an American forest are described by +every one as gorgeous; yet no one supposes that these tints are of the +least advantage to the trees. Bearing in mind how many substances +closely analogous to natural organic compounds have been recently formed +by chemists, and which exhibit the most splendid colours, it would have +been a strange fact if substances similarly coloured had not often +originated, independently of any useful end being thus gained, in the +complex laboratory of living organisms. + + +_The sub-kingdom of the Mollusca._--Throughout this great division +(taken in its largest acceptation) of the animal kingdom, secondary +sexual characters, such as we are here considering, never, as far as I +can discover, occur. Nor could they be expected in the three lowest +classes, namely in the Ascidians, Polyzoa, and Brachiopods (constituting +the Molluscoida of Huxley), for most of these animals are permanently +affixed to a support or have their sexes united in the same individual. +In the Lamellibranchiata, or bivalve shells, hermaphroditism is not +rare. In the next higher class of the Gasteropoda, or marine univalve +shells, the sexes are either united or separate. But in this latter case +the males never possess special organs for finding, securing, or +charming the females, or for fighting with other males. The sole +external difference between the sexes consists, as I am informed by Mr. +Gwyn Jeffreys, in the shell sometimes differing a little in form; for +instance, the shell of the male periwinkle (_Littorina littorea_) is +narrower and has a more elongated spire than that of the female. But +differences of this nature, it may be presumed, are directly connected +with the act of reproduction or with the development of the ova. + +The Gasteropoda, though capable of locomotion and furnished with +imperfect eyes, do not appear to be endowed with sufficient mental +powers for the members of the same sex to struggle together in rivalry, +and thus to acquire secondary sexual characters. Nevertheless with the +pulmoniferous gasteropods, or land-snails, the pairing is preceded by +courtship; for these animals, though hermaphrodites, are compelled by +their structure to pair together. Agassiz remarks,[410] "Quiconque a eu +l'occasion d'observer les amours des limacons, ne saurait mettre en +doute la seduction deployee dans les mouvements et les allures qui +preparent et accomplissent le double embrassement de ces +hermaphrodites." These animals appear also susceptible of some degree of +permanent attachment: an accurate observer, Mr. Lonsdale, informs me +that he placed a pair of land-shells (_Helix pomatia_), one of which was +weakly, into a small and ill-provided garden. After a short time the +strong and healthy individual disappeared, and was traced by its track +of slime over a wall into an adjoining well-stocked garden. Mr. Lonsdale +concluded that it had deserted its sickly mate; but after an absence of +twenty-four hours it returned, and apparently communicated the result of +its successful exploration, for both then started along the same track +and disappeared over the wall. + +Even in the highest class of the Mollusca, namely the Cephalopoda or +cuttle-fishes, in which the sexes are separate, secondary sexual +characters of the kind which we are here considering, do not, as far as +I can discover, occur. This is a surprising circumstance, as these +animals possess highly-developed sense-organs and have considerable +mental powers, as will be admitted by every one who has watched their +artful endeavours to escape from an enemy.[411] Certain Cephalopoda, +however, are characterised by one extraordinary sexual character, +namely, that the male element collects within one of the arms or +tentacles, which is then cast off, and, clinging by its sucking-discs to +the female, lives for a time an independent life. So completely does the +cast-off arm resemble a separate animal, that it was described by Cuvier +as a parasitic worm under the name of Hectocotyle. But this marvellous +structure may be classed as a primary rather than as a secondary sexual +character. + +Although with the Mollusca sexual selection does not seem to have come +into play; yet many univalve and bivalve shells, such as volutes, cones, +scallops, &c., are beautifully coloured and shaped. The colours do not +appear in most cases to be of any use as a protection; they are probably +the direct result, as in the lowest classes, of the nature of the +tissues; the patterns and the sculpture of the shell depending on its +manner of growth. The amount of light seems to a certain extent to be +influential; for although, as repeatedly stated by Mr. Gwyn Jeffreys, +the shells of some species living at a profound depth are brightly +coloured, yet we generally see the lower surfaces and the parts covered +by the mantle less highly coloured than the upper and exposed +surfaces.[412] In some cases, as with shells living amongst corals or +brightly-tinted sea-weeds, the bright colours may serve as a protection. +But many of the nudibranch mollusca, or sea-slugs, are as beautifully +coloured as any shells, as may be seen in Messrs. Alder and Hancock's +magnificent work; and from information kindly given me by Mr. Hancock, +it is extremely doubtful whether these colours usually serve as a +protection. With some species this may be the case, as with one which +lives on the green leaves of algae, and is itself bright-green. But many +brightly-coloured, white or otherwise conspicuous species, do not seek +concealment; whilst again some equally conspicuous species, as well as +other dull-coloured kinds, live under stones and in dark recesses. So +that with these nudibranch molluscs, colour apparently does not stand in +any close relation to the nature of the places which they inhabit. + +These naked sea-slugs are hermaphrodites, yet they pair together, as do +land-snails, many of which have extremely pretty shells. It is +conceivable that two hermaphrodites, attracted by each others' greater +beauty, might unite and leave offspring which would inherit their +parents' greater beauty. But with such lowly-organised creatures this is +extremely improbable. Nor is it at all obvious how the offspring from +the more beautiful pairs of hermaphrodites would have any advantage, so +as to increase in numbers, over the offspring of the less beautiful, +unless indeed vigour and beauty generally coincided. We have not here a +number of males becoming mature before the females, and the more +beautiful ones selected by the more vigorous females. If, indeed, +brilliant colours were beneficial to an hermaphrodite animal in relation +to its general habits of life, the more brightly-tinted individuals +would succeed best and would increase in number; but this would be a +case of natural and not of sexual selection. + + +_Sub-kingdom of the Vermes or Annulosa_: Class, _Annelida (or +Sea-worms)_.--In this class, although the sexes (when separate) +sometimes differ from each other in characters of such importance that +they have been placed under distinct genera or even families, yet the +differences do not seem of the kind which can be safely attributed to +sexual selection. These animals, like those in the preceding classes, +apparently stand too low in the scale, for the individuals of either sex +to exert any choice in selecting a partner, or for the individuals of +the same sex to struggle together in rivalry. + + +_Sub-kingdom of the Arthropoda_: Class, _Crustacea_.--In this great +class we first meet with undoubted secondary sexual characters, often +developed in a remarkable manner. Unfortunately the habits of +crustaceans are very imperfectly known, and we cannot explain the uses +of many structures peculiar to one sex. With the lower parasitic species +the males are of small size, and they alone are furnished with perfect +swimming-legs, antennae and sense-organs; the females being destitute of +these organs, with their bodies often consisting of a mere distorted +mass. But these extraordinary differences between the two sexes are no +doubt related to their widely different habits of life, and consequently +do not concern us. In various crustaceans, belonging to distinct +families, the anterior antennae are furnished with peculiar thread-like +bodies, which are believed to act as smelling-organs, and these are much +more numerous in the males than in the females. As the males, without +any unusual development of their olfactory organs, would almost +certainly be able sooner or later to find the females, the increased +number of the smelling-threads has probably been acquired through sexual +selection, by the better provided males having been the most successful +in finding partners and in leaving offspring. Fritz Mueller has described +a remarkable dimorphic species of Tanais, in which the male is +represented by two distinct forms, never graduating into each other. In +the one form the male is furnished with more numerous smelling-threads, +and in the other form with more powerful and more elongated chelae or +pincers which serve to hold the female. Fritz Mueller suggests that these +differences between the two male forms of the same species must have +originated in certain individuals having varied in the number of the +smelling-threads, whilst other individuals varied in the shape and size +of their chelae; so that of the former, those which were best able to +find the female, and of the latter, those which were best able to hold +her when found, have left the greater number of progeny to inherit their +respective advantages.[413] + +[Illustration: Fig. 3. Labidocera Darwinii, (from Lubbock). + +_a._ Part of right-hand anterior antenna of male, forming a prehensile +organ. + +_b._ Posterior pair of thoracic legs of male. + +_c._ Ditto of female.] + +In some of the lower crustaceans, the right-hand anterior antenna of the +male differs greatly in structure from the left-hand one, the latter +resembling in its simple tapering joints the antennae of the female. In +the male the modified antenna is either swollen in the middle or +angularly bent, or converted (fig. 3) into an elegant, and sometimes +wonderfully complex, prehensile organ.[414] It serves, as I hear from +Sir J. Lubbock, to hold the female, and for this same purpose one of the +two posterior legs (_b_) on the same side of the body is converted into +a forceps. In another family the inferior or posterior antennae are +"curiously zigzagged" in the males alone. + +[Illustration: Fig. 4. Anterior part of body of Callianassa (from +Milne-Edwards), showing the unequal and differently-constructed right +and left-hand chelae of the male. + +N.B.--The artist by mistake has reversed the drawing, and made the +left-hand chela the largest.] + +[Illustration: Fig. 5. Second leg of male Orchestia Tucuratinga (from +Fritz Mueller).] + +[Illustration: Fig. 6. Ditto of female.] + +In the higher crustaceans the anterior legs form a pair of chelae or +pincers, and these are generally larger in the male than in the female. +In many species the chelae on the opposite sides of the body are of +unequal size, the right-hand one being, as I am informed by Mr. C. +Spence Bate, generally, though not invariably, the largest. This +inequality is often much greater in the male than in the female. The two +chelae also often differ in structure (figs. 4, 5, and 6), the smaller one +resembling those of the female. What advantage is gained by their +inequality in size on the opposite sides of the body, and by the +inequality being much greater in the male than in the female; and why, +when they are of equal size, both are often much larger in the male than +in the female, is not known. The chelae are sometimes of such length and +size that they cannot possibly be used, as I hear from Mr. Spence Bate, +for carrying food to the mouth. In the males of certain freshwater +prawns (Palaemon) the right leg is actually longer than the whole +body.[415] It is probable that the great size of one leg with its chelae +may aid the male in fighting with his rivals; but this use will not +account for their inequality in the female on the opposite sides +of the body. In Gelasimus, according to a statement quoted by +Milne-Edwards,[416] the male and female live in the same burrow, which +is worth notice, as shewing that they pair, and the male closes the +mouth of the burrow with one of its chelae, which is enormously +developed; so that here it indirectly serves as a means of defence. +Their main use, however, probably is to seize and to secure the female, +and this in some instances, as with Gammarus, is known to be the case. +The sexes, however, of the common shore-crab (_Carcinus maenas_), as Mr. +Spence Bate informs me, unite directly after the female has moulted her +hard shell, and when she is so soft that she would be injured if seized +by the strong pincers of the male; but as she is caught and carried +about by the male previously to the act of moulting, she could then be +seized with impunity. + +Fritz Mueller states that certain species of Melita are distinguished +from all other amphipods by the females having "the coxal lamellae of the +penultimate pair of feet produced into hook-like processes, of which the +males lay hold with the hands of the first pair." The development of +these hook-like processes probably resulted from those females which +were the most securely held during the act of reproduction, having left +the largest number of offspring. Another Brazilian amphipod (_Orchestia +Darwinii_, fig. 7) is described by Fritz Mueller, as presenting a case of +dimorphism, like that of Tanais; for there are two male forms, which +differ in the structure of their chelae.[417] As chelae of either shape +would certainly have sufficed to hold the female, for both are now used +for this purpose, the two male forms probably originated, by some having +varied in one manner and some in another; both forms having derived +certain special, but nearly equal advantages, from their differently +shaped organs. + +It is not known that male crustaceans fight together for the possession +of the females, but this is probable; for with most animals when the +male is larger than the female, he seems to have acquired his greater +size by having conquered during many generations other males. Now Mr. +Spence Bate informs me that in most of the crustacean orders, especially +in the highest or the Brachyura, the male is larger than the female; the +parasitic genera, however, in which the sexes follow different habits of +life, and most of the Entomostraca must be excepted. The chelae of many +crustaceans are weapons well adapted for fighting. Thus a Devil-crab +(_Portunus puber_) was seen by a son of Mr. Bate fighting with a +_Carcinus maenas_, and the latter was soon thrown on its back, and had +every limb torn from its body. When several males of a Brazilian +Gelasimus, a species furnished with immense pincers, were placed +together by Fritz Mueller in a glass vessel, they mutilated and killed +each other. Mr. Bate put a large male _Carcinus maenas_ into a pan of +water, inhabited by a female paired with a smaller male; the latter was +soon dispossessed, but, as Mr. Bate adds, "if they fought, the victory +was a bloodless one, for I saw no wounds." This same naturalist +separated a male sand-skipper (so common on our sea-shores), _Gammarus +marinus_, from its female, both of which were imprisoned in the same +vessel with many individuals of the same species. The female being thus +divorced joined her comrades. After an interval the male was again put +into the same vessel and he then, after swimming about for a time, +dashed into the crowd, and without any fighting at once took away his +wife. This fact shews that in the Amphipoda, an order low in the scale, +the males and females recognise each other, and are mutually attached. + +[Illustration: Fig. 7. Orchestia Darwinii (from Fritz Mueller), showing +the differently-constructed chelae of the two male forms.] + +The mental powers of the Crustacea are probably higher than might have +been expected. Any one who has tried to catch one of the shore-crabs, so +numerous on many tropical coasts, will have perceived how wary and alert +they are. There is a large crab (_Birgus latro_), found on coral +islands, which makes at the bottom of a deep burrow a thick bed of the +picked fibres of the cocoa-nut. It feeds on the fallen fruit of this +tree by tearing off the husk, fibre by fibre; and it always begins at +that end where the three eye-like depressions are situated. It then +breaks through one of these eyes by hammering with its heavy front +pincers, and turning round, extracts the albuminous core with its narrow +posterior pincers. But these actions are probably instinctive, so that +they would be performed as well by a young as by an old animal. The +following case, however, can hardly be so considered: a trustworthy +naturalist, Mr. Gardner,[418] whilst watching a shore-crab (Gelasimus) +making its burrow, threw some shells towards the hole. One rolled in, +and three other shells remained within a few inches of the mouth. In +about five minutes the crab brought out the shell which had fallen in, +and carried it away to the distance of a foot; it then saw the three +other shells lying near, and evidently thinking that they might likewise +roll in, carried them to the spot where it had laid the first. It would, +I think, be difficult to distinguish this act from one performed by man +by the aid of reason. + +With respect to colour which so often differs in the two sexes of +animals belonging to the higher classes, Mr. Spence Bate does not know +of any well-marked instances with our British crustaceans. In some +cases, however, the male and female differ slightly in tint, but Mr. +Bate thinks not more than may be accounted for by their different habits +of life, such as by the male wandering more about and being thus more +exposed to the light. In a curious Bornean crab, which inhabits sponges, +Mr. Bate could always distinguish the sexes by the male not having the +epidermis so much rubbed off. Dr. Power tried to distinguish by colour +the sexes of the species which inhabit the Mauritius, but always failed, +except with one species of Squilla, probably the _S. stylifera_, the +male of which is described as being "of a beautiful blueish-green," with +some of the appendages cherry-red, whilst the female is clouded with +brown and grey, "with the red about her much less vivid than in the +male."[419] In this case, we may suspect the agency of sexual selection. +With Saphirina (an oceanic genus of Entomostraca, and therefore low in +the scale) the males are furnished with minute shields or cell-like +bodies, which exhibit beautiful changing colours; these being absent in +the females, and in the case of one species in both sexes.[420] It +would, however, be extremely rash to conclude that these curious organs +serve merely to attract the females. In the female of a Brazilian +species of Gelasimus, the whole body, as I am informed by Fritz Mueller, +is of a nearly uniform greyish-brown. In the male the posterior part of +the cephalo-thorax is pure white, with the anterior part of a rich +green, shading into dark brown; and it is remarkable that these colours +are liable to change in the course of a few minutes--the white becoming +dirty grey or even black, the green "losing much of its brilliancy." The +males apparently are much more numerous than the females. It deserves +especial notice that they do not acquire their bright colours until they +become mature. They differ also from the females in the larger size of +their chelae. In some species of the genus, probably in all, the sexes +pair and inhabit the same burrow. They are also, as we have seen, highly +intelligent animals. From these various considerations it seems highly +probable that the male in this species has become gaily ornamented in +order to attract or excite the female. + +It has just been stated that the male Gelasimus does not acquire his +conspicuous colours until mature and nearly ready to breed. This seems +the general rule in the whole class with the many remarkable differences +in structure between the two sexes. We shall hereafter find the same law +prevailing throughout the great sub-kingdom of the Vertebrata, and in +all cases it is eminently distinctive of characters which have been +acquired through sexual selection. Fritz Mueller[421] gives some +striking instances of this law; thus the male sand-hopper (Orchestia) +does not acquire his large claspers, which are very differently +constructed from those of the female, until nearly full-grown; whilst +young his claspers resemble those of the female. Thus, again, the male +Brachyscelus possesses, like all other amphipods, a pair of posterior +antennae; the female, and this is a most extraordinary circumstance, is +destitute of them, and so is the male as long as he remains immature. + + +Class, _Arachnida_ (Spiders).--The males are often darker, but sometimes +lighter than the females, as may be seen in Mr. Blackwall's magnificent +work.[422] In some species the sexes differ conspicuously from each +other in colour; thus the female of _Sparassus smaragdulus_ is +dullish-green; whilst the adult male has the abdomen of a fine yellow, +with three longitudinal stripes of rich red. In some species of Thomisus +the two sexes closely resemble each other; in others they differ much; +thus in _T. citreus_ the legs and body of the female are pale-yellow or +green, whilst the front legs of the male are reddish-brown: in _T. +floricolens_, the legs of the female are pale-green, those of the male +being ringed in a conspicuous manner with various tints. Numerous +analogous cases could be given in the genera Epeira, Nephila, +Philodromus, Theridion, Linyphia, &c. It is often difficult to say which +of the two sexes departs most from the ordinary coloration of the genus +to which the species belong; but Mr. Blackwall thinks that, as a +general rule, it is the male. Both sexes whilst young, as I am informed +by the same author, usually resemble each other; and both often undergo +great changes in colour during their successive moults before arriving +at maturity. In other cases the male alone appears to change colour. +Thus the male of the above-mentioned brightly-coloured Sparassus at +first resembles the female and acquires his peculiar tints only when +nearly adult. Spiders are possessed of acute senses, and exhibit much +intelligence. The females often shew, as is well known, the strongest +affection for their eggs, which they carry about enveloped in a silken +web. On the whole it appears probable that well-marked differences in +colour between the sexes have generally resulted from sexual selection, +either on the male or female side. But doubts may be entertained on this +head from the extreme variability in colour of some species, for +instance of _Theridion lineatum_, the sexes of which differ when adult; +this great variability indicates that their colours have not been +subjected to any form of selection. + +Mr. Blackwall does not remember to have seen the males of any species +fighting together for the possession of the female. Nor, judging from +analogy, is this probable; for the males are generally much smaller than +the females, sometimes to an extraordinary degree.[423] Had the males +been in the habit of fighting together, they would, it is probable, have +gradually acquired greater size and strength. Mr. Blackwall has +sometimes seen two or more males on the same web with a single female; +but their courtship is too tedious and prolonged an affair to be easily +observed. The male is extremely cautious in making his advances, as the +female carries her coyness to a dangerous pitch. De Geer saw a male that +"in the midst of his preparatory caresses was seized by the object of +his attentions, enveloped by her in a web and then devoured, a sight +which, as he adds, filled him with horror and indignation."[424] + +Westring has made the interesting discovery that the males of several +species of Theridion[425] have the power of making a stridulating sound +(like that made by many beetles and other insects, but feebler), whilst +the females are quite mute. The apparatus consists of a serrated ridge +at the base of the abdomen, against which the hard hinder part of the +thorax is rubbed; and of this structure not a trace could be detected in +the females. From the analogy of the Orthoptera and Homoptera, to be +described in the next chapter, we may feel almost sure that the +stridulation serves, as Westring remarks, either to call or to excite +the female; and this is the first case in the ascending scale of the +animal kingdom, known to me, of sounds emitted for this purpose. + + +Class, _Myriapoda_.--In neither of the two orders in this class, +including the millipedes and centipedes, can I find any well-marked +instances of sexual differences such as more particularly concern us. In +_Glomeris limbata_, however, and perhaps in some few other species, the +males differ slightly in colour from the females; but this Glomeris is a +highly variable species. In the males of the Diplopoda, the legs +belonging to one of the anterior segments of the body, or to the +posterior segment, are modified into prehensile hooks which serve to +secure the female. In some species of Iulus the tarsi of the male are +furnished with membranous suckers for the same purpose. It is a much +more unusual circumstance, as we shall see when we treat of Insects, +that it is the female in Lithobius which is furnished with prehensile +appendages at the extremity of the body for holding the male.[426] + + + + +CHAPTER X. + +SECONDARY SEXUAL CHARACTERS OF INSECTS. + + + Diversified structures possessed by the males for seizing the + females--Differences between the sexes, of which the meaning is + not understood--Difference in size between the + sexes--Thysanura--Diptera--Hemiptera--Homoptera, musical powers + possessed by the males alone--Orthoptera, musical instruments + of the males, much diversified in structure; pugnacity; + colours--Neuroptera, sexual differences in colour--Hymenoptera, + pugnacity and colours--Coleoptera, colours; furnished with + great horns, apparently as an ornament; battles; stridulating + organs generally common to both sexes. + + +In the immense class of insects the sexes sometimes differ in their +organs for locomotion, and often in their sense-organs, as in the +pectinated and beautifully plumose antennae of the males of many species. +In one of the Ephemerae, namely Chloeon, the male has great pillared +eyes, of which the female is entirely destitute.[427] The ocelli are +absent in the females of certain other insects, as in the Mutillidae, +which are likewise destitute of wings. But we are chiefly concerned with +structures by which one male is enabled to conquer another, either in +battle or courtship, through his strength, pugnacity, ornaments, or +music. The innumerable contrivances, therefore, by which the male is +able to seize the female, may be briefly passed over. Besides the +complex structures at the apex of the abdomen, which ought perhaps to be +ranked as primary organs,[428] "it is astonishing," as Mr. B. D. +Walsh[429] has remarked, "how many different organs are worked in by +nature, for the seemingly insignificant object of enabling the male to +grasp the female firmly." The mandibles or jaws are sometimes used for +this purpose; thus the male _Corydalis cornutus_ (a neuropterous insect +in some degree allied to the Dragon-flies, &c.) has immense curved jaws, +many times longer than those of the female; and they are smooth instead +of being toothed, by which means he is enabled to seize her without +injury.[430] One of the stag-beetles of North America (_Lucanus +elaphus_) uses his jaws, which are much larger than those of the female, +for the same purpose, but probably likewise for fighting. In one of the +sand-wasps (_Ammophila_) the jaws in the two sexes are closely alike, +but are used for widely different purposes; the males, as Professor +Westwood observes, "are exceedingly ardent, seizing their partners round +the neck with their sickle-shaped jaws;"[431] whilst the females use +these organs for burrowing in sand-banks and making their nests. + +[Illustration: Fig. 8. Crabro cribrarius. Upper figure, male: lower +figure, female.] + +The tarsi of the front-legs are dilated in many male beetles, or are +furnished with broad cushions of hairs; and in many genera of +water-beetles they are armed with a round flat sucker, so that the male +may adhere to the slippery body of the female. It is a much more unusual +circumstance that the females of some water-beetles (Dytiscus) have +their elytra deeply grooved, and in _Acilius sulcatus_ thickly set with +hairs, as an aid to the male. The females of some other water-beetles +(Hydroporus) have their elytra punctured for the same object.[432] In +the male of _Crabro cribrarius_ (fig. 8.), it is the tibia which is +dilated into a broad horny plate, with minute membraneous dots, giving +to it a singular appearance like that of a riddle.[433] In the male of +Penthe (a genus of beetles) a few of the middle joints of the antennae +are dilated and furnished on the inferior surface with cushions of +hair, exactly like those on the tarsi of the Carabidae, "and obviously +for the same end." In male dragon-flies, "the appendages at the tip of +the tail are modified in an almost infinite variety of curious patterns +to enable them to embrace the neck of the female." Lastly in the males +of many insects, the legs are furnished with peculiar spines, knobs or +spurs; or the whole leg is bowed or thickened, but this is by no means +invariably a sexual character; or one pair, or all three pairs are +elongated, sometimes to an extravagant length.[434] + +[Illustration: Fig. 9. Taphroderes distortus (much enlarged). Upper +figure, male; lower figure, female.] + +In all the orders, the sexes of many species present differences, of +which the meaning is not understood. One curious case is that of a +beetle (fig. 9), the male of which has the left mandible much enlarged; +so that the mouth is greatly distorted. In another Carabidous beetle, +the Eurygnathus,[435] we have the unique case, as far as known to Mr. +Wollaston, of the head of the female being much broader and larger, +though in a variable degree, than that of the male. Any number of such +cases could be given. They abound in the Lepidoptera: one of the most +extraordinary is that certain male butterflies have their fore-legs more +or less atrophied, with the tibiae and tarsi reduced to mere rudimentary +knobs. The wings, also, in the two sexes often differ in neuration,[436] +and sometimes considerably in outline, as in the _Aricoris epitus_, +which was shown to me in the British Museum by Mr. A. Butler. The males +of certain South American butterflies have tufts of hair on the margins +of the wings, and horny excrescences on the discs of the posterior +pair.[437] In several British butterflies, the males alone, as shewn by +Mr. Wonfor, are in parts clothed with peculiar scales. + +The purpose of the luminosity in the female glow-worm is likewise not +understood; for it is very doubtful whether the primary use of the light +is to guide the male to the female. It is no serious objection to this +latter belief that the males emit a feeble light; for secondary sexual +characters proper to one sex are often developed in a slight degree in +the other sex. It is a more valid objection that the larvae shine, and in +some species brilliantly: Fritz Mueller informs me that the most luminous +insect which he ever beheld in Brazil, was the larva of some beetle. +Both sexes of certain luminous species of Elater emit light. Kirby and +Spence suspect that the phosphorescence serves to frighten and drive +away enemies. + + +_Difference in Size between the Sexes._--With insects of all kinds the +males are commonly smaller than the females;[438] and this difference +can often be detected even in the larval state. So considerable is the +difference between the male and female cocoons of the silk-moth +(_Bombyx mori_), that in France they are separated by a particular mode +of weighing.