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| author | Roger Frank <rfrank@pglaf.org> | 2025-10-14 20:01:22 -0700 |
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| committer | Roger Frank <rfrank@pglaf.org> | 2025-10-14 20:01:22 -0700 |
| commit | 40de937284b64499031d4662124b1f2d717b2c45 (patch) | |
| tree | 282cb49d68fcc1c172026b2664b137d93afd5c51 | |
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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/34279-0.txt b/34279-0.txt new file mode 100644 index 0000000..cf56647 --- /dev/null +++ b/34279-0.txt @@ -0,0 +1,16499 @@ +The Project Gutenberg EBook of Handbook of Medical Entomology, by +William Albert Riley and Oskar Augustus Johanssen + +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: Handbook of Medical Entomology + +Author: William Albert Riley + Oskar Augustus Johanssen + +Release Date: November 11, 2010 [EBook #34279] + +Language: English + +Character set encoding: UTF-8 + +*** START OF THIS PROJECT GUTENBERG EBOOK HANDBOOK OF MEDICAL ENTOMOLOGY *** + + + + +Produced by Bryan Ness, Iris Schimandle, Brownfox and the +Online Distributed Proofreading Team at http://www.pgdp.net +(This file was produced from images generously made +available by The Internet Archive/American Libraries.) + + + + + +Transcriber's Note: Barring some obvious typos, the text has been left +as printed. Discrepancies identified are listed at the end of the text. + + + + +[Illustration: Some early medical entomology. Athanasius Kircher's +illustration of the Italian tarantula and the music prescribed as an +antidote for the poison of its bite. (1643).] + + + + +HANDBOOK OF MEDICAL + +ENTOMOLOGY + +WM. A. RILEY, PH.D. + +Professor of Insect Morphology and Parasitology, Cornell University + +and + +O. A. JOHANNSEN, PH.D. + +Professor of Biology, Cornell University + +[Illustration] + +ITHACA, NEW YORK + +THE COMSTOCK PUBLISHING COMPANY + +1915 + + +COPYRIGHT, 1915 + +BY THE COMSTOCK PUBLISHING COMPANY, + +ITHACA, N. Y. + + +Press of W. F. Humphrey +Geneva, N. Y. + + + + +PREFACE + + +The Handbook of Medical Entomology is the outgrowth of a course of +lectures along the lines of insect transmission and dissemination of +diseases of man given by the senior author in the Department of +Entomology of Cornell University during the past six years. More +specifically it is an illustrated revision and elaboration of his "Notes +on the Relation of Insects to Disease" published January, 1912. + +Its object is to afford a general survey of the field, and primarily to +put the student of medicine and entomology in touch with the discoveries +and theories which underlie some of the most important modern work in +preventive medicine. At the same time the older phases of the +subject--the consideration of poisonous and parasitic forms--have not +been ignored. + +Considering the rapid shifts in viewpoint, and the development of the +subject within recent years, the authors do not indulge in any hopes +that the present text will exactly meet the needs of every one +specializing in the field,--still less do they regard it as complete or +final. The fact that the enormous literature of isolated articles is to +be found principally in foreign periodicals and is therefore difficult +of access to many American workers, has led the authors to hope that a +summary of the important advances, in the form of a reference book may +not prove unwelcome to physicians, sanitarians and working +entomologists, and to teachers as a text supplementing lecture work in +the subject. + +Lengthy as is the bibliography, it covers but a very small fraction of +the important contributions to the subject. It will serve only to put +those interested in touch with original sources and to open up the +field. Of the more general works, special acknowledgment should be made +to those of Banks, Brumpt, Castellani and Chalmers, Comstock, Hewitt, +Howard, Manson, Mense, Neveau-Lemaire, Nuttall, and Stiles. + +To the many who have aided the authors in the years past, by suggestions +and by sending specimens and other materials, sincerest thanks is +tendered. This is especially due to their colleagues in the Department +of Entomology of Cornell University, and to Professor Charles W. Howard, +Dr. John Uri Lloyd, Mr. A. H. Ritchie, Dr. I. M. Unger, and Dr. Luzerne +Coville. + +They wish to express indebtedness to the authors and publishers who have +so willingly given permission to use certain illustrations. Especially +is this acknowledgment due to Professor John Henry Comstock, Dr. L. O. +Howard, Dr. Graham-Smith, and Professor G. H. T. Nuttall. Professor +Comstock not only authorized the use of departmental negatives by the +late Professor M. V. Slingerland (credited as M. V. S.), but generously +put at their disposal the illustrations from the MANUAL FOR THE STUDY OF +INSECTS and from the SPIDER BOOK. Figures 5 and 111 are from Peter's +"Der Arzt und die Heilkunst in der deutschen Vergangenheit." It should +be noted that on examining the original, it is found that Gottfried's +figure relates to an event antedating the typical epidemic of dancing +mania. + + WM. A. RILEY. + O. A. JOHANNSEN. + + CORNELL UNIVERSITY, + January, 1915. + + + + +ADDITIONS AND CORRECTIONS + +vi line 11, for Heilkunft read Heilkunst. + +18 line 2, for tarsi read tarsus. + +32 line 21, and legend under fig. 23, for C. (Conorhinus) abdominalis + read Melanolestes abdominalis. + +47 legend under figure for 33c read 34. + +92 line 22 and 25, for sangiusugus read sanguisugus. + +116 legend under fig. 83, for Graham-Smith read Manson. + +136 line 10, from bottom, insert "ring" after "chitin". + +137 line 3, for meditatunda read meditabunda. + +145 line 7, from bottom, for Rs read R_5. + +158 line 20, for have read has. + +212 after the chapter heading insert "continued". + +219 line 10, from bottom, for Cornohinus read Conorhinus. + +266 line 1, fig. 158j refers to the female. + +272 line 5, insert "palpus" before "and leg". + +281 line 6, for discodial read discoidal. + +281 last line, insert "from" before "the". + +284 line 5, for "tubercle of" read "tubercle or". + +305 lines 19, 28, 44, page 306 lines 1, 9, 22, 27, 30, page 307 line 7, + page 309 lines 8, 11, for R_{4+5} read M_{1+2}. + +309 legend under fig. 168 add Bureau of Entomology. + +312 line 36, for "near apex" read "of M_{1+2}". + +313 running head, for Muscidæ read Muscoidea. + +314 line 29, for "distal section" read "distally M_{1+2}". + +315 legend under fig. 172, for Pseudopyrellia read Orthellia, for + Lyperosia read Hæmatobia, for Umbana read urbana. + +323 and 325 legends under the figures, add "After Dr. J. H. Stokes". + +328 line 7 from bottom for Apiochæta read Aphiochæta. + + + + +CONTENTS + + +CHAPTER I + +INTRODUCTION 1-5 + + Early suggestions regarding the transmission of disease by + insects. + The ways in which arthropods may affect the health of man. + + +CHAPTER II + +ARTHROPODS WHICH ARE DIRECTLY POISONOUS 6-56 + + The Araneida, or Spiders. + The tarantulas. Bird spiders. Spiders of the genus + Latrodectus. Other venomous spiders. Summary. + The Pedipalpida, or whip-scorpions. + The Scorpionida, or true scorpions. + The Solpugida, or solpugids. + The Acarina, or mites and ticks. + The Myriapoda, or centipedes and millipedes. + The Hexapoda, or true insects. + Piercing or biting insects poisonous to man. + Hemiptera, or true bugs. + The Notonectidæ or back-swimmers. Belostomidæ or giant + water-bugs. Reduviidæ, or assassin bugs. Other + Hemiptera reported as poisonous to man. + Diptera; the midges, mosquitoes and flies. + Stinging insects. + Apis mellifica, the honey bee. Other stinging forms. + Nettling insects. + Lepidoptera, or butterflies and moths. Relief from + poisoning by nettling larvæ. + Vescicating insects and those possessing other poisons + in their blood plasma. The blister beetles. Other + cryptotoxic insects. + + +CHAPTER III + +PARASITIC ARTHROPODS AFFECTING MAN 57-130 + + Acarina, or mites. + The Trombidiidæ, or harvest mites. + The Ixodoidea, or ticks. + Argasidæ. Ixodidæ. Treatment of tick bites. + The mites. + Dermanyssidæ. Tarsonemidæ. Sarcoptidæ, the itch mites. + Demodecidæ, the follicle mites. + Hexapoda, or true insects. + Siphunculata, or sucking lice. + Hemiptera. + The bed-bug. Other bed-bugs. + Parasitic Diptera, or flies. + Psychodidæ, or moth flies. Phlebotominæ. Culicidæ, or + mosquitoes. Simuliidæ, or black-flies. Chironomidæ, or + midges. Tabanidæ, or horse-flies. Leptidæ or + snipe-flies. Oestridæ, or bot-flies. Muscidæ, the + stable-fly and others. + Siphonaptera, or fleas. + The fleas affecting man, the dog, cat, and rat. + The true chiggers, or chigoes. + +CHAPTER IV + +ACCIDENTAL OR FACULTATIVE PARASITES 131-143 + + Acarina, or mites. + Myriapoda, or centipedes and millipedes. + Lepidopterous larvæ. + Coleoptera, or beetles. + Dipterous larvæ causing myiasis. + Piophila casei, the cheese skipper. Chrysomyia macellaria, + the screw-worm fly. Calliphorinæ, the bluebottles. + Muscinæ, the house or typhoid fly, and others. + Anthomyiidæ, the lesser house-fly and others. + Sarcophagidæ, the flesh-flies. + + +CHAPTER V + +ARTHROPODS AS SIMPLE CARRIERS OF DISEASE 144-163 + + The house or typhoid fly as a carrier of disease. + Stomoxys calcitrans, the stable-fly. + Other arthropods which may serve as simple carriers of + pathogenic organisms. + + +CHAPTER VI + +ARTHROPODS AS DIRECT INOCULATORS OF DISEASE GERMS 164-174 + + Some illustrations of direct inoculations of disease germs + by arthropods. + The rôle of fleas in the transmission of the plague. + + +CHAPTER VII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC ORGANISMS 175-185 + + Insects as intermediate hosts of tape-worms. + Arthropods as intermediate hosts of nematode worms. + Filariasis and mosquitoes. + Other nematode parasites of man and animals. + + +CHAPTER VIII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA 186-211 + + Mosquitoes and malaria. + Mosquitoes and yellow fever. + + +CHAPTER IX + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA 212-229 + + Insects and trypanosomiases. + Fleas and lice as carriers of Trypanosoma lewisi. + Tsetse-flies and nagana. + Tsetse-flies and sleeping sickness in man. + South American trypanosomiasis. + Leishmanioses and insects. + Ticks and diseases of man and animals. + Cattle tick and Texas fever. + Ticks and Rocky Mountain Spotted fever of man. + + +CHAPTER X + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA +(CONTINUED) 230-240 + + Arthropods and Spirochætoses of man and animals. + African relapsing fever of man. + European relapsing fever. + North African relapsing fever of man. + Other types of relapsing fever of man. + Spirochætosis of fowls. + Other spirochæte diseases of animals. + Typhus fever and lice. + + +CHAPTER XI + +SOME POSSIBLE, BUT IMPERFECTLY KNOWN CASES OF +ARTHROPOD TRANSMISSION OF DISEASE 241-256 + + Infantile paralysis, or acute anterior poliomyelitis. + Pellagra. Leprosy. Verruga peruviana. Cancer. + + +CHAPTER XII + +KEYS TO THE ARTHROPODS NOXIOUS TO MAN 257-317 + + Crustacea. + Myriapoda, or centipedes and millipedes. + Arachnida (Orders of). + Acarina or ticks. + Hexapoda (Insecta). + Siphunculata and Hemiptera (lice and true bugs). + Diptera (mosquitoes, midges, and flies). + Siphonaptera (fleas). + + +APPENDIX + +Hydrocyanic acid gas against household insects 318-320 + Proportion of ingredients. A single room as an example. + Fumigating a large house. Precautions. + +Lesions produced by the bite of the black-fly 321-326 + + +BIBLIOGRAPHY 327-340 + + +INDEX 341-348 + + + + +CHAPTER I. + +INTRODUCTION + +EARLY SUGGESTIONS REGARDING THE TRANSMISSION OF DISEASE BY INSECTS + + +Until very recent years insects and their allies have been considered as +of economic importance merely in so far as they are an annoyance or +direct menace to man, or his flocks and herds, or are injurious to his +crops. It is only within the past fifteen years that there has sprung +into prominence the knowledge that in another and much more insiduous +manner, they may be the enemy of mankind, that they may be among the +most important of the disseminators of disease. In this brief period, +such knowledge has completely revolutionized our methods of control of +certain diseases, and has become an important weapon in the fight for +the conservation of health. + +It is nowhere truer than in the case under consideration that however +abrupt may be their coming into prominence, great movements and great +discoveries do not arise suddenly. Centuries ago there was suggested the +possibility that insects were concerned with the spread of disease, and +from time to time there have appeared keen suggestions and logical +hypotheses along this line, that lead us to marvel that the +establishment of the truths should have been so long delayed. + +One of the earliest of these references is by the Italian physician, +Mercurialis, who lived from 1530 to 1607, during a period when Europe +was being ravaged by the dread "black death", or plague. Concerning its +transmission he wrote: "There can be no doubt that flies feed on the +internal secretions of the diseased and dying, then, flying away, they +deposit their excretions on the food in neighboring dwellings, and +persons who eat of it are thus infected." + +It would be difficult to formulate more clearly this aspect of the facts +as we know them to-day, though it must always be borne in mind that we +are prone to interpret such statements in the light of present-day +knowledge. Mercurialis had no conception of the animate nature of +contagion, and his statement was little more than a lucky guess. + +Much more worthy of consideration is the approval which was given to his +view by the German Jesuit, Athanasius Kircher in 1658. One cannot read +carefully his works without believing that long before Leeuwenhook's +discovery, Kircher had seen the larger species of bacteria. Moreover, he +attributed the production of disease to these organisms and formulated, +vaguely, to be sure, a theory of the animate nature of contagion. It has +taken two and a half centuries to accumulate the facts to prove his +hypothesis. + +The theory of Mercurialis was not wholly lost sight of, for in the +medical literature of the eighteenth century there are scattered +references to flies as carriers of disease. Such a view seems even to +have been more or less popularly accepted, in some cases. Gudger (1910), +has pointed out that, as far back as 1769, Edward Bancroft, in "An Essay +on the Natural History of Guiana in South America," wrote concerning the +contagious skin-disease known as "Yaws": "It is usually believed that +this disorder is communicated by the flies who have been feasting on a +diseased object, to those persons who have sores, or scratches, which +are uncovered; and from many observations, I think this is not +improbable, as none ever receive this disorder whose skins are whole." + +Approaching more closely the present epoch, we find that in 1848, Dr. +Josiah Nott, of Mobile, Alabama, published a remarkable article on the +cause of yellow fever, in which he presented "reasons for supposing its +specific cause to exist in some form of insect life." As a matter of +fact, the bearing of Nott's work on present day ideas of the insect +transmission of disease has been very curiously overrated. The common +interpretation of his theory has been deduced from a few isolated +sentences, but his argument appears quite differently when the entire +article is studied. It must be remembered that he wrote at a period +before the epoch-making discoveries of Pasteur and before the +recognition of micro-organisms as factors in the cause of disease. His +article is a masterly refutation of the theory of "malarial" origin of +"all the fevers of hot climates," but he uses the term "insect" as +applicable to the lower forms of life, and specific references to +"mosquitoes," "aphids," "cotton-worms," and others, are merely in the +way of similes. + +But, while Nott's ideas regarding the relation of insects to yellow +fever were vague and indefinite, it was almost contemporaneously that +the French physician, Louis Daniel Beauperthuy argued in the most +explicit possible manner, that yellow fever and various others are +transmitted by mosquitoes. In the light of the data which were available +when he wrote, in 1853, it is not surprising that he erred by thinking +that the source of the virus was decomposing matter which the mosquito +took up and accidentally inoculated into man. Beauperthuy not only +discussed the rôle of mosquitoes in the transmission of disease, but he +taught, less clearly, that house-flies scatter pathogenic organisms. It +seems that Boyce (1909) who quotes extensively from this pioneer work, +does not go too far when he says "It is Dr. Beauperthuy whom we must +regard as the father of the doctrine of insect-borne disease." + +In this connection, mention must be made of the scholarly article by the +American physician, A. F. A. King who, in 1883, brought together an all +but conclusive mass of argument in support of his belief that malaria +was caused by mosquitoes. At about the same time, Finley, of Havana, was +forcefully presenting his view that the mosquito played the chief rôle +in the spread of yellow fever. + +To enter more fully into the general historical discussion is beyond the +scope of this book. We shall have occasion to make more explicit +references in considering various insect-borne diseases. Enough has been +said here to emphasize that the recognition of insects as factors in the +spread of disease was long presaged, and that there were not wanting +keen thinkers who, with a background of present-day conceptions of the +nature of disease, might have been in the front rank of investigators +along these lines. + + +THE WAYS IN WHICH ARTHROPODS MAY AFFECT THE HEALTH OF MAN + +When we consider the ways in which insects and their allies may affect +the health of man, we find that we may treat them under three main +groups: + +A. They may be directly poisonous. Such, for example, are the scorpions, +certain spiders and mites, some of the predaceous bugs, and stinging +insects. Even such forms as the mosquito deserve some consideration from +this viewpoint. + +B. They may be parasitic, living more or less permanently on or in the +body and deriving their sustenance from it. + +Of the parasitic arthropods we may distinguish, first, the _true +parasites_, those which have adopted and become confirmed in the +parasitic habit. Such are the itch mites, the lice, fleas, and the +majority of the forms to be considered as parasitic. + +In addition to these, we may distinguish a group of _accidental_, or +_facultative parasites_, species which are normally free-living, feeding +on decaying substances, but which when accidentally introduced into the +alimentary canal or other cavities of man, may exist there for a greater +or less period. For example, certain fly larvæ, or maggots, normally +feeding in putrifying meat, have been known to occur as accidental or +facultative parasites in the stomach of man. + +C. Finally, and most important, arthropods may be transmitters and +disseminators of disease. In this capacity they may function in one of +three ways; as _simple carriers_, as _direct inoculators_, or as +_essential hosts_ of disease germs. + +As simple carriers, they may, in a wholly incidental manner, transport +from the diseased to the healthy, or from filth to food, pathogenic +germs which cling to their bodies or appendages. Such, for instance, is +the relation of the house-fly to the dissemination of typhoid. + +As direct inoculators, biting or piercing species may take up from a +diseased man or animal, germs which, clinging to the mouth parts, are +inoculated directly into the blood of the insect's next victim. It it +thus that horse-flies may occasionally transmit anthrax. Similarly, +species of spiders and other forms which are ordinarily perfectly +harmless, may accidentally convey and inoculate pyogenic bacteria. + +It is as essential hosts of disease germs that arthropods play their +most important rôle. In such cases an essential part of the life cycle +of the pathogenic organism is undergone in the insect. In other words, +without the arthropod host the disease-producing organism cannot +complete its development. As illustrations may be cited the relation of +the Anopheles mosquito to the malarial parasite, and the relation of the +cattle tick to Texas fever. + +A little consideration will show that this is the most important of the +group. Typhoid fever is carried by water or by contaminated milk, and in +various other ways, as well as by the house-fly. Kill all the +house-flies and typhoid would still exist. On the other hand, malaria is +carried only by the mosquito, because an essential part of the +development of the malarial parasite is undergone in this insect. +Exterminate all of the mosquitoes of certain species and the +dissemination of human malaria is absolutely prevented. + +Once an arthropod becomes an essential host for a given parasite it may +disseminate infection in three different ways: + +1. By infecting man or animals who ingest it. It is thus, for example, +that man, dog, or cat, becomes infected with the double-pored dog +tapeworm, _Dipylidium caninum_. The cysticercoid stage occurs in the +dog louse, or in the dog or cat fleas, and by accidentally ingesting the +infested insect the vertebrate becomes infested. Similarly, _Hymenolepis +diminuta_, a common tapeworm of rats and mice, and occasional in man, +undergoes part of its life cycle in various meal-infesting insects, and +is accidentally taken up by its definitive host. It is very probable +that man becomes infested with _Dracunculus (Filaria) medinensis_ +through swallowing in drinking water, the crustacean, _Cyclops_, +containing the larvæ of this worm. + +2. By infecting man or animals on whose skin or mucous membranes the +insect host may be crushed or may deposit its excrement. The pathogenic +organism may then actively penetrate, or may be inoculated by +scratching. The causative organism of typhus fever is thus transmitted +by the body louse. + +3. By direct inoculation by its bite, the insect host may transfer the +parasite which has undergone development within it. The malarial +parasite is thus transferred by mosquitoes; the Texas fever parasite by +cattle ticks. + + + + +CHAPTER II. + +ARTHROPODS WHICH ARE DIRECTLY POISONOUS + + +Of all the myriads of insects and related forms, a very few are of +direct use to man, some few others have forced his approbation on +account of their wonderful beauty, but the great hordes of them are +loathed or regarded as directly dangerous. As a matter of fact, only a +very small number are in the slightest degree poisonous to man or to the +higher animals. The result is that entomologists and lovers of nature, +intent upon dissipating the foolish dread of insects, are sometimes +inclined to go to the extreme of discrediting all statements of serious +injury from the bites or stings of any species. + +Nevertheless, it must not be overlooked that poisonous forms do exist, +and they must receive attention in a consideration of the ways in which +arthropods may affect the health of man. Moreover, it must be recognized +that "what is one man's meat, is another man's poison," and that in +considering the possibilities of injury we must not ignore individual +idiosyncrasies. Just as certain individuals may be poisoned by what, for +others, are common articles of food, so some persons may be abnormally +susceptible to insect poison. Thus, the poison of a bee sting may be of +varying severity, but there are individuals who are made seriously sick +by a single sting, regardless of the point of entry. Some individuals +scarcely notice a mosquito bite, others find it very painful, and so +illustrations of this difference in individuals might be multiplied. + +In considering the poisonous arthropods, we shall take them up by +groups. The reader who is unacquainted with the systematic relationship +of insects and their allies is referred to Chapter XII. No attempt will +be made to make the lists under the various headings exhaustive, but +typical forms will be discussed. + + +ARANEIDA OR SPIDERS + +Of all the arthropods there are none which are more universally feared +than are the spiders. It is commonly supposed that the majority, if not +all the species are poisonous and that they are aggressive enemies of +man and the higher animals, as well as of lower forms. + +That they really secrete a poison may be readily inferred from the +effect of their bite upon insects and other small forms. Moreover, the +presence of definite and well-developed poison glands can easily be +shown. They occur as a pair of pouches (fig. 1) lying within the +cephalothorax and connected by a delicate duct with a pore on the claw +of the chelicera, or so-called "mandible" on the convex surface of the +claw in such a position that it is not plugged and closed by the flesh +of the victim. + +[Illustration: 1. Head of a spider showing poison gland (_c_) and its +relation to the chelicera (_a_).] + +The glands may be demonstrated by slowly and carefully twisting off a +chelicera and pushing aside the stumps of muscles at its base. By +exercising care, the chitinous wall of the chelicera and its claw may be +broken away and the duct traced from the gland to its outlet. The inner +lining of the sac is constituted by a highly developed glandular +epithelium, supported by a basement membrane of connective tissue and +covered by a muscular layer, (fig. 2). The muscles, which are striated, +are spirally arranged (fig. 1), and are doubtless under control of the +spider, so that the amount of poison to be injected into a wound may be +varied. + +[Illustration: 2. Section through a venom gland of Latrodectus +13-guttatus showing the peritoneal, muscular and epithelial layers. +After Bordas.] + +The poison itself, according to Kobert (1901), is a clear, colorless +fluid, of oily consistency, acid reaction, and very bitter taste. After +the spider has bitten two or three times, its supply is exhausted and +therefore, as in the case of snakes, the poison of the bite decreases +quickly with use, until it is null. To what extent the content of the +poison sacs may contain blood serum or, at least, active principles of +serum, in addition to a specific poison formed by the poison glands +themselves, Kobert regards as an open question. He believes that the +acid part of the poison, if really present, is formed by the glands and +that, in the case of some spiders, the ferment-like, or better, active +toxine, comes from the blood. + +[Illustration: 3. Chelicera of a spider.] + +But there is a wide difference between a poison which may kill an insect +and one which is harmful to men. Certain it is that there is no lack of +popular belief and newspaper records of fatal cases, but the evidence +regarding the possibility of fatal or even very serious results for man +is most contradictory. For some years, we have attempted to trace the +more circumstantial newspaper accounts, which have come to our notice, +of injury by North American species. The results have served, mainly, to +emphasize the straits to which reporters are sometimes driven when there +is a dearth of news. The accounts are usually vague and lacking in any +definite clue for locating the supposed victim. In the comparatively few +cases where the patient, or his physician, could be located, there was +either no claim that the injury was due to spider venom, or there was no +evidence to support the belief. Rarely, there was evidence that a +secondary blood poisoning, such as might be brought about by the prick +of a pin, or by any mechanical injury, had followed the bite of a +spider. Such instances have no bearing on the question of the venomous +nature of these forms. + +[Illustration: 4. The Italian tarantula (Lycosa tarantula). After +Kobert.] + +The extreme to which unreasonable fear of the bites of spiders +influenced the popular mind was evidenced by the accepted explanation of +the remarkable dancing mania, or tarantism, of Italy during the Middle +Ages. This was a nervous disorder, supposed to be due to the bite of a +spider, the European tarantula (fig. 4), though it was also, at times, +attributed to the bite of the scorpion. In its typical form, it was +characterized by so great a sensibility to music that under its +influence the victims indulged in the wildest and most frenzied dancing, +until they sank to the ground utterly exhausted and almost lifeless. The +profuse perspiring resulting from these exertions was supposed to be the +only efficacious remedy for the disease. Certain forms of music were +regarded as of especial value in treating this tarantism, and hence the +name of "tarantella" was applied to them. Our frontispiece, taken from +Athanasius Kircher's _Magnes sive de Arte Magnetica_, 1643 ed., +represents the most commonly implicated spider and illustrates some of +what Fabre has aptly designated as "medical choreography." + +The disease was, in reality, a form of hysteria, spreading by sympathy +until whole communities were involved, and was paralleled by the +outbreaks of the so-called St. Vitus's or St. John's dance, which swept +Germany at about the same time (fig. 5). The evidence that the spider +was the cause of the first is about as conclusive as is that of the +demoniacal origin of the latter. The true explanation of the outbreaks +is doubtless to be found in the depleted physical and mental condition +of the people, resulting from the wars and the frightful plagues which +devastated all Europe previous to, and during these times. An +interesting discussion of these aspects of the question is to be found +in Hecker. + +[Illustration: 5. Dancing mania. Illustration from Johann Ludwig +Gottfried's Chronik. 1632.] + +So gross has been the exaggeration and so baseless the popular fear +regarding spiders that entomologists have been inclined to discredit all +accounts of serious injury from their bites. Not only have the most +circumstantial of newspaper accounts proved to be without foundation but +there are on record a number of cases where the bite of many of the +commoner species have been intentionally provoked and where the effect +has been insignificant. Some years ago the senior author personally +experimented with a number of the largest of our northern species, and +with unexpected results. The first surprise was that the spiders were +very unwilling to bite and that it required a considerable effort to get +them to attempt to do so. In the second place, most of those +experimented with were unable to pierce the skin of the palm or the back +of the hand, but had to be applied to the thin skin between the fingers +before they were able to draw blood. Unfortunately, no special attempt +was made to determine, at the time, the species experimented with, but +among them were _Theridion tepidariorum_, _Miranda aurantia_ +(_Argiopa_), _Metargiope trifasciata_, _Marxia stellata_, _Aranea +trifolium_, _Misumena vatia_, and _Agelena nævia_. In no case was the +bite more severe than a pin prick and though in some cases the sensation +seemed to last longer, it was probably due to the fact that the mind was +intent upon the experiment. + +[Illustration: 6. An American tarantula (Eurypelma hentzii). Natural +size. After Comstock.] + +Similar experiments were carried out by Blackwell (1855), who believed +that in the case of insects bitten, death did not result any more +promptly than it would have from a purely mechanical injury of equal +extent. He was inclined to regard all accounts of serious injury to man +as baseless. The question cannot be so summarily dismissed, and we shall +now consider some of the groups which have been more explicitly +implicated. + + +THE TARANTULAS.--In popular usage, the term "tarantula" is loosely +applied to any one of a number of large spiders. The famous tarantulas +of southern Europe, whose bites were supposed to cause the dancing +mania, were Lycosidæ, or wolf-spiders. Though various species of this +group were doubtless so designated, the one which seems to have been +most implicated was _Lycosa tarantula_ (L.), (fig. 4). On the other +hand, in this country, though there are many Lycosidæ, the term +"tarantula" has been applied to members of the superfamily Avicularoidea +(fig. 6), including the bird-spiders. + +Of the Old World Lycosidæ there is no doubt that several species were +implicated as the supposed cause of the tarantism. In fact, as we have +already noted, the blame was sometimes attached to a scorpion. However, +there seems to be no doubt that most of the accounts refer to the spider +known as _Lycosa tarantula_. + +There is no need to enter into further details here regarding the +supposed virulence of these forms, popular and the older medical +literature abound in circumstantial accounts of the terrible effects of +the bite. Fortunately, there is direct experimental evidence which bears +on the question. + +Fabre induced a common south European wolf-spider, _Lycosa narbonensis_, +to bite the leg of a young sparrow, ready to leave the nest. The leg +seemed paralyzed as a result of the bite, and though the bird seemed +lively and clamored for food the next day, on the third day it died. A +mole, bitten on the nose, succumbed after thirty-six hours. From these +experiments Fabre seemed justified in his conclusion that the bite of +this spider is not an accident which man can afford to treat lightly. +Unfortunately, there is nothing in the experiments, or in the symptoms +detailed, to exclude the probability that the death of the animals was +the result of secondary infection. + +As far back as 1693, as we learn from the valuable account of Kobert, +(1901), the Italian physician, Sanguinetti allowed himself to be bitten +on the arm by two tarantulas, in the presence of witnesses. The +sensation was equivalent to that from an ant or a mosquito bite and +there were no other phenomena the first day. On the second day the wound +was inflamed and there was slight ulceration. It is clear that these +later symptoms were due to a secondary infection. These experiments have +been repeated by various observers, among whom may be mentioned Leon +Dufour, Josef Erker and Heinzel, and with the similar conclusion that +the bite of the Italian tarantula ordinarily causes no severe symptoms. +In this conclusion, Kobert, though firmly convinced of the poisonous +nature of some spiders, coincides. He also believes that striking +symptoms may be simulated or artificially induced by patients in order +to attract interest, or because they have been assured that the bite, +under all circumstances, caused tarantism. + +The so-called Russian tarantula, _Trochosa singoriensis_ (fig. 7), is +much larger than the Italian species, and is much feared. Kobert carried +out a series of careful experiments with this species and his results +have such an important bearing on the question of the venomous nature of +the tarantula that we quote his summary. Experimenting first on nearly a +hundred living specimens of _Trochosa singoriensis_ from Crimea he says +that: + +"The tarantulas, no matter how often they were placed on the skin, +handled, and irritated, could not be induced to bite either myself, the +janitor, or the ordinary experimental animals. The objection that the +tarantulas were weak and indifferent cannot stand, for as soon as I +placed two of them on the shaved skin of a rabbit, instead of an attack +on the animal, there began a furious battle between the two spiders, +which did not cease until one of the two was killed." + +[Illustration: 7. Trochosa singoriensis. After Kobert.] + +"Since the spiders would not bite, I carefully ground up the fresh +animals in physiological salt solution, preparing an extract which must +have contained, in solution, all of the poisonous substance of their +bodies. While in the case of _Latrodectus_, as we shall see, less than +one specimen sufficed to yield an active extract, I have injected the +filtered extract of six fresh Russian tarantulas, of which each one was +much heavier than an average _Latrodectus_, subcutaneously and into the +jugular vein of various cats without the animals dying or showing any +special symptoms. On the basis of my experiments I can therefore only +say that the quantity of the poison soluble in physiological salt +solution, even when the spiders are perfectly fresh and well nourished, +is very insignificant. That the poison of the Russian tarantula is not +soluble in physiological salt solution, is exceedingly improbable. +Moreover, I have prepared alcoholic extracts and was unable to find them +active. Since the Russian spider exceeds the Italian in size and in +intensity of the bite, it seems very improbable to me that the +pharmacological test of the Italian tarantula would yield essentially +other results than those from the Russian species." + +To the AVICULAROIDEA belong the largest and most formidable appearing of +the spiders and it is not strange that in the New World they have fallen +heir to the bad reputation, as well as to the name of the tarantula of +Europe. In this country they occur only in the South or in the far West, +but occasionally living specimens are brought to our northern ports in +shipments of bananas and other tropical produce, and are the source of +much alarm. It should be mentioned, however, that the large spider most +frequently found under such circumstances is not a tarantula at all, but +one of the Heteropodidæ, or giant crab-spiders, (fig. 8). + +[Illustration: 8. The giant crab-spider or banana spider (Heteropoda +venatoria). Natural size. After Comstock.] + +In spite of their prominence and the fear which they arouse there are +few accurate data regarding these American tarantulas. It has often been +shown experimentally that they can kill small birds and mammals, though +it is doubtful if these form the normal prey of any of the species, as +has been claimed. There is no question but that the mere mechanical +injury which they may inflict, and the consequent chances of secondary +infection, justify, in part, their bad reputation. In addition to the +injury from their bite, it is claimed that the body hairs of several of +the South American species are readily detached and are urticating. + +Recently, Phisalix (1912) has made a study of the physiological effects +of the venom of two Avicularoidea, _Phormictopus carcerides_ Pocock, +from Haiti and _Cteniza sauvagei_ Rossi, from Corsica. The glands were +removed aseptically and ground up with fine, sterilized sand in +distilled water. The resultant liquid was somewhat viscid, colorless, +and feebly alkaline. Injected into sparrows and mice the extract of +_Phormictopus_ proved very actively poisonous, that from a single spider +being sufficient to kill ten sparrows or twenty mice. It manifested +itself first and, above all, as a narcotic, slightly lowering the +temperature and paralyzing the respiration. Muscular and cardiac +weakening, loss of general sensibility, and the disappearance of +reflexes did not occur until near the end. The extract from _Cteniza_ +was less active and, curiously enough, the comparative effect on +sparrows and on mice was just reversed. + + +SPIDERS OF THE GENUS LATRODECTUS.--While most of the popular accounts of +evil effects from the bites of spiders will not stand investigation, it +is a significant fact that, the world over, the best authenticated +records refer to a group of small and comparatively insignificant +spiders belonging to the genus _Latrodectus_, of the family Theridiidæ. +The dread "Malmigniatte" of Corsica and South Europe, the "Karakurte" of +southeastern Russia, the "Katipo" of New Zealand, the "Mena-vodi" and +"Vancoho" of Madagascar, and our own _Latrodectus mactans_, all belong +to this genus, and concerning all of these the most circumstantial +accounts of their venomous nature are given. These accounts are not mere +fantastic stories by uneducated natives but in many cases are reports +from thoroughly trained medical men. + +The symptoms produced are general, rather than local. As summarized by +Kobert (1901) from a study of twenty-two cases treated in 1888, in the +Kherson (Russia) Government Hospital and Berislaw (Kherson) District +Hospital the typical case, aside from complications, exhibits the +following symptoms. The victim suddenly feels the bite, like the sting +of a bee. Swelling of the barely reddened spot seldom follows. The +shooting pains, which quickly set in, are not manifested at the point of +injury but localized at the joints of the lower limb and in the region +of the hip. The severity of the pain forces the victim to the hospital, +in spite of the fact that they otherwise have a great abhorrence of it. +The patient is unable to reach the hospital afoot, or, at least, not +without help, for there is usually inability to walk. The patient, even +if he has ridden, reaches the hospital covered with cold sweat and +continues to perspire for a considerable period. His expression +indicates great suffering. The respiration may be somewhat dyspnÅ“ic, +and a feeling of oppression in the region of the heart is common. There +is great aversion to solid food, but increasing thirst for milk and tea. +Retention of urine, and constipation occur. Cathartics and, at night, +strong narcotics are desired. Warm baths give great relief. After three +days, there is marked improvement and usually the patient is dismissed +after the fifth. This summary of symptoms agrees well with other +trustworthy records. + +It would seem, then, that Riley and Howard (1889), who discussed a +number of accounts in the entomological literature, were fully justified +in their statement that "It must be admitted that certain spiders of the +genus _Latrodectus_ have the power to inflict poisonous bites, which may +(probably exceptionally and depending upon exceptional conditions) bring +about the death of a human being." + +And yet, until recently the evidence bearing on the question has been +most conflicting. The eminent arachnologist, Lucas, (1843) states that +he himself, had been repeatedly bitten by the Malmigniatte without any +bad effects. Dr. Marx, in 1890, gave before the Entomological Society of +Washington, an account of a series of experiments to determine whether +the bite of _Latrodectus mactans_ is poisonous or not. He described the +poison glands as remarkably small[A] and stated that he had introduced +the poison in various ways into guinea-pigs and rabbits without +obtaining any satisfactory results. Obviously, carefully conducted +experiments with the supposed venom were needed and fortunately they +have been carried out in the greatest detail by Kobert (1901). + +This investigator pointed out that there were two factors which might +account for the discrepancies in the earlier experiments. In the first +place, the poison of spiders, as of snakes, might be so exhausted after +two or three bites that further bites, following directly, might be +without visible effect. Secondly, the application of the poison by means +of the bite, is exceedingly inexact, since even after the most careful +selection of the point of application, the poison might in one instance +enter a little vein or lymph vessel, and in another case fail to do so. +Besides, there would always remain an incalculable and very large amount +externally, in the nonabsorptive epithelium. While all of these factors +enter into the question of the effect of the bite in specific instances, +they must be as nearly as possible obviated in considering the question +of whether the spiders really secrete a venom harmful to man. + +Kobert therefore sought to prepare extracts which would contain the +active principles of the poison and which could be injected in definite +quantities directly into the blood of the experimental animal. For this +purpose various parts of the spiders were rubbed up in a mortar with +distilled water, or physiological salt solution, allowed to stand for an +hour, filtered, and then carefully washed, by adding water drop by drop +for twenty-four hours. The filtrate and the wash-water were then united, +well mixed and, if necessary, cleared by centrifuging or by exposure to +cold. The mixture was again filtered, measured, and used, in part, for +injection and, in part, for the determination of the organic materials. + +Such an extract was prepared from the cephalothoraces of eight dried +specimens of the Russian _Latrodectus_ and three cubic centimeters of +this, containing 4.29 mg. of organic material, were injected into the +jugular vein of a cat weighing 2450 grams. The previously very active +animal was paralyzed and lay in whatever position it was placed. The +sensibility of the skin of the extremities and the rump was so reduced +that there was no reaction from cutting or sticking. There quickly +followed dyspnÅ“a, convulsions, paralysis of the respiratory muscles +and of the heart. In twenty-eight minutes the cat was dead, after having +exhibited exactly the symptoms observed in severe cases of poisoning of +man from the bite of this spider. + +These experiments were continued on cats, dogs, guinea pigs and various +other animals. Not only extracts from the cephalothorax, but from other +parts of the body, from newly hatched spiders, and from the eggs were +used and all showed a similar virulence. Every effort was made to avoid +sources of error and the experiments, conducted by such a recognized +authority in the field of toxicology, must be accepted as conclusively +showing that this spider and, presumably, other species of the genus +_Latrodectus_ against which the clinical evidence is quite parallel, +possess a poison which paralyzes the heart and central nervous system, +with or without preliminary stimulus of the motor center. If the +quantity of the poison which comes into direct contact with the blood is +large, there may occur hæmolysis and thrombosis of the vessels. + +On the other hand, check experiments were carried out, using similar +extracts of many common European spiders of the genera _Tegenaria_, +_Drassus_, _Agelena_, _Eucharia_ and _Argyroneta_, as well as the +Russian tarantula, _Lycosa singoriensis_. In no other case was the +effect on experimental animals comparable to the _Latrodectus_ extract. + +Kobert concludes that in its chemical nature the poison is neither an +alkaloid, nor a glycoside, nor an acid, but a toxalbumen, or poisonous +enzyme which is very similar to certain other animal poisons, notably +that of the scorpion. + +[Illustration: 9. Latrodectus mactans; (_a_) female, x 3; (_b_) venter +of female; (_c_) dorsum of male. After Comstock.] + +The genus _Latrodectus_ is represented in the United States by at least +two species, _L. mactans_ and _L. geometricus_. Concerning _L. mactans_ +there are very circumstantial accounts of serious injury and even death +in man[B]. _Latrodectus mactans_ is coal black, marked with red or +yellow or both. It has eight eyes, which are dissimilar in color and +are distinctly in front of the middle of the thorax, the lateral eyes of +each side widely separate. The tarsus of the fourth pair of legs has a +number of curved setæ in a single series. It has on the ventral side of +its abdomen an hour-glass shaped spot. The full-grown female is about +half an inch in length. Its globose abdomen is usually marked with one +or more red spots dorsally along the middle line. The male is about half +as long but has in addition to the dorsal spots, four pairs of stripes +along the sides. Immature females resemble the male in coloring (fig. +9). + +Regarding the distribution of _Latrodectus mactans_, Comstock states +that: "Although it is essentially a Southern species, it occurs in +Indiana, Ohio, Pennsylvania, New Hampshire, and doubtless other of the +Northern States." _L. geometricus_ has been reported from California. + + +OTHER VENOMOUS SPIDERS--While conclusive evidence regarding the venomous +nature of spiders is meager and relates almost wholly to that of the +genus _Latrodectus_, the group is a large one and we are not justified +in dismissing arbitrarily, all accounts of injury from their bites. +Several species stand out as especially needing more detailed +investigation. + +_Chiracanthium nutrix_ is a common European species of the family +Clubionidæ, concerning which there is much conflicting testimony. Among +the reports are two by distinguished scientists whose accounts of +personal experiences cannot be ignored. A. Forel allowed a spider of +this species to bite him and not only was the pain extreme, but the +general symptoms were so severe that he had to be helped to his house. +The distinguished arachnologist, Bertkau reports that he, himself, was +bitten and that an extreme, burning pain spread almost instantaneously +over the arm and into the breast. There were slight chills the same day +and throbbing pain at the wound lasted for days. While this particular +species is not found in the United States, there are two other +representatives of the genus and it is possible that they possess the +same properties. We are unaware of any direct experimental work on the +poison. + +_Epeira diadema_, of Europe, belongs to a wholly different group, that +of the orb-weavers, but has long been reputed venomous. Kobert was able +to prepare from it an extract whose effects were very similar to that +prepared from _Latrodectus_, though feebler in its action. Under +ordinary circumstances this spider is unable to pierce the skin of man +and though Kobert's results seem conclusive, the spider is little to be +feared. + +_Phidippus audax_ (_P. tripunctatus_) is one of our largest Attids, or +jumping spiders. The late Dr. O. Lugger describes a case of severe +poisoning from the bite of this spider and though details are lacking, +it is quite possible that this and other large species of the same +group, which stalk their prey, may possess a more active poison than +that of web-building species. + + +SUMMARY--It is clearly established that our common spiders are not to be +feared and that the stories regarding their virulence are almost wholly +without foundation. On the other hand, the chances of secondary +infection from the bites of some of the more powerful species are not to +be ignored. + +Probably all species possess a toxin secreted by the poison gland, +virulent for insects and other normal prey of the spiders, but with +little or no effect on man. + +There are a very few species, notably