[439] In the lower classes of the animal kingdom, the +greater size of the females seems generally to depend on their +developing an enormous number of ova; and this may to a certain extent +hold good with insects. But Dr. Wallace has suggested a much more +probable explanation. He finds, after carefully attending to the +development of the caterpillars of _Bombyx cynthia_ and _yamamai_, and +especially of some dwarfed caterpillars reared from a second brood on +unnatural food, "that in proportion as the individual moth is finer, so +is the time required for its metamorphosis longer; and for this reason +the female, which is the larger and heavier insect, from having to carry +her numerous eggs, will be preceded by the male, which is smaller and +has less to mature."[440] Now as most insects are short-lived, and as +they are exposed to many dangers, it would manifestly be advantageous to +the female to be impregnated as soon as possible. This end would be +gained by the males being first matured in large numbers ready for the +advent of the females; and this again would naturally follow, as Mr. A. +E. Wallace has remarked,[441] through natural selection; for the smaller +males would be first matured, and thus would procreate a large number of +offspring which would inherit the reduced size of their male parents, +whilst the larger males from being matured later would leave fewer +offspring. + +There are, however, exceptions to the rule of male insects being smaller +than the females; and some of these exceptions are intelligible. Size +and strength would be an advantage to the males, which fight for the +possession of the female; and in these cases the males, as with the +stag-beetle (Lucanus), are larger than the females. There are, however, +other beetles which are not known to fight together, of which the males +exceed the females in size; and the meaning of this fact is not known; +but in some of these cases, as with the huge Dynastes and Megasoma, we +can at least see that there would be no necessity for the males to be +smaller than the females, in order to be matured before them, for these +beetles are not short-lived, and there would be ample time for the +pairing of the sexes. So, again, male dragon-flies (Libellulidae) are +sometimes sensibly larger, and never smaller, than the females;[442] and +they do not, as Mr. MacLachlan believes, generally pair with the +females, until a week or fortnight has elapsed, and until they have +assumed their proper masculine colours. But the most curious case, +shewing on what complex and easily-overlooked relations, so trifling a +character as a difference in size between the sexes may depend, is that +of the aculeate Hymenoptera; for Mr. F. Smith informs me that throughout +nearly the whole of this large group the males, in accordance with the +general rule, are smaller than the females and emerge about a week +before them; but amongst the Bees, the males of _Apis mellifica_, +_Anthidium manicatum_ and _Anthophora acervorum_, and amongst the +Fossores, the males of the _Methoca ichneumonides_, are larger than the +females. The explanation of this anomaly is that a marriage-flight is +absolutely necessary with these species, and the males require great +strength and size in order to carry the females through the air. +Increased size has here been acquired in opposition to the usual +relation between size and the period of development, for the males, +though larger, emerge before the smaller females. + +We will now review the several Orders, selecting such facts as more +particularly concern us. The Lepidoptera (Butterflies and Moths) will be +retained for a separate chapter. + + +Order, _Thysanura_.--The members of this Order are lowly organised for +their class. They are wingless, dull-coloured, minute insects, with +ugly, almost misshapen heads and bodies. The sexes do not differ; but +they offer one interesting fact, by showing that the males pay sedulous +court to their females even low down in the animal scale. Sir J. +Lubbock[443] in describing the _Smynthurus luteus_, says: "it is very +amusing to see these little creatures coquetting together. The male, +which is much smaller than the female, runs round her, and they butt one +another, standing face to face, and moving backward and forward like two +playful lambs. Then the female pretends to run away and the male runs +after her with a queer appearance of anger, gets in front and stands +facing her again; then she turns coyly round, but he, quicker and more +active, scuttles round too, and seems to whip her with his antennae; then +for a bit they stand face to face, play with their antennae, and seem to +be all in all to one another." + + +Order, _Diptera_ (Flies).--The sexes differ little in colour. The +greatest difference, known to Mr. F. Walker, is in the genus Bibio, in +which the males are blackish or quite black, and the females obscure +brownish-orange. The genus Elaphomyia, discovered by Mr. Wallace[444] in +New Guinea, is highly remarkable, as the males are furnished with horns, +of which the females are quite destitute. The horns spring from beneath +the eyes, and curiously resemble those of stags, being either branched +or palmated. They equal in length the whole of the body in one of the +species. They might be thought to serve for fighting, but as in one +species they are of a beautiful pink colour, edged with black, with a +pale central stripe, and as these insects have altogether a very elegant +appearance, it is perhaps more probable that the horns serve as +ornaments. That the males of some Diptera fight together is certain; for +Prof. Westwood[445] has several times seen this with some species of +Tipula or Harry-long-legs. Many observers believe that when gnats +(Culicidae) dance in the air in a body, alternately rising and falling, +the males are courting the females. The mental faculties of the Diptera +are probably fairly well developed, for their nervous system is more +highly developed than in most other Orders of insects.[446] + + +Order, _Hemiptera_ (Field-Bugs).--Mr. J. W. Douglas, who has +particularly attended to the British species, has kindly given me an +account of their sexual differences. The males of some species are +furnished with wings, whilst the females are wingless; the sexes differ +in the form of the body and elytra; in the second joints of their +antennae and in their tarsi; but as the signification of these +differences is quite unknown, they may be here passed over. The females +are generally larger and more robust than the males. With British, and, +as far as Mr. Douglas knows, with exotic species, the sexes do not +commonly differ much in colour; but in about six British species the +male is considerably darker than the female, and in about four other +species the female is darker than the male. Both sexes of some species +are beautifully marked with vermilion and black. It is doubtful whether +these colours serve as a protection. If in any species the males had +differed from the females in an analogous manner, we might have been +justified in attributing such conspicuous colours to sexual selection +with transference to both sexes. + +Some species of Reduvidae make a stridulating noise; and, in the case of +_Pirates stridulus_, this is said[447] to be effected by the movement of +the neck within the pro-thoracic cavity. According to Westring, +_Reduvius personatus_ also stridulates. But I have not been able to +learn any particulars about these insects; nor have I any reason to +suppose that they differ sexually in this respect. + + +Order, _Homoptera._--Every one who has wandered in a tropical forest +must have been astonished at the din made by the male Cicadae. The +females are mute; as the Grecian poet Xenarchus says, "Happy the Cicadas +live, since they all have voiceless wives." The noise thus made could be +plainly heard on board the "Beagle," when anchored at a quarter of a +mile from the shore of Brazil; and Captain Hancock says it can be heard +at the distance of a mile. The Greeks formerly kept, and the Chinese now +keep, these insects in cages for the sake of their song, so that it +must be pleasing to the ears of some men.[448] The Cicadidae usually sing +during the day; whilst the Fulgoridae appear to be night-songsters. The +sound, according to Landois,[449] who has recently studied the subject, +is produced by the vibration of the lips of the spiracles, which are set +into motion by a current of air emitted from the tracheae. It is +increased by a wonderfully complex resounding apparatus, consisting of +two cavities covered by scales. Hence the sound may truly be called a +voice. In the female the musical apparatus is present, but very much +less developed than in the male, and is never used for producing sound. + +With respect to the object of the music, Dr. Hartman in speaking of the +_Cicada septemdecim_ of the United States, says,[450] "the drums are now +(June 6th and 7th, 1851) heard in all directions. This I believe to be +the marital summons from the males. Standing in thick chestnut sprouts +about as high as my head, where hundreds were around me, I observed the +females coming around the drumming males." He adds, "this season (Aug. +1868) a dwarf pear-tree in my garden produced about fifty larvae of _Cic. +pruinosa_; and I several times noticed the females to alight near a male +while he was uttering his clanging notes." Fritz Mueller writes to me +from S. Brazil that he has often listened to a musical contest between +two or three males of a Cicada, having a particularly loud voice, and +seated at a considerable distance from each other. As soon as the first +had finished his song, a second immediately began; and after he had +concluded, another began, and so on. As there is so much rivalry between +the males, it is probable that the females not only discover them by the +sounds emitted, but that, like female birds, they are excited or allured +by the male with the most attractive voice. + +I have not found any well-marked cases of ornamental differences between +the sexes of the Homoptera. Mr. Douglas informs me that there are three +British species, in which the male is black or marked with black bands, +whilst the females are pale-coloured or obscure. + + +Order, _Orthoptera_.--The males in the three saltatorial families +belonging to this Order are remarkable for their musical powers, namely +the Achetidae or crickets, the Locustidae for which there is no exact +equivalent name in English, and the Acridiidae or grasshoppers. The +stridulation produced by some of the Locustidae is so loud that it can be +heard during the night at the distance of a mile;[451] and that made by +certain species is not unmusical even to the human ear, so that the +Indians on the Amazons keep them in wicker cages. All observers agree +that the sounds serve either to call or excite the mute females. But it +has been noticed[452] that the male migratory locust of Russia (one of +the Acridiidae) whilst coupled with the female, stridulates from anger or +jealousy when approached by another male. The house-cricket when +surprised at night uses its voice to warn its fellows.[453] In North +America the Katy-did (_Platyphyllum concavum_, one of the Locustidae) is +described[454] as mounting on the upper branches of a tree, and in the +evening beginning "his noisy babble, while rival notes issue from the +neighbouring trees, and the groves resound with the call of +_Katy-did-she-did_, the live-long night." Mr. Bates, in speaking of the +European field-cricket (one of the Achetidae), says, "the male has been +observed to place itself in the evening at the entrance of its burrow, +and stridulate until a female approaches, when the louder notes are +succeeded by a more subdued tone, whilst the successful musician +caresses with his antennae the mate he has won."[455] Dr. Scudder was +able to excite one of these insects to answer him, by rubbing on a file +with a quill.[456] In both sexes a remarkable auditory apparatus has +been discovered by Von Siebold, situated in the front legs.[457] + +[Illustration: Fig. 10. Gryllus campestris (from Landois). + +Right-hand figure, under side of part of the wing-nervure, much +magnified, showing the teeth, _st_. + +Left-hand figure, upper surface of wing-cover, with the projecting, +smooth nervure, _r._, across which the teeth (_st_) are scraped.] + +In the three Families the sounds are differently produced. In the males +Of the Achetidae both wing-covers have the same structure; and this in +the field-cricket (_Gryllus campestris_, fig. 10) consists, as +described by Landois,[458] of from 131 to 138 sharp, transverse ridges +or teeth (_st_) on the under side of one of the nervures of the +wing-cover. This toothed nervure is rapidly scraped across a projecting, +smooth, hard nervure (_r_) on the upper surface of the opposite wing. +First one wing is rubbed over the other, and then the movement is +reversed. Both wings are raised a little at the same time, so as to +increase the resonance. In some species the wing-covers of the males are +furnished at the base with a talc-like plate.[459] I have here given a +drawing (fig. 11) of the teeth on the under side of the nervure of +another species of Gryllus, viz. _G. domesticus_. + +[Illustration: Fig. 11. Teeth of Nervure of Gryllus domesticus (from +Landois).] + +In the Locustidae the opposite wing-covers differ in structure (fig. 12), +and cannot, as in the last family, be indifferently used in a reversed +manner. The left wing, which acts as the bow of the fiddle, lies over +the right wing which serves as the fiddle itself. One of the nervures +(_a_) on the under surface of the former is finely serrated, and is +scraped across the prominent nervures on the upper surface of the +opposite or right wing. In our British _Phasgonura viridissima_ it +appeared to me that the serrated nervure is rubbed against the rounded +hind corner of the opposite wing, the edge of which is thickened, +coloured brown, and very sharp. In the right wing, but not in the left, +there is a little plate, as transparent as talc, surrounded by nervures, +and called the speculum. In _Ephippiger vitium_, a member of this same +family, we have a curious subordinate modification; for the wing-covers +are greatly reduced in size, but "the posterior part of the pro-thorax +is elevated into a kind of dome over the wing-covers, and which has +probably the effect of increasing the sound."[460] + +[Illustration: Fig. 12. Chlorocoelus Tanana (from Bates). _a_, _b_. +Lobes of opposite wing-covers.] + +We thus see that the musical apparatus is more differentiated or +specialised in the Locustidae, which includes I believe the most powerful +performers in the Order, than in the Achetidae, in which both wing-covers +have the same structure and the same function.[461] Landois, however, +detected in one of the Locustidae, namely in Decticus, a short and narrow +row of small teeth, mere rudiments, on the inferior surface of the +right wing-cover, which underlies the other and is never used as the +bow. I observed the same rudimentary structure on the under side of the +right wing-cover in _Phasgonura viridissima_. Hence we may with +confidence infer that the Locustidae are descended from a form, in which, +as in the existing Achetidae, both wing-covers had serrated nervures on +the under surface, and could be indifferently used as the bow; but that +in the Locustidae the two wing-covers gradually became differentiated and +perfected, on the principle of the division of labour, the one to act +exclusively as the bow and the other as the fiddle. By what steps the +more simple apparatus in the Achetidae originated, we do not know, but it +is probable that the basal portions of the wing-covers overlapped each +other formerly as at present, and that the friction of the nervures +produced a grating sound, as I find is now the case with the wing-covers +of the females.[462] A grating sound thus occasionally and accidentally +made by the males, if it served them ever so little as a love-call to +the females, might readily have been intensified through sexual +selection by fitting variations in the roughness of the nervures having +been continually preserved. + +[Illustration: Fig. 13, Hind-leg of Stenobothrus pratorum: _r_, the +stridulating ridge; lower figure, the teeth, forming the ridge, much +magnified (from Landois).] + +In the last and third Family, namely the Acridiidae or grasshoppers, the +stridulation is produced in a very different manner, and is not so +shrill, according to Dr. Scudder, as in the preceding Families. The +inner surface of the femur (fig. 13, _r_) is furnished with a +longitudinal row of minute, elegant, lancet-shaped, elastic teeth, from +85 to 93 in number;[463] and these are scraped across the sharp, +projecting nervures on the wing-covers, which, are thus made to vibrate +and resound. Harris[464] says that when one of the males begins to play, +he first "bends the shank of the hind-leg beneath, the thigh, where it +is lodged in a furrow designed to receive it, and then draws the leg +briskly up and down. He does not play both fiddles together, but +alternately first upon one and then on the other." In many species, the +base of the abdomen is hollowed out into a great cavity which is +believed to act as a resounding board. In Pneumora (fig. 14), a S. +African genus belonging to this same family, we meet with a new and +remarkable modification: in the males a small notched ridge projects +obliquely from each side of the abdomen, against which the hind femora +are rubbed.[465] As the male is furnished with wings, the female being +wingless, it is remarkable that the thighs are not rubbed in the usual +manner against the wing-covers; but this may perhaps be accounted for by +the unusually small size of the hind-legs. I have not been able to +examine the inner surface of the thighs, which, judging from analogy, +would be finely serrated. The species of Pneumora have been more +profoundly modified for the sake of stridulation than any other +orthopterous insect; for in the male the whole body has been converted +into a musical instrument, being distended with air, like a great +pellucid bladder, so as to increase the resonance. Mr. Trimen informs me +that at the Cape of Good Hope these insects make a wonderful noise +during the night There is one exception to the rule that the females in +these three Families are destitute of an efficient musical apparatus; +for both sexes of Ephippiger (Locustidae) are said[466] to be thus +provided. This case may be compared with that of the reindeer, in which +species alone both sexes possess horns. Although the female orthoptera +are thus almost invariably mute, yet Landois[467] found rudiments of the +stridulating organs on the femora of the female Acridiidae, and similar +rudiments on the under surface of the wing-covers of the female +Achetidae; but he failed to find any rudiments in the females of +Decticus, one of the Locustidae. In the Homoptera the mute females of +Cicada, have the proper musical apparatus in an undeveloped state; and +we shall hereafter meet in other divisions of the animal kingdom with +innumerable instances of structures proper to the male being present in +a rudimentary condition in the female. Such cases appear at first sight +to indicate that both sexes were primordially constructed in the same +manner, but that certain organs were subsequently lost by the females. +It is, however, a more probable view, as previously explained, that the +organs in question were acquired by the males and partially transferred +to the females. + +[Illustration: Fig. 14. Pneumora (from specimens in the British Museum). +Upper figure, male; lower figure, female.] + +Landois has observed another interesting fact, namely that in the +females of the Acridiidae, the stridulating teeth on the femora remain +throughout life in the same condition in which they first appear in both +sexes during the larval state. In the males, on the other hand, they +become fully developed and acquire their perfect structure at the last +moult, when the insect is mature and ready to breed. + +From the facts now given, we see that the means by which the males +produce their sounds are extremely diversified in the Orthoptera, and +are altogether different from those employed by the Homoptera. But +throughout the animal kingdom we incessantly find the same object +gained by the most diversified means; this being due to the whole +organisation undergoing in the course of ages multifarious changes; and +as part after part varies, different variations are taken advantage of +for the same general purpose. The diversification of the means for +producing sound in the three families of the Orthoptera and in the +Homoptera, impresses the mind with the high importance of these +structures to the males, for the sake of calling or alluring the +females. We need feel no surprise at the amount of modification which +the Orthoptera have undergone in this respect, as we now know, from Dr. +Scudder's remarkable discovery,[468] that there has been more than ample +time. This naturalist has lately found a fossil insect in the Devonian +formation of New Brunswick, which is furnished with "the well-known +tympanum or stridulating apparatus of the male Locustidae." This insect, +though in most respects related to the Neuroptera, appears to connect, +as is so often the case with very ancient forms, the two Orders of the +Neuroptera and Orthoptera which are now generally ranked as quite +distinct. + +I have but little more to say on the Orthoptera. Some of the species are +very pugnacious: when two male field-crickets (_Gryllus campestris_) are +confined together, they fight till one kills the other; and the species +of Mantis are described as manoeuvring with their sword-like +front-limbs, like hussars with their sabres. The Chinese keep these +insects in little bamboo cages and match them like game-cocks.[469] With +respect to colour, some exotic locusts are beautifully ornamented; the +posterior wings being marked with red, blue, and black; but as +throughout the Order the two sexes rarely differ much in colour, it is +doubtful whether they owe these bright tints to sexual selection. +Conspicuous colours may be of use to these insects as a protection, on +the principle to be explained in the next chapter, by giving notice to +their enemies that they are unpalatable. Thus it has been observed[470] +that an Indian brightly-coloured locust was invariably rejected when +offered to birds and lizards. Some cases, however, of sexual differences +in colour in this Order are known. The male of an American cricket[471] +is described as being as white as ivory, whilst the female varies from +almost white to greenish-yellow or dusky. Mr. Walsh informs me that the +adult male of _Spectrum femoratum_ (one of the Phasmidae) "is of a +shining brownish-yellow colour; the adult female being of a dull, +opaque, cinereous-brown; the young of both sexes being green." Lastly, I +may mention that the male of one curious kind of cricket[472] is +furnished with "a long membranous appendage, which falls over the face +like a veil;" but whether this serves as an ornament is not known. + + +Order, _Neuroptera_.--Little need here be said, except in regard to +colour. In the Ephemeridae the sexes often differ slightly in their +obscure tints;[473] but it is not probable that the males are thus +rendered attractive to the females. The Libellulidae or dragon-flies are +ornamented with splendid green, blue, yellow, and vermilion metallic +tints; and the sexes often differ. Thus, the males of some of the +Agrionidae, as Prof. Westwood remarks[474] "are of a rich blue with black +wings, whilst the females are fine green with colourless wings." But in +_Agrion Ramburii_ these colours are exactly reversed in the two +sexes.[475] In the extensive N. American genus of Hetaerina, the males +alone have a beautiful carmine spot at the base of each wing. In _Anax +junius_ the basal part of the abdomen in the male is a vivid +ultra-marine blue, and in the female grass-green. In the allied genus +Gomphus, on the other hand, and in some other genera, the sexes differ +but little in colour. Throughout the animal kingdom, similar cases of +the sexes of closely-allied forms either differing greatly, or very +little, or not at all, are of frequent occurrence. Although with many +Libellulidae there is so wide a difference in colour between the sexes, +it is often difficult to say which is the most brilliant; and the +ordinary coloration of the two sexes is exactly reversed, as we have +just seen, in one species of Agrion. It is not probable that their +colours in any case have been gained as a protection. As Mr. MacLachlan, +who has closely attended to this family, writes to me, dragon-flies--the +tyrants of the insect-world--are the least liable of any insect to be +attacked by birds or other enemies. He believes that their bright +colours serve as a sexual attraction. It deserves notice, as bearing on +this subject, that certain dragon-flies appear to be attracted by +particular colours: Mr. Patterson observed[476] that the species of +Agrionidae, of which the males are blue, settled in numbers on the blue +float of a fishing line; whilst two other species were attracted by +shining white colours. + +It is an interesting fact, first observed by Schelver, that the males, +in several genera belonging to two sub-families, when they first emerge +from the pupal state are coloured exactly like the females; but that +their bodies in a short time assume a conspicuous milky-blue tint, owing +to the exudation of a kind of oil, soluble in ether and alcohol. Mr. +MacLachlan believes that in the male of _Libellula depressa_ this change +of colour does not occur until nearly a fortnight after the +metamorphosis, when the sexes are ready to pair. + +Certain species of Neurothemis present, according to Brauer[477] a +curious case of dimorphism, some of the females having their wings +netted in the usual manner; whilst other females have them "very richly +netted as in the males of the same species." Brauer "explains the +phenomenon on Darwinian principles by the supposition that the close +netting of the veins is a secondary sexual character in the males." This +latter character is generally developed in the males alone, but being, +like every other masculine character, latent in the female, is +occasionally developed in them. We have here an illustration of the +manner in which the two sexes of many animals have probably come to +resemble each other, namely by variations first appearing in the males, +being preserved in them, and then transmitted to and developed in the +females; but in this particular genus a complete transference is +occasionally and abruptly effected. Mr. MacLachlan informs me of another +case of dimorphism occurring in several species of Agrion in which a +certain number of individuals are found of an orange colour, and these +are invariably females. This is probably a case of reversion, for in +the true Libellulae, when the sexes differ in colour, the females are +always orange or yellow, so that supposing Agrion to be descended from +some primordial form having the characteristic sexual colours of the +typical Libellulae, it would not be surprising that a tendency to vary in +this manner should occur in the females alone. + +Although many dragon-flies are such large, powerful, and fierce insects, +the males have not been observed by Mr. MacLachlan to fight together, +except, as he believes, in the case of some of the smaller species of +Agrion. In another very distinct group in this Order, namely in the +Termites or white ants, both sexes at the time of swarming may be seen +running about, "the male after the female, sometimes two chasing one +female, and contending with great eagerness who shall win the +prize."[478] + + +Order, _Hymenoptera_.--That inimitable observer, M. Fabre,[479] in +describing the habits of Cerceris, a wasp-like insect, remarks that +"fights frequently ensue between the males for the possession of some +particular female, who sits an apparently unconcerned beholder of the +struggle for supremacy, and when the victory is decided, quietly flies +away in company with the conqueror." Westwood[480] says that the males +of one of the saw-flies (Tenthredinae) "have been found fighting +together, with their mandibles locked." As M. Fabre speaks of the males +of Cerceris striving to obtain a particular female, it may be well to +bear in mind that insects belonging to this Order have the power of +recognising each other after long intervals of time, and are deeply +attached. For instance, Pierre Huber, whose accuracy no one doubts, +separated some ants, and when after an interval of four months they met +others which had formerly belonged to the same community, they mutually +recognised and caressed each other with their antennae. Had they been +strangers they would have fought together. Again, when two communities +engage in a battle, the ants on the same side in the general confusion +sometimes attack each other, but they soon perceive their mistake, and +the one ant soothes the other.[481] + +In this Order slight differences in colour, according to sex, are +common, but conspicuous differences are rare except in the family of +Bees; yet both sexes of certain groups are so brilliantly coloured--for +instance in Chrysis, in which vermilion and metallic greens +prevail--that we are tempted to attribute the result to sexual +selection. In the Ichneumonidae, according to Mr. Walsh,[482] the males +are almost universally lighter coloured than the females. On the other +hand, in the Tenthredinidae the males are generally darker than the +females. In the Siricidae the sexes frequently differ; thus the male of +_Sirex juvencus_ is banded with orange, whilst the female is dark +purple; but it is difficult to say which sex is the most ornamented. In +_Tremex columbae_ the female is much brighter coloured than the male. +With ants, as I am informed by Mr. F. Smith, the males of several +species are black, the females being testaceous. In the family of Bees, +especially in the solitary species, as I hear from the same +distinguished entomologist, the sexes often differ in colour. The males +are generally the brightest, and in Bombus as well as in Apathus, much +more variable in colour than the females. In _Anthophora retusa_ the +male is of a rich fulvous-brown, whilst the female is quite black: so +are the females of several species of Xylocopa, the males being bright +yellow. In an Australian bee (_Lestis bombylans_), the female is of an +extremely brilliant steel-blue, sometimes tinted with vivid green; the +male being of a bright brassy colour clothed with rich fulvous +pubescence. As in this group the females are provided with excellent +defensive weapons in their stings, it is not probable that they have +come to differ in colour from the males for the sake of protection. + +_Mutilla Europaea_ emits a stridulating noise; and according to +Goureau[483] both sexes have this power. He attributes the sound to the +friction of the third and preceding abdominal segments; and I find that +these surfaces are marked with very fine concentric ridges, but so is +the projecting thoracic collar, on which the head articulates; and this +collar, when scratched with the point of a needle, emits the proper +sound. It is rather surprising that both sexes should have the power of +stridulating, as the male is winged and the female wingless. It is +notorious that Bees express certain emotions, as of anger, by the tone +of their humming, as do some dipterous insects; but I have not referred +to these sounds, as they are not known to be in any way connected with +the act of courtship. + + +Order, _Coleoptera_ (Beetles).