of the genus _Latrodectus_, and +possibly including the European _Chiracanthium nutrix_ and _Epeira +diadema_, which possess, in addition, a toxalbumen derived from the +general body tissue, which is of great virulence and may even cause +death in man and the higher animals. + +[Illustration: 10. A whip-scorpion (Mastigoproctus giganteus). Half +natural size. After Comstock.] + + +THE PEDIPALPIDA OR WHIP-SCORPIONS + +The tailed whip-scorpions, belonging to the family Thelyphonidæ, are +represented in the United States by the giant whip-scorpion +_Mastigoproctus giganteus_ (fig. 10), which is common in Florida, Texas +and some other parts of the South. In Florida, it is locally known as +the "grampus" or "mule-killer" and is very greatly feared. There is no +evidence that these fears have any foundation, and Dr. Marx states that +there is neither a poison gland nor a pore in the claw of the +chelicera. + + +THE SCORPIONIDA, OR TRUE SCORPIONS + +The true scorpions are widely distributed throughout warm countries and +everywhere bear an evil reputation. According to Comstock (1912), about +a score of species occur in the Southern United States. These are +comparatively small forms but in the tropics members of this group may +reach a length of seven or eight inches. They are pre-eminently +predaceous forms, which lie hidden during the day and seek their prey by +night. + +[Illustration: 11. A true scorpion. After Comstock.] + +The scorpions (fig. 11) possess large pedipalpi, terminated by strongly +developed claws, or chelæ. They may be distinguished from all other +Arachnids by the fact that the distinctly segmented abdomen is divided +into a broad basal region of seven segments and a terminal, slender, +tail-like division of five distinct segments. + +The last segment of the abdomen, or telson, terminates in a +ventrally-directed, sharp spine, and contains a pair of highly developed +poison glands. These glands open by two small pores near the tip of the +spine. Most of the species when running carry the tip of the abdomen +bent upward over the back, and the prey, caught and held by the +pedipalpi, is stung by inserting the spine of the telson and allowing it +to remain for a time in the wound. + +The glands themselves have been studied in _Prionurus citrinus_ by +Wilson (1904). He found that each gland is covered by a sheet of muscle +on its mesal and dorsal aspects, which may be described as the +_compressor muscle_. The muscle of each side is inserted by its edge +along the ventral inner surface of the chitinous wall of the telson, +close to the middle line, and by a broader insertion laterally. A layer +of fine connective tissue completely envelops each gland and forms the +basis upon which the secreting cells rest. The secreting epithelium is +columnar; and apparently of three different types of cells. + +1. The most numerous have the appearance of mucous cells, resembling the +goblet cells of columnar mucous membranes. The nucleus, surrounded by a +small quantity of protoplasm staining with hæmatoxylin, lies close to +the base of the cell. + +2. Cells present in considerable numbers, the peripheral portions of +which are filled with very numerous fine granules, staining with acid +dyes such as methyl orange. + +3. Cells few in number, filled with very large granules, or irregular +masses of a substance staining with hæmatoxylin. + +The poison, according to Kobert (1893), is a limpid, acid-reacting +fluid, soluble in water but insoluble in absolute alcohol and ether. +There are few data relative to its chemical nature. Wilson (1901) states +that a common Egyptian species, _Buthus quinquestriatus_, has a specific +gravity of 1.092, and contains 20.3% of solids and 8.4% ash. + +The venom of different species appears to differ not only quantitatively +but qualitatively. The effects of the bite of the smaller species of the +Southern United States may be painful but there is no satisfactory +evidence that it is ever fatal. On the other hand, certain tropical +species are exceedingly virulent and cases of death of man from the bite +are common. + +In the case of _Buthus quinquestriatus_, Wilson (1904) found the +symptoms in animals to be hypersecretion, salivation and lachrymation, +especially marked, convulsions followed by prolonged muscular spasm; +death from asphyxia. The temperature shows a slight, rarely +considerable, rise. Rapid and considerable increase of blood-pressure +(observed in dogs) is followed by a gradual fall with slowing of the +heart-beat. The coagulability of the blood is not affected. + +An interesting phase of Wilson's work was the experiments on desert +mammals. The condition under which these animals exist must frequently +bring them in contact with scorpions, and he found that they possess a +degree of immunity to the venom sufficient at least to protect them from +the fatal effects of the sting. + +As far as concerns its effect on man, Wilson found that much depended +upon the age. As high as 60 per cent of the cases of children under +five, resulted fatally. Caroroz (1865), states that in a Mexican state +of 15,000 inhabitants, the scorpions were so abundant and so much feared +that the authorities offered a bounty for their destruction. A result +was a large number of fatalities, over two hundred per year. Most of the +victims were children who had attempted to collect the scorpions. + +The treatment usually employed in the case of bites by the more +poisonous forms is similar to that for the bite of venomous snakes. +First, a tight ligature is applied above the wound so as to stop the +flow of blood and lymph from that region. The wound is then freely +excised and treated with a strong solution of permanganate of potash, or +with lead and opium lotion. + +In recent years there have been many attempts to prepare an antivenom, +or antiserum comparable to what has been used so effectively in the case +of snake bites. The most promising of these is that of Todd (1909), +produced by the immunization of suitable animals. This antivenom proved +capable of neutralizing the venom when mixed _in vitro_ and also acts +both prophylactically and curatively in animals. Employed curatively in +man, it appears to have a very marked effect on the intense pain +following the sting, and the evidence so far indicates that its prompt +use greatly reduces the chance of fatal results. + + +THE SOLPUGIDA, OR SOLPUGIDS + +The SOLPUGIDA are peculiar spider-like forms which are distinguished +from nearly all other arachnids by the fact that they possess no true +cephalothorax, the last two leg-bearing segments being distinct, +resembling those of the abdomen in this respect. The first pair of legs +is not used in locomotion but seemingly functions as a second pair of +pedipalpi. Figure 12 illustrates the striking peculiarities of the +group. They are primarily desert forms and occur in the warm zones of +all countries. Of the two hundred or more species, Comstock lists twelve +as occurring in our fauna. These occur primarily in the southwest. + +[Illustration: 12. A solpugid (Eremobates cinerea). After Comstock.] + +The Solpugida have long borne a bad reputation and, regarding virulence, +have been classed with the scorpions. Among the effects of their bites +have been described painful swelling, gangrene, loss of speech, cramps, +delirium, unconsciousness and even death. Opposed to the numerous loose +accounts of poisoning, there are a number of careful records by +physicians and zoölogists which indicate clearly that the effects are +local and though they may be severe, they show not the slightest symptom +of direct poisoning. + +More important in the consideration of the question is the fact that +there are neither poison glands nor pores in the fangs for the exit of +any poisonous secretion. This is the testimony of a number of prominent +zoölogists, among whom is Dr. A. Walter, who wrote to Kobert at length +on the subject and whose conclusions are presented by him. + +However, it should be noted that the fangs are very powerful and are +used in such a manner that they may inflict especially severe wounds. +Thus, there may be more opportunity for secondary infection than is +usual in the case of insect wounds. + +The treatment of the bite of the Solpugida is, therefore, a matter of +preventing infection. The wound should be allowed to bleed freely and +then washed out with a 1:3000 solution of corrosive sublimate, and, if +severe, a wet dressing of this should be applied. If infection takes +place, it should be treated in the usual manner, regardless of its +origin. + + +THE ACARINA, OR MITES AND TICKS + +A number of the parasitic Acarina evidently secrete a specific poison, +presumably carried by the saliva, but in most cases its effect on man is +insignificant. There is an abundant literature dealing with the +poisonous effect of the bite of these forms, especially the ticks, but +until recently it has been confused by failure to recognize that various +species may transmit diseases of man, rather than produce injury through +direct poisoning. We shall therefore discuss the Acarina more especially +in subsequent chapters, dealing with parasitism and with disease +transmission. + +Nevertheless, after the evidence is sifted, there can be no doubt that +the bites of certain ticks may occasionally be followed by a direct +poisoning, which may be either local or general in its effects. Nuttall +(1908) was unable to determine the cause of the toxic effect, for, in +_Argas persicus_, the species most often implicated, he failed to get +the slightest local or general effect on experimental animals, from the +injection of an emulsion prepared by crushing three of the ticks. + +It seems clearly established that the bite of certain ticks may cause a +temporary paralysis, or even complete paralysis, involving the organs of +respiration or the heart, and causing death. In 1912, Dr. I. U. Temple, +of Pendleton, Oregon, reported several cases of what he called "acute +ascending paralysis" associated with the occurrence of ticks on the head +or the back of the neck. A typical severe case was that of a six year +old child, who had retired in her usual normal health. The following +morning upon arising she was unable to stand on her feet. She exhibited +paralysis extending to the knees, slight temperature, no pain, sensory +nerves normal, motor nerves completely paralyzed, reflexes absent. The +following day the paralysis had extended to the upper limbs, and before +night of the third day the nerves of the throat (hypoglossal) were +affected. The thorax and larynx were involved, breathing was labored, +she was unable to swallow liquids, phonation was impossible and she +could only make low, guttural sounds. At this stage, two ticks, fully +distended with blood, were found over the junction of the spinal column +with the occipital bones in the hollow depression. They were removed by +the application of undiluted creoline. Though the child's life was +despaired of, by the following morning she was very much improved. By +evening she was able to speak. The paralysis gradually receded, +remaining longest in the feet, and at the end of one week the patient +was able to go home. + +There was some doubt as to the exact species of tick implicated in the +cases which Dr. Temple reported, although the evidence pointed strongly +to _Dermacentor venustus_.[C] Somewhat later, Hadwen (1913) reported +that "tick paralysis" occurs in British Columbia, where it affects not +only man, but sheep and probably other animals. It is caused by the +bites of _Dermacentor venustus_ and was experimentally produced in lambs +and a dog (Hadwen and Nuttall, 1913). It is only when the tick begins to +engorge or feed rapidly, some days after it has become attached, that +its saliva produces pathogenic effects. + +Ulceration following tick bite is not uncommon. In many of the instances +it is due to the file-like hypostome, with its recurved teeth, being +left in the wound when the tick is forcibly pulled off. + + +THE MYRIAPODA, OR CENTIPEDES AND MILLIPEDES + +The old class, Myriapoda includes the DIPLOPODA, or millipedes, and the +CHILOPODA, or centipedes. The present tendency is to raise these groups +to the rank of classes. + + +The Diplopoda + +The DIPLOPODA, or millipedes (fig. 13), are characterized by the +presence of two pairs of legs to a segment. The largest of our local +myriapods belong to this group. They live in moist places, feeding +primarily on decaying vegetable matter, though a few species +occasionally attack growing plants. + +[Illustration: 13. A millipede. After Comstock.] + +The millipedes are inoffensive and harmless. _Julus terrestris_, and +related species, when irritated pour out over the entire body a +yellowish secretion which escapes from cutaneous glands. It is volatile, +with a pungent odor, and Phisalix (1900) has shown that it is an active +poison when injected into the blood of experimental animals. This, +however, does not entitle them to be considered as poisonous arthropods, +in the sense of this chapter, any more than the toad can be considered +poisonous to man because it secretes a venom from its cutaneous glands. + + +The Chilopoda + +[Illustration: 14. Two common centipedes. + +(_a_) Lithobius forficatus. After Comstock. + +(_b_) Scutigera forceps. Natural size; after Howard.] + +The CHILOPODA, or centipedes (fig. 14), unlike the millipedes, are +predaceous forms, and possess well developed poison glands for killing +their prey. These glands are at the base of the first pair of legs +(fig. 15), which are bent forward so as to be used in holding their +prey. The legs terminate in a powerful claw, at the tip of which is the +outlet of the poison glands. + +The poison is a limpid, homogeneous, slightly acid fluid, which +precipitates in distilled water. Briot (1904) extracted it from the +glands of _Scolopendra morsitans_, a species common in central France, +and found that it was actively venomous for the ordinary experimental +animals. A rabbit of two kilograms weight received an injection of three +cubic centimeters in the vein of the ear and died in a minute. A white +rat, weighing forty-eight grams, received one and a half cubic +centimeters in the hind leg. There was an almost immediate paralysis of +the leg and marked necrosis of the tissues. + +[Illustration: 15. Mandible of Scolopendra cingulata showing venom +gland. After Dubosq.] + +As for the effect on man, there is little foundation for the fear with +which centipedes are regarded. Our native species produce, at most, +local symptoms,--sometimes severe local pain and swelling,--but there is +no authentic record of fatal results. In the tropics, some of the +species attain a large size, _Scolopendra gigantea_ reaching a length of +nearly a foot. These forms are justly feared, and there is good evidence +that death sometimes, though rarely, results from their bite. + +One of the most careful accounts of death from the sting of the scorpion +is that of Linnell, (1914), which relates to a comparatively small +Malayan species, unfortunately undetermined. The patient, a coolie, aged +twenty, was admitted to a hospital after having been stung two days +previously on the left heel. For cure, the other coolies had made him +eat the head of the scorpion. On admission, the patient complained of +"things creeping all over the body". Temp. 102.8°. On the fourth day he +had paralysis of the legs, and on the fifth day motor paralysis to the +umbilicus, sensation being unaltered. On the sixth day there was +retention of the urine and on the ninth day (first test after third day) +sugar was present. On the thirteenth day the patient became comatose, +but could be roused to eat and drink. The temperature on the following +day fell below 95° and the patient was still comatose. Death fifteenth +day. + +Examination of the spinal (lumbar) cord showed acute disseminated +myelitis. In one part there was an acute destruction of the anterior +horn and an infiltration of round cells. In another portion Clarke's +column had been destroyed. The perivascular sheaths were crowded with +small round cells and the meninges were congested. Some of the cells of +the anterior horn were swollen and the nuclei eccentric; chromatolysis +had occurred in many of them. + +As for treatment, Castellani and Chalmers (1910), recommend bathing the +part well with a solution of ammonia (one in five, or one in ten). After +bathing, apply a dressing of the same alkali or, if there is much +swelling and redness, an ice-bag. If necessary, hypodermic injections of +morphine may be given to relieve the pain. At a later period +fomentations may be required to reduce the local inflammation. + + +THE HEXAPODA OR TRUE INSECTS + +There are a number of HEXAPODA, or true insects, which are, in one way +or another, poisonous to man. These belong primarily to the orders +Hemiptera, or true bugs; Lepidoptera, or butterflies and moths (larval +forms); Diptera, or flies; Coleoptera, or beetles; and Hymenoptera, or +ants, bees, and wasps. There are various ways in which they may be +poisonous. + +1. _Piercing_ or _biting_ forms may inject an irritating or poisonous +saliva into the wound caused by their mouth-parts. + +2. _Stinging forms_ may inject a poison, from glands at the caudal end +of the abdomen, into wounds produced by a specially modified ovipositer, +the _sting_. + +3. _Nettling_ properties may be possessed by the hairs of the insect. + +4. _Vescicating_, or _poisonous blood plasma_, or _body fluids_ are +known to exist in a large number of species and may, under exceptional +circumstances, affect man. + +For convenience of discussion, we shall consider poisonous insects under +these various headings. In this, as in the preceding discussion, no +attempt will be made to give an exhaustive list of the poisonous forms. +Typical instances will be selected and these will be chosen largely from +North American species. + + +PIERCING OR BITING INSECTS POISONOUS TO MAN + + +HEMIPTERA + +Several families of the true bugs include forms which, while normally +inoffensive, are capable of inflicting painful wounds on man. In these, +as in all of the Hemiptera, the mouth-parts are modified to form an +organ for piercing and sucking. This is well shown by the accompanying +illustration (fig. 16). + +The upper lip, or _labrum_, is much reduced and immovable, the lower +lip, or _labium_, is elongated to form a jointed sheath, within which +the lance-like mandibles and maxillæ are enclosed. The mandibles are +more or less deeply serrate, depending on the species concerned. + +[Illustration: 16. Beak of hemipteron.] + +The poison is elaborated by the salivary glands, excepting, possibly, in +_Belostoma_ where Locy is inclined to believe that it is secreted by the +maxillary glands. The salivary glands of the Hemiptera have been the +subject of much study but the most recent, comprehensive work has been +done by Bugnion and Popoff, (1908 and 1910) to whose text the reader is +referred for details. + +The Hemiptera have two pairs of salivary glands: the _primary gland_, of +which the efferent duct leads to the salivary syringe, and the +_accessory gland_, of which the very long and flexuous duct empties into +the primary duct at its point of insertion. Thus, when one observes the +isolated primary gland it appears as though it had efferent ducts +inserted at the same point. In _Nepa_ and the _Fulgoridæ_ there are two +accessory glands and therefore apparently three ducts at the same point +on the primary gland. The _ensemble_ differs greatly in appearance in +different species but we shall show here Bugnion and Popoff's figure of +the apparatus of _Notonecta maculata_, a species capable of inflicting a +painful bite on man (fig. 17). + +[Illustration: 17. Salivary glands of Notonecta maculata. After Bugnion +and Popoff.] + +[Illustration: 18. Pharyngeal syringe or salivary pump of Fulgora +maculata. After Bugnion and Popoff.] + +[Illustration: 19. Heteroptera, (_a_) Melanolestes picipes; (_b_) +Notonecta undulata; (_c_, _d_) Aradus robustus (_c_) adult, (_d_) nymph, +much enlarged; (_e_) Arilus cristatus; (_f_) Belostoma americana; (_g_) +Nabis (Coriscus) subcoleoptratus, enlarged; (_h_) Cimex lectularius, +(_i_) Oeciacus vicarius, much enlarged; (_j_) Lyctocoris fitchii, much +enlarged. After Lugger.] + +Accessory to the salivary apparatus there is on the ventral side of the +head, underneath the pharynx, a peculiar organ which the Germans have +called the "Wanzenspritze," or syringe. The accompanying figure of the +structure in _Fulgora maculata_ (fig. 18) shows its relation to the +ducts of the salivary glands and to the beak. It is made up of a +dilatation forming the body of the pump, in which there is a chitinous +piston. Attached to the piston is a strong retractor muscle. The +function of the salivary pump is to suck up the saliva from the salivary +ducts and to force it out through the beak. + +Of the Hemiptera reported as attacking man, we shall consider briefly +the forms most frequently noted. + + +The NOTONECTIDÆ, or _back swimmers_, (fig. 19_b_) are small, aquatic +bugs that differ from all others in that they always swim on their +backs. They are predaceous; feeding on insects and other small forms. +When handled carelessly they are able to inflict a painful bite, which +is sometimes as severe as the sting of a bee. In fact, they are known in +Germany as "Wasserbienen." + + +The BELOSTOMATIDÆ, or _giant water bugs_, (fig. 19_f_) include the +largest living Hemiptera. They are attracted to lights and on account of +the large numbers which swarm about the electric street lamps in some +localities they have received the popular name "electric light bugs." +Our largest representatives in the northern United States belong to the +two genera _Belostoma_ and _Banacus_, distinguished from each other by +the fact that _Belostoma_ has a groove on the under side of the femur of +the front leg, for the reception of the tibia. + +The salivary glands of Belostoma were figured by Leidy (1847) and later +were studied in more detail by Locy (1884). There are two pairs of the +glands, those of one pair being long and extending back as far as the +beginning of the abdomen, while the others are about one-fourth as long. +They lie on either side of the Å“sophagus. On each side of the +Å“sophagus there is a slender tube with a sigmoid swelling which may +serve as a poison reservoir. In addition to this salivary system, there +is a pair of very prominent glands on the ventral side of the head, +opening just above the base of the beak. These Locy has called the +"cephalic glands" and he suggests that they are the source of the +poison. They are the homologues of the maxillary glands described for +other Hemiptera, and it is by no means clear that they are concerned +with the production of venom. It seems more probable that in +_Belostoma_, as in other Hemiptera, it is produced by the salivary +glands, though the question is an open one. + +The Belostomatidæ feed not only on insects, but on small frogs, fish, +salamanders and the like. Matheson (1907) has recorded the killing of a +good-sized bird by _Belostoma americana_. A woodpecker, or flicker, was +heard to utter cries of distress, and fluttered and fell from a tree. On +examination it was found that a bug of this species had inserted its +beak into the back part of the skull and was apparently busily engaged +in sucking the blood or brains of the bird. Various species of +_Belostoma_ have been cited as causing painful bites in man. We can +testify from personal experience that the bite of _Belostoma americana_ +may almost immediately cause severe, shooting pains that may extend +throughout the arm and that they may be felt for several days. + +[Illustration: 20. Reduvius (OpsicÅ“tus) personatus. (×2).] + +Relief from the pain may be obtained by the use of dilute ammonia, or a +menthol ointment. In the not uncommon case of secondary infection the +usual treatment for that should be adopted. + +[Illustration: 21. (_a_) Reduvius personatus, nymph. Photograph by M. V. +S.] + + +The REDUVIIDÆ, or _assassin-bugs_ are capable of inflicting very painful +wounds, as most collectors of Hemiptera know to their sorrow. Some +species are frequently to be found in houses and outhouses and Dr. +Howard suggests that many of the stories of painful spider bites relate +to the attack of these forms. + +[Illustration: 21. (_b_) Reduvius personatus, adult (×2) Photograph by +M. V. S.] + +An interesting psychological study was afforded in the summer of 1899, +by the "kissing-bug" scare which swept over the country. It was reported +in the daily papers that a new and deadly bug had made its appearance, +which had the unpleasant habit of choosing the lips or cheeks for its +point of attack on man. So widespread were the stories regarding this +supposedly new insect that station entomologists all over the country +began to receive suspected specimens for identification. At Cornell +there were received, among others, specimens of stone-flies, may-flies +and even small moths, with inquiries as to whether they were +"kissing-bugs." + +[Illustration: 22. Rasahus biguttatus. (×2). After Howard.] + +Dr. L. O. Howard has shown that the scare had its origin in newspaper +reports of some instances of bites by either _Melanolestes picipes_ +(fig. 19a) or _Opsicoetes personatus_ (fig. 20), in the vicinity of +Washington, D. C. He then discusses in considerable detail the more +prominent of the Reduviidæ which, with greater or less frequency pierce +the skin of human beings. These are _Opsicoetes personatus_, +_Melanolestes picipes_, _Coriscus subcoleoptratus_ (fig. 19_g_), +_Rasahus thoracicus_, _Rasahus biguttatus_ (fig. 22), _Conorhinus +sanguisugus_ (fig. 71), and _Melanolestes abdominalis_ (fig. 23). + +[Illustration: 23. Melanolestes abdominalis (×2). After Marlatt.] + +One of the most interesting of these species is _Reduvius personatus_, +(= _OpsicÅ“tus personatus_), which is popularly known as the "masked +bed-bug hunter." It owes this name to the fact that the immature nymphs +(fig. 21) have their bodies and legs completely covered by dust and +lint, and that they are supposed to prey upon bed-bugs. LeConte is +quoted by Howard as stating that "This species is remarkable for the +intense pain caused by its bite. I do not know whether it ever willingly +plunges its rostrum into any person, but when caught, or unskilfully +handled it always stings. In this case the pain is almost equal to the +bite of a snake, and the swelling and irritation which result from it +will sometimes last for a week." + +A species which very commonly attacks man is _Conorhinus sanguisugus_, +the so-called "big bed-bug" of the south and southern United States. It +is frequently found in houses and is known to inflict an exceedingly +painful bite. As in the case of a number of other predaceous Hemiptera, +the salivary glands of these forms are highly developed. The effect of +the bite on their prey and, as Marlatt has pointed out, the constant and +uniform character of the symptoms in nearly all cases of bites in man, +clearly indicate that their saliva contains a specific substance. No +satisfactory studies of the secretions have been made. On the other +hand, Dr. Howard is doubtless right in maintaining that the very serious +results which sometimes follow the bite are due to the introduction of +extraneous poison germs. This is borne out by the symptoms of most of +the cases cited in literature and also by the fact that treatment with +corrosive sublimate, locally applied to the wound, has yielded favorable +results. + + +OTHER HEMIPTERA REPORTED AS POISONOUS TO MAN--A large number of other +Hemiptera have been reported as attacking man. Of these, there are +several species of Lygæidæ, Coreidæ, and Capsidæ. Of the latter, _Lygus +pratensis_, the tarnished plant-bug, is reported by Professor Crosby as +sucking blood. _Orthotylus flavosparsus_ is another Capsid which has +been implicated. _Empoasca mali_ and _Platymetopius acutus_ of the +Jassidæ have also been reported as having similar habits. + +Whenever the periodical cicada or "seventeen-year locust" becomes +abundant, the newspapers contain accounts of serious results from its +bites. The senior author has made scores of attempts to induce this +species to bite and only once successfully. At that time the bite was in +no wise more severe than a pin-prick. A student in our department +reports a similar experience. There is no case on record which bears +evidence of being worthy of any credence, whatsoever. + +Under the heading of poisonous Hemiptera we might consider the bed-bugs +and the lice. These will be discussed later, as parasites and as +carriers of disease, and therefore need only be mentioned here. + + +DIPTERA + +Several species of blood-sucking Diptera undoubtedly secrete a saliva +possessing poisonous properties. Chief among these are the Culicidæ, or +mosquitoes, and the Simuliidæ, or black-flies. As we shall consider +these forms in detail under the heading of parasitic species and +insects transmitting disease, we shall discuss here only the poison of +the mosquitoes. + +It is well known that mosquitoes, when they bite, inject into the wound +a minute quantity of poison. The effect of this varies according to the +species of mosquito and also depends very much on the susceptibility of +the individual. Soon after the bite a sensation of itching is noticed +and often a wheal, or eminence, is produced on the skin, which may +increase to a considerable swelling. The scratching which is induced may +cause a secondary infection and thus lead to serious results. Some +people seem to acquire an immunity against the poison. + +The purpose of this irritating fluid may be, as Reaumur suggested, to +prevent the coagulation of the blood and thus not only to cause it to +flow freely when the insect bites but to prevent its rapid coagulation +in the stomach. Obviously, it is not developed as a protective fluid, +and its presence subjects the group to the additional handicap of the +vengeance of man. + +[Illustration: 24. Diagram of a longitudinal section of a mosquito.] + +As to the origin of the poison, there has been little question, until +recent years, that it was a secretion from the salivary glands. +Macloskie (1888) showed that each gland is subdivided into three lobes, +the middle of which differs from the others in having evenly granulated +contents and staining more deeply than the others (fig. 24). This middle +lobe he regarded as the source of the poison. Bruck, (1911), by the use +of water, glycerine, chloroform, and other fluids, extracted from the +bodies of a large number of mosquitoes a toxine which he calls +_culicin_. This he assumes comes from the salivary glands. Animal +experimentation showed that this extract possessed hemolytic powers. +Inoculated into the experimenter's own skin it produced lesions which +behaved exactly as do those of mosquito bites. + +Similarly, most writers on the subject have concurred with the view that +the salivary glands are the source of the poison. However, recent work, +especially that of Nuttall and Shipley (1903), and Schaudinn (1904), has +shown that the evidence is by no means conclusive. Nuttall dissected out +six sets (thirty-six acini) of glands from freshly killed _Culex +pipiens_ and placed them in a drop of salt solution. The drop was +allowed to dry, it being thought that the salt crystals would facilitate +the grinding up of the glands with the end of a small glass rod, this +being done under microscopic control. After grinding up, a small drop of +water was added of the size of the original drop of saline, and an equal +volume of human blood taken from the clean finger-tip was quickly mixed +therewith, and the whole drawn up into a capillary tube. Clotting was +not prevented and no hemolysis occurred. Salivary gland emulsion added +to a dilute suspension of corpuscles did not lead to hemolysis. This +experiment was repeated a number of times, with slight modification, but +with similar results. The data obtained from the series "do not support +the hypothesis that the salivary glands, at any rate in _Culex pipiens_, +contain a substance which prevents coagulation." + +Much more detailed, and the more important experiments made along this +line, are those of Schaudinn (1904). The results of these experiments +were published in connection with a technical paper on the alternation +of generations and of hosts in _Trypanosoma_ and _Spirochæta_, and for +this reason seem to have largely escaped the notice of entomologists. +They are so suggestive that we shall refer to them in some detail. + +Schaudinn observed that the three Å“sophageal diverticula (commonly, +but incorrectly, known as the "sucking stomach") (fig. 24) usually +contain large bubbles of gas and in addition, he always found yeast +cells. On the ground of numerous observations, Schaudinn was convinced +that these yeast plants are normal and constant commensals of the +insect. He regarded them as the cause of the gas bubbles to be found in +diverticula. It was found that as the insect fed, from time to time the +abdomen underwent convulsive contractions which resulted in the emptying +of the Å“sophageal diverticula and the salivary glands through blood +pressure. + +In order to test the supposed toxic action of the salivary glands, +Schaudinn repeatedly introduced them under his skin and that of his +assistant, in a drop of salt solution, and never obtained a suggestion +of the irritation following a bite of the insect, even though the glands +were carefully rubbed to fragments after their implantation. Like +Nuttall, he failed to get satisfactory evidence that the secretion of +the salivary glands retarded coagulation of the blood. + +He then carefully removed the Å“sophageal diverticula with their +content of yeast and introduced them into an opening in the skin of the +hand. Within a few seconds there was noticeable the characteristic +itching irritation of the mosquito bite; and in a short time there +appeared reddening and typical swelling. This was usually much more +severe than after the usual mosquito bite, and the swelling persisted +and itched longer. This was because by the ordinary bite of the mosquito +most of the yeast cells are again sucked up, while in these experiments +they remained in the wound. These experiments were repeated a number of +times on himself, his assistant and others, and always with the same +result. From them Schaudinn decided that the poisonous action of the +mosquito bite is caused by an enzyme from a commensal fungus. These +conclusions have not, as yet, been satisfactorily tested. + +Relief from the effect of the mosquito bite may be obtained by bathing +the swellings with weak ammonia or, according to Howard, by using moist +soap. The latter is to be rubbed gently on the puncture and is said to +speedily allay the irritation. Howard also quotes from the _Journal of +Tropical Medicine and Hygiene_ to the effect that a few drops of a +solution of thirty to forty grains of iodine to an ounce of saponated +petroleum rubbed into the mosquito bite, or wasp sting, allay the pain +instantaneously. + +Methods of mosquito control will be discussed later, in considering +these insects as parasites and as carriers of disease. + + +STINGING INSECTS + +The stinging insects all belong to the order HYMENOPTERA. In a number of +families of this group the ovipositor is modified to form a sting and is +connected with poison-secreting glands. We shall consider the apparatus +of the honey-bee and then make briefer reference to that of other forms. + + +APIS MELLIFICA, THE HONEY BEE--The sting of the worker honey-bee is +situated within a so-called sting chamber at the end of the abdomen. +This chamber is produced by the infolding of the greatly reduced and +modified eighth, ninth and tenth abdominal segments into the seventh.[D] +From it the dart-like sting can be quickly exserted. + +The sting (fig. 25) is made up of a central shaft, ventro-laterad of +which are the paired _lancets_, or darts, which are provided with sharp, +recurved teeth. Still further laterad lie the paired whitish, +finger-like _sting palpi_. Comparative morphological as well as +embryological studies have clearly established that these three parts +correspond to the three pairs of gonopophyses of the ovipositor of more +generalized insects. + +[Illustration: 25. Sting of a honey bee. _Psn Sc_, base of acid poison +gland; _B Gl_, alkaline poison gland; _Stn Plp_, sting palpi; _Sh B_, +bulb of sting; _Sh A_, basal arm; _Lct_, lancets or darts; _Sh s_, shaft +of sting. Modified from Snodgrass.] + +[Illustration: 26. Poison apparatus of a honey bee. Modified from +Snodgrass.] + +An examination of the internal structures (fig. 26) reveals two distinct +types of poison glands, the acid-secreting and the alkaline-secreting +glands, and a prominent poison reservoir. In addition, there is a small +pair of accessory structures which have been called lubricating glands, +on account of the supposed function of their product. The acid-secreting +gland empties into the distal end of the poison reservoir which in turn +pours the secretion into the muscular bulb-like enlargement at the base +of the shaft. The alkaline secreting gland empties into the bulb ventrad +of the narrow neck of the reservoir. + +The poison is usually referred to as formic acid. That it is not so +easily explained has been repeatedly shown and is evidenced by the +presence of the two types of glands. Carlet maintains that the product +of either gland is in itself innocent,--it is only when they are +combined that the toxic properties appear. + +The most detailed study of the poison of the honey-bee is that of Josef +Langer (1897), who in the course of his work used some 25,000 bees. +Various methods of obtaining the active poison for experimental purposes +were used. For obtaining the pure secretion, bees were held in the +fingers and compressed until the sting was exserted, when a clear drop +of the poison was visible at its tip. This was then taken up in a +capillary tube or dilute solutions obtained by dipping the tip of the +sting into a definite amount of distilled water. + +An aqueous solution of the poison was more readily obtained by pulling +out the sting and poison sacs by means of forceps, and grinding them up +in water. The somewhat clouded fluid was then filtered one or more +times. For obtaining still greater quantities, advantage was taken of +the fact that while alcohol coagulates the poison, the active principle +remains soluble in water. Hence the stings with the annexed glands where +collected in 96 per cent alcohol, after filtering off of the alcohol +were dried at 40° C., then rubbed to a fine powder and this was +repeatedly extracted with water. Through filtering of this aqueous +extract there was obtained a yellowish-brown fluid which produced the +typical reactions, according to concentration of the poison. + +The freshly expelled drop of poison is limpid, of distinct acid +reaction, tastes bitter and has a delicate aromatic odor. On +evaporation, it leaves a sticky residue, which at 100 degrees becomes +fissured, and suggests dried gum arabic. The poison is readily soluble +in water and possesses a specific gravity of 1.1313. On drying at room +temperature, it leaves a residue of 30 per cent, which has not lost in +poisonous action or in solubility. In spite of extended experiments, +Langer was unable to determine the nature of the active principle. He +showed that it was not, as had been supposed, an albuminous body, but +rather an organic base. + +The pure poison, or the two per cent aqueous solution, placed on the +uninjured skin showed absolutely no irritating effect, though it +produced a marked reaction on the mucus membrane of the nose or eye. A +single drop of one-tenth per cent aqueous solution of the poison brought +about a typical irritation in the conjunctiva of the rabbit's eye. On +the other hand, the application of a drop of the poison, or its +solution, to the slightest break in the skin, or by means of a needle +piercing the skin, produced typical effects. There is produced a local +necrosis, in the neighborhood of which there is infiltration of +lymphocytes, Å“dema, and hyperæmia. + +The effect of the sting on man (fig. 27) is usually transitory but there +are some individuals who are made sick for hours, by a single sting. +Much depends, too, on the place struck. It is a common experience that +an angry bee will attempt to reach the eye of its victim and a sting on +the lid may result in severe and prolonged swelling. In the case of a +man stung on the cheek, Legiehn observed complete aphonia and a breaking +out of red blotches all over the body. A sting on the tongue has been +known to cause such collateral Å“dema as to endanger life through +suffocation. Cases of death of man from the attacks of bees are rare but +are not unknown. Such results are usually from a number of stings but, +rarely, death has been known to follow a single sting, entering a blood +vessel of a particularly susceptible individual. + +[Illustration: Effect of bee stings. After Root.] + +It is clearly established that partial immunity from the effects of the +poison may be acquired. By repeated injections of the venom, mice have +been rendered capable of bearing doses that certainly would have killed +them at first. It is a well-known fact that most bee-keepers become +gradually hardened to the stings, so that the irritation and the +swelling become less and less. Some individuals have found this immunity +a temporary one, to be reacquired each season. A striking case of +acquired immunity is related by the Roots in their "A B C and X Y Z of +Bee Culture." The evidence in the case is so clear that it should be +made more widely available and hence we quote it here. + +A young man who was determined to become a bee-keeper, was so +susceptible to the poison that he was most seriously affected by a +single sting, his body breaking out with red blotches, breathing growing +difficult, and his heart action being painfully accelerated. "We finally +suggested taking a live bee and pressing it on the back of his hand +until it merely pierced his skin with the sting, then immediately +brushing off both bee and sting. This was done and since no serious +effect followed, it was repeated inside of four or five days. This was +continued for some three or four weeks, when the patient began to have a +sort of itching sensation all over his body. The hypodermic injections +of bee-sting poison were then discontinued. At the end of a month they +were repeated at intervals of four or five days. Again, after two or +three weeks the itching sensation came on, but it was less pronounced. +The patient was given a rest of about a month, when the doses were +repeated as before." By this course of treatment the young man became so +thoroughly immunized that neither unpleasant results nor swelling +followed the attacks of the insects and he is able to handle bees with +the same freedom that any experienced bee-keeper does. + +In an interesting article in the _Entomological News_ for November, +1914, J. H. Lovell calls attention to the fact that "There has been a +widespread belief among apiarists that a beekeeper will receive more +stings when dressed in black than when wearing white clothing. A large +amount of evidence has been published in the various bee journals +showing beyond question that honey-bees under certain conditions +discriminate against black. A few instances may be cited in +illustration. Of a flock of twelve chickens running in a bee-yard seven +black ones were stung to death, while five light colored ones escaped +uninjured. A white dog ran among the bee-hives without attracting much +attention, while at the same time a black dog was furiously assailed by +the bees. Mr. J. D. Byer, a prominent Canadian beekeeper, relates that a +black and white cow, tethered about forty feet from an apiary, was one +afternoon attacked and badly stung by bees. On examination it was found +that the black spots had five or six stings to one on the white. All +noticed this fact, although no one was able to offer any explanation. A +white horse is in much less danger of being stung, when driven near an +apiary, than a black one. It has, indeed, been observed repeatedly that +domestic animals of all kinds, if wholly or partially black, are much +more liable to be attacked by bees, if they wander among the hives, than +those which are entirely white." + +In order to test the matter experimentally, the following series of +experiments was performed. In the language of the investigator: + +"On a clear, warm day in August I dressed wholly in white with the +exception of a black veil. Midway on the sleeve of my right arm there +was sewed a band of black cloth ten inches wide. I then entered the +bee-yard and, removing the cover from one of the hives, lifted a piece +of comb with both hands and gently shook it. Instantly many of the bees +flew to the black band, which they continued to attack as long as they +were disturbed. Not a single bee attempted to sting the left sleeve, +which was of course entirely white, and very few even alighted upon it." + +"This experiment was repeated a second, third and fourth time; in each +instance with similar results. I estimated the number of bees on the +band of black cloth at various moments was from thirty to forty; it was +evident from their behavior that they were extremely irritable. To the +left white sleeve and other portions of my clothing they paid very +little attention; but the black veil was very frequently attacked." + +"A few days later the experiments were repeated, but the band of black +cloth, ten inches wide, was sewed around my left arm instead of around +the right arm as before. When the bees were disturbed, after the hive +cover had been removed, they fiercely attacked the band of black cloth +as in the previous experiences; but the right white sleeve and the white +suit were scarcely noticed. At one time a part of the black cloth was +almost literally covered with furiously stinging bees, and the black +veil was assailed by hundreds. The bees behaved in a similar manner when +a second hive on the opposite side of the apiary was opened." + +"A white veil which had been procured for this purpose, was next +substituted for the black veil. The result was most surprising, for, +whereas in the previous experiments hundreds of bees had attacked the +black veil, so few flew against the white veil as to cause me no +inconvenience. Undoubtedly beekeepers will find it greatly to their +advantage to wear white clothing when working among their colonies of +bees and manipulating the frames of the hives." + +When a honey-bee stings, the tip of the abdomen, with the entire sting +apparatus, is torn off and remains in the wound. Here the muscles +continue to contract, for some minutes, forcing the barbs deeper and +deeper into the skin, and forcing out additional poison from the +reservoir. + +Treatment, therefore, first consists in removing the sting without +squeezing out additional poison. This is accomplished by lifting and +scraping it out with a knife-blade or the fingernail instead of grasping +and pulling it out. Local application of alkalines, such as weak +ammonia, are often recommended on the assumption that the poison is an +acid to be neutralized on this manner, but these are of little or no +avail. They should certainly not be rubbed in, as that would only +accelerate the absorption of the poison. The use of cloths wrung out in +hot water and applied as hot as can be borne, affords much relief in the +case of severe stings. The application of wet clay, or of the end of a +freshly cut potato is sometimes helpful. + +In extreme cases, where there is great susceptibility, or where there +may have been many stings, a physician should be called. He may find +strychnine injections or other treatment necessary, if general symptoms +develop. + +[Illustration: 28. The poison apparatus of Formica. Wheeler, after +Forel.] + + +OTHER STINGING FORMS--Of the five thousand, or more, species of bees, +most possess a sting and poison apparatus and some of the larger forms +are capable of inflicting a much more painful sting than that of the +common honey-bee. In fact, some, like the bumble bees, possess the +advantage that they do not lose the sting from once using it, but are +capable of driving it in repeatedly. In the tropics there are found many +species of stingless bees but these are noted for their united efforts +to drive away intruders by biting. Certain species possess a very +irritating saliva which they inject into the wounds. + +The ants are not ordinarily regarded as worthy of consideration under +the heading of "stinging insects" but as a matter of fact, most of them +possess well developed stings and some of them, especially in the +tropics, are very justly feared. Even those which lack the sting possess +well-developed poison glands and the parts of the entire stinging +apparatus, in so far as it is developed in the various species, may +readily be homologized with those of the honey-bee. + +The ants lacking a sting are those of the subfamily CAMPONOTINÆ, which +includes the largest of our local species. It is an interesting fact +that some of these species possess the largest poison glands and +reservoir (fig. 28) and it is found that when they attack an enemy they +bring the tip of the abdomen forward and spray the poison in such a way +that it is introduced into the wound made by the powerful mandibles. + +[Illustration: 29. A harmless, but much feared larva, the "tomato worm." +Natural size. Photograph by M. V. S.] + +More feared than any of the other Hymenoptera are the hornets and wasps. +Of these there are many species, some of which attain a large size and +are truly formidable. Phisalix (1897), has made a study of the venom of +the common hornet and finds that, like the poison of the honey-bee, it +is neither an albuminoid nor an alkaloid. Its toxic properties are +destroyed at 120° C. Phisalix also says that the venom is soluble in +alcohol. If this be true, it differs in this respect from that of the +bee. An interesting phase of the work of Phisalix is that several of her +experiments go to show that the venom of hornets acts as a vaccine +against that of vipers. + + +NETTLING INSECTS + +So far, we have considered insects which possess poison glands connected +with the mouth-parts or a special sting