--Many beetles are coloured so as to +resemble the surfaces which they habitually frequent. Other species are +ornamented with gorgeous metallic tints,--for instance, many Carabidae, +which live on the ground and have the power of defending themselves by +an intensely acrid secretion,--the splendid diamond-beetles which are +protected by an extremely hard covering,--many species of Chrysomela, +such as _C. cerealis_, a large species beautifully striped with various +colours, and in Britain confined to the bare summit of Snowdon,--and a +host of other species. These splendid colours, which are often arranged +in stripes, spots, crosses and other elegant patterns, can hardly be +beneficial, as a protection, except in the case of some flower-feeding +species; and we cannot believe that they are purposeless. Hence the +suspicion arises, that they serve as a sexual attraction; but we have no +evidence on this head, for the sexes rarely differ in colour. Blind +beetles, which cannot of course behold each other's beauty, never +exhibit, as I hear from Mr. Waterhouse, jun., bright colours, though +they often have polished coats: but the explanation of their obscurity +may be that blind insects inhabit caves and other obscure stations. + +Some Longicorns, however, especially certain Prionidae, offer an +exception to the common rule that the sexes of beetles do not differ in +colour. Most of these insects are large and splendidly coloured. The +males in the genus Pyrodes,[484] as I saw in Mr. Bates' collection, are +generally redder but rather duller than the females, the latter being +coloured of a more or less splendid golden green. On the other hand, in +one species the male is golden-green, the female being richly tinted +with red and purple. In the genus Esmeralda the sexes differ so greatly +in colour that they have been ranked as distinct species: in one species +both are of a beautiful shining green, but the male has a red thorax. On +the whole, as far as I could judge, the females of those Prionidae, in +which the sexes differ, are coloured more richly than the males; and +this does not accord with the common rule in regard to colour when +acquired through sexual selection. + +[Illustration: Fig. 15. Chalcosoma atlas. Upper figure, male (reduced); +lower figure, female (nat. size).] + +[Illustration: Fig. 16. Copris isidis. (Left-hand figures, males.)] + +[Illustration: Fig. 17. Phanaeus faunus.] + +[Illustration: Fig. 18. Dipelicus cantori.] + +[Illustration: Fig. 19. Onthophagus rangifer, enlarged.] + +A most remarkable distinction between the sexes of many beetles is +presented by the great horns which rise from the head, thorax, or +clypeus of the males; and in some few cases from the under surface of +the body. These horns, in the great family of the Lamellicorns, resemble +those of various quadrupeds, such as stags, rhinoceroses, &c., and are +wonderful both from their size and diversified shapes. Instead of +describing them, I have given figures of the males and females of some +of the more remarkable forms. (Figs. 15 to 19.) The females generally +exhibit rudiments of the horns in the form of small knobs or ridges; but +some are destitute of even a rudiment. On the other hand, the horns are +nearly as well developed in the female as in the male of _Phanaeus +lancifer_; and only a little less well developed in the females of some +other species of the same genus and of Copris. In the several +subdivisions of the family, the differences in structure of the horns do +not run parallel, as I am informed by Mr. Bates, with their more +important and characteristic differences; thus within the same natural +section of the genus Onthophagus, there are species which have either a +single cephalic horn, or two distinct horns. + +In almost all cases, the horns are remarkable from their excessive +variability; so that a graduated series can be formed, from the most +highly developed males to others so degenerate that they can barely be +distinguished from the females. Mr. Walsh[485] found that in _Phanaeus +carnifex_ the horns were thrice as long in some males as in others. Mr. +Bates, after examining above a hundred males of _Onthophagus rangifer_ +(fig. 19), thought that he had at last discovered a species in which +the horns did not vary; but further research proved the contrary. + +The extraordinary size of the horns, and their widely different +structure in closely-allied forms, indicate that they have been formed +for some important purpose; but their excessive variability in the males +of the same species leads to the inference that this purpose cannot be +of a definite nature. The horns do not show marks of friction, as if +used for any ordinary work. Some authors suppose[486] that as the males +wander much more than the females, they require horns as a defence +against their enemies; but in many cases the horns do not seem well +adapted for defence, as they are not sharp. The most obvious conjecture +is that they are used by the males for fighting together; but they have +never been observed to fight; nor could Mr. Bates, after a careful +examination of numerous species, find any sufficient evidence in their +mutilated or broken condition of their having been thus used. If the +males had been habitual fighters, their size would probably have been +increased through sexual selection, so as to have exceeded that of the +female; but Mr. Bates, after comparing the two sexes in above a hundred +species of the Copridae, does not find in well-developed individuals any +marked difference in this respect. There is, moreover, one beetle, +belonging to the same great division of the Lamellicorns, namely +Lethrus, the males of which are known to fight, but they are not +provided with horns, though their mandibles are much larger than those +of the female. + +The conclusion, which best agrees with the fact of the horns having been +so immensely yet not fixedly developed,--as shewn by their extreme +variability in the same species and by their extreme diversity in +closely-allied species--is that they have been acquired as ornaments. +This view will at first appear extremely improbable; but we shall +hereafter find with many animals, standing much higher in the scale, +namely fishes, amphibians, reptiles and birds, that various kinds of +crests, knobs, horns and combs have been developed apparently for this +sole purpose. + +[Illustration: Fig. 20. Onitis furcifer, male, viewed from beneath.] + +[Illustration: Fig. 21. Left-hand figure, male of Onitis furcifer, +viewed laterally. Right-hand figure, female. _a._ Rudiment of cephalic +horn. _b._ Trace of thoracic horn or crest.] + +The males of _Onitis furcifer_ (fig. 20) are furnished with singular +projections on their anterior femora, and with a great fork or pair of +horns on the lower surface of the thorax. This situation seems extremely +ill adapted for the display of these projections, and they may be of +some real service; but no use can at present be assigned to them. It is +a highly remarkable fact, that although the males do not exhibit even a +trace of horns on the upper surface of the body, yet in the females a +rudiment of a single horn on the head (fig. 21, _a_), and of a crest +(_b_) on the thorax, are plainly visible. That the slight thoracic crest +in the female is a rudiment of a projection proper to the male, though +entirely absent in the male of this particular species, is clear: for +the female of _Bubas bison_ (a form which comes next to _Onitis_) has a +similar slight crest on the thorax, and the male has in the same +situation a great projection. So again there can be no doubt that the +little point (_a_) on the head of the female _Onitis furcifer_, as well +of the females of two or three allied species, is a rudimentary +representative of the cephalic horn, which is common to the males of so +many lamellicorn beetles, as in Phanaeus, fig. 17. The males indeed of +some unnamed beetles in the British Museum, which are believed actually +to belong to the genus Onitis, are furnished with a similar horn. The +remarkable nature of this case will be best perceived by an +illustration: the Ruminant quadrupeds run parallel with the lamellicorn +beetles, in some females possessing horns as large as those of the male, +in others having them much smaller, or existing as mere rudiments +(though this is as rare with ruminants as it is common with +Lamellicorns), or in having none at all. Now if a new species of deer or +sheep were discovered with the female bearing distinct rudiments of +horns, whilst the head of the male was absolutely smooth, we should have +a case like that of _Onitis furcifer_. + +In this case the old belief of rudiments having been created to complete +the scheme of nature is so far from holding good, that all ordinary +rules are completely broken through. The view which seems the most +probable is that some early progenitor of Onitis acquired, like other +Lamellicorns, horns on the head and thorax, and then transferred them, +in a rudimentary condition, as with so many existing species, to the +female, by whom they have ever since been retained. The subsequent loss +of the horns by the male may have resulted through the principle of +compensation from the development of the projections on the lower +surface, whilst the female has not been thus affected, as she is not +furnished with these projections, and consequently has retained the +rudiments of the horns on the upper surface. Although this view is +supported by the case of Bledius immediately to be given, yet the +projections on the lower surface differ greatly in structure and +development in the males of the several species of Onitis, and are even +rudimentary in some; nevertheless the upper surface in all these species +is quite destitute of horns. As secondary sexual characters are so +eminently variable, it is possible that the projections on the lower +surface may have been first acquired by some progenitor of Onitis and +produced their effect through compensation, and then have been in +certain cases almost completely lost. + +[Illustration: Fig. 22. Bledius taurus, magnified. Left-hand figure, +male; right-hand figure, female.] + +All the cases hitherto given refer to the Lamellicorns, but the males of +some few other beetles, belonging to two widely distinct groups, namely, +the Curculionidae and Staphylinidae, are furnished with horns,--in the +former on the lower surface of the body,[487] in the latter on the upper +surface of the head and thorax. In the Staphylinidae the horns of the +males in the same species are extraordinarily variable, just as we have +seen with the Lamellicorns. In Siagonium we have a case of dimorphism, +for the males can be divided into two sets, differing greatly in the +size of their bodies, and in the development of their horns, without any +intermediate gradations. In a species of Bledius (fig. 22), also +belonging to the Staphylinidae, male specimens can be found in the same +locality, as Professor Westwood states, "in which the central horn of +the thorax is very large, but the horns of the head quite rudimental; +and others, in which the thoracic horn is much shorter, whilst the +protuberances on the head are long."[488] Here, then, we apparently have +an instance of compensation of growth, which throws light on the curious +case just given of the loss of the upper horns by the males of _Onitis +furcifer_. + + +_Law of Battle._--Some male beetles, which seem ill fitted for fighting, +nevertheless engage in conflicts for the possession of the females. Mr. +Wallace[489] saw two males of _Leptorhynchus angustatus_, a linear +beetle with a much elongated rostrum, "fighting for a female, who stood +close by busy at her boring. They pushed at each other with their +rostra, and clawed and thumped, apparently in the greatest rage." The +smaller male, however, "soon ran away, acknowledging himself +vanquished." In some few cases the males are well adapted for fighting, +by possessing great toothed mandibles, much larger than those of the +females. This is the case with the common stag-beetle (_Lucanus +cervus_), the males of which emerge from the pupal state about a week +before the other sex, so that several may often be seen pursuing the +same female. At this period they engage in fierce conflicts. When Mr. A. +H. Davis[490] enclosed two males with one female in a box, the larger +male severely pinched the smaller one, until he resigned his +pretensions. A friend informs me that when a boy he often put the males +together to see them fight, and he noticed that they were much bolder +and fiercer than the females, as is well known to be the case with the +higher animals. The males would seize hold of his finger, if held in +front, but not so the females. With many of the Lucanidae, as well as +with the above-mentioned Leptorhynchus, the males are larger and more +powerful insects than the females. The two sexes of _Lethrus cephalotes_ +(one of the Lamellicorns) inhabit the same burrow; and the male has +larger mandibles than the female. If, during the breeding-season, a +strange male attempts to enter the burrow, he is attacked; the female +does not remain passive, but closes the mouth of the burrow, and +encourages her mate by continually pushing him on from behind. The +action does not cease until the aggressor is killed or runs away.[491] +The two sexes of another lamellicorn beetle, the _Ateuchus cicatricosus_ +live in pairs, and seem much attached to each other; the male excites +the female to roll the balls of dung in which the ova are deposited; and +if she is removed, he becomes much agitated. If the male is removed, the +female ceases all work, and as M. Brulerie[492] believes, would remain +on the spot until she died. + +The great mandibles of the male Lucanidae are extremely variable both in +size and structure, and in this respect resemble the horns on the head +and thorax of many male Lamellicorns and Staphylinidae. A perfect series +can be formed from the best-provided to the worst-provided or degenerate +males. Although the mandibles of the common stag-beetle, and probably of +many other species, are used as efficient weapons for fighting, it is +doubtful whether their great size can thus be accounted for. We have +seen that with the _Lucanus elaphus_ of N. America they are used for +seizing the female. As they are so conspicuous and so elegantly +branched, the suspicion has sometimes crossed my mind that they may be +serviceable to the males as an ornament, in the same manner as the horns +on the head and thorax of the various above described species. The male +_Chiasognathus grantii_ of S. Chile--a splendid beetle belonging to the +same family--has enormously-developed mandibles (fig. 23); he is bold +and pugnacious; when threatened on any side he faces round, opening his +great jaws, and at the same time stridulating loudly; but the mandibles +were not strong enough to pinch my finger so as to cause actual pain. + +[Illustration: Fig. 23. Chiasognathus grantii, reduced. Upper figure, +male; lower figure, female.] + +Sexual selection, which implies the possession of considerable +perceptive powers and of strong passions, seems to have been more +effective with the Lamellicorns than with any other family of the +Coleoptera or beetles. With some species the males are provided with +weapons for fighting; some live in pairs and show mutual affection; +many have the power of stridulating when excited; many are furnished +with the most extraordinary horns, apparently for the sake of ornament; +some which are diurnal in their habits are gorgeously coloured; and, +lastly, several of the largest beetles in the world belong to this +family, which was placed by Linnaeus and Fabricius at the head of the +Order of the Coleoptera.[493] + + +_Stridulating organs._--Beetles belonging to many and widely distinct +families possess these organs. The sound can sometimes be heard at the +distance of several feet or even yards,[494] but is not comparable with +that produced by the Orthoptera. The part which may be called the rasp +generally consists of a narrow slightly-raised surface, crossed by very +fine, parallel ribs, sometimes so fine as to cause iridescent colours, +and having a very elegant appearance under the microscope. In some +cases, for instance, with Typhaeus, it could be plainly seen that +extremely minute, bristly, scale-like prominences, which cover the whole +surrounding surface in approximately parallel lines, give rise to the +ribs of the rasp by becoming confluent and straight, and at the same +time more prominent and smooth. A hard ridge on any adjoining part of +the body, which in some cases is specially modified for the purpose, +serves as the scraper for the rasp. The scraper is rapidly moved across +the rasp, or conversely the rasp across the scraper. + +[Illustration: Fig. 24. Necrophorus (from Landois). _r._ The two rasps. +Left-hand figure, part of the rasp highly magnified.] + +These organs are situated in widely different positions. In the +carrion-beetles (Necrophorus) two parallel rasps (_r_, fig. 24) stand on +the dorsal surface of the fifth abdominal segment, each rasp being +crossed, as described by Landois,[495] by from 126 to 140 fine ribs. +These ribs are scraped by the posterior margins of the elytra, a small +portion of which projects beyond the general outline. In many +Crioceridae, and in _Clythra 4-punctata_ (one of the Chrysomelidae), and +in some Tenebrionidae, &c.,[496] the rasp is seated on the dorsal apex of +the abdomen, on the pygidium or pro-pygidium, and is scraped as above by +the elytra. In Heterocerus, which belongs to another family, the rasps +are placed on the sides of the first abdominal segment, and are scraped +by ridges on the femora.[497] In certain Curculionidae and Carabidae,[498] +the parts are completely reversed in position, for the rasps are seated +on the inferior surface of the elytra, near their apices, or along their +outer margins, and the edges of the abdominal segments serve as the +scrapers. In _Pelobius hermanni_ (one of Dytiscidae or water-beetles) a +strong ridge runs parallel and near to the sutural margin of the elytra, +and is crossed by ribs, coarse in the middle part, but becoming +gradually finer at both ends, especially at the upper end; when this +insect is held under water or in the air, a stridulating noise is +produced by scraping the extreme horny margin of the abdomen against the +rasp. In a great number of long-horned beetles (Longicornia) the organs +are altogether differently situated, the rasp being on the meso-thorax, +which is rubbed against the pro-thorax; Landois counted 238 very fine +ribs on the rasp of _Cerambyx heros_. + +[Illustration: Fig. 25. Hind-leg of Geotrupes stercorarius (from +Landois). _r._ Rasp. _c._ Coxa. _f._ Femur. _t._ Tibia. _tr._ Tarsi.] + +Many Lamellicorns have the power of stridulating, and the organs differ +greatly in position. Some species stridulate very loudly, so that when +Mr. F. Smith caught a _Trox sabulosus_, a gamekeeper who stood by +thought that he had caught a mouse; but I failed to discover the proper +organs in this beetle. In Geotrupes and Typhaeus a narrow ridge runs +obliquely across (_r_, fig. 25) the coxa of each hind-leg, having in +_G. stercorarius_ 84 ribs, which are scraped by a specially-projecting +part of one of the abdominal segments. In the nearly allied _Copris +lunaris_, an excessively narrow fine rasp runs along the sutural margin +of the elytra, with another short rasp near the basal outer margin; but +in some other Coprini the rasp is seated, according to Leconte,[499] on +the dorsal surface of the abdomen. In Oryctes it is seated on the +pro-pygidium, and in some other Dynastini, according to the same +entomologist, on the under surface of the elytra. Lastly, Westring +states that in _Omaloplia brunnea_ the rasp is placed on the +pro-sternum, and the scraper on the meta-sternum, the parts thus +occupying the under surface of the body, instead of the upper surface as +in the Longicorns. + +We thus see that the stridulating organs in the different coleopterous +families are wonderfully diversified in position, but not much in +structure. Within the same family some species are provided with these +organs, and some are quite destitute of them. This diversity is +intelligible, if we suppose that originally various species made a +shuffling or hissing noise by the rubbing together of the hard and rough +parts of their bodies which were in contact; and that from the noise +thus produced being in some way useful, the rough surfaces were +gradually developed into regular stridulating organs. Some beetles as +they move, now produce, either intentionally or unintentionally, a +shuffling noise, without possessing any proper organs for the purpose. +Mr. Wallace informs me that the _Euchirus longimanus_ (a Lamellicorn, +with the anterior legs wonderfully elongated in the male) "makes, whilst +moving, a low hissing sound by the protrusion and contraction of the +abdomen; and when seized it produces a grating sound by rubbing its +hind-legs against the edges of the elytra." The hissing sound is clearly +due to a narrow rasp running along the sutural margin of each elytron; +and I could likewise make the grating sound by rubbing the shagreened +surface of the femur against the granulated margin of the corresponding +elytron; but I could not here detect any proper rasp; nor is it likely +that I could have overlooked it in so large an insect. After examining +Cychrus and reading what Westring has written in his two papers about +this beetle, it seems very doubtful whether it possesses any true rasp, +though it has the power of emitting a sound. + +From the analogy of the Orthoptera and Homoptera, I expected to find +that the stridulating organs in the Coleoptera differed according to +sex; but Landois, who has carefully examined several species, observed +no such difference; nor did Westring; nor did Mr. G. R. Crotch in +preparing the numerous specimens which he had the kindness to send me +for examination. Any slight sexual difference, however, would be +difficult to detect, on account of the great variability of these +organs. Thus in the first pair of the _Necrophorus humator_ and of the +_Pelobius_ which I examined, the rasp was considerably larger in the +male than in the female; but not so with succeeding specimens. In +_Geotrupes stercorarius_ the rasp appeared to me thicker, opaquer, and +more prominent in three males than in the same number of females; +consequently my son, Mr. F. Darwin, in order to discover whether the +sexes differed in their power of stridulating, collected 57 living +specimens, which he separated into two lots, according as they made, +when held in the same manner, a greater or lesser noise. He then +examined their sexes, but found that the males were very nearly in the +same proportion to the females in both lots. Mr. F. Smith has kept alive +numerous specimens of _Mononychus pseudacori_ (Curculionidae), and is +satisfied that both sexes stridulate, and apparently in an equal degree. + +Nevertheless the power of stridulating is certainly a sexual character +in some few Coleoptera. Mr. Crotch has discovered that the males alone +of two species of Heliopathes (Tenebrionidae) possess stridulating +organs. I examined five males of _H. gibbus_, and in all these there was +a well-developed rasp, partially divided into two, on the dorsal surface +of the terminal abdominal segment; whilst in the same number of females +there was not even a rudiment of the rasp, the membrane of this segment +being transparent and much thinner than in the male. In _H. +cribratostriatus_ the male has a similar rasp, excepting that it is not +partially divided into two portions, and the female is completely +destitute of this organ; but in addition the male has on the apical +margins of the elytra, on each side of the suture, three or four short +longitudinal ridges, which are crossed by extremely fine ribs, parallel +to and resembling those on the abdominal rasp; whether these ridges +serve as an independent rasp, or as a scraper for the abdominal rasp, I +could not decide: the female exhibits no trace of this latter structure. + +Again, in three species of the Lamellicorn genus Oryctes, we have a +nearly parallel case. In the females of _O. gryphus_ and _nasicornis_ +the ribs on the rasp of the pro-pygidium are less continuous and less +distinct than in the males; but the chief difference is that the whole +upper surface of this segment, when held in the proper light, is seen to +be clothed with hairs, which are absent or are represented by +excessively fine down in the males. It should be noticed that in all +Coleoptera the effective part of the rasp is destitute of hairs. In _O. +senegalensis_ the difference between the sexes is more strongly marked, +and this is best seen when the proper segment is cleaned and viewed as a +transparent object. In the female the whole surface is covered with +little separate crests, bearing spines; whilst in the male these crests +become, in proceeding towards the apex, more and more confluent, +regular, and naked; so that three-fourths of the segment is covered with +extremely fine parallel ribs, which are quite absent in the female. In +the females, however, of all three species of Oryctes, when the abdomen +of a softened specimen is pushed backwards and forwards, a slight +grating or stridulating sound can be produced. + +In the case of the Heliopathes and Oryctes there can hardly be a doubt +that the males stridulate in order to call or to excite the females; but +with most beetles the stridulation apparently serves both sexes as a +mutual call. This view is not rendered improbable from beetles +stridulating under various emotions; we know that birds use their voices +for many purposes besides singing to their mates. The great +Chiasognathus stridulates in anger or defiance; many species do the same +from distress or fear, when held so that they cannot escape; Messrs. +Wollaston and Crotch were able, by striking the hollow stems of trees in +the Canary Islands, to discover the presence of beetles belonging to the +genus Acalles by their stridulation. Lastly the male Ateuchus +stridulates to encourage the female in her work, and from distress when +she is removed.[500] Some naturalists believe that beetles make this +noise to frighten away their enemies; but I cannot think that the +quadrupeds and birds which are able to devour the larger beetles with +their extremely hard coats, would be frightened by so slight a grating +sound. The belief that the stridulation serves as a sexual call is +supported by the fact that death-ticks (_Anobium tessellatum_) are well +known to answer each other's ticking, or, as I have myself observed, a +tapping noise artificially made; and Mr. Doubleday informs me that he +has twice or thrice observed a female ticking,[501] and in the course of +an hour or two has found her united with a male, and on one occasion +surrounded by several males. Finally, it seems probable that the two +sexes of many kinds of beetles were at first enabled to find each other +by the slight shuffling noise produced by the rubbing together of the +adjoining parts of their hard bodies; and that as the males or females +which made the greatest noise succeeded best in finding partners, the +rugosities on various parts of their bodies were gradually developed by +means of sexual selection into true stridulating organs. + + + + +CHAPTER XI. + +INSECTS, _continued_.--ORDER LEPIDOPTERA. + + + Courtship of butterflies--Battles--Ticking noise--Colours + common to both sexes, or more brilliant in the + males--Examples--Not due to the direct action of the conditions + of life--Colours adapted for protection--Colours of + moths--Display--Perceptive powers of the + Lepidoptera--Variability--Causes of the difference in colour + between the males and females--Mimickry, female butterflies + more brilliantly coloured than the males--Bright colours of + caterpillars--Summary and concluding remarks on the secondary + sexual characters of insects--Birds and insects compared. + + +In this great Order the most interesting point for us is the difference +in colour between the sexes of the same species, and between the +distinct species of the same genus. Nearly the whole of the following +chapter will be devoted to this subject; but I will first make a few +remarks on one or two other points. Several males may often be seen +pursuing and crowding round the same female. Their courtship appears to +be a prolonged affair, for I have frequently watched one or more males +pirouetting round a female until I became tired, without seeing the end +of the courtship. Although butterflies are such weak and fragile +creatures, they are pugnacious, and an Emperor butterfly[502] has been +captured with the tips of its wings broken from a conflict with another +male. Mr. Collingwood in speaking of the frequent battles between the +butterflies of Borneo says, "They whirl round each other with the +greatest rapidity, and appear to be incited by the greatest ferocity." +One case is known of a butterfly, namely the _Ageronia feronia_, which +makes a noise like that produced by a toothed wheel passing under a +spring catch, and which could be heard at the distance of several yards. +At Rio de Janeiro this sound was noticed by me, only when two were +chasing each other in an irregular course, so that it is probably made +during the courtship of the sexes; but I neglected to attend to this +point.