and which actively inject their +poison into man. There remain to be considered those insects which +possess poisonous hairs or body fluids which, under favorable +circumstances, may act as poisons. To the first of these belong +primarily the larvæ of certain Lepidoptera. + + +LEPIDOPTERA + +[Illustration: 30. Another innocent but much maligned caterpillar, the +larva of the Regal moth. Photograph by M. V. S.] + +When we consider the reputedly poisonous larvæ of moths and butterflies, +one of the first things to impress us is that we cannot judge by mere +appearance. Various species of Sphingid, or hawk-moth larvæ, bear at the +end of the body a chitinous horn, which is often referred to as a +"sting" and regarded as capable of inflicting dangerous wounds. It would +seem unnecessary to refer to this absurd belief if it were not that each +summer the newspapers contain supposed accounts of injury from the +"tomato worm" (fig. 29) and others of this group. The grotesque, spiny +larva (fig. 30) of one of our largest moths, _Citheronia regalis_ is +much feared though perfectly harmless, and similar instances could be +multiplied. + +[Illustration: 31. The flannel moth (Lagoa crispata). (_a_) Poisonous +larva.] + +[Illustration: 31. (_b_) Adult. Enlarged. Photographs by M. V. S.] + +But if the larvæ are often misjudged on account of their ferocious +appearance, the reverse may be true. A group of most innocent looking +and attractive caterpillars is that of the flannel-moth larvæ, of which +_Lagoa crispata_ may be taken as an example. Its larva (fig. 31) has a +very short and thick body, which is fleshy and completely covered and +hidden by long silken hairs of a tawny or brown color, giving a convex +form to the upper side. Interspersed among these long hairs are +numerous short spines connected with underlying hypodermal poison +glands. These hairs are capable of producing a marked nettling effect +when they come in contact with the skin. This species is found in our +Atlantic and Southern States. Satisfactory studies of its poisonous +hairs and their glands have not yet been made. + +[Illustration: 32 The poisonous saddle back caterpillar. Empretia +(Sibine) stimulea. Photograph by M. V. S.] + +[Illustration: 33_a_. Io moth larvæ on willow. Photograph by M. V. S.] + +_Sibine stimulea (Empretia stimulea)_, or the saddle-back caterpillar +(fig. 32), is another which possesses nettling hairs. This species +belongs to the group of Eucleidæ, or slug caterpillars. It can be +readily recognized by its flattened form, lateral, bristling spines and +by the large green patch on the back resembling a saddle-cloth, while +the saddle is represented by an oval, purplish-brown spot. The small +spines are venomous and affect some persons very painfully. The larva +feeds on the leaves of a large variety of forest trees and also on +cherry, plum, and even corn leaves. It is to be found throughout the +Eastern and Southern United States. + +[Illustration: 33_b_. Io moth. Full grown larva. Photograph by M. V. S.] + +[Illustration: 34. Io moth. Adult. Photograph by M. V. S.] + +_Automeris io_ is the best known of the nettling caterpillars. It is the +larva of the Io moth, one of the Saturniidæ. The mature caterpillar, +(fig. 33), which reaches a length of two and one-half inches, is of a +beautiful pale green with sublateral stripes of cream and red color and +a few black spines among the green ones. The green radiating spines give +the body a mossy appearance. They are tipped with a slender chitinous +hair whose tip is readily broken off in the skin and whose poisonous +content causes great irritation. Some individuals are very susceptible +to the poison, while others are able to handle the larvæ freely without +any discomfort. The larvæ feed on a wide range of food plants. They are +most commonly encountered on corn and on willow, because of the +opportunities for coming in contact with them. + +[Illustration: 35. Larva of brown-tail moth. (Natural size). Photograph +by M. V. S.] + +The larvæ of the brown-tail moth (_Euproctis chrysorrhÅ“a_) (fig. 35 +and 36), where they occur in this country, are, on account of their +great numbers, the most serious of all poisonous caterpillars. It is not +necessary here, to go into details regarding the introduction of this +species from Europe into the New England States. This is all available +in the literature from the United States Bureau of Entomology and from +that of the various states which are fighting the species. Suffice to +say, there is every prospect that the pest will continue to spread +throughout the Eastern United States and Canada and that wherever it +goes it will prove a direct pest to man as well as to his plants. + +Very soon after the introduction of the species there occurred in the +region where it had gained a foothold, a mysterious dermatitis of man. +The breaking out which usually occurred on the neck or other exposed +part of the body was always accompanied by an intense itching. It was +soon found that this dermatitis was caused by certain short, barbed +hairs of the brown-tail caterpillars and that not only the caterpillars +but their cocoons and even the adult female moths might harbor these +nettling hairs and thus give rise to the irritation. In many cases the +hairs were wafted to clothing on the line and when this was worn it +might cause the same trouble. Still worse, it was found that very +serious internal injury was often caused by breathing or swallowing the +poisonous hairs. + +[Illustration: 36. Browntail moths. One male and two females. Photograph +by M. V. S.] + +The earlier studies seemed to indicate that the irritation was purely +mechanical in origin, the result of the minute barbed hairs working into +the skin in large numbers. Subsequently, however, Dr. Tyzzer (1907) +demonstrated beyond question that the trouble was due to a poison +contained in the hairs. In the first place, it is only the peculiar +short barbed hairs which will produce the dermatitis when rubbed on the +skin, although most of the other hairs are sharply barbed. Moreover, it +was found that in various ways the nettling properties could be +destroyed without modifying the structure of the hairs. This was +accomplished by baking for one hour at 110° C, by warming to 60° C in +distilled water, or by soaking in one per cent. or in one-tenth per +cent. of potassium hydrate or sodium hydrate. The most significant part +of his work was the demonstration of the fact that if the nettling +hairs are mingled with blood, they immediately produce a change in the +red corpuscles. These at once become coarsely crenated, and the roleaux +are broken up in the vicinity of the hair (fig. 37_b_). The corpuscles +decrease in size, the coarse crenations are transformed into slender +spines which rapidly disappear, leaving the corpuscles in the form of +spheres, the light refraction of which contrasts them sharply with the +normal corpuscles. The reaction always begins at the basal sharp point +of the hair. It could not be produced by purely mechanical means, such +as the mingling of minute particles of glass wool, the barbed hairs of a +tussock moth, or the other coarser hairs of the brown-tail, with the +blood. + +[Illustration: 37. (_a_) Ordinary hairs and three poison hairs of +subdorsal and lateral tubercles of the larva of the browntail moth. +Drawing by Miss Kephart.] + +The question of the source of the poison has been studied in our +laboratory by Miss Cornelia Kephart. She first confirmed Dr. Tyzzer's +general results and then studied carefully fixed specimens of the larvæ +to determine the distribution of the hairs and their relation to the +underlying tissues. + +[Illustration: 37. (_b_) Effect of the poison on the blood corpuscles of +man. After Tyzzer.] + +The poison hairs (fig. 37), are found on the subdorsal and lateral +tubercles (fig. 38), in bunches of from three to twelve on the minute +papillæ with which the tubercles are thickly covered. The underlying +hypodermis is very greatly thickened, the cells being three or four +times the length of the ordinary hypodermal cells and being closely +crowded together. Instead of a pore canal through the cuticula for each +individual hair, there is a single pore for each papillæ on a tubercle, +all the hairs of the papilla being connected with the underlying cells +through the same pore canal, (figs. 39 and 40). + +[Illustration: 38. Cross section of the larva of the browntail moth +showing the tubercles bearing the poison hairs. Drawing by Miss +Kephart.] + +The hypodermis of this region is of two distinct types of cells. First, +there is a group of slender fusiform cells, one for each poison hair on +the papilla, which are the trichogen, or hair-formative cells. They are +crowded to one side and towards the basement membrane by a series of +much larger, and more prominent cells (fig. 40), of which there is a +single one for each papilla. These larger cells have a granular +protoplasm with large nuclei and are obviously actively secreting. They +are so characteristic in appearance as to leave no question but that +they are the true poison glands. + +[Illustration: 39. Epithelium underlying poison hairs of the larva of +the browntail moth. Drawing by Miss Kephart.] + +Poisonous larvæ of many other species have been reported from Europe and +especially from the tropics but the above-mentioned species are the more +important of those occurring in the United States and will serve as +types. It should be noted in this connection that through some curious +misunderstanding GÅ“ldi (1913) has featured the larva of _Orgyia +leucostigma_, the white-marked tussock moth, as the most important of +the poisonous caterpillars of this country. Though there are occasional +reports of irritation from its hairs such cases are rare and there is no +evidence that there is any poison present. Indeed, subcutaneous +implantation of the hairs leads to no poisoning, but merely to temporary +irritation. + +[Illustration: 40. Same as figure 39, on larger scale.] + +Occasionally, the hairs of certain species of caterpillars find +lodgement in the conjunctiva, cornea, or iris of the eye of man and give +rise to the condition known as _opthalmia nodosa_. The essential feature +of this trouble is a nodular conjunctivitis which simulates tuberculosis +of the conjunctiva and hence has been called _pseudo-tubercular_. It may +be distinguished microscopically by the presence of the hairs. + +[Illustration: 41. (_a_) Nodular conjunctivitis in the eye of a child. +De Schweinitz and Shumway.] + +Numerous cases of opthalmia nodosa are on record. Of those from this +country, one of the most interesting is reported by de Schweinitz and +Shumway (1904). It is that of a child of fifteen years whose eye had +become inflamed owing to the presence of some foreign body. Downward and +inward on the bulbar conjunctiva were a number of flattened, +grayish-yellow nodules, between which was a marked congestion of the +conjunctival and episcleral vessels (fig. 41_a_). Twenty-seven nodules +could be differentiated, those directly in the center of the collection +being somewhat confluent and assuming a crescentic and circular +appearance. The nodules were excised and, on sectioning, were found to +be composed of a layer of spindle cells and round cells, outside of +which the tissue was condensed into a capsule. The interior consisted of +epithelioid cells, between which was a considerable intercellular +substance. Directly in the center of a certain number of nodules was +found the section of a hair (fig. 41_b_). The evidence indicated that +the injury had resulted from playing with caterpillars of one of the +Arctiid moths, _Spilosoma virginica_. Other reported cases have been +caused by the hairs of larvæ of _Lasiocampa rubi_, _L. pini_, +_Porthetria dispar_, _Psilura monacha_ and _Cnethocampa processionea_. + +[Illustration: 41_b_. Section through one of the nodules showing the +caterpillar hair. De Schweinitz and Shumway.] + + +RELIEF FROM POISONING BY NETTLING LARVÆ--The irritation from nettling +larvæ is often severe and, especially in regions where the brown-tail +abounds, inquiries as to treatment arise. In general, it may be said +that cooling lotions afford relief, and that scratching, with the +possibilities of secondary infection, should be avoided, in so far as +possible. + +Among the remedies usually at hand, weak solutions of ammonia, or a +paste of ordinary baking soda are helpful. Castellani and Chalmers +recommend cleaning away the hairs by bathing the region with an alkaline +lotion, such as two per cent solution of bicarbonate of soda, and then +applying an ointment of ichthyol (10%). + +In the brown-tail district, there are many proprietary remedies of which +the best ones are essentially the following, as recommended by Kirkland +(1907): + + Carbolic acid ½ drachm. + Zinc oxide ½ oz. + Lime water 8 oz. + +Shake thoroughly and rub well into the affected parts. + +In some cases, and especially where there is danger of secondary +infection, the use of a weak solution of creoline (one teaspoonful to a +quart of water), is to be advised. + + +VESCICATING INSECTS AND THOSE POSSESSING OTHER POISONS IN THEIR BLOOD +PLASMA + +We have seen that certain forms, for example, the poisonous spiders, not +only secrete a toxine in their poison glands, but that such a substance +may be extracted from other parts of their body, or even their eggs. +There are many insects which likewise possess a poisonous blood plasma. +Such forms have been well designated by Taschenberg as _cryptotoxic_ +(κÏυπτοσ = hidden). We shall consider a few representative +forms. + +[Illustration: 42_a_. Blister beetle.] + +[Illustration: 42_b_. An American blister beetle. Meloe angusticollis. +Photograph by M. V. S.] + + +THE BLISTER BEETLES--Foremost among the cryptotoxic insects are the +_Meloidæ_ or "blister beetles," to which the well-known "Spanish fly" +(fig. 42_a_), formerly very generally used in medical practice, belongs. +The vescicating property is due to the presence in the blood plasma of a +peculiar, volatile, crystalline substance known as _cantharidin_, which +is especially abundant in the reproductive organs of the beetle. +According to Kobert, the amount of this varies in different species from +.4 or .5% to 2.57% of the dry weight of the beetle. + +While blister beetles have been especially used for external +application, they are also at times used internally as a stimulant and a +diuretic. The powder or extract was formerly much in vogue as an +aphrodisiac, and formed the essential constituent of various philters, +or "love powders". It is now known that its effects on the reproductive +organs appear primarily after the kidneys have been affected to such an +extent as to endanger life, and that many cases of fatal poison have +been due to its ignorant use. + +There are many cases on record of poisoning and death due to internal +use, and in some instances from merely external application. There are +not rarely cases of poisoning of cattle from feeding on herbage bearing +a large number of the beetles and authentic cases are known of human +beings who have been poisoned by eating the flesh of such cattle. Kobert +states that the beetles are not poisonous to birds but that the flesh of +birds which have fed on them is poisonous to man, and that if the flesh +of chickens or frogs which have fed on the cantharidin be fed to cats it +causes in them the same symptoms as does the cantharidin. + +Treatment of cases of cantharidin poison is a matter for a skilled +physician. Until he can be obtained, emetics should be administered and +these should be followed by white of egg in water. Oils should be +avoided, as they hasten the absorption of the poison. + + +OTHER CRYPTOTOXIC INSECTS--Though the blister beetles are the best known +of the insects with poisonous blood plasma, various others have been +reported and we shall refer to a few of the best authenticated. + +One of the most famous is the Chrysomelid beetle, _Diamphidia simplex_, +the body fluids of whose larvæ are used by certain South African bushmen +as an arrow poison. Its action is due to the presence of a toxalbumin +which exerts a hæmolytic action on the blood, and produces inflammation +of the subcutaneous connective tissue and mucous membranes. Death +results from general paralysis. Krause (1907) has surmised that the +active principle may be a bacterial toxin arising from decomposition of +the tissues of the larva, but he presents no support of this view and it +is opposed by all the available evidence. + +In China, a bug, _Heuchis sanguinea_, belonging to the family Cicadidæ, +is used like the Meloidæ, to produce blistering, and often causes +poisoning. It has been assumed that its vescicating properties are due +to cantharidin, but the presence of this substance has not been +demonstrated. + +Certain Aphididæ contain a strongly irritating substance which produces, +not merely on mucous membranes but on outer skin, a characteristic +inflammation. + +It has been frequently reported that the larvæ of the European cabbage +butterfly, _Pieris brassicæ_, accidentally eaten by cows, horses, ducks, +and other domestic animals, cause severe colic, attempts to vomit, +paralysis of the hind legs, salivation, and stomatitis. On _postmortem_ +there are to be found hæmorrhagic gastro-enteritis, splenitis, and +nephritis. Kobert has recently investigated the subject and has found a +poisonous substance in the blood of not only the larvæ but also the +pupæ. + + +FOOTNOTES: + +[A] This is diametrically opposed to the findings of Bordas (1905) in +the case of the European _Latrodectus 13-guttatus_, whose glands are +"much larger than those of other spiders." From a considerable +comparative study, we should also unhesitatingly make this statement +regarding the glands of our American species, _L. mactans_. + +[B] Dr. E. H. Coleman (Kellogg, 1915) has demonstrated its virulence by +a series of experiments comparable with those of Kobert. + +[C] According to Stiles, the species occurring in the Northwest which is +commonly identified as _D. venustus_ should be called _D. andersoni_ +(see footnote, chapter 12). + +[D] It should be remembered that in all the higher Hymenoptera the first +abdominal segment is fused with the thorax and that what is apparently +the sixth segment is, in reality, the seventh. + + + + +CHAPTER III + +PARASITIC ARTHROPODA AFFECTING MAN + + +The relation of insects to man as simple parasites has long been +studied, and until very recent years the bulk of the literature of +medical entomology referred to this phase of the subject. This is now +completely overshadowed by the fact that so many of these parasitic +forms are more than simple parasites, they are transmitters of other +microscopic parasites which are pathogenic to man. Yet the importance of +insects as parasites still remains and must be considered in a +discussion of the relation of insects to the health of man. In taking up +the subject we shall first consider some general features of the +phenomenon of animal parasitism. + +Parasitism is an adaptation which has originated very often among living +organisms and in widely separated groups. It would seem simple to define +what is meant by a "parasite" but, in reality, the term is not easily +limited. It is often stated that a parasite is "An organism which lives +at the expense of another," but this definition is applicable to a +predatory species or, in its broadest sense, to all organisms. For our +purpose we may say with Braun: "A parasite is an organism which, for the +purpose of obtaining food, takes up its abode, temporarily or +permanently, on or within another living organism". + +Thus, parasitism is a phase of the broad biological phenomenon of +_symbiosis_, or living together of organisms. It is distinguished from +_mutualism_, or symbiosis in the narrow sense, by the fact that only one +party to the arrangement obtains any advantage, while the other is to a +greater or less extent injured. + +Of parasites we may distinguish on the basis of their location on or in +the host, _ecto-parasites_, which live outside of the body; and +_endo-parasites_, which live within the body. On account of their method +of breathing the parasitic arthropods belong almost exclusively to the +first of these groups. + +On the basis of relation to their host, we find _temporary parasites_, +those which seek the host only occasionally, to obtain food; and the +_stationary_ or _permanent_ parasites which, at least during certain +stages, do not leave their host. + +_Facultative parasites_ are forms which are not normally parasitic, but +which, when accidentally ingested, or otherwise brought into the body, +are able to exist for a greater or less period of time in their unusual +environment. These are generally called in the medical literature +"pseudoparasites" but the term is an unfortunate one. + +We shall now take up the different groups of arthropods, discussing the +more important of the parasitic forms attacking man. The systematic +relationship of these forms, and key for determining important species +will be found in Chapter XII. + + +ACARINA OR MITES + +The ACARINA, or _mites_, form a fairly natural group of arachnids, +characterized, in general, by a sac-like, unsegmented body which is +generally fused with the cephalothorax. The mouth-parts have been united +to form a beak or rostrum. + +The representatives of this group undergo a marked metamorphosis. +Commonly, the larvæ on hatching from the egg, possess but three pairs of +legs, and hence are called _hexapod larvæ_. After a molt, they transform +into nymphs which, like the adult, have four pairs of legs and are +called _octopod nymphs_. These after a period of growth, molt one or +more times and, acquiring external sexual organs, become adult. + +Most of the mites are free-living, but there are many parasitic species +and as these have originated in widely separated families, the Acarina +form an especially favorable group for study of the origin of +parasitism. Such a study has been made by Ewing (1911), who has reached +the following conclusions: + +"We have strong evidence indicating that the parasitic habit has +originated independently at least eleven times in the phylogeny of the +Ararina. Among the zoophagous parasites, the parasitic habit has been +developed from three different types of free-living Acarina: (a) +predaceous forms, (b) scavengers, (c) forms living upon the juices of +plants." + +Ewing also showed that among the living forms of Acarina we can trace +out all the stages of advancing parasitism, semiparasitism, facultative +parasitism, even to the fixed and permanent type, and finally to +endoparasitism. + +Of the many parasitic forms, there are several species which are serious +parasites of man and we shall consider the more important of these. +Infestation by mites is technically known as _acariasis_. + +[Illustration: 43. Effect of the harvest mites on the skin of man. +Photograph by J. C. Bradley.] + + +The Trombidiidæ, or Harvest Mites + +In many parts of this country it is impossible for a visitor to go into +the fields and, particularly, into berry patches and among tall weeds +and grass in the summer or early fall without being affected by an +intolerable itching, which is followed, later, by a breaking out of +wheals, or papules, surrounded by a bright red or violaceous aureola, +(fig. 43). It is often regarded as a urticaria or eczema, produced by +change of climate, an error in diet, or some condition of general +health. + +Sooner or later, the victim finds that it is due to none of these, but +to the attacks of an almost microscopic red mite, usually called +"jigger" or "chigger" in this country. As the term "chigger" is applied +to one of the true fleas, _Dermatophilus penetrans_, of the tropics, +these forms are more correctly known as "harvest mites." Natives of an +infested region may be so immune or accustomed to its attacks as to be +unaware of its presence, though such immunity is by no means possessed +by all who have been long exposed to the annoyance. + +[Illustration: 44. Harvest mites. (Larvæ of Trombidium). After C. V. +Riley.] + +The harvest mites, or chiggers, attacking man are larval forms, +possessing three pairs of legs (fig. 44). Their systematic position was +at first unknown and they were classed under a special genus _Leptus_, a +name which is very commonly still retained in the medical literature. It +is now known that they are the larval forms of various species of the +genus _Trombidium_, a group of predaceous forms, the adults of which +feed primarily on insects and their eggs. In this country the species +best known are those to be found late in summer, as larvæ at the base of +the wings of houseflies or grasshoppers. + +There is much uncertainty as to the species of the larvæ attacking man +but it is clear that several are implicated. Bruyant has shown that in +France the larvæ _Trombidium inapinatum_ and _Trombidium holosericeum_ +are those most frequently found. The habit of attacking man is abnormal +and the larvæ die after entering the skin. Normally they are parasitic +on various insects. + +Most recent writers agree that, on man, they do not bore into the skin, +as is generally supposed, but enter a hair follicle or sebaceous gland +and from the bottom of this, pierce the cutis with their elongate +hypopharynx. According to Braun, there arises about the inserted +hypopharynx a fibrous secretion--the so-called "beak" which is, in +reality, a product of the host. Dr. J. C. Bradley, however, has made +careful observations on their method of attack, and he assures us that +the mite ordinarily remains for a long time feeding on the surface of +the skin, where it produces the erythema above described. During this +time it is not buried in the skin but is able to retreat rapidly into it +through a hair follicle or sweat gland. The irritation from the mites +ceases after a few days, but not infrequently the intolerable itching +leads to so much scratching that secondary infection follows. + +Relief from the irritation may be afforded by taking a warm salt bath as +soon as possible after exposure or by killing the mites by application +of benzine, sulphur ointment or carbolized vaseline. When they are few +in number, they can be picked out with a sterile needle. + +Much may be done in the way of warding off their attacks by wearing +gaiters or close-woven stockings extending from ankle to the knee. Still +more efficacious is the sprinkling of flowers of sulphur in the +stockings and the underclothes from a little above the knee, down. The +writers have known this to make it possible for persons who were +especially susceptible to work with perfect comfort in badly infested +regions. Powdered naphthalene is successfully used in the same way and +as Chittenden (1906) points out, is a safeguard against various forms of +man-infesting tropical insect pests. + +The question of the destruction of the mites in the field is sometimes +an important one, and under some conditions, is feasible. Chittenden +states that much can be accomplished by keeping the grass, weeds, and +useless herbage mowed closely, so as to expose the mites to the sun. He +believes that in some cases good may be done by dusting the grass and +other plants, after cutting, with flowers of sulphur or by spraying with +dilute kerosene emulsion in which sulphur has been mixed. More recently +(1914) he calls attention to the value of cattle, and more especially +sheep, in destroying the pests by tramping on them and by keeping the +grass and herbage closely cropped. + + +IXODOIDEA OR TICKS + +Until recently, the ticks attracted comparatively little attention from +entomologists. Since their importance as carriers of disease has been +established, interest in the group has been enormously stimulated and +now they rank second only to the mosquitoes in the amount of detailed +study that has been devoted to them. + +[Illustration: 45_a_. Argus persicus. Capitulum of male. After Nuttall +and Warburton.] + +The ticks are the largest of the Acarina. They are characterized by the +fact that the hypostome, or "tongue" (fig. 45) is large and file-like, +roughened by sharp teeth. They possess a breathing pore on each side of +the body, above the third or fourth coxæ (fig. 45_b_). + +[Illustration: 45_b_. Left spiracle of nymph of _Argas persicus_. After +Nuttall and Warburton.] + +There are two distinct families of the IXODOIDEA, differing greatly in +structure, life-history and habits. These are the ARGASIDÆ and the +IXODIDÆ. We shall follow Nuttall (1908) in characterizing these two +families and in pointing out their biological differences, and shall +discuss briefly the more important species which attack man. The +consideration of the ticks as carriers of disease will be reserved for a +later chapter. + + +Argasidæ + +In the ticks belonging to the family ARGASIDÆ, there is comparatively +little sexual dimorphism, while this is very marked in the Ixodidæ. The +capitulum, or so-called "head" is ventral, instead of terminal; the +palpi are leg-like, with the segments subequal; the scutum, or dorsal +shield, is absent; eyes, when present, are lateral, on supracoxal folds. +The spiracles are very small; coxæ unarmed; tarsi without ventral +spurs, and the pulvilli are absent or rudimentary. + +In habits and life history the Argasidæ present striking +characteristics. In the first place, they are long-lived, a factor which +counts for much in the maintenance of the species. They are intermittent +feeders, being comparable with the bed-bug in this respect. There are +two or more nymphal stages, and they may molt after attaining maturity. +The female lays comparatively few eggs in several small batches. + +Nuttall (1911) concludes that "The Argasidæ represent the relatively +primitive type of ticks because they are less constantly parasitic than +are the Ixodidæ. Their nymphs and adults are rapid feeders and chiefly +infest the habitat of their hosts. * * * Owing to the Argasidæ infesting +the habitats of their hosts, their resistance to prolonged starvation +and their rapid feeding habits, they do not need to bring forth a large +progeny, because there is less loss of life in the various stages, as +compared with the Ixodidæ, prior to their attaining maturity." + +[Illustration: 46. Argus persicus. Dorsal and ventral aspects. (×4). +After Hassell.] + +Of the Argasidæ, we have in the United States, several species which +have been reported as attacking man. + +_Argas persicus_, the famous "Miana bug" (fig. 46), is a very widely +distributed species, being reported from Europe, Asia, Africa, and +Australia. It is everywhere preeminently a parasite of fowls. According +to Nuttall it is specifically identical with _Argas americanus_ Packard +or _Argas miniatus_ Koch, which is commonly found on fowls in the United +States, in the South and Southwest. Its habits are comparable to those +of the bed-bug. It feeds intermittently, primarily at night, and instead +of remaining on its host, it then retreats to cracks and crevices. +Hunter and Hooker (1908) record that they have found the larva to remain +attached for five or eight days before dropping. Unlike the Ixodidæ, the +adults oviposit frequently. + +[Illustration: 47. Otiobius (Ornithodoros) megnini, head of nymph. After +Stiles.] + +[Illustration: 48. Otiobius (Ornithodoros) megnini, male. (_a_) dorsal, +(_b_) ventral aspect. After Nuttall and Warburton.] + +The most remarkable feature of the biology of this species is the great +longevity, especially of the adult. Hunter and Hooker report keeping +larvæ confined in summer in pill boxes immediately after hatching for +about two months while under similar conditions those of the Ixodid, +_Boophilus annulatus_ lived for but two or three days. Many writers have +recorded keeping adults for long periods without food. We have kept +specimens in a tin box for over a year and a half and at the end of that +time a number were still alive. Laboulliene kept unfed adults for over +three years. In view of the effectiveness of sulphur in warding off the +attacks of Trombidiidæ, it is astonishing to find that Lounsbury has +kept adults of _Argas persicus_ for three months in a box nearly filled +with flowers of sulphur, with no apparent effect on them. + +We have already called attention to the occasional serious effects of +the bites of this species. While such reports have been frequently +discredited there can be no doubt that they have foundation in fact. The +readiness with which this tick attacks man, and the extent to which old +huts may be infested makes it especially troublesome. + +_Otiobius (Ornithodoros) megnini_, the "spinose ear-tick" (figs. 47, +48), first described from Mexico, as occurring in the ears of horses, is +a common species in our Southwestern States and is recorded by Banks as +occurring as far north as Iowa. + +The species is remarkable for the great difference between the spiny +nymph stage and the adult. The life history has been worked out by +Hooker (1908). Seed ticks, having gained entrance to the ear, attach +deeply down in the folds, engorge, and in about five days, molt; as +nymphs with their spinose body they appear entirely unlike the larvæ. As +nymphs they continue feeding sometimes for months. Finally the nymph +leaves the host, molts to form the unspined adult, and without further +feeding is fertilized and commences oviposition. + +The common name is due to the fact that in the young stage the ticks +occur in the ear of their hosts, usually horses or cattle. Not +uncommonly it has been reported as occurring in the ear of man and +causing very severe pain. Stiles recommends that it be removed by +pouring some bland oil into the ear. + +Banks (1908) reports three species of _Ornithodoros_--_O. turicata_, +_coriaceus_ and _talaje_--as occurring in the United States. All of +these attack man and are capable of inflicting very painful bites. + + +Ixodidæ + +The ticks belonging to the family IXODIDÆ (figs. 49 and 50) exhibit a +marked sexual dimorphism. The capitulum is anterior, terminal, instead +of ventral as in the Argasidæ; the palpi are relatively rigid (except in +the subfamily Ixodinæ), with rudimentary fourth segment; scutum present; +eyes, when present, dorsal, on side of scutum. The spiracles are +generally large, situated well behind the fourth coxæ; coxæ generally +with spurs; pulvilli always present. + +In habits and life history the typical Ixodidæ differ greatly from the +Argasidæ. They are relatively short-lived, though some recent work +indicates that their longevity has been considerably under-estimated. +Typically, they are permanent feeders, remaining on the host, or hosts, +during the greater part of their life. They molt twice only, on leaving +the larval and the nymphal stages. The adult female deposits a single, +large batch of eggs. Contrasting the habits of the Ixodidæ to those of +the Argasidæ, Nuttall (1911) emphasizes that the Ixodidæ are more highly +specialized parasites. "The majority are parasitic on hosts having no +fixed habitat and consequently all stages, as a rule, occur upon the +host." + +[Illustration: 49. Ixodes ricinus; male, ventral aspect. After Braun and +Luehe.] + +As mere parasites of man, apart from their power to transmit disease, +the Ixodidæ are much less important than the Argasidæ. Many are reported +as occasionally attacking man and of these the following native species +may be mentioned. + +[Illustration: 50. Ixodes ricinus, var. scapularis, female. Capitulum +and scutum; ventral aspect of capitulum; coxæ; tarsus 4; spiracle; +genital and anal grooves. After Nuttall and Warburton.] + +_Ixodes ricinus_, the European castor bean tick (figs. 49, 50), is a +species which has been often reported from this country but Banks (1908) +has shown that, though it does occur, practically all of the records +apply to _Ixodes scapularis_ or _Ixodes cookei_. In Europe, _Ixodes +ricinus_ is very abundant and very commonly attacks man. At the point +of penetration of the hypostome there is more or less inflammation but +serious injury does not occur unless there have been introduced +pathogenic bacteria or, unless the tick has been abruptly removed, +leaving the capitulum in the wound. Under the latter circumstances, +there may be an abscess formed about the foreign body and occasionally, +serious results have followed. Under certain conditions the tick, in +various stages, may penetrate under the skin and produce a tumor, within +which it may survive for a considerable period of time. + +_Ixodes cookei_ is given by Banks as "common on mammals in the Eastern +States as far west as the Rockies." It is said to affect man severely. + +_Amblyomma americanum_, (fig. 158c), the "lone star tick," is widely +distributed in the United States. Its common name is derived from the +single silvery spot on the scutum of the female. Hunter and Hooker +regard this species as, next to _Boophilus annulatus_, the most +important tick in the United States. Though more common on cattle, it +appears to attack mammals generally, and "in portions of Louisiana and +Texas it becomes a pest of considerable importance to moss gatherers and +other persons who spend much time in the forests." + +_Amblyomma cajennense_, noted as a pest of man in central and tropical +America, is reported from various places in the south and southwestern +United States. + +_Dermacentor variabilis_ is a common dog tick of the eastern United +States. It frequently attacks man, but the direct effects of its bite +are negligible. + +The "Rocky Mountain spotted fever tick" (_Dermacentor andersoni_ +according to Stiles, _D. venustus_ according to Banks) is, from the +viewpoint of its effects on man, the most important of the ticks of the +United States. This is because, as has been clearly established, it +transmits the so-called "spotted fever" of man in our northwestern +states. This phase of the subject will be discussed later and it need +merely be mentioned here, that this species has been reported as causing +painful injuries by its bites. Dr. Stiles states that he has seen cases +of rather severe lymphangitis and various sores and swellings developing +from this cause. In one case, of an individual bitten near the elbow, +the arm became very much swollen and the patient was confined in bed for +several days. The so-called tick paralysis produced by this species is +discussed in a preceding chapter. + +There are many other records of various species of ticks attacking man, +but the above-mentioned will serve as typical and it is not necessary to +enter into greater detail. + + +TREATMENT OF TICK BITES--When a tick attaches to man the first thing to +be done is to remove it without leaving the hypostome in the wound to +fester and bring about secondary effects. This is best accomplished by +applying to the tick's body some substance which will cause it to more +readily loosen its hold. Gasoline or petroleum, oil or vaseline will +serve. For removing the spinose ear-tick, Stiles recommends pouring some +bland oil into the ear. Others have used effectively a pledget of cotton +soaked in chloroform. + +In general, the treatment recommended by Wellman for the bites of +_Ornithodoros moubata_ will prove helpful. It consists of prolonged +bathing in very hot water, followed by the application of a strong +solution of bicarbonate of soda, which is allowed to dry upon the skin. +He states that this treatment is comforting. For severe itching he +advises smearing the bites with vaseline, which is slightly impregnated +with camphor or menthol. Medical aid should be sought when complications +arise. + +The DERMANYSSIDÆ are Gamasid mites which differ from others of the group +in that they are parasitic on vertebrates. None of the species normally +attack man, but certain of them, especially the poultry mite, may be +accidental annoyances. + +[Illustration: 51. Dermanyssus gallinæ, female. After Delafond.] + +_Dermanyssus gallinæ_ (fig. 51), the red mite of poultry, is an +exceedingly common and widespread parasite of fowls. During the day it +lives in cracks and crevices of poultry houses, under supports of +roosts, and in litter of the food and nests, coming out at night to +feed. They often attack people working in poultry houses or handling +and plucking infested fowls. They may cause an intense pruritis, but +they do not produce a true dermatosis, for they do not find conditions +favorable for multiplication on the skin of man. + + +Tarsonemidæ + +The representatives of the family TARSONEMIDÆ are minute mites, with the +body divided into cephalothorax and abdomen. There is marked sexual +dimorphism. The females possess stigmata at the anterior part of the +body, at the base of the rostrum, and differ from all other mites in +having on each side, a prominent clavate organ between the first and +second legs. The larva, when it exists, is hexapodous and resembles the +adult. A number of the species are true parasites on insects, while +others attack plants. Several of them may be accidental parasites of +man. + +[Illustration: 52. Pediculoides ventricosus, female. After Webster.] + +[Illustration: 53. Pediculoides ventricosus, gravid female. (×80). +After Webster.] + +_Pediculoides ventricosus_ (fig. 52 and 53) is, of all the Tarsonemidæ +reported, the one which has proved most troublesome to man. It is a +predaceous species which attacks a large number of insects but which has +most commonly been met with by man through its fondness for certain +grain-infesting insects, notably the Angoumois grain moth, _Sitotroga +cerealella_, and the wheat straw-worm, _Isosoma grande_. In recent years +it has attracted much attention in the United States and its +distribution and habits have been the object of detailed study by +Webster (1901). + +[Illustration: 54. Pediculoides ventricosus, male. After Braun.] + +There is a very striking sexual dimorphism in this species. The +non-gravid female is elongate, about 200µ by 70µ (fig. 52), with the +abdomen slightly striated longitudinally. The gravid female (fig. 53) +has the abdomen enormously swollen, so that it is from twenty to a +hundred times greater than the rest of the body. The species is +viviparous and the larvæ undergo their entire growth in the body of the +mother. They emerge as sexually mature males and females which soon +pair. The male (fig. 54) is much smaller, reaching a length of only 320µ +but is relatively broad, 80µ, and angular. Its abdomen is very greatly +reduced. + +As far back as 1850 it was noted as causing serious outbreaks of +peculiar dermatitis among men handling infested grain. For some time the +true source of the difficulty was unknown and it was even believed that +the grain had been poisoned. Webster has shown that in this country (and +probably in Europe as well) its attacks have been mistaken for those of +the red bugs or "chiggers" (larval Trombiidæ). More recently a number of +outbreaks of a mysterious "skin disease" were traced to the use of straw +mattresses, which were found to be swarming with these almost +microscopic forms which had turned their attentions to the occupants of +the beds. Other cases cited were those of farmers running wheat through +a fanning mill, and of thrashers engaged in feeding unthrashed grain +into the cylinder of the machine. + +[Illustration: 55. Lesions produced by the attacks of Pediculoides +ventricosus. After Webster.] + +The medical aspects of the question have been studied especially by +Schamberg and Goldberger and from the latter's summary (1910) we derive +the following data. Within twelve to sixteen hours after exposure, +itching appears and in severe cases, especially where exposure is +continued night after night by sleeping on an infested bed, the itching +may become almost intolerable. Simultaneously, there appears an eruption +which characteristically consists of wheals surrounded by a vesicle +(fig. 55). The vesicle as a rule does not exceed a pin head in size but +may become as large as a pea. Its contents rapidly become turbid and in +a few hours it is converted into a pustule. The eruption is most +abundant on the trunk, slight on the face and extremities and almost +absent on the feet and hands. In severe cases there may be +constitutional disturbances marked, at the outset, by chilliness, +nausea, and vomiting, followed for a few days by a slight elevation of +temperature, with the appearance of albumin in the urine. In some cases +the eruption may simulate that of chicken-pox or small-pox. + +Treatment for the purpose of killing the mites is hardly necessary as +they attach feebly to the surface and are readily brushed off by +friction of the clothes. "Antipruritic treatment is always called for; +warm, mildly alkaline baths or some soothing ointment, such as zinc +oxide will be found to fulfil this indication." Of course, reinfestation +must be guarded against, by discarding, or thoroughly fumigating +infested mattresses, or by avoiding other sources. Goldberger suggests +that farm laborers who must work with infested wheat or straw might +protect themselves by anointing the body freely with some bland oil or +grease, followed by a change of clothes and bath as soon as their work +is done. We are not aware of any experiments to determine the effect of +flowers of sulphur, but their efficiency in the case of "red bugs" +suggests that they are worth a trial against _Pediculoides_. + +Various species of TYROGLYPHIDÆ (fig. 150_f_) may abound on dried fruits +and other products and attacking persons handling them, may cause a +severe dermatitis, comparable to that described above for _Pediculoides +ventricosus_. Many instances of their occurrence as such temporary +ectoparasites are on record. Thus, workers who handle vanilla pods are +subject to a severe dermatitis, known as vanillism, which is due to the +attacks of _Tyroglyphus siro_, or a closely related species. The +so-called "grocer's itch" is similarly caused by mites infesting various +products. Castellani has shown that in Ceylon, workers employed in the +copra mills, where dried cocoanut is ground up for export, are much +annoyed by mites, which produce the so-called "copra itch." The skin of +the hands, arms and legs, and sometimes of the whole body, except the +face, is covered by fairly numerous, very pruriginous papules, often +covered by small, bloody crusts due to scratching. The condition is +readily mistaken for scabies. It is due to the attacks of _Tyroglyphus +longior castellanii_ which occur in enormous numbers in some samples of +the copra. + + +Sarcoptidæ + +The SARCOPTIDÆ are minute whitish mites, semi-globular in shape, with a +delicate transversely striated cuticula. They lack eyes and tracheæ. The +mouth-parts are fused at the base to form a cone which is usually +designated as the head. The legs are short and stout, and composed of +five segments. The tarsi may or may not possess a claw and may terminate +in a pedunculated sucker, or simple long bristle, or both. The presence +or absence of these structures and their distribution are much used in +classification. The mites live on or under the skin of mammals and +birds, where they produce the disease known as scabies, mange, or itch. +Several species of the Sarcoptidæ attack man but the most important of +these, and the one pre-eminent as the "itch mite" is _Sarcoptes +scabiei_. + +The female of _Sarcoptes scabiei_, of man, is oval and yellowish white; +the male more rounded and of a somewhat reddish tinge, and much smaller. +The body is marked by transverse striæ which are partly interrupted on +the back. There are transverse rows of scales, or pointed spines, and +scattered bristles on the dorsum. + +[Illustration: 56_a_. Sarcoptes scabiei, male. (×100). After +Fürstenberg.] + +The male (fig. 56) which is from 200-240µ in length, and 150-200µ in +breadth, possesses pedunculated suckers on each pair of legs except the +third, which bears, instead, a long bristle. The female (fig. 56) +300-450µ in length and 250-350µ in breadth, has the pedunculated suckers +on the first and second pairs of legs, only, the third and fourth +terminating in bristles. + +[Illustration: 56_b_. Sarcoptes scabiei, female. (×100.) After +Fürstenberg.] + +The mite lives in irregular galleries from a few millimeters to several +centimeters in length, which it excavates in the epidermis (fig. 57). It +works especially where the skin is thin, such as between the fingers, in +the bend of the elbows and knees, and in the groin, but it is by no +means restricted to these localities. The female, alone, tunnels into +the skin; the males remain under the superficial epidermal scales, and +seldom are found, as they die soon after mating. + +As she burrows into the skin the female deposits her eggs, which measure +about 150 × 100µ. Fürstenberg says that each deposits an average of +twenty-two to twenty-four eggs, though Gudden reports a single burrow as +containing fifty-one. From these there develop after about seven days, +the hexapod larvæ. These molt on the sixteenth day to form an octopod +nymph, which molts again the twenty-first day. At the end of the fourth +week the nymphs molt to form the sexually mature males and the so-called +pubescent females. These pair, the males die, and the females again cast +their skin, and become the oviparous females. Thus the life cycle is +completed in about twenty-eight days. + +The external temperature exercises a great influence on the development +of the mites and thus, during the winter, the areas of infestation not +only do not spread, but they become restricted. As soon as the +temperature rises, the mites increase and the infestation becomes much +more extensive. + +[Illustration: 57. Sarcoptes scabiei. Diagrammatic representation of the +course in the skin of man.] + +In considering the possible sources of infestation, and the chances of +reinfestation after treatment, the question of the ability of the mite +to live apart from its host is a very important one. Unfortunately there +are few reliable data on this subject. Gerlach found that, exposed in +the dry, warm air of a room they became very inactive within twenty-four +hours, that after two days they showed only slight movement, and that +after three or four days they could not be revived by moisture and +warming. The important fact was brought out that in moist air, in folded +soiled underwear, they survived as long as ten days. Bourguignon found +that under the most favorable conditions the mites of _Sarcoptes scabiei +equi_ would live for sixteen days. + +The disease designated the "itch" or "scabies," in man has been known +from time immemorial, but until within less than a hundred years it was +almost universally attributed to malnutrition, errors of diet, or "bad +blood." This was in spite of the fact that the mite was known to Mouffet +and that Bonomo had figured both the adult and the egg and had declared +the mite the sole cause of the disease. In 1834 the Corsican medical +student, Francis Renucci, demonstrated the mite before a clinic in Saint +Louis Hospital in Paris and soon thereafter there followed detailed +studies of the life history of the various itch mites of man and +animals. + +[Illustration: 58. Scabies on the hand. From portfolio of Dermochromes +by permission of Rebman & Co., of New York. Publishers.] + +The disease is a cosmopolitan one, being exceedingly abundant in some +localities. Its spread is much favored where large numbers of people are +crowded together under insanitary conditions and hence it increases +greatly during wars and is widely disseminated and abundant immediately +afterwards. Though more commonly to be met with among the lower classes, +it not infrequently appears among those of the most cleanly, careful +habits, and it is such cases that are most liable to wrong diagnosis by +the physician. + +Infection occurs solely through the passage, direct or indirect, of the +young fertilized females to the skin of a healthy individual. The adult, +oviparous females do not quit their galleries and hence do not serve to +spread the disease. The young females move about more or less at night +and thus the principal source of infestation is through sleeping in the +same bed with an infested person, or indirectly through bedclothes, or +even towels or clothing. Diurnal infestation through contact or clothing +is exceptional. Many cases are known of the disease being contracted +from animals suffering from scabies, or mange. + +When a person is exposed to infestation, the trouble manifests itself +after eight or ten days, though there usually elapses a period of twenty +to thirty days before there is a suspicion of anything serious. The +first symptom is an intense itching which increases when the patient is +in bed. When the point of irritation is examined the galleries may +usually be seen as characteristic sinuous lines, at first whitish in +color but soon becoming blackish because of the contained eggs and +excrement. The galleries, which may not be very distinct in some cases, +may measure as much as four centimeters in length. Little vesicles, of +the size of a pin head are produced by the secretions of the feeding +mite; they are firm, and projecting, and contain a limpid fluid. Figures +58 and 59 show the typical appearance of scabies on the hands, while +figure 60 shows a severe general infestation. The intolerable itching +induces scratching and through this various complications may arise. The +lesions are not normally found on the face and scalp, and are rare on +the back. + +[Illustration: 59. Scabies on the hand. After Duhring.] + +[Illustration: 60. Generalized infection of Scabies. After Morrow.] + +Formerly, scabies was considered a very serious disease, for its cause +and method of treatment were unknown, and potentially it may continue +indefinitely. Generation after generation of the mites may develop and +finally their number become so great that the general health of the +individual is seriously affected. Now that the true cause of the disease +is known, it is easily controlled. + +Treatment usually consists in softening the skin by friction with soap +and warm water, followed by a warm bath, and then applying some +substance to kill the mites. Stiles gives the following directions, +modified from Bourguignon's, as "a rather radical guide, to be modified +according to facilities and according to the delicacy of the skin or +condition of the patient": + +1. The patient, stripped naked, is energetically rubbed all over (except +the head) for twenty minutes, with green soap and warm water. 2. He is +then placed in a warm bath for thirty minutes, during which time the +rubbing is continued. 3. The parasiticide is next rubbed in for twenty +minutes and is allowed to remain on the body for four or five hours; in +the meantime the patient's clothes are sterilized, to kill the eggs or +mites attached to them. 4. A final bath is taken to remove the +parasiticide. + +The parasiticide usually relied on is the officinal sulphur ointment of +the United States pharmacopÅ“ia. When infestation is severe it is +necessary to repeat treatment after three or four days in order to kill +mites which have hatched from the eggs. + +The above treatment is too severe for some individuals and may, of +itself, produce a troublesome dermatitis. We have seen cases where the +treatment was persisted in and aggravated the condition because it was +supposed to be due to the parasite. For delicate-skinned patients the +use of balsam of Peru is very satisfactory, and usually causes no +irritation whatever. Of course, sources of reinfection should be +carefully guarded against. + +_Sarcoptes scabiei crustosæ_, which is a distinct variety, if not +species, of the human itch mite, is the cause of so-called Norwegian +itch. This disease is very contagious, and is much more resistant than +the ordinary scabies. Unlike the latter, it may occur on the face and +scalp. + +_Sarcoptes scabiei_ not only attacks man but also occurs on a large +number of mammals. Many species, based on choice of host, and minute +differences in size and secondary characters, have been established, but +most students of the subject relegate these to varietal rank. Many of +them readily attack man, but they have become sufficiently adapted to +their normal host so that they are usually less persistent on man. + +_Notoedres cati_ (usually known as _Sarcoptes minor_) is a species of +itch mites which produce an often fatal disease of cats. The body is +rounded and it is considerably smaller than _Sarcoptes scabiei_, the +female (fig. 61) measuring 215-230µ long and 165-175µ wide; the males +145-150µ by 120-125µ. The most important character separating +_Notoedres_ from _Sarcoptes_ is the position of the anus, which is +dorsal instead of terminal. The mite readily transfers to man but does +not persist, the infestation usually disappearing spontaneously in about +two weeks. Infested cats are very difficult to cure, unless treatment is +begun at the very inception of the outbreak, and under ordinary +circumstances it is better to kill them promptly, to avoid spread of the +disease to children and others who may be exposed. + +[Illustration: 61. Notoedres cati, male and female. After Railliet.] + + +Demodecidæ + +The DEMODECIDÆ are small, elongate, vermiform mites which live in the +hair follicles of mammals. The family characteristics will be brought +out in the discussion of the species infesting man, _Demodex +folliculorum_. + +[Illustration: 62. Demodex folliculorum. (×200) After Blanchard.] + +_Demodex folliculorum_ (fig. 62) is to be found very commonly in the +hair follicles and sebaceous glands of man. It is vermiform in +appearance, and with the elongate abdomen transversely striated so as to +give it the appearance of segmentation. The female is 380-400µ long by +45µ; the male 300µ by 40µ. The three-jointed legs, eight in number, are +reduced to mere stubs in the adult. The larval form is hexopod. These +mites thus show in their form a striking adaptation to their +environment. In the sebaceous glands and hair follicles they lie with +their heads down (fig. 63). Usually there are only a few in a gland, but +Gruby has counted as many as two hundred. + +[Illustration: 63. Demodex folliculorum. Section through skin showing +the mites in situ. Magnification of Nos. 1, 2, 6 and 7, ×150; Nos, 3, +4, 5, ×450. After Megnin.] + +The frequency with which they occur in man is surprising. According to +European statistics they are found in 50 per cent to 60 per cent or even +more. Gruby found them in forty out of sixty persons examined. These +figures are very commonly quoted, but reliable data for the United +States seem to be lacking. Our studies indicate that it is very much +less common in this country than is generally assumed. + +The Demodex in man does not, as a rule, cause the slightest +inconvenience to its host. It is often stated that they give rise to +comedons or "black-heads" but there is no clear evidence that they are +ever implicated. Certain it is that they are not the usual cause. A +variety of the same, or a very closely related species of _Demodex_, on +the dog gives rise to the very resistant and often fatal follicular +mange. + + +HEXAPODA OR TRUE INSECTS + +The HEXAPODA, or true insects, are characterized by the fact that the +adult possesses three pairs of legs. The body is distinctly segmented +and is divided into head, thorax, and abdomen. + +The mouth-parts in a generalized form, consist of an upper lip, or +_labrum_, which is a part of the head capsule, and a central unpaired +_hypopharynx_, two _mandibles_, two _maxillæ_ and a lower lip, or +_labium_, made up of the fused pair of second maxillæ. These parts may +be greatly modified, dependent upon whether they are used for biting, +sucking, piercing and sucking, or a combination of biting and sucking. + +Roughly speaking, insects may be grouped into those which undergo +_complete metamorphosis_ and those which have _incomplete +metamorphosis_. They are said to undergo complete metamorphosis when the +young form, as it leaves the egg, bears no resemblance to the adult. For +example, the maggot changes to a quiescent pupa and from this emerges +the winged active fly. They undergo incomplete metamorphosis, when the +young insect, as it leaves the egg, resembles the adult to a greater or +less extent, and after undergoing a certain number of molts becomes +sexually mature. + +Representatives of several orders have been reported as accidental or +faculative parasites of man, but the true parasites are restricted to +four orders. These are the Siphunculata; the Hemiptera, the Diptera and +the Siphonaptera. + + +SIPHUNCULATA + +The order SIPHUNCULATA was established by Meinert to include the true +sucking lice. These are small wingless insects, with reduced +mouth-parts, adapted for sucking; thorax apparently a single piece due +to indistinct separation of its three segments: the compound eyes +reduced to a single ommatidium on each side. The short, powerful legs +are terminated by a single long claw. Metamorphosis incomplete. + +There has been a great deal of discussion regarding the structure of the +mouth-parts, and the relationships of the sucking lice, and the +questions cannot yet be regarded as settled. The conflicting views are +well represented by Cholodkovsky (1904 and 1905) and by Enderlein +(1904). + +[Illustration: 64. Pediculus showing the blind sac (_b_) containing the +mouth parts (_a_) beneath the alimentary canal (_p_). After Pawlowsky.] + +Following Graber, it is generally stated that the mouth-parts consist of +a short tube furnished with hooks in front, which constitutes the lower +lip, and that within this is a delicate sucking tube derived from the +fusion of the labrum and the mandibles. Opposed to this, Cholodkovsky +and, more recently, Pawlowsky, (1906), have shown that the piercing +apparatus lies in a blind sac under the pharynx and opening into the +mouth cavity (fig. 64). It does not form a true tube but a furrow with +its open surface uppermost. Eysell has shown that, in addition, there is +a pair of chitinous rods which he regards as the homologues of the +maxillæ. + +When the louse feeds, it everts the anterior part of the mouth cavity, +with its circle of hooks. The latter serve for anchoring the bug, and +the piercing apparatus is then pushed out. + +[Illustration: 65. Pediculus humanus, ventral aspect of male. (×10)] + +Most writers have classed the sucking lice as a sub-order of the +Hemiptera, but the more recent anatomical and developmental studies +render this grouping untenable. An important fact, bearing on the +question, is that, as shown by Gross, (1905), the structure of the +ovaries is radically different from that of the Hemiptera. + +Lice infestation and its effects are known medically as _pediculosis_. +Though their continued presence is the result of the grossest neglect +and filthiness, the original infestation may be innocently obtained and +by people of the most careful habits. + +Three species commonly attack man. Strangely enough, there are very few +accurate data regarding their life history. + +_Pediculus humanus_ (fig. 65), the head louse, is the most widely +distributed. It is usually referred to in medical literature as +_Pediculus capitis_, but the Linnean specific name has priority. In +color it is of a pale gray, blackish on the margins. It is claimed by +some authors that the color varies according to the color of the skin of +the host. The abdomen is composed of seven distinct segments, bearing +spiracles laterally. There is considerable variation in size. The males +average 1.8 mm. and the females 2.7 mm. in length. + +The eggs, fifty to sixty in number, stick firmly to the hairs of the +host and are known as nits. They are large and conspicuous, especially +on dark hair and are provided with an operculum, or cap, at the free +end, where the nymphs emerge. They hatch in about six days and about the +eighteenth day the young lice are sexually mature. + +[Illustration: 66. Pediculosis of the head. The illustration shows the +characteristic indications of the presence of lice, viz: the occipital +eczema gluing the hairs together, the swollen cervical glands, and the +porrigo, or eruption of contagious pustules upon the neck. After Fox.] + +The head lice live by preference on the scalp of their host but +occasionally they are found on the eyelashes and beard, or in the pubic +region. They may also occur elsewhere on the body. The penetration of +the rostrum into the skin and the discharge of an irritating saliva +produce a severe itching, accompanied by the formation of an eczema-like +eruption (fig. 66). When the infestation is severe, the discharge from +the pustules mats down the hair, and scabs are formed, under which the +insects swarm. "If allowed to run, a regular carapace may form, called +_trichoma_, and the head exudes a fÅ“tid odor. Various low plants may +grow in the trichoma, the whole being known as _plica +palonica_."--Stiles. + +Sources of infestation are various. School children may obtain the lice +from seatmates, by wearing the hats or caps of infested mates, or by the +use, in common, of brushes and combs. They may be obtained from infested +beds or sleeper berths. Stiles reports an instance in which a large +number of girls in a fashionable boarding school developed lousiness a +short time after traveling in a sleeping car. + +Treatment is simple, for the parasites may readily be controlled by +cleanliness and washing the head with a two per cent solution of +carbolic acid or even kerosene. The latter is better used mixed with +equal parts of olive oil, to avoid irritation. The treatment should be +applied at night and followed the next morning by a shampoo with soap +and warm water. It is necessary to repeat the operation in a few days. +Xylol, used pure, or with the addition of five per cent of vaseline, is +also very efficacious. Of course, the patient must be cautioned to stay +away from a lighted lamp or fire while using either the kerosene or +xylol. While these treatments will kill the eggs or nits, they will not +remove them from the hairs. Pusey recommends repeated washings with +vinegar or 25 per cent of acetic acid in water, for the purpose of +loosening and removing the nits. + +Treatment of severe infestations in females is often troublesome on +account of long hair. For such cases the following method recommended by +Whitfield (1912) is especially applicable: + +The patient is laid on her back on the bed with her head over the edge, +and beneath the head is placed a basin on a chair so that the hair lies +in the basin. A solution of 1 in 40 carbolic acid is then poured over +the hair into the basin and sluiced backwards and forwards until the +whole of the hair is thoroughly soaked with it. It is especially +necessary that care should be taken to secure thorough saturation of the +hair over the ears and at the nape of the neck, since these parts are +not only the sites of predilection of the parasites but they are apt to +escape the solution. This sluicing is carried out for ten minutes by the +clock. At the end of ten minutes the hair is lifted from the basin and +allowed to drain, but is not dried or even thoroughly wrung out. The +whole head is then swathed with a thick towel or better, a large piece +of common house flannel, which is fastened up to form a sort of turban, +and is allowed to remain thus for an hour. It can then be washed or +simply allowed to dry, as the carbolic quickly disperses. At the end of +this period every pediculus and what is better, every ovum is dead and +no relapse will occur unless there is exposure to fresh contagion. +Whitfield states that there seem to be no disadvantages in this method, +which he has used for years. He has never seen carboluria result from +it, but would advise first cutting the hair of children under five years +of age. + +_Pediculus corporis_ (= _P. vestimenti_) the body louse, is larger than +the preceding species, the female measuring 3.3 mm., and the male 3 mm. +in length. The color is a dirty white, or grayish. _P. corporis_ has +been regarded by some authorities as merely a variety of _P. humanus_ +but Piaget maintains there are good characters separating the two +species. + +The body louse lives in the folds and seams of the clothing of its host, +passing to the skin only when it wishes to feed. Brumpt states that he +has found enormous numbers of them in the collars of glass-ware or +grains worn by certain naked tribes in Africa. + +Exact data regarding the life-history of this species have been +supplied, in part, by the work of Warburton (1910), cited by Nuttall. He +found that _Pediculus corporis_ lives longer than _P. humanus_ under +adverse conditions. This is doubtless due to its living habitually on +the clothing, whereas _humanus_ lives upon the head, where it has more +frequent opportunities of feeding. He reared a single female upon his +own person, keeping the louse enclosed in a cotton-plugged tube with a +particle of cloth to which it could cling. The tube was kept next to his +body, thus simulating the natural conditions of warmth and moisture +under which the lice thrive. The specimen was fed twice daily, while it +clung to the cloth upon which it rested. Under these conditions she +lived for one month. Copulation commenced five days after the female had +hatched and was repeated a number of times, sexual union lasting for +hours. The female laid one hundred and twenty-four eggs within +twenty-five days. + +The eggs hatched after eight days, under favorable conditions, such as +those under which the female was kept. They did not hatch in the cold. +Eggs kept near the person during the day and hung in clothing by the +bedside at night, during the winter, in a cold room, did not hatch until +the thirty-fifth day. When the nymphs emerge from the eggs, they feed at +once, if given a chance to do so. They are prone to scatter about the +person and abandon the fragment of cloth to which the adult clings. + +The adult stage is reached on the eleventh day, after three molts, about +four days apart. Adults enter into copulation about the fifth day and as +the eggs require eight days for development, the total cycle, under +favorable conditions, is about twenty-four days. Warburton's data differ +considerably from those commonly quoted and serve to emphasize the +necessity for detailed studies of some of the commonest of parasitic +insects. + +Body lice are voracious feeders, producing by their bites and the +irritating saliva which they inject, rosy elevations and papules which +become covered with a brownish crust. The intense itching provokes +scratching, and characteristic white scars (fig. 67) surrounded by +brownish pigment (fig. 68) are formed. The skin may become thickened and +take on a bronze tinge. This melanoderma is especially marked in the +region between the shoulders but it may become generalized, a prominent +characteristic of "vagabond's disease." According to Dubre and Beille, +this melanoderma is due to a toxic substance secreted by the lice, which +indirectly provokes the formation of pigment. + +[Illustration: 67. Pediculosis in man caused by the body louse. After +Morrow.] + +Control measures, in the case of the body louse, consist in boiling or +steaming the clothes or in some cases, sterilizing by dry heat. The +dermatitis may be relieved by the use of zinc-oxide ointment, to which +Pusey recommends that there be added, on account of their parasiticidal +properties, sulphur and balsam of Peru, equal parts, 15 to 30 grains to +the ounce. + +_Phthirius pubis_ (= _P. inguinalis_), the pubic louse, or so-called +"crab louse," differs greatly from the preceding in appearance. It is +characterized by its relatively short head which fits into a broad +depression in the thorax. The latter is broad and flat and merges into +the abdomen. The first pair of legs is slender and terminated by a +straight claw. The second and third pairs of legs are thicker and are +provided with powerful claws fitted for clinging to hairs. The females +(fig. 69) measure 1.5 to 2 mm. in length by 1.5 mm. in breadth. The male +averages a little over half as large. The eggs, or nits, are fixed at +the base of the hairs. Only a few, ten to fifteen are deposited by a +single female, and they hatch in about a week's time. The young lice +mature in two weeks. + +The pubic louse usually infests the hairs of the pubis and the perineal +region. It may pass to the arm pits or even to the beard or moustache. +Rarely, it occurs on the eyelids, and it has even been found, in a very +few instances, occurring in all stages, on the scalp. Infestation may be +contracted from beds or even from badly infested persons in a crowd. We +have seen several cases which undoubtedly were due to the use of public +water closets. It produces papular eruption and an intense pruritis. +When abundant, there occurs a grayish discoloration of the skin which +Duguet has shown is due to a poisonous saliva injected by the louse, as +is the melanoderma caused by the body louse. + +[Illustration: 68. Melanoderma caused by the body louse. From Portfolio +of Dermochromes, by permission of Rebman & Co., New York, Publishers.] + +The pubic louse may be exterminated by the measures recommended for the +head louse, or by the use of officinal mercurial ointment. + +[Illustration: 69. Phthirius pubis. Ventral aspect of female. (×12).] + + +HEMIPTERA + +Several species of HEMIPTERA-HETEROPTERA are habitual parasites of man, +and others occur as occasional or accidental parasites. Of all these, +the most important and widespread are the bed-bugs, belonging to the +genus _Cimex_ (= _Acanthia_). + + +THE BED-BUGS--The bed-bugs are characterized by a much flattened oval +body, with the short, broad head unconstricted behind, and fitting into +the strongly excavated anterior margin of the thorax. The compound eyes +are prominent, simple eyes lacking. Antennæ four-jointed, the first +segment short, the second long and thick, and the third and fourth +slender. The tarsi are short and three segmented. + +It is often assumed in the literature of the subject that there is but a +single species of _Cimex_ attacking man, but several such species are to +be recognized. These are distinguishable by the characters given in +Chapter XII. We shall consider especially _Cimex lectularius_, the most +common and widespread species. + +_Cimex lectularius_ (= _Acanthia lectularia_, _Clinocoris +lectularius_), is one of the most cosmopolitan of human parasites but, +like the lice, it has been comparatively little studied until recent +years, when the possibility that it may be concerned with the +transmission of various diseases has awakened interest in the details of +its life-history and habits. + +[Illustration: 70. Cimex lectularius adult and eggs. Photograph by M. V. +S.] + +The adult insect (fig. 70) is 4-5 mm. long by 3 mm. broad, reddish brown +in color, with the beak and body appendages lighter in color. The short, +broad and somewhat rectangular head has no neck-like constriction but +fits into the broadly semilunar prothorax. The four segmented labium or +proboscis encloses the lancet-like maxillæ and mandibles. The distal of +the four antennal segments is slightly club-shaped. The prothorax is +characteristic of the species, being deeply incised anteriorly and with +its thin lateral margins somewhat turned up. The mesothorax is +triangular, with the apex posteriorly, and bears the greatly atrophied +first pair of wings. There is no trace of the metathoracic pair. The +greatly flattened abdomen has eight visible segments, though in reality +the first is greatly reduced and has been disregarded by most writers. +The body is densely covered