[503] + +Every one has admired the extreme beauty of many butterflies and of some +moths; and we are led to ask, how has this beauty been acquired? Have +their colours and diversified patterns simply resulted from the direct +action of the physical conditions to which these insects have been +exposed, without any benefit being thus derived? Or have successive +variations been accumulated and determined either as a protection or for +some unknown purpose, or that one sex might be rendered attractive to +the other? And, again, what is the meaning of the colours being widely +different in the males and females of certain species, and alike in the +two sexes of other species? Before attempting to answer these questions +a body of facts must be given. + +With most of our English butterflies, both those which are beautiful, +such as the admiral, peacock, and painted lady (Vanessae), and those +which are plain-coloured, such as the meadow-browns (Hipparchiae), the +sexes are alike. This is also the case with the magnificent Heliconidae +and Danaidae of the tropics. But in certain other tropical groups, and +with some of our English butterflies, as the purple emperor, orange-tip, +&c. (_Apatura Iris_ and _Anthocharis cardamines_), the sexes differ +either greatly or slightly in colour. No language suffices to describe +the splendour of the males of some tropical species. Even within the +same genus we often find species presenting an extraordinary difference +between the sexes, whilst others have their sexes closely alike. Thus in +the South American genus Epicalia, Mr. Bates, to whom I am much indebted +for most of the following facts and for looking over this whole +discussion, informs me that he knows twelve species, the two sexes of +which haunt the same stations (and this is not always the case with +butterflies), and therefore cannot have been differently affected by +external conditions[504]. In nine of these species the males rank +amongst the most brilliant of all butterflies, and differ so greatly +from the comparatively plain females that they were formerly placed in +distinct genera. The females of these nine species resemble each other +in their general type of coloration, and likewise resemble both sexes in +several allied genera, found in various parts of the world. Hence in +accordance with the descent-theory we may infer that these nine species, +and probably all the others of the genus, are descended from an +ancestral form which was coloured in nearly the same manner. In the +tenth species the female still retains the same general colouring, but +the male resembles her, so that he is coloured in a much less gaudy and +contrasted manner than the males of the previous species. In the +eleventh and twelfth species, the females depart from the type of +colouring which is usual with their sex in this genus, for they are +gaily decorated in nearly the same manner as the males, but in a +somewhat less degree. Hence in these two species the bright colours of +the males seem to have been transferred to the females; whilst the male +of the tenth species has either retained or recovered the plain colours +of the female as well as of the parent-form of the genus; the two sexes +being thus rendered in both cases, though in an opposite manner, nearly +alike. In the allied genus Eubagis, both sexes of some of the species +are plain-coloured and nearly alike; whilst with the greater number the +males are decorated with beautiful metallic tints, in a diversified +manner, and differ much from their females. The females throughout the +genus retain the same general style of colouring, so that they commonly +resemble each other much more closely than they resemble their own +proper males. + +In the genus Papilio, all the species of the AEneas group are remarkable +for their conspicuous and strongly contrasted colours, and they +illustrate the frequent tendency to gradation in the amount of +difference between the sexes. In a few species, for instance in _P. +ascanius_, the males and females are alike; in others the males are a +little or very much more superbly coloured than the females. The genus +Junonia allied to our Vanessae offers a nearly parallel case, for +although the sexes of most of the species resemble each other and are +destitute of rich colours, yet in certain species, as in _J. oenone_, +the male is rather more brightly coloured than the female, and in a few +(for instance _J. andremiaja_) the male is so different from the female +that he might be mistaken for an entirely distinct species. + +Another striking case was pointed out to me in the British museum by Mr. +A. Butler, namely one of the Tropical American Theclae, in which both +sexes are nearly alike and wonderfully splendid; in another, the male +is coloured in a similarly gorgeous manner, whilst the whole upper +surface of the female is of a dull uniform brown. Our common little +English blue butterflies of the genus Lycaena, illustrate the various +differences in colour between the sexes, almost as well, though not in +so striking a manner, as the above exotic genera. In _Lycaena agestis_ +both sexes have wings of a brown colour, bordered with small ocellated +orange spots, and are consequently alike. In _L. oegon_ the wings of +the male are of a fine blue, bordered with black; whilst the wings of +the female are brown, with a similar border, and closely resemble those +of _L. agestis_. Lastly, in _L. arion_ both sexes are of a blue colour +and nearly alike, though in the female the edges of the wings are rather +duskier, with the black spots plainer; and in a bright blue Indian +species both sexes are still more closely alike. + +I have given the foregoing cases in some detail in order to shew, in the +first place, that when the sexes of butterflies differ, the male as a +general rule is the most beautiful, and departs most from the usual type +of colouring of the group to which the species belongs. Hence in most +groups the females of the several species resemble each other much more +closely than do the males. In some exceptional cases, however, to which +I shall hereafter allude, the females are coloured more splendidly than +the males. In the second place these cases have been given to bring +clearly before the mind that within the same genus, the two sexes +frequently present every gradation from no difference in colour to so +great a difference that it was long before the two were placed by +entomologists in the same genus. In the third place, we have seen that +when the sexes nearly resemble each other, this apparently may be due +either to the male having transferred his colours to the female, or to +the male having retained, or perhaps recovered, the primordial colours +of the genus to which the species belongs. It also deserves notice that +in those groups in which the sexes present any difference of colour, the +females usually resemble the males to a certain extent, so that when the +males are beautiful to an extraordinary degree, the females almost +invariably exhibit some degree of beauty. From the numerous cases of +gradation in the amount of difference between the sexes, and from the +prevalence of the same general type of coloration throughout the whole +of the same group, we may conclude that the causes, whatever they may +be, which have determined the brilliant colouring of the males alone of +some species, and of both sexes in a more or less equal degree of other +species, have generally been the same. + +As so many gorgeous butterflies inhabit the tropics, it has often been +supposed that they owe their colours to the great heat and moisture of +these zones; but Mr. Bates[505] has shewn by the comparison of various +closely-allied groups of insects from the temperate and tropical +regions, that this view cannot be maintained; and the evidence becomes +conclusive when brilliantly-coloured males and plain-coloured females of +the same species inhabit the same district, feed on the same food, and +follow exactly the same habits of life. Even when the sexes resemble +each other, we can hardly believe that their brilliant and +beautifully-arranged colours are the purposeless result of the nature of +the tissues, and the action of the surrounding conditions. + +With animals of all kinds, whenever colour has been modified for some +special purpose, this has been, as far as we can judge, either for +protection or as an attraction between the sexes. With many species of +butterflies the upper surfaces of the wings are obscurely coloured, and +this in all probability leads to their escaping observation and danger. +But butterflies when at rest would be particularly liable to be attacked +by their enemies; and almost all the kinds when resting raise their +wings vertically over their backs, so that the lower sides alone are +exposed to view. Hence it is this side which in many cases is obviously +coloured so as to imitate the surfaces on which these insects commonly +rest. Dr. Roessler, I believe, first noticed the similarity of the closed +wings of certain Vanessae and other butterflies to the bark of trees. +Many analogous and striking facts could be given. The most interesting +one is that recorded by Mr. Wallace[506] of a common Indian and Sumatran +butterfly (Kallima), which disappears like magic when it settles in a +bush; for it hides its head and antennae between its closed wings, and +these in form, colour, and veining cannot be distinguished from a +withered leaf together with the footstalk. In some other cases the lower +surfaces of the wings are brilliantly coloured, and yet are protective; +thus in _Thecla rubi_ the wings when closed are of an emerald green and +resemble the young leaves of the bramble, on which this butterfly in the +spring may often be seen seated. + +Although the obscure tints of the upper or under surface of many +butterflies no doubt serve to conceal them, yet we cannot possibly +extend this view to the brilliant and conspicuous colours of many kinds, +such as our admiral and peacock Vanessae, our white cabbage-butterflies +(Pieris), or the great swallow-tail Papilio which haunts the open +fens--for these butterflies are thus rendered visible to every living +creature. With these species both sexes are alike; but in the common +brimstone butterfly (_Gonepteryx rhamni_), the male is of an intense +yellow, whilst the female is much paler; and in the orange-tip +(_Anthocharis cardamines_) the males alone have the bright orange tips +to their wings. In these cases the males and females are equally +conspicuous, and it is not credible that their difference in colour +stands in any relation to ordinary protection. Nevertheless it is +possible that the conspicuous colours of many species may be in an +indirect manner beneficial, as will hereafter be explained, by leading +their enemies at once to recognise them as unpalatable. Even in this +case it does not certainly follow that their bright colours and +beautiful patterns were acquired for this special purpose. In some other +remarkable cases, beauty has been gained for the sake of protection, +through the imitation of other beautiful species, which inhabit the same +district and enjoy an immunity from attack by being in some way +offensive to their enemies. + +The female of our orange-tip butterfly, above referred to, and of an +American species (_Anth. genutia_) probably shew us, as Mr. Walsh has +remarked to me, the primordial colours of the parent-species of the +genus; for both sexes of four or five widely-distributed species are +coloured in nearly the same manner. We may infer here, as in several +previous cases, that it is the males of _Anth. cardamines_ and _genutia_ +which have departed from the usual type of colouring of their genus. In +the _Anth. sara_ from California, the orange-tips have become partially +developed in the female; for her wings are tipped with reddish-orange, +but paler than in the male, and slightly different in some other +respects. In an allied Indian form, the _Iphias glaucippe_, the +orange-tips are fully developed in both sexes. In this Iphias the under +surface of the wings marvellously resembles, as pointed out to me by Mr. +A. Butler, a pale-coloured leaf; and in our English orange-tip, the +under surface resembles the flower-head of the wild parsley, on which it +may be seen going to rest at night.[507] The same reasoning power which +compels us to believe that the lower surfaces have here been coloured +for the sake of protection, leads us to deny that the wings have been +tipped, especially when this character is confined to the males, with +bright orange for the same purpose. + + +Turning now to Moths: most of these rest motionless with their wings +depressed during the whole or greater part of the day; and the upper +surfaces of their wings are often shaded and coloured in an admirable +manner, as Mr. Wallace has remarked, for escaping detection. With most +of the Bombycidae and Noctuidae,[508] when at rest, the front-wings +overlap and conceal the hind-wings; so that the latter might be brightly +coloured without much risk; and they are thus coloured in many species +of both families. During the act of flight, moths would often be able to +escape from their enemies; nevertheless, as the hind-wings are then +fully exposed to view, their bright colours must generally have been +acquired at the cost of some little risk. But the following fact shews +us how cautious we ought to be in drawing conclusions on this head. The +common yellow under-wings (Triphaena) often fly about during the day or +early evening, and are then conspicuous from the colour of their +hind-wings. It would naturally be thought that this would be a source of +danger; but Mr. J. Jenner Weir believes that it actually serves them as +a means of escape, for birds strike at these brightly coloured and +fragile surfaces, instead of at the body. For instance, Mr. Weir turned +into his aviary a vigorous specimen of _Triphaena pronuba_, which was +instantly pursued by a robin; but the bird's attention being caught by +the coloured wings, the moth was not captured until after about fifty +attempts, and small portions of the wings were repeatedly broken off. He +tried the same experiment, in the open air, with a _T. fimbria_ and +swallow; but the large size of this moth probably interfered with its +capture.[509] We are thus reminded of a statement made by Mr. +Wallace,[510] namely, that in the Brazilian forests and Malayan islands, +many common and highly-decorated butterflies are weak flyers, though +furnished with a broad expanse of wings; and they "are often captured +with pierced and broken wings, as if they had been seized by birds, from +which they had escaped: if the wings had been much smaller in proportion +to the body, it seems probable that the insect would more frequently +have been struck or pierced in a vital part, and thus the increased +expanse of the wings may have been indirectly beneficial." + + +_Display._--The bright colours of butterflies and of some moths are +specially arranged for display, whether or not they serve in addition as +a protection. Bright colours would not be visible during the night; and +there can be no doubt that moths, taken as a body, are much less gaily +decorated than butterflies, all of which are diurnal in their habits. +But the moths in certain families, such as the Zygaenidae, various +Sphingidae, Uraniidae, some Arctiidae and Saturniidae, fly about during the +day or early evening, and many of these are extremely beautiful, being +far more brightly coloured than the strictly nocturnal kinds. A few +exceptional cases, however, of brightly-coloured nocturnal species have +been recorded.[511] + +There is evidence of another kind in regard to display. Butterflies, as +before remarked, elevate their wings when at rest, and whilst basking in +the sunshine often alternately raise and depress them, thus exposing to +full view both surfaces; and although the lower surface is often +coloured in an obscure manner as a protection, yet in many species it is +as highly coloured as the upper surface, and sometimes in a very +different manner. In some tropical species the lower surface is even +more brilliantly coloured than the upper.[512] In one English +fritillary, the _Argynnis aglaia_, the lower surface alone is ornamented +with shining silver discs. Nevertheless, as a general rule, the upper +surface, which is probably the most fully exposed, is coloured more +brightly and in a more diversified manner than the lower. Hence the +lower surface generally affords to entomologists the most useful +character for detecting the affinities of the various species. + +Now if we turn to the enormous group of moths, which do not habitually +expose to full view the under surface of their wings, this side is very +rarely, as I hear from Mr. Stainton, coloured more brightly than the +upper side, or even with equal brightness. Some exceptions to the rule, +either real or apparent, must be noticed, as that of Hypopyra, specified +by Mr. Wormald.[513] Mr. R. Trimen informs me that in Guenee's great +work, three moths are figured, in which the under surface is much the +most brilliant. For instance, in the Australian Gastrophora the upper +surface of the fore-wing is pale greyish-ochreous, while the lower +surface is magnificently ornamented by an ocellus of cobalt-blue, placed +in the midst of a black mark, surrounded by orange-yellow, and this by +bluish-white. But the habits of these three moths are unknown; so that +no explanation can be given of their unusual style of colouring. Mr. +Trimen also informs me that the lower surface of the wings in certain +other Geometrae[514] and quadrifid Noctuae are either more variegated or +more brightly-coloured than the upper surface; but some of these species +have the habit of "holding their wings quite erect over their backs, +retaining them in this position for a considerable time," and thus +exposing to view the under surface. Other species when settled on the +ground or herbage have the habit of now and then suddenly and slightly +lifting up their wings. Hence the lower surface of the wings being more +brightly-coloured than the upper surface in certain moths is not so +anomalous a circumstance as it at first appears. The Saturniidae include +some of the most beautiful of all moths, their wings being decorated, as +in our British Emperor moth, with fine ocelli; and Mr. T. W. Wood[515] +observes that they resemble butterflies in some of their movements; "for +instance, in the gentle waving up and down of the wings, as if for +display, which is more characteristic of diurnal than of nocturnal +Lepidoptera." + +It is a singular fact that no British moths, nor as far as I can +discover hardly any foreign species, which are brilliantly coloured, +differ much in colour according to sex; though this is the case with +many brilliant butterflies. The male, however, of one American moth, the +_Saturnia Io_, is described as having its fore-wings deep yellow, +curiously marked with purplish-red spots; whilst the wings of the female +are purple-brown, marked with grey lines.[516] The British moths which +differ sexually in colour are all brown, or various tints of dull +yellow, or nearly white. In several species the males are much darker +than the females,[517] and these belong to groups which generally fly +about during the afternoon. On the other hand, in many genera, as Mr. +Stainton informs me, the males have the hind-wings whiter than those of +the female--of which, fact _Agrotis exclamationis_ offers a good +instance. The males are thus rendered more conspicuous than the females, +whilst flying about in the dusk. In the Ghost Moth (_Hepialus humuli_) +the difference is more strongly marked; the males being white and the +females yellow with darker markings. It is difficult to conjecture what +the meaning can be of these differences between the sexes in the shades +of darkness or lightness; but we can hardly suppose that they are the +result of mere variability with sexually-limited inheritance, +independently of any benefit thus derived. + +From the foregoing statements it is impossible to admit that the +brilliant colours of butterflies and of some few moths, have commonly +been acquired for the sake of protection. We have seen that their +colours and elegant patterns are arranged and exhibited as if for +display. Hence I am led to suppose that the females generally prefer, or +are most excited by the more brilliant males; for on any other +supposition the males would be ornamented, as far as we can see, for no +purpose. We know that ants and certain lamellicorn beetles are capable +of feeling an attachment for each other, and that ants recognise their +fellows after an interval of several months. Hence there is no abstract +improbability in the Lepidoptera, which probably stand nearly or quite +as high in the scale as these insects, having sufficient mental capacity +to admire bright colours. They certainly discover flowers by colour, +and, as I have elsewhere shewn, the plants which are fertilised +exclusively by the wind never have a conspicuously-coloured corolla. The +Humming-bird Sphinx may often be seen to swoop down from a distance on a +bunch of flowers in the midst of green foliage; and I have been assured +by a friend, that these moths repeatedly visited flowers painted on the +walls of a room in the South of France. The common white butterfly, as I +hear from Mr. Doubleday, often flies down to a bit of paper on the +ground, no doubt mistaking it for one of its own species. Mr. +Collingwood[518] in speaking of the difficulty of collecting certain +butterflies in the Malay Archipelago, states that "a dead specimen +pinned upon a conspicuous twig will often arrest an insect of the same +species in its headlong flight, and bring it down within easy reach of +the net, especially if it be of the opposite sex." + +The courtship of butterflies is a prolonged affair. The males sometimes +fight together in rivalry; and many may be seen pursuing or crowding +round the same female. If, then, the females do not prefer one male to +another, the pairing must be left to mere chance, and this does not +appear to me a probable event. If, on the other hand, the females +habitually, or even occasionally, prefer the more beautiful males, the +colours of the latter will have been rendered brighter by degrees, and +will have been transmitted to both sexes or to one sex, according to +which law of inheritance prevailed. The process of sexual selection will +have been much facilitated, if the conclusions arrived at from various +kinds of evidence in the supplement to the ninth chapter can be trusted; +namely that the males of many Lepidoptera, at least in the imago state, +greatly exceed in number the females. + +Some facts, however, are opposed to the belief that female butterflies +prefer the more beautiful males; thus, as I have been assured by several +observers, fresh females may frequently be seen paired with battered, +faded or dingy males; but this is a circumstance which could hardly +fail often to follow from the males emerging from their cocoons earlier +than the females. With moths of the family of the Bombycidae, the sexes +pair immediately after assuming the imago state; for they cannot feed, +owing to the rudimentary condition of their mouths. The females, as +several entomologists have remarked to me, lie in an almost torpid +state, and appear not to evince the least choice in regard to their +partners, This is the case with the common silk-moth (_B. mori_), as I +have been told by some continental and English breeders. Dr. Wallace, +who has had such immense experience in breeding _Bombyx cynthia_, is +convinced that the females evince no choice or preference. He has kept +above 300 of these moths living together, and has often found the most +vigorous females mated with stunted males. The reverse apparently seldom +occurs; for, as he believes, the more vigorous males pass over the +weakly females, being attracted by those endowed with most vitality. +Although we have been indirectly induced to believe that the females of +many species prefer the more beautiful males, I have no reason to +suspect, either with moths or butterflies, that the males are attracted +by the beauty of the females. If the more beautiful females had been +continually preferred, it is almost certain, from the colours of +butterflies being so frequently transmitted to one sex alone, that the +females would often have been rendered more beautiful than their male +partners. But this does not occur except in a few instances; and these +can be explained, as we shall presently see, on the principle of +mimickry and protection. + + +As sexual selection primarily depends on variability, a few words must +be added on this subject. In respect to colour there is no difficulty, +as any number of highly variable Lepidoptera could be named. One good +instance will suffice. Mr. Bates shewed me a whole series of specimens +of _Papilio sesostris_ and _childrenae_; in the latter the males varied +much in the extent of the beautifully enamelled green patch on the +fore-wings, and in the size of the white mark, as well as of the +splendid crimson stripe on the hind-wings; so that there was a great +contrast between the most and least gaudy males. The male of _Papilio +sesostris_, though a beautiful insect, is much less so than _P. +childrenae_. It likewise varies a little in the size of the green patch +on the fore-wings, and in the occasional appearance of a small crimson +stripe on the hind-wings, borrowed, as it would seem, from its own +female; for the females of this and of many other species in the AEneas +group possess this crimson stripe. Hence between the brightest specimens +of _P. sesostris_ and the least bright of _P. childrenae_, there was but +a small interval; and it was evident that as far as mere variability is +concerned, there would be no difficulty in permanently increasing by +means of selection the beauty of either species. The variability is here +almost confined to the male sex; but Mr. Wallace and Mr. Bates have +shewn[519] that the females of some other species are extremely +variable, the males being nearly constant. As I have before mentioned +the Ghost Moth (_Hepialus humuli_) as one of the best instances in +Britain of a difference in colour between the sexes of moths, it may be +worth adding[520] that in the Shetland Islands, males are frequently +found which closely resemble the females. In a future chapter I shall +have occasion to shew that the beautiful eye-like spots or ocelli, so +common on the wings of many Lepidoptera, are eminently variable. + + +On the whole, although many serious objections may be urged, it seems +probable that most of the species of Lepidoptera which are brilliantly +coloured, owe their colours to sexual selection, excepting in certain +cases, presently to be mentioned, in which conspicuous colours are +beneficial as a protection. From the ardour of the male throughout the +animal kingdom, he is generally willing to accept any female; and it is +the female which usually exerts a choice. Hence if sexual selection has +here acted, the male, when the sexes differ, ought to be the most +brilliantly coloured; and this undoubtedly is the ordinary rule. When +the sexes are brilliantly coloured and resemble each other, the +characters acquired by the males appear to have been transmitted to both +sexes. But will this explanation of the similarity and dissimilarity in +colour between the sexes suffice? + +The males and females of the same species of butterfly are known[521] in +several cases to inhabit different stations, the former commonly basking +in the sunshine, the latter haunting gloomy forests. It is therefore +possible that different conditions of life may have acted directly on +the two sexes; but this is not probable,[522] as in the adult state they +are exposed during a very short period to different conditions; and the +larvae of both are exposed to the same conditions. Mr. Wallace believes +that the less brilliant colours of the female have been specially +gained in all or almost all cases for the sake of protection. On the +contrary it seems to me more probable that the males alone, in the large +majority of cases, have acquired their bright colours through sexual +selection, the females having been but little modified. Consequently the +females of distinct but allied species ought to resemble each other much +more closely than do the males of the same species; and this is the +general rule. The females thus approximately show us the primordial +colouring of the parent-species of the group to which they belong. They +have, however, almost always been modified to a certain extent by some +of the successive steps of variation, through the accumulation of which +the males were rendered beautiful, having been transferred to them. The +males and females of allied though distinct species will also generally +have been exposed during their prolonged larval state to different +conditions, and may have been thus indirectly affected; though with the +males any slight change of colour thus caused will often have been +completely masked by the brilliant tints gained through sexual +selection. When we treat of Birds, I shall have to discuss the whole +question whether the differences in colour between the males and females +have been in part specially gained by the latter as a protection; so +that I will here only give unavoidable details. + +In all cases when the more common form of equal inheritance by both +sexes has prevailed, the selection of bright-coloured males would tend +to make the females bright-coloured; and the selection of dull-coloured +females would tend to make the males dull. If both processes were +carried on simultaneously, they would tend to neutralise each other. As +far as I can see, it would be extremely difficult to change through +selection the one form of inheritance into the other. But by the +selection of successive variations, which were from the first sexually +limited in their transmission, there would not be the slightest +difficulty in giving bright colours to the males alone, and at the same +time or subsequently, dull colours to the females alone. In this latter +manner female butterflies and moths may, as I fully admit, have been +rendered inconspicuous for the sake of protection, and widely different +from their males. + +Mr. Wallace[523] has argued with much force in favour of his view that +when the sexes differ, the female has been specially modified for the +sake of protection; and that this has been effected by one form of +inheritance, namely, the transmission of characters to both sexes, +having been changed through the agency of natural selection into the +other form, namely, transmission to one sex. I was at first strongly +inclined to accept this view; but the more I have studied the various +classes throughout the animal kingdom, the less probable it has +appeared. Mr. Wallace urges that both sexes of the _Heliconidae_, +_Danaidae_, _Acroeidae_ are equally brilliant because both are protected +from the attacks of birds and other enemies, by their offensive odour; +but that in other groups, which do not possess this immunity, the +females have been rendered inconspicuous, from having more need of +protection than the males. This supposed difference in the "need of +protection by the two sexes" is rather deceptive, and requires some +discussion. It is obvious that brightly-coloured individuals, whether +males or females, would equally attract, and obscurely-coloured +individuals equally escape, the attention of their enemies. But we are +concerned with the effects of the destruction or preservation of certain +individuals of either sex, on the character of the race. With insects, +after the male has fertilised the female, and after the latter has laid +her eggs, the greater or less immunity from danger of either sex could +not possibly have any effect on the offspring. Before the sexes have +performed their proper functions, if they existed in equal numbers and +if they strictly paired (all other circumstances being the same), the +preservation of the males and females would be equally important for the +existence of the species and for the character of the offspring. But +with most animals, as is known to be the case with the domestic +silk-moth, the male can fertilise two or three females; so that the +destruction of the males would not be so injurious to the species as +that of the females. On the other hand, Dr. Wallace believes that with +moths the progeny from a second or third fertilisation is apt to be +weakly, and therefore would not have so good chance of surviving. When +the males exist in much greater numbers than the females, no doubt many +males might be destroyed with impunity to the species; but I cannot see +that the results of ordinary selection for the sake of protection would +be influenced by the sexes existing in unequal numbers; for the same +proportion of the more conspicuous individuals, whether males or +females, would probably be destroyed. If indeed the males presented a +greater range of variation in colour, the result would be different; but +we need not here follow out such complex details. On the whole I cannot +perceive that an inequality in the numbers of the two sexes would +influence in any marked manner the effects of ordinary selection on the +character of the offspring. + +Female Lepidoptera require, as Mr. Wallace insists, some days to +deposit their fertilised ova and to search for a proper place; during +this period (whilst the life of the male was of no importance) the +brighter-coloured females would be exposed to danger and would be liable +to be destroyed. The duller-coloured females on the other hand would +survive, and thus would influence, it might be thought, in a marked +manner the character of the species,--either of both sexes or of one +sex, according to which form of inheritance prevailed. But it must not +be forgotten that the males emerge from the cocoon-state some days +before the females, and during this period, whilst the unborn females +were safe, the brighter-coloured males would be exposed to danger; so +that ultimately both sexes would probably be exposed during a nearly +equal length of time to danger, and the elimination of conspicuous +colours would not be much more effective in the one than the other sex. + +It is a more important consideration that female Lepidoptera, as Mr. +Wallace remarks, and as is known to every collector, are generally +slower flyers than the males. Consequently the latter, if exposed to +greater danger from being conspicuously coloured, might be able to +escape from their enemies, whilst the similarly-coloured females would +be destroyed; and thus the females would have the most influence in +modifying the colour of their progeny. + +There is one other consideration: bright colours, as far as sexual +selection is concerned, are commonly of no service to the females; so +that if the latter varied in brightness, and the variations were +sexually limited in their transmission, it would depend on mere chance +whether the females had their bright colours increased; and this would +tend throughout the Order to diminish the number of species with +brightly-coloured females in comparison with the species having +brightly-coloured males. On the other hand, as bright colours are +supposed to be highly serviceable to the males in their love-struggles, +the brighter males (as we shall see in the chapter on Birds) although +exposed to rather greater danger, would on an average procreate a +greater number of offspring than the duller males. In this case, if the +variations were limited in their transmission to the male sex, the males +alone would be rendered more brilliantly coloured; but if the variations +were not thus limited, the preservation and augmentation of such +variations would depend on whether more evil was caused to the species +by the females being rendered conspicuous, than good to the males by +certain individuals being successful over their rivals. + +As there can hardly be a doubt that both sexes of many butterflies and +moths have been rendered dull-coloured for the sake of protection, so +it may have been with the females alone of some species in which +successive variations towards dullness first appeared in the female sex +and were from the first limited in their transmission to the same sex. +If not thus limited, both sexes would become dull-coloured. We shall +immediately see, when we treat of mimickry, that the females alone of +certain butterflies have been rendered extremely beautiful for the sake +of protection, without any of the successive protective variations +having been transferred to the male, to whom they could not possibly +have been in the least degree injurious, and therefore could not have +been eliminated through natural selection. Whether in each particular +species, in which the sexes differ in colour, it is the female which has +been specially modified for the sake of protection; or whether it is the +male which has been specially modified for the sake of sexual +attraction, the female having retained her primordial colouring only +slightly changed through the agencies before alluded to; or whether +again both sexes have been modified, the female for protection and the +male for sexual attraction, can only be definitely decided when we know +the life-history of each species. + +Without distinct evidence, I am unwilling to admit that a double process +of selection has long been going on with a multitude of species,--the +males having been rendered more brilliant by beating their rivals; and +the females more dull-coloured by having escaped from their enemies. We +may take as an instance the common brimstone butterfly (Gonepteryx), +which appears early in the spring before any other kind. The male of +this species is of a far more intense yellow than the female, though she +is almost equally conspicuous; and in this case it does not seem +probable that she specially acquired her pale tints as a protection, +though it is probable that the male acquired his bright colours as a +sexual attraction. The female of _Anthocharis cardamines_ does not +possess the beautiful orange tips to her wings with which the male is +ornamented; consequently she closely resembles the white butterflies +(Pieris) so common in our gardens; but we have no evidence that this +resemblance is beneficial. On the contrary, as she resembles both sexes +of several species of the same genus inhabiting various quarters of the +world, it is more probable that she has simply retained to a large +extent her primordial colours. + +Various facts support the conclusion that with the greater number of +brilliantly-coloured Lepidoptera, it is the male which has been +modified; the two sexes having come to differ from each other, or to +resemble each other, according to which form of inheritance has +prevailed. Inheritance is governed by so many unknown laws or +conditions, that they seem to us to be most capricious in their +action;[524] and we can so far understand how it is that with +closely-allied species the sexes of some differ to an astonishing +degree, whilst the sexes of others are identical in colour. As the +successive steps in the process of variation are necessarily all +transmitted through the female, a greater or less number of such steps +might readily become developed in her; and thus we can understand the +frequent gradations from an extreme difference to no difference at all +between the sexes of the species within the same group. These cases of +gradation are much too common to favour the supposition that we here see +females actually undergoing the process of transition and losing their +brightness for the sake of protection; for we have every reason to +conclude that at any one time the greater number of species are in a +fixed condition. With respect to the differences between the females of +the species in the same genus or family, we can perceive that they +depend, at least in part, on the females partaking of the colours of +their respective males. This is well illustrated in those groups in +which the males are ornamented to an extraordinary degree; for the +females in these groups generally partake to a certain extent of the +splendour of their male partners. Lastly, we continually find, as +already remarked, that the females of almost all the species in the same +genus, or even family, resemble each other much more closely in colour +than do the males; and this indicates that the males have undergone a +greater amount of modification than the females. + + +_Mimickry._--This principle was first made clear in an admirable paper +by Mr. Bates,[525] who thus threw a flood of light on many obscure +problems. It had previously been observed that certain butterflies in S. +America belonging to quite distinct families, resembled the Heliconidae +so closely in every stripe and shade of colour that they could not be +distinguished except by an experienced entomologist. As the Heliconidae +are coloured in their usual manner, whilst the others depart from the +usual colouring of the groups to which they belong, it is clear that the +latter are the imitators, and the Heliconidae the imitated. Mr. Bates +further observed that the imitating species are comparatively rare, +whilst the imitated swarm in large numbers; the two sets living mingled +together. From the fact of the Heliconidae being conspicuous and +beautiful insects, yet so numerous in individuals and species, he +concluded that they must be protected from the attacks of birds by some +secretion or odour; and this hypothesis has now been confirmed by a +considerable body of curious evidence.[526] From these considerations +Mr. Bates inferred that the butterflies which imitate the protected +species had acquired their present marvellously deceptive appearance, +through variation and natural selection, in order to be mistaken for the +protected kinds and thus to escape being devoured. No explanation is +here attempted of the brilliant colours of the imitated, but only of the +imitating butterflies. We must account for the colours of the former in +the same general manner, as in the cases previously discussed in this +chapter. Since the publication of Mr. Bates' paper, similar and equally +striking facts have been observed by Mr. Wallace[527] in the Malayan +region, and by Mr. Trimen in South Africa. + +As some writers[528] have felt much difficulty in understanding how the +first steps in the process of mimickry could have been effected through +natural selection, it may be well to remark that the process probably +has never commenced with forms widely dissimilar in colour. But with two +species moderately like each other, the closest resemblance if +beneficial to either form could readily be thus gained; and if the +imitated form was subsequently and gradually modified through sexual +selection or any other means, the imitating form would be led along the +same track, and thus be modified to almost any extent, so that it might +ultimately assume an appearance or colouring wholly unlike that of the +other members of the group to which it belonged. As extremely slight +variations in colour would not in many cases suffice to render a species +so like another protected species as to lead to its preservation, it +should be remembered that many species of Lepidoptera are liable to +considerable and abrupt variations in colour. A few instances have been +given in this chapter; but under this point of view Mr. Bates' original +paper on mimickry, as well as Mr. Wallace's papers, should be consulted. + +In the foregoing cases both sexes of the imitating species +resemble the imitated; but occasionally the female alone mocks a +brilliantly-coloured and protected species inhabiting the same district. +Consequently the female differs in colour from her own male, and, which +is a rare and anomalous circumstance, is the more brightly-coloured of +the two. In all the few species of Pieridae, in which the female is more +conspicuously coloured than the male, she imitates, as I am informed by +Mr. Wallace, some protected species inhabiting the same region. The +female of _Diadema anomala_ is rich purple-brown with almost the whole +surface glossed with satiny blue, and she closely imitates the +_Euploea midamus_, "one of the commonest butterflies of the East;" +whilst the male is bronzy or olive-brown, with only a slight blue gloss +on the outer parts of the wings.[529] Both sexes of this Diadema and of +_D. bolina_ follow the same habits of life, so that the differences in +colour between the sexes cannot be accounted for by exposure to +different conditions;[530] even if this explanation were admissible in +other instances.[531] + +The above cases of female butterflies which are more brightly-coloured +than the males, shew us, firstly, that variations have arisen in a state +of nature in the female sex, and have been transmitted exclusively, or +almost exclusively, to the same sex; and, secondly, that this form of +inheritance has not been determined through natural selection. For if we +assume that the females, before they became brightly coloured in +imitation of some protected kind, were exposed during each season for a +longer period to danger than the males; or if we assume that they could +not escape so swiftly from their enemies, we can understand how they +alone might originally have acquired through natural selection and +sexually-limited inheritance their present protective colours. But +except on the principle of these variations having been transmitted +exclusively to the female offspring, we cannot understand why the males +should have remained dull-coloured; for it would surely not have been in +any way injurious to each individual male to have partaken by +inheritance of the protective colours of the female, and thus to have +had a better chance of escaping destruction. In a group in which +brilliant colours are so common as with butterflies, it cannot be +supposed that the males have been kept dull-coloured through sexual +selection by the females rejecting the individuals which were rendered +as beautiful as themselves. We may, therefore, conclude that in these +cases inheritance by one sex is not due to the modification through +natural selection of a tendency to equal inheritance by both sexes. + +It may be well here to give an analogous case in another Order, of +characters acquired only by the female, though not in the least +injurious, as far as we can judge, to the male. Amongst the Phasmidae, or +spectre-insects, Mr. Wallace states that "it is often the females alone +that so strikingly resemble leaves, while the males show only a rude +approximation." Now, whatever may be the habits of these insects, it is +highly improbable that it could be disadvantageous to the males to +escape detection by resembling leaves.[532] Hence we may conclude that +the females alone in this latter as in the previous cases originally +varied in certain characters; these characters having been preserved and +augmented through ordinary selection for the sake of protection and from +the first transmitted to the female offspring alone. + + +_Bright Colours of Caterpillars._--Whilst reflecting on the beauty of +many butterflies, it occurred to me that some caterpillars were +splendidly coloured, and as sexual selection could not possibly have +here acted, it appeared rash to attribute the beauty of the mature +insect to this agency, unless the bright colours of their larvae could +be in some manner explained. In the first place it may be observed that +the colours of caterpillars do not stand in any close correlation with +those of the mature insect. Secondly, their bright colours do not serve +in any ordinary manner as a protection. As an instance of this, Mr. +Bates informs me that the most conspicuous caterpillar which he ever +beheld (that of a Sphinx) lived on the large green leaves of a tree on +the open llanos of South America; it was about four inches in length, +transversely banded with black and yellow, and with its head, legs, and +tail of a bright red. Hence it caught the eye of any man who passed by +at the distance of many yards, and no doubt of every passing bird. + +I then applied to Mr. Wallace, who has an innate genius for solving +difficulties. After some consideration he replied: "Most caterpillars +require protection, as may be inferred from some kinds being furnished +with spines or irritating hairs, and from many being coloured green like +the leaves on which they feed, or curiously like the twigs of the trees +on which they live." I may add as another instance of protection, that +there is a caterpillar of a moth, as I am informed by Mr. J. Mansel +Weale, which lives on the mimosas in South Africa, and fabricates for +itself a case, quite undistinguishable from the surrounding thorns. From +such considerations Mr. Wallace thought it probable that +conspicuously-coloured caterpillars were protected by having a nauseous +taste; but as their skin is extremely tender, and as their intestines +readily protrude from a wound, a slight peck from the beak of a bird +would be as fatal to them as if they had been devoured. Hence, as Mr. +Wallace remarks, "distastefulness alone would be insufficient to protect +a caterpillar unless some outward sign indicated to its would-be +destroyer that its prey was a disgusting morsel." Under these +circumstances it would be highly advantageous to a caterpillar to be +instantaneously and certainly recognised as unpalatable by all birds and +other animals. Thus the most gaudy colours would be serviceable, and +might have been gained by variation and the survival of the most +easily-recognised individuals. + +This hypothesis appears at first sight very bold; but when it was +brought before the Entomological Society[533] it was supported by +various statements; and Mr. J. Jenner Weir, who keeps a large number of +birds in an aviary, has made, as he informs me, numerous trials, and +finds no exception to the rule, that all caterpillars of nocturnal and +retiring habits with smooth skins, all of a green colour, and all which +imitate twigs, are greedily devoured by his birds. The hairy and spinose +kinds are invariably rejected, as were four conspicuously-coloured +species. When the birds rejected a caterpillar, they plainly shewed, by +shaking their heads and cleansing their beaks, that they were disgusted +by the taste.[534] Three conspicuous kinds of caterpillars and moths +were also given by Mr. A. Butler to some lizards and frogs, and were +rejected; though other kinds were eagerly eaten. Thus the probable truth +of Mr. Wallace's view is confirmed, namely, that certain caterpillars +have been made conspicuous for their own good, so as to be easily +recognised by their enemies, on nearly the same principle that certain +poisons are coloured by druggists for the good of man. This view will, +it is probable, be hereafter extended to many animals, which are +coloured in a conspicuous manner. + + +_Summary and Concluding Remarks on Insects._--Looking back to the +several Orders, we have seen that the sexes often differ in various +characters, the meaning of which is not understood. The sexes, also, +often differ in their organs of sense or locomotion, so that the males +may quickly discover or reach the females, and still oftener in the +males possessing diversified contrivances for retaining the females when +found. But we are not here much concerned with sexual differences of +these kinds. + +In almost all the Orders, the males of some species, even of weak and +delicate kinds, are known to be highly pugnacious; and some few are +furnished with special weapons for fighting with their rivals. But the +law of battle does not prevail nearly so widely with insects as with the +higher animals. Hence probably it is that the males have not often been +rendered larger and stronger than the females. On the contrary they are +usually smaller, in order that they may be developed within a shorter +time, so as to be ready in large numbers for the emergence of the +females. + +In two families of the Homoptera the males alone possess, in an +efficient state, organs which may be called vocal; and in three families +of the Orthoptera the males alone possess stridulating organs. In both +cases these organs are incessantly used during the breeding-season, not +only for calling the females, but for charming or exciting them in +rivalry with other males. No one who admits the agency of natural +selection, will dispute that these musical instruments have been +acquired through sexual selection. In four other Orders the members of +one sex, or more commonly of both sexes, are provided with organs for +producing various sounds, which apparently serve merely as call-notes. +Even when both sexes are thus provided, the individuals which were able +to make the loudest or most continuous noise would gain partners before +those which were less noisy, so that their organs have probably been +gained through sexual selection. It is instructive to reflect on the +wonderful diversity of the means for producing sound, possessed by the +males alone or by both sexes in no less than six Orders, and which were +possessed by at least one insect at an extremely remote geological +epoch. We thus learn how effectual sexual selection has been in leading +to modifications of structure, which sometimes, as with the Homoptera, +are of an important nature. + +From the reasons assigned in the last chapter, it is probable that the +great horns of the males of many lamellicorn, and some other beetles, +have been acquired as ornaments. So perhaps it may be with certain other +peculiarities confined to the male sex. From the small size of insects, +we are apt to undervalue their appearance. If we could imagine a male +Chalcosoma (fig. 15) with its polished, bronzed coat of mail, and vast +complex horns, magnified to the size of a horse or even of a dog, it +would be one of the most imposing animals in the world. + +The colouring of insects is a complex and obscure subject. When the male +differs slightly from the female, and neither are brilliantly coloured, +it is probable that the two sexes have varied in a slightly different +manner, with the variations transmitted to the same sex, without any +benefit having been thus derived or evil suffered. When the male is +brilliantly coloured and differs conspicuously from the female, as with +some dragon-flies and many butterflies, it is probable that he alone has +been modified, and that he owes his colours to sexual selection; whilst +the female has retained a primordial or very ancient type of colouring, +slightly modified by the agencies before explained, and has therefore +not been rendered obscure, at least in most cases, for the sake of +protection. But the female alone has sometimes been coloured +brilliantly so as to imitate other protected species inhabiting the same +district. When the sexes resemble each other and both are obscurely +coloured, there is no doubt that they have been in a multitude of cases +coloured for the sake of protection. So it is in some instances when +both are brightly coloured, causing them to resemble surrounding objects +such as flowers, or other protected species, or indirectly by giving +notice to their enemies that they are of an unpalatable nature. In many +other cases in which the sexes resemble each other and are brilliantly +coloured, especially when the colours are arranged for display, we may +conclude that they have been gained by the male sex as an attraction, +and have been transferred to both sexes. We are more especially led to +this conclusion whenever the same type of coloration prevails throughout +a group, and we find that the males of some species differ widely in +colour from the females, whilst both sexes of other species are quite +alike, with intermediate gradations connecting these extreme states. + +In the same manner as bright colours have often been partially +transferred from the males to the females, so it has been with the +extraordinary horns of many lamellicorn and some other beetles. So, +again, the vocal or instrumental organs proper to the males of the +Homoptera and Orthoptera have generally been transferred in a +rudimentary, or even in a nearly perfect condition to the females; yet +not sufficiently perfect to be used for producing sound. It is also an +interesting fact, as bearing on sexual selection, that the stridulating +organs of certain male Orthoptera are not fully developed until the last +moult; and that the colours of certain male dragon-flies are not fully +developed until some little time after their emergence from the pupal +state, and when they are ready to breed. + +Sexual selection implies that the more attractive individuals are +preferred by the opposite sex; and as with insects, when the sexes +differ, it is the male which, with rare exceptions, is the most +ornamented and departs most from the type to which the species +belongs;--and as it is the male which searches eagerly for the female, +we must suppose that the females habitually or occasionally prefer the +more beautiful males, and that these have thus acquired their beauty. +That in most or all the orders the females have the power of rejecting +any particular male, we may safely infer from the many singular +contrivances possessed by the males, such as great jaws, adhesive +cushions, spines, elongated legs, &c., for seizing the female; for these +contrivances shew that there is some difficulty in the act. In the case +of unions between distinct species, of which many instances have been +recorded, the female must have been a consenting party. Judging from +what we know of the perceptive powers and affections of various insects, +there is no antecedent improbability in sexual selection having come +largely into action; but we have as yet no direct evidence on this head, +and some facts are opposed to the belief. Nevertheless, when we see many +males pursuing the same female, we can hardly believe that the pairing +is left to blind chance--that the female exerts no choice, and is not +influenced by the gorgeous colours or other ornaments, with which the +male alone is decorated. + +If we admit that the females of the Homoptera and Orthoptera appreciate +the musical tones emitted by their male partners, and that the various +instruments for this purpose have been perfected through sexual +selection, there is little improbability in the females of other insects +appreciating beauty in form or colour, and consequently in such +characters having been thus gained by the males. But from the +circumstance of colour being so variable, and from its having been so +often modified for the sake of protection, it is extremely difficult to +decide in how large a proportion of cases sexual selection has come into +play. This is more especially difficult in those Orders, such as the +Orthoptera, Hymenoptera, and Coleoptera, in which the two sexes rarely +differ much in colour; for we are thus cut off from our best evidence of +some relation between the reproduction of the species and colour. With +the Coleoptera, however, as before remarked, it is in the great +lamellicorn group, placed by some authors at the head of the Order, and +in which we sometimes see a mutual attachment between the sexes, that we +find the males of some species possessing weapons for sexual strife, +others furnished with wonderful horns, many with stridulating organs, +and others ornamented with splendid metallic tints. Hence it seems +probable that all these characters have been gained through the same +means, namely sexual selection. + +When we treat of Birds, we shall see that they present in their +secondary sexual characters the closest analogy with insects. Thus, many +male birds are highly pugnacious, and some are furnished with special +weapons for fighting with their rivals. They possess organs which are +used during the breeding-season for producing vocal and instrumental +music. They are frequently ornamented with combs, horns, wattles and +plumes of the most diversified kinds, and are decorated with beautiful +colours, all evidently for the sake of display. We shall find that, as +with insects, both sexes, in certain groups, are equally beautiful, and +are equally provided with ornaments which are usually confined to the +male sex. In other groups both sexes are equally plain-coloured and +unornamented. Lastly, in some few anomalous cases, the females are more +beautiful than the males. We shall often find, in the same group of +birds, every gradation from no difference between the sexes, to an +extreme difference. In the latter case we shall see that the females, +like female insects, often possess more or less plain traces of the +characters which properly belong to the males. The analogy, indeed, in +all these respects between birds and insects, is curiously close. +Whatever explanation applies to the one class probably applies to the +other; and this explanation, as we shall hereafter attempt to shew, is +almost certainly sexual selection. + + + FOOTNOTES: + + [1] As the works of the first-named authors are so well known, + I need not give the titles; but as those of the latter are less + well known in England, I will give them:--'Sechs Vorlesungen + ueber die Darwin'sche Theorie:' zweite Auflage, 1868, von Dr. L. + Buechner; translated into French under the title 'Conferences + sur la Theorie Darwinienne,' 1869. 'Der Mensch, im Lichte der + Darwin'sche Lehre,' 1865, von Dr. F. Rolle. I will not attempt + to give references to all the authors who have taken the same + side of the question. Thus G. Canestrini has published + ('Annuario della Soc. d. Nat.,' Modena, 1867, p. 81) a very + curious paper on rudimentary characters, as bearing on the + origin of man. Another work has (1869) been published by Dr. + Barrago Francesco, bearing in Italian the title of "Man, made + in the image of God, was also made in the image of the ape." + + [2] Prof. Haeckel is the sole author who, since the publication + of the 'Origin,' has discussed, in his various works, in a very + able manner, the subject of sexual selection, and has seen its + full importance. + + [3] 'Grosshirnwindungen des Menschen,' 1868, s. 96. + + [4] 'Lec. sur la Phys.' 1866, p. 890, as quoted by M. Dally, + 'L'Ordre des Primates et le Transformisme,' 1868, p. 29. + + [5] 'Naturgeschichte der Saeugethiere von Paraguay,' 1830, s. + 50. + + [6] Brehm, 'Thierleben,' B. i. 1864, s. 75, 86. On the Ateles, + s. 105. For other analogous statements, see s. 25, 107. + + [7] With respect to insects see Dr. Laycock 'On a General Law + of Vital Periodicity,' British Association, 1842. Dr. + Macculloch, 'Silliman's North American Journal of Science,' + vol. xvii. p. 305, has seen a dog suffering from tertian ague. + + [8] I have given the evidence on this head in my 'Variation of + Animals and Plants under Domestication,' vol. ii. p. 15. + + [9] "Mares e diversis generibus Quadrumanorum sine dubio + dignoscunt feminas humanas a maribus. Primum, credo, odoratu, + postea aspectu. Mr. Youatt, qui diu in Hortis Zoologicis + (Bestiariis) medicus animalium erat, vir in rebus observandis + cautus et sagax, hoc mihi certissime probavit, et curatores + ejusdem loci et alii e ministris confirmaverunt. Sir Andrew + Smith et Brehm notabant idem in Cynocephalo. Illustrissimus + Cuvier etiam narrat multa de hac re qua ut opinor nihil turpius + potest indicari inter omnia hominibus et Quadrumanis communia. + Narrat enim Cynocephalum quendam in furorem incidere aspectu + feminarum aliquarum, sed nequaquam accendi tanto furore ab + omnibus. Semper eligebat juniores, et dignoscebat in turba, et + advocabat voce gestuque." + + [10] This remark is made with respect to Cynocephalus and the + anthropomorphous apes by Geoffroy Saint-Hilaire and F. Cuvier, + 'Hist. Nat. des Mammiferes,' tom. i. 1824. + + [11] Huxley, 'Man's Place in Nature,' 1863, p. 34. + + [12] 'Man's Place in Nature,' 1863, p. 67. + + [13] The human embryo (upper fig.) is from Ecker, 'Icones + Phys.,' 1851-1859, tab. xxx. fig. 2. This embryo was ten lines + in length, so that the drawing is much magnified. The embryo of + the dog is from Bischoff, 'Entwicklungsgeschichte des + Hunde-Eies,' 1845, tab. xi. fig. 42 B. This drawing is five + times magnified, the embryo being 25 days old. The internal + viscera have been omitted, and the uterine appendages in both + drawings removed. I was directed to these figures by Prof. + Huxley, from whose work, 'Man's Place in Nature.' the idea of + giving them was taken. Haeckel has also given analogous drawings + in his 'Schoepfungsgeschichte.' + + [14] Prof. Wyman in 'Proc. of American Acad. of Sciences,' vol. + iv. 1860, p. 17. + + [15] Owen, 'Anatomy of Vertebrates,' vol. i. p. 533. + + [16] 'Die Grosshirnwindungen des Menschen,' 1868, s. 95. + + [17] 'Anatomy of Vertebrates,' vol. ii. p. 553. + + [18] 'Proc. Soc. Nat. Hist.' Boston, 1863, vol. ix. p. 185. + + [19] 'Man's Place in Nature,' p. 65. + + [20] I had written a rough copy of this chapter before reading + a valuable paper, "Caratteri rudimentali in ordine all'origine + del uomo" ('Annuario della Soc. d. Nat.,' Modena, 1867, p. 81), + by G. Canestrini, to which paper I am considerably indebted. + Haeckel has given admirable discussions on this whole subject, + under the title of Dysteleology, in his 'Generelle Morphologie' + and 'Schoepfungsgeschichte.' + + [21] Some good criticisms on this subject have been given by + Messrs. Murie and Mivart, in 'Transact. Zoolog. Soc.' 1869, + vol. vii. p. 92. + + [22] 'Variation of Animals and Plants under Domestication,' + vol. ii. pp. 317 and 397. See also 'Origin of Species,' 5th + edit. p. 535. + + [23] For instance M. Richard ('Annales des Sciences Nat.' 3rd + series, Zoolog. 1852, tom. xviii. p. 13) describes and figures + rudiments of what he calls the "muscle pedieux de la main," + which he says is sometimes "infiniment petit." Another muscle, + called "le tibial posterieur," is generally quite absent in the + hand, but appears from time to time in a more or less + rudimentary condition. + + [24] Prof. W. Turner, 'Proc. Royal Soc. Edinburgh,' 1866-67, p. + 65. + + [25] Canestrini quotes Hyrt. ('Annuario della Soc. dei + Naturalisti,' Modena, 1867, p. 97) to the same effect. + + [26] 'The Diseases of the Ear,' by J. Toynbee, F.R.S., 1860, p. + 12. + + [27] See also some remarks, and the drawings of the ears of the + Lemuroidea, in Messrs. Murie and Mivart's excellent paper in + 'Transact. Zoolog. Soc.' vol. vii. 1869, pp. 6 and 90. + + [28] Mueller's 'Elements of Physiology,' Eng. translat., 1842, + vol. ii. p. 1117. Owen, 'Anatomy of Vertebrates,' vol. iii. p. + 260; ibid. on the Walrus, 'Proc. Zoolog. Soc.' November 8th, + 1854. See also R. Knox, 'Great Artists and Anatomists,' p. 106. + This rudiment apparently is somewhat larger in Negroes and + Australians than in Europeans, see Carl Vogt, 'Lectures on + Man,' Eng. translat. p. 129. + + [29] 'The Physiology and Pathology of Mind,' 2nd edit. 1868, p. + 134. + + [30] Eschricht, Ueber die Richtung der Haare am menschlichen + Koerper 'Muellers Archiv fuer Anat. und Phys.' 1837, s. 47. I + shall often have to refer to this very curious paper. + + [31] Paget, 'Lectures on Surgical Pathology,' 1853, vol. i. p. + 71. + + [32] Eschricht, ibid. s. 40, 47. + + [33] Dr. Webb, 'Teeth in Man and the Anthropoid Apes,' as + quoted by Dr. C. Carter Blake in 'Anthropological Review,' + July, 1867, p. 299. + + [34] Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 320, 321, + and 325. + + [35] 'On the Primitive Form of the Skull,' Eng. translat. in + 'Anthropological Review,' Oct. 1868, p. 426. + + [36] Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 416, 434, + 441. + + [37] 'Annuario della Soc. d. Nat.' Modena, 1867, p. 94. + + [38] M. C. Martins ("De l'Unite Organique," in 'Revue des Deux + Mondes,' June 15, 1862, p. 16), and Haeckel ('Generelle + Morphologie,' B. ii. s. 278), have both remarked on the + singular fact of this rudiment sometimes causing death. + + [39] 'The Lancet,' Jan. 24, 1863, p. 83. Dr. Knox, 'Great + Artists and Anatomists,' p. 63. See also an important memoir on + this process by Dr. Grube, in the 'Bulletin de l'Acad. Imp. de + St. Petersbourg,' tom. xii. 1867, p. 448. + + [40] "On the Caves of Gibraltar," 'Transact. Internat. Congress + of Prehist. Arch.' Third Session, 1869, p. 54. + + [41] Quatrefages has lately collected the evidence on this + subject. 'Revue des Cours Scientifiques,' 1867-1868, p. 625. + + [42] Owen, 'On the Nature of Limbs,' 1849, p. 114. + + [43] Leuckart, in Todd's 'Cyclop. of Anat.' 1849-52, vol. iv. + p. 1415. In man this organ is only from three to six lines in + length, but, like so many other rudimentary parts, it is + variable in development as well as in other characters. + + [44] See, on this subject, Owen, 'Anatomy of Vertebrates,' vol. + iii. pp. 675, 676, 706. + + [45] See the evidence on these points, as given by Lubbock, + 'Prehistoric Times,' p. 354, &c. + + [46] 'L'Instinct chez les Insectes.' 'Revue des Deux Mondes,' + Feb. 1870, p. 690. + + [47] 'The American Beaver and his Works,' 1868. + + [48] 'The Principles of Psychology,' 2nd edit. 1870, pp. + 418-443. + + [49] 'Contributions to the Theory of Natural Selection,' 1870, + p. 212 + + [50] 'Recherches sur les Moeurs des Fourmis,' 1810, p. 173. + + [51] All the following statements, given on the authority of + these two naturalists, are taken from Rengger's 'Naturges. der + Saeugethiere von Paraguay,' 1830, s. 41-57, and from Brehm's + 'Thierleben,' B. i. s. 10-87. + + [52] 'Bridgewater Treatise,' p. 263. + + [53] W. C. L. Martin, 'Nat. Hist. of Mammalia,' 1841, p. 405. + + [54] Quoted by Vogt, 'Memoire sur les Microcephales,' 1867, p. + 168. + + [55] 'The Variation of Animals and Plants under Domestication,' + vol. i. p. 27. + + [56] 'Les Moeurs des Fourmis,' 1810, p. 150. + + [57] Quoted in Dr. Maudsley's 'Physiology and Pathology of + Mind,' 1868, pp. 19, 220. + + [58] Dr. Jerdon, 'Birds of India,' vol. i. 1862, p. xxi. + + [59] Mr. L. H. Morgan's work on 'The American Beaver,' 1868, + offers a good illustration of this remark. I cannot, however, + avoid thinking that he goes too far in underrating the power of + Instinct. + + [60] 'The Moor and the Loch,' p. 45. Col. Hutchinson on 'Dog + Breaking,' 1850, p. 46. + + [61] 'Personal Narrative,' Eng. translat., vol. iii. p. 106. + + [62] Quoted by Sir C. Lyell, 'Antiquity of Man,' p. 497. + + [63] 'Journal of Researches during the Voyage of the "Beagle,"' + 1845, p. 398. 'Origin of Species,' 5th edit. p. 260. + + [64] 'Lettres Phil. sur l'Intelligence des Animaux,' nouvelle + edit. 1802, p. 86. + + [65] See the evidence on this head in chap. i. vol. i. 'On the + Variation of Animals and Plants under Domestication.' + + [66] 'Proc. Zoolog. Soc.' 1864, p. 186. + + [67] Savage and Wyman in 'Boston Journal of Nat. Hist.' vol. + iv. 1843-44, p. 383. + + [68] 'Saeugethiere von Paraguay,' 1830, s. 51-56. + + [69] 'Thierleben,' B. i. s. 79, 82. + + [70] 'The Malay Archipelago,' vol. i. 1869, p. 87. + + [71] 'Primeval Man,' 1869, pp. 145, 147. + + [72] 'Prehistoric Times,' 1865, p. 473, &c. + + [73] Quoted in 'Anthropological Review,' 1864, p. 158. + + [74] Rengger, ibid. s. 45. + + [75] See my 'Variation of Animals and Plants under + Domestication,' vol. i. p. 27. + + [76] See a discussion on this subject in Mr. E. B. Tylor's very + interesting work, 'Researches into the Early History of + Mankind,' 1865, chaps. ii. to iv. + + [77] Hon. Daines Barrington in 'Philosoph. Transactions,' 1773, + p. 262. See also Dureau de la Malle, in 'Ann. des Sc. Nat.' 3rd + series, Zoolog. tom. x. p. 119. + + [78] 'On the Origin of Language,' by H. Wedgwood, 1866. + 'Chapters on Language,' by the Rev. F. W. Farrar, 1865. These + works are most interesting. See also 'De la Phys. et de + Parole,' par Albert Lemoine, 1865, p. 190. The work on this + subject, by the late Prof. Aug. Schleicher, has been translated + by Dr. Bikkers into English, under the title of 'Darwinism + tested by the Science of Language,' 1869. + + [79] Vogt, 'Memoire sur les Microcephales,' 1867, p. 169. With + respect to savages, I have given some facts in my 'Journal of + Researches,' &c., 1845, p. 206. + + [80] See clear evidence on this head in the two works so often + quoted, by Brehm and Rengger. + + [81] See remarks on this head by Dr. Maudsley, 'The Physiology + and Pathology of Mind,' 2nd edit. 1868, p. 199. + + [82] Many curious cases have been recorded. See, for instance, + 'Inquiries Concerning the Intellectual Powers,' by Dr. + Abercrombie, 1838, p. 150. + + [83] 'The Variation of Animals and Plants under Domestication,' + vol. ii. p. 6. + + [84] See some good remarks to this effect by Dr. Maudsley, 'The + Physiology and Pathology of Mind,' 1808, p. 199. + + [85] Macgillivray, 'Hist. of British Birds,' vol. ii. 1839, p. + 29. An excellent observer, Mr. Blackwall, remarks that the + magpie learns to pronounce single words, and even short + sentences, more readily than almost any other British bird; + yet, as he adds, after long and closely investigating its + habits, he has never known it, in a state of nature, display + any unusual capacity for imitation. 'Researches in Zoology,' + 1834, p. 158. + + [86] See the very interesting parallelism between the + development of species and languages, given by Sir C. Lyell in + 'The Geolog. Evidences of the Antiquity of Man,' 1863, chap. + xxiii. + + [87] See remarks to this effect by the Rev. F. W. Farrar, in an + interesting article, entitled "Philology and Darwinism" in + 'Nature,' March 24th, 1870, p. 528. + + [88] 'Nature,' Jan. 6th, 1870, p. 257. + + [89] Quoted by C. S. Wake, 'Chapters on Man,' 1868, p. 101. + + [90] Buckland, 'Bridgewater Treatise,' p. 411. + + [91] See some good remarks on the simplification of languages, + by Sir J. Lubbock, 'Origin of Civilisation,' 1870, p. 278. + + [92] 'Conferences sur la Theorie Darwinienne,' French + translat., 1869, p. 132. + + [93] The Rev. Dr. J. M'Cann, 'Anti-Darwinism,' 1869, p. 13. + + [94] 'The Spectator,' Dec. 4th, 1869, p. 1430. + + [95] See an excellent article on this subject by the Rev. F. W. + Farrar, in the 'Anthropological Review,' Aug. 1864, p. ccxvii. + For further facts see Sir J. Lubbock, 'Prehistoric Times,' 2nd + edit. 1869. p. 564; and especially the chapters on Religion in + his 'Origin of Civilisation,' 1870. + + [96] The Worship of Animals and Plants, in the 'Fortnightly + Review,' Oct. 1, 1869, p. 422. + + [97] Tylor, 'Early History of Mankind,' 1865, p. 6. See also + the three striking chapters on the Development of Religion, in + Lubbock's 'Origin of Civilisation,' 1870. In a like manner Mr. + Herbert Spencer, in his ingenious essay in the 'Fortnightly + Review' (May 1st, 1870, p. 535), accounts for the earliest + forms of religious belief throughout the world, by man being + led through dreams, shadows, and other causes, to look at + himself as a double essence, corporeal and spiritual. As the + spiritual being is supposed to exist after death and to be + powerful, it is propitiated by various gifts and ceremonies, + and its aid invoked. He then further shews that names or + nicknames given from some animal or other object to the early + progenitors or founders of a tribe, are supposed after a long + interval to represent the real progenitor of the tribe; and + such animal or object is then naturally believed still to exist + as a spirit, is held sacred, and worshipped as a god. + Nevertheless I cannot but suspect that there is a still earlier + and ruder stage, when anything which manifests power or + movement is thought to be endowed with some form of life, and + with mental faculties analogous to our own. + + [98] See an able article on the Psychical Elements of Religion, + by Mr. L. Owen Pike, in 'Anthropolog. Review,' April, 1870, p. + lxiii. + + [99] 'Religion, Moral, &c., der Darwin'schen Art-Lehre,' 1869, + s. 53. + + [100] 'Prehistoric Times,' 2nd edit. p. 571. In this work (at + p. 553) there will be found an excellent account of the many + strange and capricious customs of savages. + + [101] See, for instance, on this subject, Quatrefages, 'Unite + de l'Espece Humaine,' 1861, p. 21, &c. + + [102] 'Dissertation on Ethical Philosophy,' 1837, p. 231, &c. + + [103] 'Metaphysics of Ethics,' translated by J. W. Semple, + Edinburgh, 1836, p. 136. + + [104] Mr. Bain gives a list ('Mental and Moral Science,' 1868, + p. 543-725) of twenty-six British authors who have written on + this subject, and whose names are familiar to every reader; to + these, Mr. Bain's own name, and those of Mr. Lecky, Mr. + Shadworth Hodgson, and Sir J. Lubbock, as well as of others, + may be added. + + [105] Sir B. Brodie, after observing that man is a social + animal ('Psychological Enquiries,' 1854, p. 192), asks the + pregnant question, "ought not this to settle the disputed + question as to the existence of a moral sense?" Similar ideas + have probably occurred to many persons, as they did long ago to + Marcus Aurelius. Mr. J. S. Mill speaks, in his celebrated work, + 'Utilitarianism,' (1864, p. 46), of the social feelings as a + "powerful natural sentiment," and as "the natural basis of + sentiment for utilitarian morality;" but on the previous page + he says, "if, as is my own belief, the moral feelings are not + innate, but acquired, they are not for that reason less + natural." It is with hesitation that I venture to differ from + so profound a thinker, but it can hardly be disputed that the + social feelings are instinctive or innate in the lower animals; + and why should they not be so in man? Mr. Bain (see, for + instance, 'The Emotions and the Will,' 1865, p. 481) and others + believe that the moral sense is acquired by each individual + during his lifetime. On the general theory of evolution this is + at least extremely improbable. + + [106] 'Die Darwin'sche Theorie,' s. 101. + + [107] Mr. R. Brown in 'Proc. Zoolog. Soc.' 1868, p. 409. + + [108] Brehm, 'Thierleben,' B. i. 1864, s. 52, 79. For the case + of the monkeys extracting thorns from each other, see s. 54. + With respect to the Hamadryas turning over stones, the fact is + given (s. 76) on the evidence of Alvarez, whose observations + Brehm thinks quite trustworthy. For the cases of the old male + baboons attacking the dogs, see s. 79; and with respect to the + eagle, s. 56. + + [109] 'Annals and Mag. of Nat. Hist.' November, 1868, p. 382. + + [110] Sir J. Lubbock, 'Prehistoric Times,' 2nd edit. p. 446. + + [111] As quoted by Mr. L. H. Morgan, 'The American Beaver,' + 1868, p. 272. Capt. Stansbury also gives an interesting account + of the manner in which a very young pelican, carried away by a + strong stream, was guided and encouraged in its attempts to + reach the shore by half a dozen old birds. + + [112] As Mr. Bain states, "effective aid to a sufferer springs + from sympathy proper:" 'Mental and Moral Science,' 1868, p. + 245. + + [113] 'Thierleben,' B. i. s. 85. + + [114] 'De l'Espece et de la Class.' 1869, p. 97. + + [115] 'Der Darwin'schen Art-Lehre,' 1869, s. 54. + + [116] Brehm, 'Thierleben,' B. i. s. 76. + + [117] See the first and striking chapter in Adam Smith's + 'Theory of Moral Sentiments.' Also Mr. Bain's 'Mental and Moral + Science,' 1868, p. 244, and 275-282. Mr. Bain states, that + "sympathy is, indirectly, a source of pleasure to the + sympathiser;" and he accounts for this through reciprocity. He + remarks that "the person benefited, or others in his stead, may + make up, by sympathy and good offices returned, for all the + sacrifice." But if, as appears to be the case, sympathy is + strictly an instinct, its exercise would give direct pleasure, + in the same manner as the exercise, as before remarked, of + almost every other instinct. + + [118] This fact, the Rev. L. Jenyns states (see his edition of + 'White's Nat. Hist. of Selborne,' 1853, p. 204) was first + recorded by the illustrious Jenner, in 'Phil. Transact.' 1824, + and has since been confirmed by several observers, especially + by Mr. Blackwall. This latter careful observer examined, late + in the autumn, during two years, thirty-six nests; he found + that twelve contained young dead birds, five contained eggs on + the point of being hatched, and three eggs not nearly hatched. + Many birds not yet old enough for a prolonged flight are + likewise deserted and left behind. See Blackwall, 'Researches + in Zoology,' 1834, pp. 108, 118. For some additional evidence, + although this is not wanted, see Leroy, 'Lettres Phil.' 1802, + p. 217. + + [119] Hume remarks ('An Enquiry Concerning the Principles of + Morals,' edit. of 1751, p. 132), "there seems a necessity for + confessing that the happiness and misery of others are not + spectacles altogether indifferent to us, but that the view of + the former ... communicates a secret joy; the appearance of the + latter ... throws a melancholy damp over the imagination." + + [120] 'Mental and Moral Science,' 1868, p. 254. + + [121] I have given one such case, namely of three Patagonian + Indians who preferred being shot, one after the other, to + betraying the plans of their companions in war ('Journal of + Researches,' 1845, p. 103). + + [122] Dr. Prosper Despine, in his 'Psychologie Naturelle,' 1868 + (tom. i. p. 243; tom ii. p. 169) gives many curious cases of + the worst criminals, who apparently have been entirely + destitute of conscience. + + [123] See an able article in the 'North British Review,' 1867, + p. 395. See also Mr. W. Bagehot's articles on the Importance of + Obedience and Coherence to Primitive Man, in the 'Fortnightly + Review,' 1867, p. 529, and 1868, p. 457, &c. + + [124] The fullest account which I have met with is by Dr. + Gerland, in his 'Ueber das Aussterben der Naturvoelker,' 1868; + but I shall have to recur to the subject of infanticide in a + future chapter. + + [125] See the very interesting discussion on Suicide in Lecky's + 'History of European Morals,' vol. i. 1869, p. 223. + + [126] See, for instance, Mr. Hamilton's account of the Kaffirs, + 'Anthropological Review,' 1870, p. xv. + + [127] Mr. M'Lennan has given 'Primitive Marriage,' 1865, p. + 176, a good collection of facts on this head. + + [128] Lecky, 'History of European Morals,' vol. i. 1869, p. + 109. + + [129] 'Embassy to China,' vol. ii. p. 348. + + [130] See on this subject copious evidence in Chap. vii. of Sir + J. Lubbock, 'Origin of Civilisation,' 1870. + + [131] For instance Lecky, 'Hist. European Morals,' vol. i. p. + 124. + + [132] This term is used in an able article in the 'Westminster + Review,' Oct. 1869, p. 498. For the Greatest Happiness + principle, see J. S. Mill, 'Utilitarianism,' p. 17. + + [133] Good instances are given by Mr. Wallace in 'Scientific + Opinion,' Sept. 15, 1869; and more fully in his 'Contributions + to the Theory of Natural Selection,' 1870, p. 353. + + [134] Tennyson, 'Idylls of the King,' p. 244. + + [135] 'The Thoughts of the Emperor M. Aurelius Antoninus,' Eng. + translat., 2nd edit., 1869, p. 112. Marcus Aurelius was born + A.D. 121. + + [136] Letter to Mr. Mill in Bain's 'Mental and Moral Science,' + 1868, p. 722. + + [137] A writer in the 'North British Review' (July, 1869, p. + 531), well capable of forming a sound judgment, expresses + himself strongly to this effect. Mr. Lecky ('Hist. of Morals,' + vol. i. p. 143) seems to a certain extent to coincide. + + [138] See his remarkable work on 'Hereditary Genius,' 1869, p. + 349. The Duke of Argyll ('Primeval Man,' 1869, p. 188) has some + good remarks on the contest in man's nature between right and + wrong. + + [139] 'The Thoughts of Marcus Aurelius,' &c., p. 139. + + [140] 'Investigations in Military and Anthropolog. Statistics + of American Soldiers,' by B. A. Gould, 1869, p. 256. + + [141] With respect to the "Cranial forms of the American + aborigines," see Dr. Aitken Meigs in 'Proc. Acad. Nat. Sci.' + Philadelphia, May, 1866. On the Australians, see Huxley, in + Lyell's 'Antiquity of Man,' 1863, p. 87. On the Sandwich + Islanders, Prof. J. Wyman, 'Observations on Crania,' Boston, + 1868, p. 18. + + [142] 'Anatomy of the Arteries,' by R. Quain. + + [143] 'Transact. Royal Soc.' Edinburgh, vol. xxiv. p. 175, 189. + + [144] 'Proc. Royal Soc.' 1867, p. 544; also 1868, p. 483, 524. + There is a previous paper, 1866, p. 229. + + [145] 'Proc. R. Irish Academy,' vol. x. 1868, p. 141. + + [146] 'Act. Acad.,' St. Petersburg, 1778, part ii. p. 217. + + [147] Brehm, 'Thierleben,' B. i. s. 58, 87. Rengger, + 'Saeugethiere von Paraguay,' s. 57. + + [148] 'Variation of Animals and Plants under Domestication,' + vol. ii. chap. xii. + + [149] 'Hereditary Genius: an Inquiry into its Laws and + Consequences,' 1869. + + [150] Mr. Bates remarks ('The Naturalist on the Amazons,' 1863, + vol. ii. p. 159), with respect to the Indians of the same S. + American tribe, "no two of them were at all similar in the + shape of the head; one man had an oval visage with fine + features, and another was quite Mongolian in breadth and + prominence of cheek, spread of nostrils, and obliquity of + eyes." + + [151] Blumenbach, 'Treatises on Anthropolog.' Eng. translat., + 1865, p. 205. + + [152] Godron, 'De l'Espece,' 1859, tom. ii. livre 3. + Quatrefages, 'Unite de l'Espece Humaine,' 1861. Also Lectures + on Anthropology, given in the 'Revue des Cours Scientifiques,' + 1866-1868. + + [153] 'Hist. Gen. et Part. des Anomalies de l'Organisation,' in + three volumes, tom. i. 1832. + + [154] I have fully discussed these laws in my 'Variation of + Animals and Plants under Domestication,' vol. ii. chap. xxii. + and xxiii. M. J. P. Durand has lately 1868; published a + valuable essay 'De l'Influence des Milieux, &c.' He lays much + stress on the nature of the soil. + + [155] 'Investigations in Military and Anthrop. Statistics,' &c. + 1869, by B. A. Gould, p. 93, 107, 126, 131, 134. + + [156] For the Polynesians, see Prichard's 'Physical Hist. of + Mankind,' vol. v. 1847, p. 145, 283. Also Godron, 'De + l'Espece,' tom. ii. p. 289. There is also a remarkable + difference in appearance between the closely-allied Hindoos + inhabiting the Upper Ganges and Bengal; see Elphinstone's + 'History of India,' vol. i. p. 324. + + [157] 'Memoirs, Anthropolog. Soc.' vol. iii. 1867-69, p. 561, + 565, 567. + + [158] Dr. Brakenridge, 'Theory of Diathesis,' 'Medical Times,' + June 19 and July 17, 1869. + + [159] I have given authorities for these several statements in + my 'Variation of Animals under Domestication,' vol. ii. p. + 297-300. Dr. Jaeger, "Ueber das Laengenwachsthum der Knochen," + 'Jenaischen Zeitschrift,' B. v. Heft i. + + [160] 'Investigations,' &c. By B. A. Gould, 1869, p. 288. + + [161] 'Saeugethiere von Paraguay,' 1830, s. 4. + + [162] 'History of Greenland,' Eng. translat. 1767, vol. i. p. + 230. + + [163] 'Intermarriage.' By Alex. Walker, 1838. p. 377. + + [164] 'The Variation of Animals under Domestication,' vol. i. + p. 173. + + [165] 'Principles of Biology,' vol. i. p. 455. + + [166] Paget, 'Lectures on Surgical Pathology,' vol. i. 1853, p. + 209. + + [167] 'The Variation of Animals under Domestication,' vol. ii. + p. 8. + + [168] 'Saeugethiere von Paraguay,' s. 8, 10. I have had good + opportunities for observing the extraordinary power of eyesight + in the Fuegians.' See also Lawrence ('Lectures on Physiology,' + &c., 1822, p. 404) on this same subject. M. Giraud-Teulon has + recently collected ('Revue des Cours Scientifiques,' 1870, p. + 625) a large and valuable body of evidence proving that the + cause of short-sight, "_C'est le travail assidu, de pres._" + + [169] Prichard, 'Phys. Hist. of Mankind,' on the authority of + Blumenbach, vol. i. 1851, p. 311; for the statement by Pallas, + vol. iv. 1844, p. 407. + + [170] Quoted by Prichard, 'Researches into the Phys. Hist. of + Mankind,' vol. v. p. 463. + + [171] Mr. Forbes' valuable paper is now published in the + 'Journal of the Ethnological Soc. of London,' new series, vol. + ii. 1870, p. 193. + + [172] Dr. Wilckens ('Landwirthschaft. Wochenblatt,' No. 10, + 1869) has lately published an interesting essay shewing how + domestic animals, which live in mountainous regions, have their + frames modified. + + [173] 'Memoire sur les Microcephales,' 1867, p. 50, 125, 169, + 171, 184-198. + + [174] See Dr. A. Farre's well-known article in the 'Cyclop. of + Anat. and Phys.' vol. v. 1859, p. 642. Owen 'Anatomy of + Vertebrates,' vol. iii. 1868, p. 687. Prof. Turner in + 'Edinburgh Medical Journal,' Feb. 1865. + + [175] 'Annuario della Soc. dei Naturalisti in Modena,' 1867, p. + 83. Prof. Canestrini gives extracts on this subject from + various authorities. Laurillard remarks, that as he has found a + complete similarity in the form, proportions, and connexion of + the two malar bones in several human subjects and in certain + apes, he cannot consider this disposition of the parts as + simply accidental. + + [176] A whole series of cases is given by Isid. Geoffroy + St.-Hilaire, 'Hist. des Anomalies,' tom. iii. p. 437. + + [177] In my 'Variation of Animals under Domestication' (vol. + ii. p. 57) I attributed the not very rare cases of + supernumerary mammae in women to reversion. I was led to this as + a _probable_ conclusion, by the additional mammae being + generally placed symmetrically on the breast, and more + especially from one case, in which a single efficient mamma + occurred in the inguinal region of a woman, the daughter of + another woman with supernumerary mammae. But Prof. Preyer ('Der + Kampf um das Dasein,' 1869, s. 45) states that _mammae erraticae_ + have been known to occur in other situations, even on the back; + so that the force of my argument is greatly weakened or perhaps + quite destroyed. + + With much hesitation I, in the same work (vol. ii. p. 12), + attributed the frequent cases of polydactylism in men to + reversion. I was partly led to this through Prof. Owen's + statement, that some of the Ichthyopterygia possess more than + five digits, and therefore, as I supposed, had retained a + primordial condition; but after reading Prof. Gegenbaur's paper + ('Jenaischen Zeitschrift,' B. v. Heft 3, s. 341), who is the + highest authority in Europe on such a point, and who disputes + Owen's conclusion, I see that it is extremely doubtful whether + supernumerary digits can thus be accounted for. It was the fact + that such digits not only frequently occur and are strongly + inherited, but have the power of regrowth after amputation, + like the normal digits of the lower vertebrata, that chiefly + led me to the above conclusion. This extraordinary fact of + their regrowth remains inexplicable, if the belief in reversion + to some extremely remote progenitor must be rejected. I cannot, + however, follow Prof. Gegenbaur in supposing that additional + digits could not reappear through reversion, without at the + same time other parts of the skeleton being simultaneously and + similarly modified; for single characters often reappear + through reversion. + + [178] 'Anatomy of Vertebrates,' vol. iii. 1868, p. 323. + + [179] 'Generelle Morphologie,' 1866, B. ii. s. clv. + + [180] Carl Vogt's 'Lectures on Man,' Eng. translat. 1864, p. + 151. + + [181] C. Carter Blake, on a jaw from La Naulette, 'Anthropolog. + Review,' 1867, p. 295. Schaaffhausen, ibid. 1868, p. 426. + + [182] 'The Anatomy of Expression,' 1844, p. 110, 131. + + [183] Quoted by Prof. Canestrini in the 'Annuario,' &c., 1867, + p. 90. + + [184] These papers deserve careful study by any one who desires + to learn how frequently our muscles vary, and in varying come + to resemble those of the Quadrumana. The following references + relate to the few points touched on in my text: Proc. Royal + Soc. vol. xiv. 1865, p. 379-384; vol. xv. 1866, p. 241, 242; + vol. xv. 1867, p. 544; vol. xvi. 1868, p. 524. I may here add + that Dr. Murie and Mr. St. George Mivart have shewn in their + Memoir on the Lemuroidea ('Transact. Zoolog. Soc.' vol. vii. + 1869, p. 96), how extraordinarily variable some of the muscles + are in these animals, the lowest members of the Primates. + Gradations, also, in the muscles leading to structures found in + animals still lower in the scale, are numerous in the + Lemuroidea. + + [185] Prof. Macalister in 'Proc. R. Irish Academy,' vol. x. + 1868, p. 124. + + [186] Prof. Macalister (ibid. p. 121) has tabulated his + observations, and finds that muscular abnormalities are most + frequent in the fore-arms, secondly in the face, thirdly in the + foot, &c. + + [187] The Rev. Dr. Haughton, after giving ('Proc. R. Irish + Academy,' June 27, 1864, p. 715) a remarkable case of variation + in the human _flexor pollicis longus_, adds, "This remarkable + example shews that man may sometimes possess the arrangement of + tendons of thumb and fingers characteristic of the macaque; but + whether such a case should be regarded as a macaque passing + upwards into a man, or a man passing downwards into a macaque, + or as a congenital freak of nature, I cannot undertake to say." + It is satisfactory to hear so capable an anatomist, and so + embittered an opponent of evolutionism, admitting even the + possibility of either of his first propositions. Prof. + Macalister has also described ('Proc. R. Irish Acad.' vol. x. + 1864, p. 138) variations in the _flexor pollicis longus_, + remarkable from their relations to the same muscle in the + Quadrumana. + + [188] The authorities for these several statements are given in + my 'Variation of Animals under Domestication,' vol. ii. p. + 320-335. + + [189] This whole subject has been discussed in chap. xxiii. + vol. ii. of my 'Variation of Animals and Plants under + Domestication.' + + [190] See the ever memorable 'Essay on the Principle of + Population,' by the Rev. T. Malthus, vol. i. 1826, p. 6, 517. + + [191] 'Variation of Animals and Plants under Domestication,' + vol. ii. p. 111-113, 163. + + [192] Mr. Sedgwick, 'British and Foreign Medico-Chirurg. + Review,' July, 1863, p. 170. + + [193] 'The Annals of Rural Bengal,' by W. W. Hunter, 1868, p. + 259. + + [194] 'Primitive Marriage,' 1865. + + [195] See some good remarks to this effect by W. Stanley + Jevons, "A Deduction from Darwin's Theory," 'Nature,' 1869, p. + 231. + + [196] Latham, 'Man and his Migrations,' 1851, p. 135. + + [197] Messrs. Murie and Mivart in their "Anatomy of the + Lemuroidea" ('Transact. Zoolog. Soc.' vol. vii. 1869, p. 96-98) + say, "some muscles are so irregular in their distribution that + they cannot be well classed in any of the above groups." These + muscles differ even on the opposite sides of the same + individual. + + [198] 'Quarterly Review,' April, 1869, p. 392. This subject is + more fully discussed in Mr. Wallace's 'Contributions to the + Theory of Natural Selection,' 1870, in which all the essays + referred to in this work are republished. The 'Essay on Man' + has been ably criticised by Prof. Claparede, one of the most + distinguished zoologists in Europe, in an article published in + the 'Bibliotheque Universelle,' June, 1870. The remark quoted + in my text will surprise every one who has read Mr. Wallace's + celebrated paper on 'The Origin of Human Races deduced from the + Theory of Natural Selection,' originally published in the + 'Anthropological Review,' May, 1864, p. clviii. I cannot here + resist quoting a most just remark by Sir J. Lubbock + ('Prehistoric Times,' 1865, p. 479) in reference to this paper, + namely, that Mr. Wallace, "with characteristic unselfishness, + ascribes it (_i.e._ the idea of natural selection) unreservedly + to Mr. Darwin, although, as is well known, he struck out the + idea independently, and published it, though not with the same + elaboration, at the same time." + + [199] Quoted by Mr. Lawson Tait in his "Law of Natural + Selection,"--'Dublin Quarterly Journal of Medical Science,' + Feb. 1869. Dr. Keller is likewise quoted to the same effect. + + [200] Owen, 'Anatomy of Vertebrates,' vol. iii. p. 71. + + [201] 'Quarterly Review,' April, 1869, p. 392. + + [202] In _Hylobates syndactylus_, as the name expresses, two of + the digits regularly cohere; and this, as Mr. Blyth informs me, + is occasionally the case with the digits of _H. agilis_, _lar_, + and _leuciscus_. + + [203] Brehm, 'Thierleben,' B. i. s. 80. + + [204] "The Hand, its mechanism," &c. 'Bridgewater Treatise,' + 1833, p. 38. + + [205] Haeckel has an excellent discussion on the steps by which + man became a biped: 'Natuerliche Schoepfungsgeschichte,' 1868, s. + 507. Dr. Buechner ('Conferences sur la Theorie Darwinienne,' + 1869, p. 135) has given good cases of the use of the foot as a + prehensile organ by man; also on the manner of progression of + the higher apes to which I allude in the following paragraph: + see also Owen ('Anatomy of Vertebrates,' vol. iii. p. 71) on + this latter subject. + + [206] "On the Primitive Form of the Skull," translated in + 'Anthropological Review,' Oct. 1868, p. 428. Owen ('Anatomy of + Vertebrates,' vol. ii. 1866, p. 551) on the mastoid processes + in the higher apes. + + [207] 'Die Grenzen der Thierwelt, eine Betrachtung zu Darwin's + Lehre,' 1868, s. 51. + + [208] Dujardin, 'Annales des Sc. Nat.' 3rd series, Zoolog. tom. + xiv. 1850, p. 203. See also Mr. Lowne, 'Anatomy and Phys. of + the _Musca vomitoria_,' 1870, p. 14. My son, Mr. F. Darwin, + dissected for me the cerebral ganglia of the _Formica rufa_. + + [209] 'Philosophical Transactions,' 1869, p. 513. + + [210] Quoted in C. Vogt's 'Lectures on Man,' Eng. translat. + 1864, p. 88, 90. Prichard, 'Phys. Hist. of Mankind,' vol. i. + 1838, p. 305. + + [211] 'Comptes Rendus des Seances,' &c. June 1, 1868. + + [212] 'The Variation of Animals and Plants under + Domestication,' vol. ii. p. 124-129. + + [213] Schaaffhausen gives from Blumenbach and Busch, the cases + of the spasms and cicatrix, in 'Anthropolog. Review,' Oct. + 1868, p. 420. Dr. Jarrold ('Anthropologia,' 1808, p. 115, 116) + adduces from Camper and from his own observations, cases of the + modification of the skull from the head being fixed in an + unnatural position. He believes that certain trades, such as + that of a shoemaker, by causing the head to be habitually held + forward, makes the forehead more rounded and prominent. + + [214] 'Variation of Animals,' &c., vol. i. p. 117 on the + elongation of the skull; p. 119, on the effect of the lopping + of one ear. + + [215] Quoted by Schaaffhausen, in 'Anthropolog. Review,' Oct. + 1868, p. 419. + + [216] Owen, 'Anatomy of Vertebrates,' vol. iii. p. 619. + + [217] Isidore Geoffroy St.-Hilaire remarks ('Hist. Nat. + Generale,' tom. ii. 1859, p. 215-217) on the head of man being + covered with long hair; also on the upper surfaces of monkeys + and of other mammals being more thickly clothed than the lower + surfaces. This has likewise been observed by various authors. + Prof. P. Gervais ('Hist. Nat. des Mammiferes,' tom. i. 1854, p. + 28), however, states that in the Gorilla the hair is thinner on + the back, where it is partly rubbed off, than on the lower + surface. + + [218] Mr. St. George Mivart, 'Proc. Zoolog. Soc.' 1865, p. 562, + 583. Dr. J. E. Gray, 'Cat. Brit. Mus.: Skeletons.' Owen, + 'Anatomy of Vertebrates,' vol. ii. p. 517. Isidore Geoffroy, + 'Hist. Nat. Gen.' tom. ii. p. 244. + + [219] 'The Variation of Animals and Plants under + Domestication,' vol. ii. p. 280, 282. + + [220] 'Primeval Man,' 1869, p. 66. + + [221] 'Anthropological Review,' May, 1864, p. clviii. + + [222] After a time the members or tribes which are absorbed + into another tribe assume, as Mr. Maine remarks ('Ancient Law,' + 1861, p. 131), that they are the co-descendants of the same + ancestors. + + [223] Morlot, 'Soc. Vaud. Sc. Nat.' 1860, p. 294. + + [224] I have given instances in my 'Variation of Animals under + Domestication,' vol. ii. p. 196. + + [225] See a remarkable series of articles on Physics and + Politics in the 'Fortnightly Review,' Nov. 1867; April 1, 1868; + July 1, 1869. + + [226] 'Origin of Civilisation,' 1870, p. 265. + + [227] Mr. Wallace gives cases in his 'Contributions to the + Theory of Natural Selection,' 1870, p. 354. + + [228] 'Ancient Law,' 1861, p. 22. For Mr. Bagehot's remarks, + 'Fortnightly Review,' April 1, 1868, p. 452. + + [229] 'The Variation of Animals and Plants under + Domestication,' vol. i. p. 309. + + [230] 'Fraser's Magazine,' Sept. 1868, p. 353. This article + seems to have struck many persons, and has given rise to two + remarkable essays and a rejoinder in the 'Spectator,' Oct. 3rd + and 17th 1868. It has also been discussed in the 'Q. Journal of + Science,' 1869, p. 152, and by Mr. Lawson Tait in the 'Dublin + Q. Journal of Medical Science,' Feb. 1869, and by Mr. E. Ray + Lankester in his 'Comparative Longevity,' 1870, p. 128. Similar + views appeared previously in the 'Australasian,' July 13, 1867. + I have borrowed ideas from several of these writers. + + [231] For Mr. Wallace, see 'Anthropolog. Review,' as before + cited. Mr. Galton in 'Macmillan's Magazine,' Aug. 1865, p. 318; + also his great work, 'Hereditary Genius,' 1870. + + [232] 'Hereditary Genius,' 1870, p. 132-140. + + [233] See the fifth and sixth columns, compiled from good + authorities, in the table given in Mr. E. R. Lankester's + 'Comparative Longevity,' 1870, p. 115. + + [234] 'Hereditary Genius,' 1870, p. 330. + + [235] 'Origin of Species' (fifth edition, 1869), p. 104. + + [236] 'Hereditary Genius,' 1870, p. 347. + + [237] E. Ray Lankester, 'Comparative Longevity,' 1870, p. 115. + The table of the intemperate is from Nelson's 'Vital + Statistics.' In regard to profligacy, see Dr. Farr, "Influence + of Marriage on Mortality," 'Nat. Assoc. for the Promotion of + Social Science,' 1858. + + [238] 'Fraser's Magazine,' Sept. 1868, p. 353. 'Macmillan's + Magazine,' Aug. 1865, p. 318. The Rev. F. W. Farrar ('Fraser's + Mag.,' Aug. 1870, p. 264) takes a different view. + + [239] "On the Laws of the Fertility of Women," in 'Transact. + Royal Soc.' Edinburgh, vol. xxiv. p. 287. See, also, Mr. + Galton, 'Hereditary Genius,' p. 352-357, for observations to + the above effect. + + [240] 'Tenth Annual Report of Births, Deaths, &c., in + Scotland,' 1867, p. xxix. + + [241] These quotations are taken from our highest authority on + such questions, namely, Dr. Farr, in his paper "On the + Influence of Marriage on the Mortality of the French People," + read before the Nat. Assoc. for the Promotion of Social + Science, 1858. + + [242] Dr. Farr, ibid. The quotations given below are extracted + from the same striking paper. + + [243] I have taken the mean of the quinquennial means, given in + 'The Tenth Annual Report of Births, Deaths, &c., in Scotland,' + 1867. The quotation from Dr. Stark is copied from an article in + the 'Daily News,' Oct. 17th, 1868, which Dr. Farr considers + very carefully written. + + [244] See the ingenious and original argument on this subject + by Mr. Galton, 'Hereditary Genius,' p. 340-342. + + [245] Mr. Greg, 'Fraser's Magazine,' Sept. 1868, p. 357. + + [246] 'Hereditary Genius,' 1870, p. 357-359. The Rev. F. H. + Farrar ('Fraser's Mag.' , Aug. 1870, p. 257) advances arguments + on the other side. Sir C. Lyell had already ('Principles of + Geology,' vol. ii. 1868, p. 489) called attention, in a + striking passage, to the evil influence of the Holy Inquisition + in having lowered, through selection, the general standard of + intelligence in Europe. + + [247] Mr. Galton, 'Macmillan's Magazine,' August, 1865, p. 325. + See, also, 'Nature,' "On Darwinism and National Life," Dec. + 1869, p. 184. + + [248] 'Last Winter in the United States,' 1868, p. 29. + + [249] 'On the Origin of Civilisation,' 'Proc. Ethnological + Soc.' Nov. 26, 1867. + + [250] 'Primeval Man,' 1869. + + [251] 'Royal Institution of Great Britain,' March 15, 1867. + Also, 'Researches into the Early History of Mankind,' 1865. + + [252] 'Primitive Marriage,' 1865. See, likewise, an excellent + article, evidently by the same author, in the 'North British + Review,' July, 1869. Also, Mr. L. H. Morgan, "A Conjectural + Solution of the Origin of the Class. System of Relationship," + in 'Proc. American Acad. of Sciences,' vol. vii. Feb. 1868. + Prof. Schaaffhausen ('Anthropolog. Review,' Oct. 1869, p. 373) + remarks on "the vestiges of human sacrifices found both in + Homer and the Old Testament." + + [253] Sir J. Lubbock, 'Prehistoric Times,' 2nd edit. 1869, + chap. xv. and xvi. _et passim_. + + [254] Dr. F. Mueller has made some good remarks to this effect + in the 'Reise der Novara: Anthropolog. Theil,' Abtheil. iii. + 1868, s. 127. + + [255] Isidore Geoffroy St.-Hilaire gives a detailed account of + the position assigned to man by various naturalists in their + classifications: 'Hist. Nat. Gen.' tom. ii. 1859, p. 170-189. + + [256] See the very interesting article, "L'Instinct chez les + Insectes," by M. George Pouchet, 'Revue des Deux Mondes,' Feb. + 1870, p. 682. + + [257] Westwood, 'Modern Class. of Insects,' vol. ii. 1840, p. + 87. + + [258] 'Proc. Zoolog. Soc.' 1869, p. 4. + + [259] 'Evidence as to Man's Place in Nature,' 1863, p. 70, _et + passim_. + + [260] Isid. Geoffroy, 'Hist. Nat. Gen.' tom. ii. 1859, p. 217. + + [261] "Ueber die Richtung der Haare," &c., Mueller's 'Archiv fuer + Anat. und Phys.' 1837, s. 51. + + [262] On the hair in Hylobates, see 'Nat. Hist. of Mammals,' by + C. L. Martin, 1841, p. 415. Also, Isid. Geoffroy on the + American monkeys and other kinds, 'Hist. Nat. Gen.' vol. ii. + 1859, p. 216, 243. Eschricht, ibid., s. 46, 55, 61. Owen, 'Anat. + of Vertebrates,' vol. iii. p. 619. Wallace, 'Contributions to + the Theory of Natural Selection,' 1870. p. 344. + + [263] 'Origin of Species,' 5th edit. 1869, p. 194. 'The + Variation of Animals and Plants under Domestication,' vol. ii. + 1868, p. 348. + + [264] 'An Introduction to the Classification of Animals,' 1869, + p. 99. + + [265] This is nearly the same classification as that + provisionally adopted by Mr. St. George Mivart ('Transact. + Philosoph. Soc.' 1867, p. 300), who, after separating the + Lemuridae, divides the remainder of the Primates into the + Hominidae, the Simiadae answering to the Catarhines, the Cebidae, + and the Hapalidae,--these two latter groups answering to the + Platyrhines. + + [266] 'Transact. Zoolog. Soc.' vol. vi. 1867, p. 214. + + [267] Mr. St. G. Mivart, 'Transact. Phil. Soc.' 1867, p. 410. + + [268] Messrs. Murie and Mivart on the Lemuroidea. 'Transact. + Zoolog. Soc.' vol. vii. 1869, p. 5. + + [269] Haeckel has come to this same conclusion. See 'Ueber die + Entstehung des Menschengeschlechts,' in Virchow's 'Sammlung. + gemein. wissen. Vortraege,' 1868, s. 61. Also his 'Natuerliche + Schoepfungsgeschichte,' 1868, in which he gives in detail his + views on the genealogy of man. + + [270] 'Anthropological Review,' April, 1867, p. 236. + + [271] 'Elements of Geology,' 1865, p. 583-585. 'Antiquity of + Man', 1863; p. 145. + + [272] 'Man's Place in Nature,' p. 105. + + [273] Elaborate tables are given in his 'Generelle Morphologie' + (B. ii. s. cliii. and s. 425); and with more especial reference + to man in his 'Natuerliche Schoepfungsgeschichte,' 1868. Prof. + Huxley, in reviewing this latter work ('The Academy,' 1869, p. + 42) says, that he considers the phylum or lines of descent of + the Vertebrata to be admirably discussed by Haeckel, although he + differs on some points. He expresses, also, his high estimate + of the value of the general tenor and spirit of the whole work. + + [274] 'Palaeontology,' 1860, p. 199. + + [275] I had the satisfaction of seeing, at the Falkland + Islands, in April, 1833, and therefore some years before any + other naturalist, the locomotive larvae of a compound Ascidian, + closely allied to, but apparently generically distinct from, + Synoicum. The tail was about five times as long as the oblong + head, and terminated in a very fine filament. It was plainly + divided, as sketched by me under a simple microscope, by + transverse opaque partitions, which I presume represent the + great cells figured by Kowalevsky. At an early stage of + development the tail was closely coiled round the head of the + larva. + + [276] 'Memoires de l'Acad. des Sciences de St. Petersbourg,' + tom. x. No. 15, 1866. + + [277] This is the conclusion of one of the highest authorities + in comparative anatomy, namely, Prof. Gegenbaur: 'Grundzuege der + vergleich. Anat.' 1870, s. 876. The result has been arrived at + chiefly from the study of the Amphibia; but it appears from the + researches of Waldeyer (as quoted in Humphry's 'Journal of + Anat. and Phys.' 1869, p. 161), that the sexual organs of even + "the higher vertebrata are, in their early condition, + hermaphrodite." Similar views have long been held by some + authors, though until recently not well based. + + [278] The male Thylacinus offers the best instance. Owen, + 'Anatomy of Vertebrates,' vol. iii. p. 771. + + [279] Serranus is well known often to be in an hermaphrodite + condition; but Dr. Guenther informs me that he is convinced that + this is not its normal state. Descent from an ancient + androgynous prototype would, however, naturally favour and + explain, to a certain extent, the recurrence of this condition + in these fishes. + + [280] Mr. Lockwood believes (as quoted in 'Quart. Journal of + Science,' April, 1868, p. 269), from what he has observed of + the development of Hippocampus, that the walls of the abdominal + pouch of the male in some way afford nourishment. On male + fishes hatching the ova in their mouths, see a very interesting + paper by Prof. Wyman, in 'Proc. Boston Soc. of Nat. Hist.' + Sept. 15, 1857; also Prof. Turner, in 'Journal of Anat. and + Phys.' Nov. 1, 1866, p. 78. Dr. Guenther has likewise described + similar cases. + + [281] All vital functions tend to run their course in fixed and + recurrent periods, and with tidal animals the periods would + probably be lunar; for such animals must have been left dry or + covered deep with water,--supplied with copious food or + stinted,--during endless generations, at regular lunar + intervals. If then the Vertebrata are descended from an animal + allied to the existing tidal Ascidians, the mysterious fact, + that with the higher and now terrestrial Vertebrata, not to + mention other classes, many normal and abnormal vital processes + run their course according to lunar periods, is rendered + intelligible. A recurrent period, if approximately of the right + duration, when once gained, would not, as far as we can judge, + be liable to be changed; consequently it might be thus + transmitted during almost any number of generations. This + conclusion, if it could be proved sound, would be curious; for + we should then see that the period of gestation in each mammal, + and the hatching of each bird's eggs, and many other vital + processes, still betrayed the primordial birthplace of these + animals. + + [282] 'History of India,' 1841, vol. i. p. 323. Father Ripa + makes exactly the same remark with respect to the Chinese. + + [283] A vast number of measurements of Whites, Blacks, and + Indians, are given in the 'Investigations in the Military and + Anthropolog. Statistics of American Soldiers,' by B. A. Gould, + 1869, p. 298-358; on the capacity of the lungs, p. 471. See + also the numerous and valuable tables, by Dr. Weisbach, from + the observations of Dr. Scherzer and Dr. Schwarz, in the 'Reise + der Novara: Anthropolog. Theil,' 1867. + + [284] See, for instance, Mr. Marshall's account of the brain of + a Bush-woman, in 'Phil. Transact.' 1864, p. 519. + + [285] Wallace, 'The Malay Archipelago,' vol. ii. 1869, p. 178. + + [286] With respect to the figures in the famous Egyptian caves + of Abou-Simbel, M. Pouchet says ('The Plurality of the Human + Races,' Eng. translat. 1864, p. 50), that he was far from + finding recognisable representations of the dozen or more + nations which some authors believe that they can recognise. + Even some of the most strongly-marked races cannot be + identified with that degree of unanimity which might have been + expected from what has been written on the subject. Thus + Messrs. Nott and Gliddon ('Types of Mankind,' p. 148) state + that Rameses II., or the Great, has features superbly European; + whereas Knox, another firm believer in the specific distinction + of the races of man ('Races of Man,' 1850, p. 201), speaking of + young Memnon (the same person with Rameses II., as I am + informed by Mr. Birch) insists in the strongest manner that he + is identical in character with the Jews of Antwerp. Again, + whilst looking in the British Museum with two competent judges, + officers of the establishment, at the statue of Amunoph III., + we agreed that he had a strongly negro cast of features; but + Messrs. Nott and Gliddon (ibid. p. 146, fig. 53) describe him + as "a hybrid, but not of negro intermixture." + + [287] As quoted by Nott and Gliddon, 'Types of Mankind,' 1854, + p. 439. They give also corroborative evidence; but C. Vogt + thinks that the subject requires further investigation. + + [288] "Diversity of Origin of the Human Races," in the + 'Christian Examiner,' July, 1850. + + [289] 'Transact. B. Soc. of Edinburgh,' vol. xxii. 1861, p. + 567. + + [290] 'On the Phenomena of Hybridity in the Genus Homo,' Eng. + translat. 1864. + + [291] See the interesting letter by Mr. T. A. Murray, in the + 'Anthropolog. Review,' April, 1868, p. liii. In this letter + Count Strzelecki's statement, that Australian women who have + borne children to a white man are afterwards sterile with their + own race, is disproved. M. A. de Quatrefages has also collected + ('Revue des Cours Scientifiques,' March, 1869, p. 239) much + evidence that Australians and Europeans are not sterile when + crossed. + + [292] 'An Examination of Prof. Agassiz's Sketch of the Nat. + Provinces of the Animal World,' Charleston, 1855, p. 44. + + [293] 'Military and Anthropolog. Statistics of American + Soldiers,' by B. A. Gould, 1869, p. 319. + + [294] 'The Variation of Animals and Plants under + Domestication,' vol. ii. p. 109. I may here remind the reader + that the sterility of species when crossed is not a + specially-acquired quality; but, like the incapacity of certain + trees to be grafted together, is incidental on other acquired + differences. The nature of these differences is unknown, but + they relate more especially to the reproductive system, and + much less to external structure or to ordinary differences in + constitution. One important element in the sterility of crossed + species apparently lies in one or both having been long + habituated to fixed conditions; for we know that changed + conditions have a special influence on the reproductive system, + and we have good reason to believe (as before remarked) that + the fluctuating conditions of domestication tend to eliminate + that sterility which is so general with species in a natural + state when crossed. It has elsewhere been shewn by me (ibid. + vol. ii. p. 185, and 'Origin of Species,' 5th edit. p. 317) + that the sterility of crossed species has not been acquired + through natural selection: we can see that when two forms have + already been rendered very sterile, it is scarcely possible + that their sterility should be augmented by the preservation or + survival of the more and more sterile individuals; for as the + sterility increases fewer and fewer offspring will be produced + from which to breed, and at last only single individuals will + be produced, at the rarest intervals. But there is even a + higher grade of sterility than this. Both Gaertner and Koelreuter + have proved that in genera of plants including numerous + species, a series can be formed from species which when crossed + yield fewer and fewer seeds, to species which never produce a + single seed, but yet are affected by the pollen of the other + species, for the germen swells. It is here manifestly + impossible to select the more sterile individuals, which have + already ceased to yield seeds; so that the acme of sterility, + when the germen alone is affected, cannot be gained through + selection. This acme, and no doubt the other grades of + sterility, are the incidental results of certain unknown + differences in the constitution of the reproductive system of + the species which are crossed. + + [295] 'The Variation of Animals,' &c., vol. ii. p. 92. + + [296] M. de Quatrefages has given ('Anthropolog. Review,' Jan. + 1869, p. 22) an interesting account of the success and energy + of the Paulistas in Brazil, who are a much crossed race of + Portuguese and Indians, with a mixture of the blood of other + races. + + [297] For instance with the aborigines of America and + Australia. Prof. Huxley says ('Transact. Internat. Congress of + Prehist. Arch.' 1868. p. 105) that the skulls of many South + Germans and Swiss are "as short and as broad as those of the + Tartars," &c. + + [298] See a good discussion on this subject in Waitz, + 'Introduct. to Anthropology,' Eng. translat. 1863, p. 198-208, + 227. I have taken some of the above statements from H. Tuttle's + 'Origin and Antiquity of Physical Man,' Boston, 1866, p. 35. + + [299] Prof. Naegeli has carefully described several striking + cases in his 'Botanische Mittheilungen,' B. ii. 1866, s. + 294-369. Prof. Asa Gray has made analogous remarks on some + intermediate forms in the Compositae of N. America. + + [300] 'Origin of Species,' 5th edit. p. 68. + + [301] See Prof. Huxley to this effect in the 'Fortnightly + Review,' 1865, p. 275. + + [302] 'Lectures on Man,' Eng. translat. 1864, p. 468. + + [303] 'Die Racen des Schweines,' 1860, s. 46. 'Vorstudien fuer + Geschichte, &c., Schweineschaedel,' 1864, s. 104. With respect + to cattle, see M. de Quatrefages, 'Unite de l'Espece Humaine,' + 1861, p. 119. + + [304] Tylor's 'Early History of Mankind,' 1865; for the + evidence with respect to gesture-language, see p. 54. Lubbock's + 'Prehistoric Times,' 2nd edit. 1869. + + [305] 'The Primitive Inhabitants of Scandinavia,' Eng. + translat. edited by Sir J. Lubbock, 1868, p. 104. + + [306] Hodder M. Westropp, on Cromlechs, &c., 'Journal of + Ethnological Soc.' as given in 'Scientific Opinion,' June 2nd. + 1869, p. 3. + + [307] 'Journal of Researches: Voyage of the "Beagle,"' p. 46. + + [308] 'Prehistoric Times,' 1869, p. 574. + + [309] Translation in 'Anthropological Review,' Oct. 1868, p. + 431. + + [310] 'Transact. Internat. Congress of Prehistoric Arch.' 1868, + p. 172-175. See also Broca (translation) in 'Anthropological + Review,' Oct. 1868, p. 410. + + [311] Dr. Gerland, 'Ueber das Aussterben der Naturvoelker,' + 1868, s. 82. + + [312] Gerland (ibid. s. 12) gives facts in support of this + statement. + + [313] See remarks to this effect in Sir H. Holland's 'Medical + Notes and Reflections,' 1839, p. 390. + + [314] I have collected ('Journal of Researches, Voyage of the + "Beagle,"' p. 435) a good many cases bearing on this subject: + see also Gerland, ibid. s. 8. Poeppig speaks of the "breath of + civilisation as poisonous to savages." + + [315] Sproat, 'Scenes and Studies of Savage Life,' 1868, p. + 284. + + [316] Bagehot, "Physics and Politics," 'Fortnightly Review,' + April 1, 1868, p. 455. + + [317] "On Anthropology," translation, 'Anthropolog. Review,' + Jan. 1868, p. 38. + + [318] 'The Annals of Rural Bengal,' 1868, p. 134. + + [319] 'The Variation of Animals and Plants under + Domestication,' vol. ii. p. 95. + + [320] Pallas, 'Act. Acad. St. Petersburgh,' 1780, part ii. p. + 69. He was followed by Rudolphi, in his 'Beytraege zur + Anthropologie,' 1812. An excellent summary of the evidence is + given by Godron, 'De l'Espece,' 1859, vol. ii. p. 246, &c. + + [321] Sir Andrew Smith, as quoted by Knox, 'Races of Man,' + 1850, p. 473. + + [322] See De Quatrefages on this head, 'Revue des Cours + Scientifiques,' Oct. 17, 1868, p. 731. + + [323] Livingstone's 'Travels and Researches in S. Africa,' + 1857, p. 338, 329. D'Orbigny, as quoted by Godron, 'De + l'Espece,' vol. ii. p. 266. + + [324] See a paper read before the Royal Soc. in 1813, and + published in his Essays in 1818. I have given an account of Dr. + Wells' views in the Historical Sketch (p. xvi) to my 'Origin of + Species.' Various cases of colour correlated with + constitutional peculiarities are given in my 'Variation of + Animals under Domestication,' vol. ii. p. 227, 335. + + [325] See, for instance, Nott and Gliddon, 'Types of Mankind,' + p. 68. + + [326] Major Tulloch, in a paper read before the Statistical + Society, April 20th, 1840, and given in the 'Athenaeum,' 1840, + p. 353. + + [327] 'The Plurality of the Human Race' (translat.), 1864, p. + 60. + + [328] Quatrefages, 'Unite de l'Espece Humaine,' 1861, p. 205. + Waitz, 'Introduct. to Anthropology,' translat. vol. i. 1863, p. + 124. Livingstone gives analogous cases in his 'Travels.' + + [329] In the spring of 1862 I obtained permission from the + Director-General of the Medical department of the Army, to + transmit to the surgeons of the various regiments on foreign + service a blank table, with the following appended remarks, but + I have received no returns. "As several well-marked cases have + been recorded with our domestic animals of a relation between + the colour of the dermal appendages and the constitution; and + it being notorious that there is some limited degree of + relation between the colour of the races of man and the climate + inhabited by them; the following investigation seems worth + consideration. Namely, whether there is any relation in + Europeans between the colour of their hair, and their liability + to the diseases of tropical countries. If the surgeons of the + several regiments, when stationed in unhealthy tropical + districts, would be so good as first to count, as a standard of + comparison, how many men, in the force whence the sick are + drawn, have dark and light-coloured hair, and hair of + intermediate or doubtful tints; and if a similar account were + kept by the same medical gentlemen, of all the men who suffered + from malarious and yellow fevers, or from dysentery, it would + soon be apparent, after some thousand cases had been tabulated, + whether there exists any relation between the colour of the + hair and constitutional liability to tropical diseases. Perhaps + no such relation would be discovered, but the investigation is + well worth making. In case any positive result were obtained, + it might be of some practical use in selecting men for any + particular service. Theoretically the result would be of high + interest, as indicating one means by which a race of men + inhabiting from a remote period an unhealthy tropical climate, + might have become dark-coloured by the better preservation of + dark-haired or dark-complexioned individuals during a long + succession of generations." + + [330] 'Anthropological Review,' Jan. 1866, p. xxi. + + [331] See, for instance, Quatrefages ('Revue des Cours + Scientifiques,' Oct. 10, 1868, p. 724) on the effects of + residence in Abyssinia and Arabia, and other analogous cases. + Dr. Rolle ('Der Mensch, seine Abstammung,' &c., 1865, s. 99) + states, on the authority of Khanikof, that the greater number + of German families settled in Georgia, have acquired in the + course of two generations dark hair and eyes. Mr. D. Forbes + informs me that the Quichuas in the Andes vary greatly in + colour, according to the position of the valleys inhabited by + them. + + [332] Harlan, 'Medical Researches,' p. 532. Quatrefages ('Unite + de l'Espece Humaine,' 1861, p. 128) has collected much evidence + on this head. + + [333] See Prof. Schaaffhausen, translat. in 'Anthropological + Review,' Oct. 1868, p. 429. + + [334] Mr. Catlin states ('N. American Indians,' 3rd edit. 1842, + vol. i. p. 49) that in the whole tribe of the Mandans, about + one in ten or twelve of the members of all ages and both sexes + have bright silvery grey hair, which is hereditary. Now this + hair is as coarse and harsh as that of a horse's mane, whilst + the hair of other colours is fine and soft. + + [335] On the odour of the skin, Godron, 'Sur l'Espece,' tom. + ii. p. 217. On the pores in the skin, Dr. Wilckens, 'Die + Aufgaben der landwirth. Zootechnik,' 1869, s. 7. + + [336] Westwood, 'Modern Class. of Insects,' vol. ii. 1810, p. + 541. In regard to the statement about Tanais, mentioned below, + I am indebted to Fritz Mueller. + + [337] Kirby and Spence, 'Introduction to Entomology,' vol. iii. + 1826, p. 309. + + [338] Even with those of plants in which the sexes are + separate, the male flowers are generally mature before the + female. Many hermaphrodite plants are, as first shewn by C. K. + Sprengel, dichogamous; that is, their male and female organs + are not ready at the same time, so that they cannot be + self-fertilised. Now with such plants the pollen is generally + mature in the same flower before the stigma, though there are + some exceptional species in which the female organs are mature + before the male. + + [339] I have received information, hereafter to be given, to + this effect with respect to poultry. Even with birds, such as + pigeons, which pair for life, the female, as I hear from Mr. + Jenner Weir, will desert her mate if he is injured or grows + weak. + + [340] On the Gorilla, Savage and Wyman, 'Boston Journal of Nat. + Hist.' vol. v. 1845-47, p. 423. On Cynocephalus, Brehm, + 'Illust. Thierleben,' B. i. 1864, s. 77. On Mycetes, Rengger, + 'Naturgesch.: Saeugethiere von Paraguay,' 1830, s. 14, 20. On + Cebus, Brehm, ibid. s. 108. + + [341] Pallas, 'Spicilegia Zoolog.' Fasc. xii. 1777, p. 29. Sir + Andrew Smith, 'Illustrations of the Zoology of S. Africa,' + 1849, pl. 29, on the Kobus. Owen, in his 'Anatomy of + Vertebrates' (vol. iii. 1868, p. 633) gives a table + incidentally showing which species of Antelopes pair and which + are gregarious. + + [342] Dr. Campbell, in 'Proc. Zoolog. Soc.' 1869, p. 138. See + also an interesting paper, by Lieut. Johnstone, in 'Proc. + Asiatic Soc. of Bengal,' May, 1868. + + [343] 'The Ibis,' vol. iii. 1861, p. 133, on the Progne + Widow-bird. See also on the Vidua axillaris, ibid. vol. ii. + 1860, p. 211. On the polygamy of the Capercailzie and Great + Bustard, see L. Lloyd, 'Game Birds of Sweden,' 1867, p. 19, and + 182. Montagu and Selby speak of the Black Grouse as polygamous + and of the Red Grouse as monogamous. + + [344] The Rev. E. S. Dixon, however, speaks positively + ('Ornamental Poultry,' 1848, p. 76) about the eggs of the + guinea-fowl being infertile when more than one female is kept + with the same male. + + [345] Noel Humphreys, 'River Gardens,' 1857. + + [346] Kirby and Spence, 'Introduction to Entomology,' vol. iii. + 1826, p. 342. + + [347] One parasitic Hymenopterous insect (Westwood, 'Modern + Class. of Insects,' vol. ii, p. 160) forms an exception to the + rule, as the male has rudimentary wings, and never quits the + cell in which it is born, whilst the female has well-developed + wings. Audouin believes that the females are impregnated by the + males which are born in the same cells with them; but it is + much more probable that the females visit other cells, and thus + avoid close inter-breeding. We shall hereafter meet with a few + exceptional cases, in various classes, in which the female, + instead of the male, is the seeker and wooer. + + [348] 'Essays and Observations,' edited by Owen, vol. i. 1861, + p. 194. + + [349] Prof. Sachs ('Lehrbuch der Botanik,' 1870, s. 633) in + speaking of the male and female reproductive cells, remarks, + "verhaelt sich die eine bei der Vereinigung activ, ... die + andere erscheint bei der Vereinigung passiv." + + [350] 'Reise der Novara: Anthropolog. Theil,' 1867, s. 216-269. + The results were calculated by Dr. Weisbach from measurements + made by Drs. K. Scherzer and Schwarz. On the greater + variability of the males of domesticated animals, see my + 'Variation of Animals and Plants under Domestication,' vol. ii. + 1868, p. 75. + + [351] 'Proceedings Royal Soc.' vol. xvi. July, 1868, p. 519 and + 524. + + [352] 'Proc. Royal Irish Academy,' vol. x. 1868, p. 123. + + [353] 'Massachusetts Medical Soc.' vol. ii. No. 3, 1808, p. 9. + + [354] 'The Variation of Animals and Plants under + Domestication,' vol. ii. 1868, p. 75. In the last chapter but + one, the provisional hypothesis of pangenesis, above alluded + to, is fully explained. + + [355] These facts are given on the high authority of a great + breeder, Mr. Teebay, in Tegetmeier's 'Poultry Book,' 1868, p. + 158. On the characters of chickens of different breeds, and on + the breeds of the pigeon, alluded to in the above paragraph, + see 'Variation of Animals,' &c., vol. i. p. 160, 249; vol. ii. + p. 77. + + [356] 'Novae species Quadrupedum e Glirium ordine,' 1778, p. 7. + On the transmission of colour by the horse, see 'Variation of + Animals, &c. under Domestication,' vol. i. p. 21. Also vol. ii. + p. 71, for a general discussion on Inheritance as limited by + Sex. + + [357] Dr. Chapuis, 'Le Pigeon Voyageur Belge,' 1865, p. 87. + Boitard et Corbie, 'Les Pigeons de Voliere,' &c., 1824, p. 173. + + [358] References are given in my 'Variation of Animals under + Domestication,' vol. ii. p. 72. + + [359] I am much obliged to Mr. Cupples for having made + enquiries for me in regard to the Roebuck and Red Deer of + Scotland from Mr. Robertson, the experienced head-forester to + the Marquis of Breadalbane. In regard to Fallow-deer, I am + obliged to Mr. Eyton and others for information. For the + _Cervus alces_ of N. America, see 'Land and Water,' 1868, p. + 221 and 254; and for the _C. Virginianus_ and _strongyloceros_ + of the same continent, see J. D. Caton, in 'Ottawa Acad. of + Nat. Sc.' 1868, p. 13. For _Cervus Eldi_ of Pegu, see Lieut. + Beavan, 'Proc. Zoolog. Soc.' 1867, p. 762. + + [360] _Antilocapra Americana._ Owen, 'Anatomy of Vertebrates,' + vol. iii. p. 627. + + [361] I have been assured that the horns of the sheep in North + Wales can always be felt, and are sometimes even an inch in + length, at birth. With cattle Youatt says ('Cattle,' 1834, p. + 277) that the prominence of the frontal bone penetrates the + cutis at birth, and that the horny matter is soon formed over + it. + + [362] I am greatly indebted to Prof. Victor Carus for having + made inquiries for me, from the highest authorities, with + respect to the merino sheep of Saxony. On the Guinea coast of + Africa there is a breed of sheep in which, as with merinos, the + rams alone bear horns; and Mr. Winwood Reade informs me that in + the one case observed, a young ram born on Feb. 10th first + showed horns on March 6th, so that in this instance the + development of the horns occurred at a later period of life, + conformably with our rule, than in the Welsh sheep, in which + both sexes are horned. + + [363] In the common peacock (_Pavo cristatus_) the male alone + possesses spurs, whilst both sexes of the Java peacock (_P. + muticus_) offer the unusual case of being furnished with spurs. + Hence I fully expected that in the latter species they would + have been developed earlier in life than in the common peacock; + but M. Hegt of Amsterdam informs me, that with young birds of + the previous year, belonging to both species, compared on April + 23rd, 1869, there was no difference in the development of the + spurs. The spurs, however, were as yet represented merely by + slight knobs or elevations. I presume that I should have been + informed if any difference in the rate of development had + subsequently been observed. + + [364] In some other species of the Duck Family the speculum in + the two sexes differs in a greater degree; but I have not been + able to discover whether its full development occurs later in + life in the males of such species, than in the male of the + common duck, as ought to be the case according to our rule. + With the allied _Mergus cucullatus_ we have, however, a case of + this kind: the two sexes differ conspicuously in general + plumage, and to a considerable degree in the speculum, which is + pure white in the male and greyish-white in the female. Now the + young males at first resemble, in all respects, the female, and + have a greyish-white speculum, but this becomes pure white at + an earlier age than that at which the adult male acquires his + other more strongly-marked sexual differences in plumage: see + Audubon, 'Ornithological Biography,' vol. iii. 1835, p. + 249-250. + + [365] 'Das Ganze der Taubenzucht,' 1837, s. 21, 24. For the + case of the streaked pigeons, see Dr. Chapuis, 'Le Pigeon + Voyageur Belge.' 1865, p. 87. + + [366] For full particulars and references on all these points + respecting the several breeds of the Fowl, see 'Variation of + Animals and Plants under Domestication,' vol. i. p. 250, 256. + In regard to the higher animals, the sexual differences which + have arisen under domestication are described in the same work + under the head of each species. + + [367] 'Twenty-ninth Annual Report of the Registrar-General for + 1866.' In this report (p. xii) a special decennial table is + given. + + [368] For Norway and Russia, see abstract of Prof. Faye's + researches, in 'British and Foreign Medico-Chirurg. Review,' + April, 1867, p. 343, 345. For France, the 'Annuaire pour l'An + 1867.' p. 213. + + [369] In regard to the Jews, see M. Thury, 'La Loi de + Production des Sexes,' 1863, p. 25. + + [370] Babbage, 'Edinburgh Journal of Science,' 1829, vol. i. p. + 88; also p. 90, on still-born children. On illegitimate + children in England, see 'Report of Registrar-General for + 1866,' p. xv. + + [371] 'British and Foreign Medico-Chirurg. Review,' April, + 1867, p. 343. Dr. Stark also remarks ('Tenth Annual Report of + Births, Deaths, &c., in Scotland,' 1867, p. xxviii) that "These + examples may suffice to shew that, at almost every stage of + life, the males in Scotland have a greater liability to death + and a higher death-rate than the females. The fact, however, of + this peculiarity being most strongly developed at that + infantile period of life when the dress, food, and general + treatment of both sexes are alike, seems to prove that the + higher male death-rate is an impressed, natural, and + constitutional peculiarity due to sex alone." + + [372] With the savage Guaranys of Paraguay, according to the + accurate Azara ('Voyages dans l'Amerique merid.' tom. ii. 1809, + p. 60, 179), the women in proportion to the men are as 14 to + 13. + + [373] Leuckart in Wagner, 'Handwoerterbuch der Phys.' B. iv. + 1853, s. 774. + + [374] Anthropological Review, April, 1870, p. cviii. + + [375] During the last eleven years a record has been kept of + the number of mares which have proved barren or prematurely + slipped their foals; and it deserves notice, as shewing how + infertile these highly-nurtured and rather closely-interbred + animals have become, that not far from one-third of the mares + failed to produce living foals. Thus during 1866, 809 male + colts and 816 female colts were born, and 743 mares failed to + produce offspring. During 1867, 836 males and 902 females were + born, and 794 mares failed. + + [376] I am much indebted to Mr. Cupples for having procured for + me the above returns from Scotland, as well as some of the + following returns on cattle. Mr. R. Elliot, of Laighwood, first + called my attention to the premature deaths of the males,--a + statement subsequently confirmed by Mr. Aitchison and others. + To this latter gentleman, and to Mr. Payan, I owe my thanks for + the larger returns on sheep. + + [377] Bell, 'History of British Quadrupeds,' p. 100. + + [378] 'Illustrations of the Zoology of S. Africa,' 1849, pl. + 29. + + [379] Brehm ('Illust. Thierleben,' B. iv. s. 990) comes to the + same conclusion. + + [380] On the authority of L. Lloyd, 'Game Birds of Sweden,' + 1867, p. 12, 132. + + [381] 'Nat. Hist. of Selbourne,' letter xxix. edit. of 1825, + vol. i. p. 139. + + [382] Mr. Jenner Weir received similar information, on making + enquiries during the following year. To shew the number of + chaffinches caught, I may mention that in 1869 there was a + match between two experts; and one man caught in a day 62, and + another 40, male chaffinches. The greatest number ever caught + by one man in a single day was 70. + + [383] 'Ibis,' vol. ii. p. 260, as quoted in Gould's + 'Trochilidae,' 1861, p. 52. For the foregoing proportions, I am + indebted to Mr. Salvin for a table of his results. + + [384] 'Ibis,' 1860, p. 137; and 1867, p. 369. + + [385] 'Ibis,' 1862, p. 137. + + [386] Leuckart quotes Bloch (Wagner, 'Handwoerterbuch der Phys.' + B. iv. 1853, s. 775), that with fish there are twice as many + males as females. + + [387] Quoted in the 'Farmer,' March 18, 1869, p. 369. + + [388] 'The Stormontfield Piscicultural Experiments,' 1866, p. + 23. The 'Field' newspaper, June 29th, 1867. + + [389] 'Land and Water,' 1868, p. 41. + + [390] Yarrell, 'Hist. British Fishes,' vol. i. 1836, p. 307; on + the _Cyprinus carpio_, p. 331; on the _Tinca vulgaris_, p. 331; + on the _Abramis brama_, p. 336. See, for the minnow (_Leuciscus + phoxinus_), 'Loudon's Mag. of Nat. Hist.' vol. v. 1832, p. 682. + + [391] Leuckart quotes Meinecke (Wagner, 'Handwoerterbuch der + Phys.' B. iv. 1853, s. 775) that with Butterflies the males are + three or four times as numerous as the females. + + [392] 'The Naturalist on the Amazons,' vol. ii. 1863, p. 228, + 347. + + [393] Four of these cases are given by Mr. Trimen in his + 'Rhopalocera Africae Australis.' + + [394] Quoted by Trimen, 'Transact. Ent. Soc.' vol. v. part iv. + 1866, p. 330. + + [395] 'Transact. Linn. Soc.' vol. xxv. p. 37. + + [396] 'Proc. Entomolog. Soc.' Feb. 17th, 1868. + + [397] Quoted by Dr. Wallace in 'Proc. Ent. Soc.' 3rd series, + vol. v. 1867, p. 487. + + [398] Blanchard, 'Metamorphoses, Moeurs des Insectes,' 1868, p. + 225-226. + + [399] 'Lepidopteren-Doubblettren Liste,' Berlin, No. x. 1866. + + [400] This naturalist has been so kind as to send me some + results from former years, in which the females seemed to + preponderate; but so many of the figures were estimates, that I + found it impossible to tabulate them. + + [401] Guenther's 'Record of Zoological Literature,' 1867, p. + 260. On the excess of female Lucanus, ibid. p. 250. On the + males of Lucanus in England, Westwood, 'Modern Class. of + Insects,' vol. i. p. 187. On the Siagonium, ibid. p. 172. + + [402] Walsh, in 'The American Entomologist,' vol. i. 1869, p. + 103. F. Smith, 'Record of Zoological Literature,' 1867, p. 328. + + [403] 'Farm Insects,' p. 45-46. + + [404] 'Observations on N. American Neuroptera,' by H. Hagen and + B. D. Walsh, 'Proc. Ent. Soc. Philadelphia,' Oct. 1863, p. 168, + 223, 239. + + [405] 'Proc. Ent. Soc. London,' Feb. 17, 1868. + + [406] Another great authority in this class, Prof. Thorell of + Upsala ('On European Spiders,' 1869-70, part i. p. 205) speaks + as if female spiders were generally commoner than the males. + + [407] See, on this subject, Mr. Pickard-Cambridge, as quoted in + 'Quarterly Journal of Science,' 1868, p. 429. + + [408] I have often been struck with the fact, that in several + species of Primula the seeds in the capsules which contained + only a few were very much larger than the numerous seeds in the + more productive capsules. + + [409] 'Principles of Biology,' vol. ii. 1867, chaps. ii.-xi. + + [410] 'De l'Espece et de la Class.' &c., 1869, p. 106. + + [411] See, for instance, the account which I have given in my + 'Journal of Researches,' 1845, p. 7. + + [412] I have given ('Geolog. Observations on Volcanic Islands,' + 1844, p. 53) a curious instance of the influence of light on + the colours of a frondescent incrustation, deposited by the + surf on the coast-rocks of Ascension, and formed by the + solution of triturated sea-shells. + + [413] 'Facts and Arguments for Darwin,' English translat. 1869, + p. 20. See the previous discussion on the olfactory threads. + Sars has described a somewhat analogous case (as quoted in + 'Nature,' 1870, p. 455) in a Norwegian crustacean, the + _Pontoporeia affinis_. + + [414] See Sir J. Lubbock in 'Annals. and Mag. of Nat. Hist.' + vol. xi. 1853, pl. i. and x.; and vol. xii. (1853) pl. vii. See + also Lubbock in 'Transact. Ent. Soc.' vol. iv. new series, + 1856-1858, p. 8. With respect to the zigzagged antennae + mentioned below, see Fritz Mueller, 'Facts and Arguments for + Darwin' 1869, p. 40, foot-note. + + [415] See a paper by Mr. C. Spence Bate, with figures, in + 'Proc. Zoolog. Soc.' 1868, p. 363; and on the nomenclature of + the genus, ibid. p. 585. I am greatly indebted to Mr. Spence + Bate for nearly all the above statements with respect to the + chelae of the higher crustaceans. + + [416] 'Hist. Nat. des Crust.' tom. ii. 1837, p. 50. + + [417] Fritz Mueller, 'Facts and Arguments for Darwin,' 1869, p. + 25-28. + + [418] 'Travels in the Interior of Brazil,' 1846, p. 111. I have + given, in my 'Journal of Researches,' p. 463, an account of the + habits of the Birgos. + + [419] Mr. Ch. Fraser, in 'Proc. Zoolog. Soc.' 1869, p. 3. I am + indebted to Mr. Bate for the statement from Dr. Power. + + [420] Claus, 'Die freilebenden Copepoden,' 1863, s. 35. + + [421] 'Facts and Arguments,' &c., p. 79. + + [422] 'A History of the Spiders of Great Britain,' 1861-64. For + the following facts, see p. 102, 77, 88. + + [423] Aug. Vinson ('Araneides des Iles de la Reunion,' pl. vi. + figs. 1 and 2) gives a good instance of the small size of the + male in _Epeira nigra_. In this species, as I may add, the male + is testaceous and the female black with legs banded with red. + Other even more striking cases of inequality in size between + the sexes have been recorded ('Quarterly Journal of Science,' + 1868, July, p. 429); but I have not seen the original accounts. + + [424] Kirby and Spence, 'Introduction to Entomology,' vol. i. + 1818, p. 280. + + [425] Theridion (Asagena, Sund.) serratipes, 4-punctatum et + guttatum; see Westring, in Kroyer, 'Naturhist. Tidskrift,' vol. + iv. 1842-1843, p. 349; and vol. ii. 1846-1849, p. 342. See, + also, for other species, 'Araneae Svecicae,' p. 184. + + [426] Walckenaer et P. Gervais, 'Hist. Nat. des Insectes: + Apteres,' tom. iv. 1847, p. 17, 19, 68. + + [427] Sir J. Lubbock, 'Transact. Linnean Soc.' vol. xxv. 1866, + p. 484. With respect to the Mutillidae see Westwood, 'Modern + Class. of Insects,' vol. ii. p. 213. + + [428] These organs in the male often differ in closely-allied + species, and afford excellent specific characters. But their + importance, under a functional point of view, as Mr. E. + MacLachlan has remarked to me, has probably been overrated. It + has been suggested, that slight differences in these organs + would suffice to prevent the intercrossing of well-marked + varieties or incipient species, and would thus aid in their + development. That this can hardly be the case, we may infer + from the many recorded cases (see for instance, Bronn, + 'Geschichte der Natur,' B. ii. 1843, s. 164; and Westwood, + 'Transact. Ent. Soc.' vol. iii. 1842, p. 195) of distinct + species having been observed in union. Mr. MacLachlan informs + me (vide 'Stett. Ent. Zeitung,' 1867, s. 155) that when several + species of Phryganidae, which present strongly-pronounced + differences of this kind, were confined together by Dr. Aug. + Meyer, _they coupled_, and one pair produced fertile ova. + + [429] 'The Practical Entomologist,' Philadelphia, vol. ii. May, + 1867, p. 88. + + [430] Mr. Walsh, ibid. p. 107. + + [431] 'Modern Classification of Insects,' vol. ii. 1840, p. + 206, 205. Mr. Walsh, who called my attention to this double use + of the jaws, says that he has repeatedly observed this fact. + + [432] We have here a curious and inexplicable case of + dimorphism, for some of the females of four European species of + Dytiscus, and of certain species of Hydroporus, have their + elytra smooth; and no intermediate gradations between sulcated + or punctured and quite smooth elytra have been observed. See + Dr. H. Schaum, as quoted in the 'Zoologist,' vol. v.-vi. + 1847-48, p. 1896. Also Kirby and Spence, 'Introduction to + Entomology,' vol. iii. 1826, p. 305. + + [433] Westwood, 'Modern Class.' vol. ii. p. 193. The following + statement about Penthe, and others in inverted commas, are + taken from Mr. Walsh, 'Practical Entomologist,' Philadelphia, + vol. ii. p. 88. + + [434] Kirby and Spence, 'Introduct.' &c., vol. iii. p. 332-336. + + [435] 'Insecta Maderensia,' 1854, p. 20. + + [436] E. Doubleday, 'Annals and Mag. of Nat. Hist.' vol. i. + 1848, p. 379. I may add that the wings in certain Hymenoptera + (see Shuckard, 'Fossorial Hymenop.' 1837, p. 39-43) differ in + neuration according to sex. + + [437] H. W. Bates, in 'Journal of Proc. Linn. Soc.' vol. vi. + 1862, p. 74. Mr. Wonfor's observations are quoted in 'Popular + Science Review,' 1868, p. 343. + + [438] Kirby and Spence, 'Introduction to Entomology,' vol. iii. + p. 299. + + [439] Robinet, 'Vers a Soie,' 1848, p. 207. + + [440] 'Transact. Ent. Soc.' 3rd series, vol. v. p. 486. + + [441] 'Journal of Proc. Ent. Soc.' Feb. 4th, 1867, p. lxxi. + + [442] For this and other statements on the size of the sexes, + see Kirby and Spence, ibid. vol. iii. p. 300; on the duration + of life in insects, see p. 344. + + [443] 'Transact. Linnean Soc.' vol. xxvi. 1868, p. 296. + + [444] 'The Malay Archipelago,' vol. ii. 1869, p. 313. + + [445] 'Modern Classification of Insects,' vol. ii. 1840, p. + 526. + + [446] See Mr. B. T. Lowne's very interesting work, 'On the + Anatomy of the Blow-Fly, Musca vomitoria,' 1870, p. 14. + + [447] Westwood, 'Modern Class. of Insects,' vol. ii. p. 473. + + [448] These particulars are taken from Westwood's 'Modern + Class. of Insects,' vol. ii. 1840, p. 422. See, also, on the + Fulgoridae, Kirby and Spence, 'Introduct.' vol. ii. p. 401. + + [449] 'Zeitschrift fuer wissenschaft. Zoolog.' B. xvii. 1867, s. + 152-158. + + [450] I am indebted to Mr. Walsh for having sent me this + extract from a 'Journal of the Doings of Cicada septemdecim,' + by Dr. Hartman. + + [451] L. Guilding, 'Transact. Linn. Soc.' vol. xv. p. 154. + + [452] Koeppen, as quoted in the 'Zoological Record,' for 1867, + p. 460. + + [453] Gilbert White, 'Nat. Hist. of Selborne,' vol. ii. 1825, + p. 262. + + [454] Harris, 'Insects of New England,' 1842, p. 128. + + [455] 'The Naturalist on the Amazons,' vol. i. 1863, p. 252. + Mr. Bates gives a very interesting discussion on the gradations + in the musical apparatus of the three families. See also + Westwood, 'Modern Class.' vol. ii. p. 445 and 453. + + [456] 'Proc. Boston Soc. of Nat. Hist.' vol. xi. April, 1868. + + [457] 'Nouveau Manuel d'Anat. Comp.' (French translat.), tom. + i. 1850 p. 567. + + [458] 'Zeitschrift fuer wissenschaft. Zoolog.' B. xvii. 1867, s. + 117. + + [459] Westwood, 'Modern Class. of Insects,' vol. i. p. 440. + + [460] Westwood, 'Modern Class. of Insects,' vol. i. p. 453. + + [461] Landois, ibid. s. 121, 122. + + [462] Mr. Walsh also informs me that he has noticed that the + female of the _Platyphyllum concavum_, "when captured makes a + feeble grating noise by shuffling her wing-covers together." + + [463] Landois, ibid. s. 113. + + [464] 'Insects of New England,' 1842, p. 133. + + [465] Westwood, 'Modern Classification,' vol. i. p. 462. + + [466] Westwood, ibid. vol. i. p. 453. + + [467] Landois, ibid. s. 115, 116, 120, 122. + + [468] 'Transact. Ent. Soc.' 3rd series, vol. ii. ('Journal of + Proceedings, p. 117.) + + [469] Westwood, 'Modern Class. of Insects,' vol. i. p. 427; for + crickets, p. 445. + + [470] Mr. Ch. Horne, in 'Proc. Ent. Soc.' May 3, 1869, p. xii. + + [471] The _Oecanthus nivalis_, Harris, 'Insects of New + England,' 1842, p. 124. + + [472] Platyblemnus: Westwood, 'Modern. Class.' vol. i. p. 447. + + [473] B. D. Walsh, the Pseudo-neuroptera of Illinois, in 'Proc. + Ent. Soc. of Philadelphia,' 1862, p. 361. + + [474] 'Modern Class.' vol. ii. p. 37. + + [475] Walsh, ibid. p. 381. I am indebted to this naturalist for + the following facts on Hetaerina, Anax, and Gomphus. + + [476] 'Transact. Ent. Soc' vol. i. 1836, p. lxxxi. + + [477] See abstract in the 'Zoological Record' for 1867, p. 450. + + [478] Kirby and Spence, 'Introduct. to Entomology,' vol. ii. + 1818, p. 35. + + [479] See an interesting article, "The Writings of Fabre," in + 'Nat. Hist. Review,' April, 1862, p. 122. + + [480] 'Journal of Proc. of Entomolog. Soc.' Sept. 7th, 1863, + p. 169. + + [481] P. Huber, 'Recherches sur les Moeurs des Fourmis,' 1810, + p. 150, 165. + + [482] 'Proc. Entomolog. Soc. of Philadelphia,' 1866, p. + 238-239. + + [483] Quoted by Westwood, 'Modern Class. of Insects,' vol. ii. + p. 214. + + [484] Pyrodes pulcherrimus, in which the sexes differ + conspicuously, has been described by Mr. Bates in 'Transact. + Ent. Soc.' 1869, p. 50. I will specify the few other cases in + which I have heard of a difference in colour between the sexes + of beetles. Kirby and Spence ('Introduct. to Entomology,' vol. + iii. p. 301) mention a Cantharis, Meloe, Rhagium, and the + _Leptura testacea_; the male of the latter being testaceous, + with a black thorax, and the female of a dull red all over. + These two latter beetles belong to the Order of Longicorns. + Messrs. R. Trimen and Waterhouse, junr., inform me of two + Lamellicorns, viz., a Peritrichia and Trichius, the male of the + latter being more obscurely coloured than the female. In + _Tillus elongatus_ the male is black, and the female always, as + it is believed, of a dark blue colour with a red thorax. The + male, also, of _Orsodacna atra_, as I hear from Mr. Walsh, is + black, the female (the so-called _O. ruficollis_) having a + rufous thorax. + + [485] 'Proc. Entomolog. Soc. of Philadelphia,' 1864, p. 228. + + [486] Kirby and Spence, 'Introduct. Entomolog.' vol. iii. p. + 300. + + [487] Kirby and Spence, ibid. vol. iii. p. 329. + + [488] 'Modern Classification of Insects,' vol. i. p. 172. On + the same page there is an account of Siagonium. In the British + Museum I noticed one male specimen of Siagonium in an + intermediate condition, so that the dimorphism is not strict. + + [489] 'The Malay Archipelago,' vol. ii. 1869, p. 276. + + [490] 'Entomological Magazine,' vol. i. 1833, p. 82. See also + on the conflicts of this species, Kirby and Spence, ibid. vol. + iii. p. 314; and Westwood, ibid. vol. i. p. 187. + + [491] Quoted from Fischer, in 'Dict. Class. d'Hist. Nat.' tom. + x. p. 324. + + [492] 'Ann. Soc. Entomolog. France,' 1866, as quoted in + 'Journal of Travel,' by A. Murray, 1868, p. 135. + + [493] Westwood, 'Modern Class.' vol. i. p. 184. + + [494] Wollaston, On certain musical Curculionidae, 'Annals and + Mag. of Nat. Hist.' vol. vi. 1860, p. 14. + + [495] 'Zeitschrift fuer wiss. Zoolog.' B. xvii. 1867, s. 127. + + [496] I am greatly indebted to Mr. G. R. Crotch for having sent + me numerous prepared specimens of various beetles belonging to + these three families and others, as well as for valuable + information of all kinds. He believes that the power of + stridulation in the Clythra has not been previously observed. I + am also much indebted to Mr. E. W. Janson, for information and + specimens. I may add that my son, Mr. F. Darwin, finds that + _Dermestes murinus_ stridulates, but he searched in vain for + the apparatus. Scolytus has lately been described by Mr. Algen + as a stridulator, in the 'Edinburgh Monthly Magazine,' 1869, + Nov., p. 130. + + [497] Schioedte, translated in 'Annals and Mag. of Nat. Hist.' + vol. xx. 1867, p. 37. + + [498] Westring has described (Kroyer, 'Naturhist. Tidskrift,' + B. ii. 1848-49, p. 334) the stridulating organs in these two, + as well as in other families. In the Carabidae I have examined + _Elaphrus uliginosus_ and _Blethisa multipunctata_, sent to me + by Mr. Crotch. In Blethisa the transverse ridges on the + furrowed border of the abdominal segment do not come into play, + as far as I could judge, in scraping the rasps on the elytra. + + [499] I am indebted to Mr. Walsh, of Illinois, for having sent + me extracts from Leconte's 'Introduction to Entomology,' p. + 101, 143. + + [500] M. P. de la Brulerie, as quoted in 'Journal of Travel,' + A. Murray, vol. i. 1868, p. 135. + + [501] Mr. Doubleday informs me that "the noise is produced by + the insect raising itself on its legs as high as it can, and + then sinking its thorax five or six times, in rapid succession, + against the substance upon which it is sitting." For references + on this subject see Landois, 'Zeitschrift fuer wissen. Zoolog.' + B. xvii. s. 131. Olivier says (as quoted by Kirby and Spence, + 'Introduct.' vol. ii. p. 395) that the female of _Pimelia + striata_ produces a rather loud sound by striking her abdomen + against any hard substance, "and that the male, obedient to + this call, soon attends her and they pair." + + [502] Apatura Iris: 'The Entomologist's Weekly Intelligencer,' + 1859, p. 139. For the Bornean Butterflies see C. Collingwood, + 'Rambles of a Naturalist,' 1868, p. 183. + + [503] See my 'Journal of Researches,' 1845, p. 33. Mr. + Doubleday has detected ('Proc. Ent. Soc.' March 3rd, 1845, p. + 123) a peculiar membranous sac at the base of the front wings, + which is probably connected with the production of the sound. + + [504] See also Mr. Bates' paper in 'Proc. Ent. Soc. of + Philadelphia,' 1865, p. 206. Also Mr. Wallace on the same + subject, in regard to Diadema, in 'Transact. Entomolog. Soc. of + London,' 1869, p. 278. + + [505] 'The Naturalist on the Amazons,' vol. i. 1863, p. 19. + + [506] See the interesting article in the 'Westminster Review,' + July, 1867, p. 10. A woodcut of the Kallima is given by Mr. + Wallace in Hardwicke's 'Science Gossip,' Sept. 1867, p. 196. + + [507] See the interesting observations by Mr. T. W. Wood, 'The + Student,' Sept. 1868, p. 81. + + [508] Mr. Wallace in 'Hardwicke's Science Gossip,' Sept. 1867, + p. 193. + + [509] See also, on this subject, Mr. Weir's paper in 'Transact. + Ent. Soc.' 1869, p. 23. + + [510] 'Westminster Review,' July, 1867, p. 16. + + [511] For instance, Lithosia; but Prof. Westwood ('Modern + Class. of Insects,' vol. ii. p. 390) seems surprised at this + case. On the relative colours of diurnal and nocturnal + Lepidoptera, see ibid. p. 333 and 392; also Harris, 'Treatise + on the Insects of New England,' 1842, p. 315. + + [512] Such differences between the upper and lower surfaces of + the wings of several species of Papilio, may be seen in the + beautiful plates to Mr. Wallace's Memoir on the Papilionidae of + the Malayan Region, in 'Transact. Linn. Soc.' vol. xxv. part i. + 1865. + + [513] 'Proc. Ent. Soc.' March 2nd, 1868. + + [514] See also an account of the S. American genus Erateina + (one of the Geometrae) in 'Transact. Ent. Soc.' new series, vol. + v. pl. xv. and xvi. + + [515] 'Proc. Ent. Soc. of London,' July 6, 1868, p. xxvii. + + [516] Harris, 'Treatise,' &c., edited by Flint, 1862, p. 395. + + [517] For instance, I observe in my son's cabinet that the + males are darker than the females in the _Lasiocampa quercus_, + _Odonestis potatoria_, _Hypogymna dispar_, _Dasychira + pudibunda_, and _Cycnia mendica_. In this latter species the + difference in colour between the two sexes is strongly marked; + and Mr. Wallace informs me that we here have, as he believes, + an instance of protective mimickry confined to one sex, as will + hereafter be more fully explained. The white female of the + Cycnia resembles the very common _Spilosoma menthrasti_, both + sexes of which are white; and Mr. Stainton observed that this + latter moth was rejected with utter disgust by a whole brood of + young turkeys, which were fond of eating other moths; so that + if the Cycnia was commonly mistaken by British birds for the + Spilosoma, it would escape being devoured, and its white + deceptive colour would thus be highly beneficial. + + [518] 'Rambles of a Naturalist in the Chinese Seas,' 1868, p. + 182. + + [519] Wallace on the Papilionidae of the Malayan Region, in + 'Transact. Linn. Soc.' vol. xxv. 1865, p. 8, 36. A striking case + of a rare variety, strictly intermediate between two other + well-marked female varieties, is given by Mr. Wallace. See also + Mr. Bates, in 'Proc. Entomolog. Soc.' Nov. 19th, 1866, p. xl. + + [520] Mr. R. MacLachlan, 'Transact. Ent. Soc.' vol. ii. part + 6th, 3rd series, 1866, p. 459. + + [521] H. W. Bates, 'The Naturalist on the Amazons,' vol. ii. + 1863, p. 228. A. R. Wallace, in 'Transact. Linn. Soc.' vol. xxv. + 1865, p. 10. + + [522] On this whole subject see 'The Variation of Animals and + Plants under Domestication,' vol. ii. 1868, chap. xxiii. + + [523] A. R. Wallace, in 'The Journal of Travel,' vol. i. 1868, + p. 88. 'Westminster Review,' July, 1857, p. 37. See also + Messrs. Wallace and Bates in 'Proc. Ent. Soc.' Nov. 19th, 1866, + p. xxxix. + + [524] 'The Variation of Animals and Plants under + Domestication,' vol. ii. chap. xii. p. 17. + + [525] 'Transact. Linn. Soc.' vol. xxiii. 1862, p. 495. + + [526] 'Proc. Ent. Soc.' Dec. 3rd, 1866, p. xlv. + + [527] 'Transact. Linn. Soc.' vol. xxv. 1865, p. 1; also + 'Transact. Ent. Soc.' vol. iv. (3rd series), 1867, p. 301. + + [528] See an ingenious article entitled, "Difficulties of the + Theory of Natural Selection," in the 'Month,' 1869. The writer + strangely supposes that I attribute the variations in colour of + the Lepidoptera, by which certain species belonging to distinct + families have come to resemble others, to reversion to a common + progenitor; but there is no more reason to attribute these + variations to reversion than in the case of any ordinary + variation. + + [529] Wallace, "Notes on Eastern Butterflies," 'Transact. Ent. + Soc.' 1869, p. 287. + + [530] Wallace, in 'Westminster Review,' July, 1867, p. 37; and + in 'Journal of Travel and Nat. Hist.' vol. i. 1868, p. 88. + + [531] See remarks by Messrs. Bates and Wallace, in 'Proc. Ent. + Soc.' Nov. 19, 1866, p. xxxix. + + [532] See Mr. Wallace in 'Westminster Review,' July, 1867, p. + 11 and 37. The male of no butterfly, as Mr. Wallace informs me, + is known to differ in colour, as a protection, from the female; + and he asks me how I can explain this fact on the principle + that one sex alone has varied and has transmitted its + variations exclusively to the same sex, without the aid of + selection to check the variations being inherited by the other + sex. No doubt if it could be shewn that the females of very + many species had been rendered beautiful through protective + mimickry, but that this has never occurred with the males, it + would be a serious difficulty. But the number of cases as yet + known hardly suffices for a fair judgment. We can see that the + males, from having the power of flying more swiftly, and thus + escaping danger, would not be so likely as the females to have + had their colours modified for the sake of protection; but this + would not in the least have interfered with their receiving + protective colours through inheritance from the females. In the + second place, it is probable that sexual selection would + actually tend to prevent a beautiful male from becoming + obscure, for the less brilliant individuals would be less + attractive to the females. Supposing that the beauty of the + male of any species had been mainly acquired through sexual + selection, yet if this beauty likewise served as a protection, + the acquisition would have been aided by natural selection. But + it would be quite beyond our power to distinguish between the + two processes of sexual and ordinary selection. Hence it is not + likely that we should be able to adduce cases of the males + having been rendered brilliant exclusively through protective + mimickry, though this is comparatively easy with the females, + which have rarely or never been rendered beautiful, as far as + we can judge, for the sake of sexual attraction, although they + have often received beauty through inheritance from their male + parents. + + [533] 'Proc. Entomolog. Soc.' Dec. 3rd, 1866, p. xlv., and + March 4th, 1867, p. lxxx. + + [534] See Mr. J. Jenner Weir's paper on insects and + insectivorous birds, in 'Transact. Ent. Soc.' 1869, p. 21; also + Mr. Butler's paper, ibid. p. 27. + + +END OF VOL. I. + +LONDON: + +PRINTED BY WILLIAM CLOWES AND SONS, DUKE STREET, STAMFORD STREET, AND +CHARING CROSS. + + +50A, ALBEMARLE STREET, LONDON, +_January, 1871_. + + +MR. MURRAY'S + +LIST OF POPULAR WORKS. + + +_One Shilling._ + +THE PRINCIPAL SPEECHES AND ADDRESSES OF H.R.H. 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