with short bristles and hairs, the former +being peculiarly saber-shaped structures sharply toothed at the apex and +along the convex side (fig. 159_b_). + +The peculiar disagreeable odor of the adult bed-bug is due to the +secretion of the stink glands which lie on the inner surface of the +mesosternum and open by a pair of orifices in front of the metacoxæ, +near the middle line. In the nymphs, the thoracic glands are not +developed but in the abdomen there are to be found three unpaired dorsal +stink glands, which persist until the fifth molt, when they become +atrophied and replaced by the thoracic glands. The nymphal glands occupy +the median dorsal portion of the abdomen, opening by paired pores at the +anterior margin of the fourth, fifth and sixth segments. The secretion +is a clear, oily, volatile fluid, strongly acid in reaction. Similar +glands are to be found in most of the Hemiptera-Heteroptera and their +secretion is doubtless protective, through being disagreeable to the +birds. In the bed-bug, as Marlatt points out, "it is probably an +illustration of a very common phenomenon among animals, i.e., the +persistence of a characteristic which is no longer of any special value +to the possessor." In fact, its possession is a distinct disadvantage to +the bed-bug, as the odor frequently reveals the presence of the bugs, +before they are seen. + +The eggs of the bed-bug (fig. 70) are pearly white, oval in outline, +about a millimeter long, and possess a small operculum or cap at one +end, which is pushed off when the young hatches. They are laid +intermittently, for a long period, in cracks and crevices of beds and +furniture, under seams of mattresses, under loose wall paper, and +similar places of concealment of the adult bugs. Girault (1905) observed +a well-fed female deposit one hundred and eleven eggs during the +sixty-one days that she was kept in captivity. She had apparently +deposited some of her eggs before being captured. + +The eggs hatch in six to ten days, the newly emerged nymphs being about +1.5 mm. in length and of a pale yellowish white color. They grow slowly, +molting five times. At the last molt the mesathoracic wing pads appear, +characteristic of the adult. The total length of the nymphal stage +varies greatly, depending upon conditions of food supply, temperature +and possibly other factors. Marlatt (1907) found under most favorable +conditions a period averaging eight days between molting which, added to +an equal egg period, gave a total of about seven weeks from egg to adult +insect. Girault (1912) found the postembryonic period as low as +twenty-nine days and as high as seventy days under apparently similar +and normal conditions of food supply. Under optimum and normal +conditions of food supply, beginning August 27, the average nymphal life +was 69.9 days; average number of meals 8.75 and the molts 5. Under +conditions allowing about half the normal food supply the average +nymphal life was from 116.9 to 139 days. Nymphs starved from birth lived +up to 42 days. We have kept unfed nymphs, of the first stage, alive in a +bottle for 75 days. The interesting fact was brought out that under +these conditions of minimum food supply there were sometimes six molts +instead of the normal number. + +The adults are remarkable for their longevity, a factor which is of +importance in considering the spread of the insect and methods of +control. Dufour (1833) (not De Geer, as often stated) kept specimens for +a year, in a closed vial, without food. This ability, coupled with their +willingness to feed upon mice, bats, and other small mammals, and even +upon birds, accounts for the long periods that deserted houses and camps +may remain infested. There is no evidence that under such conditions +they are able to subsist on the starch of the wall paper, juices of +moistened wood, or the moisture in the accumulations of dust, as is +often stated. + +There are three or four generations a year, as Girault's breeding +experiments have conclusively shown. He found that the bed-bug does not +hibernate where the conditions are such as to allow it to breed and that +breeding is continuous unless interrupted by the lack of food or, during +the winter, by low temperature. + +Bed-bugs ordinarily crawl from their hiding places and attack the face +and neck or uncovered parts of the legs and arms of their victims. If +undisturbed, they will feed to repletion. We have found that the young +nymph would glut itself in about six minutes, though some individuals +fed continuously for nine minutes, while the adult required ten to +fifteen minutes for a full meal. When gorged, it quickly retreats to a +crack or crevice to digest its meal, a process which requires two or +three days. The effect of the bite depends very greatly on the +susceptibility of the individual attacked. Some persons are so little +affected that they may be wholly ignorant of the presence of a large +number of bugs. Usually the bite produces a small hard swelling, or +wheal, whitish in color. It may even be accompanied by an edema and a +disagreeable inflammation, and in such susceptible individuals the +restlessness and loss of sleep due to the presence of the insects may be +a matter of considerable importance. Stiles (1907) records the case of a +young man who underwent treatment for neurasthenia, the diagnosis being +agreed upon by several prominent physicians; all symptoms promptly +disappeared, however, immediately following a thorough fumigation of +his rooms, where nearly a pint of bed-bugs were collected. + +It is natural to suppose that an insect which throughout its whole life +is in such intimate relationship with man should play an important rôle +in the transmission of disease. Yet comparatively little is definitely +known regarding the importance of the bed-bug in this respect. It has +been shown that it is capable of transmitting the bubonic plague, and +South American trypanosomiasis. Nuttall succeeded in transmitting +European relapsing fever from mouse to mouse by its bite. It has been +claimed that Oriental sore, tuberculosis, and even syphilis may be so +carried. These phases of the subject will be considered later. + +The sources of infestation are many, and the invasion of a house is not +necessarily due to neglect, though the continued presence of the pests +is quite another matter. In apartments and closely placed houses they +are known to invade new quarters by migration. They are frequently to be +met with in boat and sleeper berths, and even the plush seats of day +coaches, whence a nucleus may be carried in baggage to residences. They +may be brought in the laundry or in clothes of servants. + +Usually they are a great scourge in frontier settlements and it is +generally believed that they live in nature under the bark of trees, in +lumber, and under similar conditions. This belief is founded upon the +common occurrence of bugs resembling the bed-bug, in such places. As a +matter of fact, they are no relation to bed-bugs but belong to +plant-feeding forms alone (fig. 19 _c_, _d_). + +It is also often stated that bed-bugs live in poultry houses, in +swallows nests, and on bats, and that it is from these sources that they +gain access to dwellings. These bugs are specifically distinct from the +true bed-bug, but any of them may, rarely, invade houses. Moreover, +chicken houses are sometimes thoroughly infested with the true _Cimex +lectularius_. + +Control measures consist in the use of iron bedsteads and the reduction +of hiding places for the bugs. If the infestation is slight they may be +exterminated by a vigilant and systematic hunt, and by squirting +gasoline or alcohol into cracks and crevices of the beds, and furniture. +Fumigation must be resorted to in more general infestations. + +The simplest and safest method of fumigation is by the use of flowers of +sulphur at the rate of two pounds to each one thousand cubic feet of +room space. The sulphur should be placed in a pan, a well made in the +top of the pile and a little alcohol poured in, to facilitate burning. +The whole should be placed in a larger pan and surrounded by water so as +to avoid all danger of fire. Windows should be tightly closed, beds, +closets and drawers opened, and bedding spread out over chairs in order +to expose them fully to the fumes. As metal is tarnished by the sulphur +fumes, ornaments, clocks, instruments, and the like should be removed. +When all is ready the sulphur should be fired, the room tightly closed +and left for twelve to twenty-four hours. Still more efficient in large +houses, or where many hiding places favor the bugs, is fumigation with +hydrocyanic acid gas. This is a deadly poison and must be used under +rigid precautions. Through the courtesy of Professor Herrick, who has +had much experience with this method, we give in the Appendix, the clear +and detailed directions taken from his bulletin on "Household Insects." + +Fumigation with formaldehyde gas, either from the liquid or "solid" +formalin, so efficient in the case of contagious diseases, is useless +against bed-bugs and most other insects. + + +OTHER BED-BUGS--_Cimex hemipterus_ (= _C. rotundatus_) is a tropical and +subtropical species, occurring in both the old and new world. Patton and +Cragg state that it is distributed throughout India, Burma, Assam, the +Malay Peninsula, Aden, the Island of Mauritius, Reunion, St. Vincent and +Porto Rico. "It is widely distributed in Africa, and is probably the +common species associated there with man." Brumpt also records it for +Cuba, the Antilles, Brazil, and Venezuela. + +This species, which is sometimes called the Indian bed-bug, differs from +_C. lectularius_ in being darker and in having a more elongate abdomen. +The head also is shorter and narrower, and the prothorax has rounded +borders. + +It has the same habits and practically the same life cycle as _Cimex +lectularius_. Mackie, in India, has found that it is capable of +transmitting the Asiatic type of recurrent fever. Roger suggested that +it was also capable of transmitting Kala-azar and Patton has described +in detail the developmental stages of _Leishmania_, the causative +organism of Kala-azar, in the stomach of this bug, but Brumpt declares +that the forms described are those of a common, non-pathogenic +flagellate to be found in the bug, and have nothing to do with the +human disease. Brumpt has shown experimentally that _Cimex hemipterus_ +may transmit _Trypanosoma cruzi_ in its excrement. + +_Cimex boueti_, occurring in French Guinea, is another species attacking +man. Its habits and general life history are the same as for the above +species. It is 3 to 4.5 mm. in length, has vestigial elytra, and much +elongated antennæ and legs. The extended hind legs are about as long as +the body. + +_Cimex columbarius_, a widely distributed species normally living in +poultry houses and dove cotes, _C. inodorus_, infesting poultry in +Mexico, _C. hirundinis_, occurring in the nests of swallows in Europe +and _Oeciacus vicarius_ (fig. 19i) occurring in swallows' nests in this +country, are species which occasionally infest houses and attack man. + +[Illustration: 71. Conorhinus sanguisugus.] + +_Conorhinus sanguisugus_, the cone-nosed bed-bug. We have seen in our +consideration of poisonous insects, that various species of Reduviid +bugs readily attack man. Certain of these are nocturnal and are so +commonly found in houses that they have gained the name, of "big +bed-bugs." The most noted of these, in the United States, is _Conorhinus +sanguisugus_ (fig. 71), which is widely distributed in our Southern +States. + +Like its near relatives, _Conorhinus sanguisugus_ is carnivorous in +habit and feeds upon insects as well as upon mammalian and human blood. +It is reported as often occurring in poultry houses and as attacking +horses in barns. The life history has been worked out in considerable +detail by Marlatt, (1902), from whose account we extract the following. + +[Illustration: 72. Beak of Conorhinus sanguisugus. After Marlatt.] + +The eggs are white, changing to yellow and pink before hatching. The +young hatch within twenty days and there are four nymphal stages. In all +these stages the insect is active and predaceous, the mouth-parts (fig. +72) being powerfully developed. The eggs are normally deposited, and the +early stages are undoubtedly passed, out of doors, the food of the +immature forms being other insects. Immature specimens are rarely found +indoors. It winters both in the partly grown and adult stage, often +under the bark of trees or in any similar protection, and only in its +nocturnal spring and early summer flights does it attack men. Marlatt +states that this insect seems to be decidedly on the increase in the +region which it particularly infests,--the plains region from Texas +northward and westward. In California a closely related species of +similar habits is known locally as the "monitor bug." + +The effect of the bite of the giant bed-bug on man is often very severe, +a poisonous saliva apparently being injected into the wound. We have +discussed this phase of the subject more fully under the head of +poisonous insects. + +_Conorhinus megistus_ is a Brazilian species very commonly attacking +man, and of special interest since Chagas has shown that it is the +carrier of a trypanosomiasis of man. Its habits and life history have +been studied in detail by Neiva, (1910). + +This species is now pre-eminently a household insect, depositing its +eggs in cracks and crevices in houses, though this is a relatively +recent adaptation. The nymphs emerge in from twenty to forty days, +depending upon the temperature. There are five nymphal stages, and as in +the case of true bed-bugs, the duration of these is very greatly +influenced by the availability of food and by temperature. Neiva reckons +the entire life cycle, from egg to egg, as requiring a minimum of three +hundred and twenty-four days. + +The nymphs begin to suck blood in three to five days after hatching. +They usually feed at night and in the dark, attacking especially the +face of sleeping individuals. The bite occasions but little pain. The +immature insects live in cracks and crevices in houses and invade the +beds which are in contact with walls, but the adults are active flyers +and attack people sleeping in hammocks. The males as well as the females +are blood suckers. + +Like many blood-sucking forms, _Conorhinus megistus_ can endure for long +periods without food. Neiva received a female specimen which had been +for fifty-seven days alive in a tightly closed box. They rarely feed on +two consecutive days, even on small quantities of blood, and were never +seen to feed on three consecutive days. + +Methods of control consist in screening against the adult bugs, and the +elimination of crevices and such hiding places of the nymphs. Where the +infestation is considerable, fumigation with sulphur is advisable. + + +PARASITIC DIPTERA OR FLIES + +Of the DIPTERA or two-winged flies, many species occasionally attack +man. Of these, a few are outstanding pests, many of them may also serve +to disseminate disease, a phase of our subject which will be considered +later. We shall now consider the most important of the group from the +viewpoint of their direct attacks on man. + + +Psychodidæ or Moth-Flies + +The PSYCHODIDÆ or Moth-flies, include a few species which attack man, +and at least one species, _Phlebotomus papatasii_, is known to transmit +the so-called "three-day fever" of man. Another species is supposed to +be the vector of Peruvian verruga. + +The family is made up of small, sometimes very small, nematocerous +Diptera, which are densely covered with hairs, giving them a moth-like +appearance. The wings are relatively large, oval or lanceolate in shape, +and when at rest are held in a sloping manner over the abdomen, or are +held horizontally in such a way as to give the insect a triangular +outline. Not only is the moth-like appearance characteristic, but the +venation of the wings (fig. 163, d) is very peculiar and, according to +Comstock, presents an extremely generalized form. All of the +longitudinal veins separate near the base of the wing except veins R_2 +and R_3 and veins M_1 and M_2. Cross veins are wanting in most cases. + +Comparatively little is known regarding the life-history and habits of +the Psychodidæ, but one genus, _Phlebotomus_, contains minute, +blood-sucking species, commonly known as sand-flies. The family is +divided into two subfamilies, the PSYCHODINÆ and the PHLEBOTOMINÆ. The +second of these, the PHLEBOTOMINÆ, is of interest to us. + + +THE PHLEBOTOMINÆ--The Phlebotominæ differ from the Psychodinæ in that +the radical sector branches well out into the wing rather than at the +base of the wing. They are usually less hairy than the Psychodinæ. The +ovipositor is hidden and less strongly chitinized. The species attacking +man belong to the genus _Phlebotomus_, small forms with relatively +large, hairy wings which are held upright, and with elongate proboscis. +The mandibles and maxillæ are serrated and fitted for biting. + +According to Miss Summers (1913) there are twenty-nine known species of +the genus _Phlebotomus_, five European, eleven Asiatic, seven African +and six American. One species only, _Phlebotomus vexator_, has been +reported for the United States. This was described by Coquillett, +(1907), from species taken on Plummer's Island, Maryland. It measures +only 1.5 mm. in length. As it is very probable that this species is much +more widely distributed, and that other species of these minute flies +will be found to occur in our fauna, we quote Coquillett's description. + +_Phlebotomus vexator_, Coq.: Yellow, the mesonotum brown, hairs chiefly +brown; legs in certain lights appear brown, but are covered with a white +tomentum; wings hyaline, unmarked; the first vein (R_1) terminates +opposite one-fifth of the length of the first submarginal cell (cell +R_2); this cell is slightly over twice as long as its petiole; terminal, +horny portion of male claspers slender, bearing many long hairs; the +apex terminated by two curved spines which are more than one-half as +long as the preceding part, and just in front of these are two similar +spines, while near the middle of the length of this portion is a fifth +spine similar to the others. Length 1.5 mm. + +The life-history of the Phlebotomus flies has been best worked out for +the European _Phlebotomus papatasii_ and we shall briefly summarize the +account of DÅ“rr and Russ (1913) based primarily on work on this +species. The European Phlebotomus flies appear at the beginning of the +warm season, a few weeks after the cessation of the heavy rains and +storms of springtime. They gradually become more abundant until they +reach their first maximum, which in Italy is near the end of July +(Grassi). They then become scarcer but reach a second maximum in +September. At the beginning of winter they vanish completely, +hibernating individuals not being found. + +After fertilization there is a period of eight to ten days before +oviposition. The eggs are then deposited, the majority in a single mass +covered by a slimy secretion from the sebaceous glands. The larvæ emerge +in fourteen to twenty days. There is uncertainty as to the length of +larval life, specimens kept in captivity remaining fifty or more days +without transforming. Growth may be much more rapid in nature. The larvæ +do not live in fluid media but in moist detritus in dark places. Marett +believes that they live chiefly on the excrement of pill-bugs (Oniscidæ) +and lizards. Pupation always occurs during the night. The remnants of +the larval skin remain attached to the last two segments of the +quiescent pupa and serve to attach it to the stone on which it lives. +The pupal stage lasts eleven to sixteen days, the adult escaping at +night. + +Only the females suck blood. They attack not only man but all +warm-blooded animals and, according to recent workers, also cold-blooded +forms, such as frogs, lizards, and larvæ. Indeed, Townsend (1914) +believes that there is an intimate relation between _Phlebotomus_ and +lizards, or other reptiles the world over. The Phlebotomus passes the +daylight hours within the darkened recesses of the loose stone walls and +piles of rock in order to escape wind and strong light. Lizards inhabit +the same places, and the flies, always ready to suck blood in the +absence of light and wind, have been found more prone to suck reptilian +than mammalian blood. + +On hot summer nights, when the wind is not stirring, the Phlebotomus +flies, or sand-flies, as they are popularly called, invade houses and +sleeping rooms in swarms and attack the inmates. As soon as light begins +to break the flies either escape to the breeding places, or cool, dark +places protected from the wind, or a part of them remain in the rooms, +hiding behind pictures, under garments, and in similar places. Wherever +the Phlebotomus flies occur they are an intolerable nuisance. On account +of their small size they can easily pass through the meshes of ordinary +screens and mosquito curtains. They attack silently and inflict a very +painful, stinging bite, followed by itching. The ankles, dorsum of the +feet, wrists, inner elbow, knee joint and similar places are favorite +places of attack, possibly on account of their more delicate skin. + +Special interest has been attracted to these little pests in recent +years, since it has been shown that they transmit the European +"pappatici fever" or "three day fever." More recently yet, it appears +that they are the carriers of the virus of the Peruvian "verruga." This +phase of the subject will be discussed later. + +Control measures have not been worked out. As Newstead says, "In +consideration of the facts which have so far been brought to light +regarding the economy of Phlebotomus, it is clearly evident that the +task of suppressing these insects is an almost insurmountable one. Had +we to deal with insects as large and as accessible as mosquitoes, the +adoption of prophylactic measures would be comparatively easy, but owing +to the extremely minute size and almost flea-like habits of the adult +insects, and the enormous area over which the breeding-places may occur, +we are faced with a problem which is most difficult of solution." For +these reasons, Newstead considers that the only really prophylactic +measures which can at present be taken, are those which are considered +as precautionary against the bites of the insects. + +Of repellents, he cites as one of the best a salve composed of the +following: + + Ol. Anisi 3 grs. + Ol. Eucalypti 3 grs. + Ol. Terebenth 3 grs. + Unq. Acid Borac. + +Of sprays he recommends as the least objectionable and at the same time +one of the most effective, formalin. "The dark portions and angles of +sleeping apartments should be sprayed with a one per cent. solution of +this substance every day during the season in which the flies are +prevalent. A fine spraying apparatus is necessary for its application +and an excessive amount must not be applied. It is considered an +excellent plan also to spray the mosquito curtains regularly every day +towards sunset; nets thus treated are claimed to repel the attacks of +these insects." This effectiveness of formalin is very surprising for, +as we have seen, it is almost wholly ineffective against bed-bugs, +mosquitoes, house flies and other insects, where it has been tried. + +A measure which promises to be very effective, where it can be adopted, +is the use of electric fans so placed as to produce a current of air in +the direction of the windows of sleeping apartments. On account of the +inability of the Phlebotomus flies to withstand even slight breezes, it +seems very probable that they would be unable to enter a room so +protected. + + +Culicidæ or Mosquitoes + +From the medical viewpoint, probably the most interesting and important +of the blood-sucking insects are the mosquitoes. Certainly this is true +of temperate zones, such as those of the United States. The result is +that no other group of insects has aroused such widespread interest, or +has been subjected to more detailed study than have the mosquitoes, +since their rôle as carriers of disease was made known. There is an +enormous literature dealing with the group, but fortunately for the +general student, this has been well summarized by a number of workers. +The most important and helpful of the general works are those of Howard +(1901), Smith (1904), Blanchard (1905), Mitchell (1907), and especially +of Howard, Dyar, and Knab, whose magnificent monograph is still in +course of publication. + +Aside from their importance as carriers of disease, mosquitoes are +notorious as pests of man, and the earlier literature on the group is +largely devoted to references to their enormous numbers and their +blood-thirstiness in certain regions. They are to be found in all parts +of the world, from the equator to the Arctic and Antarctic regions. +Linnæus, in the "Flora Lapponica," according to Howard, Dyar and Knab, +"dwells at some length upon the great abundance of mosquitoes in Lapland +and the torments they inflicted upon man and beast. He states that he +believes that nowhere else on earth are they found in such abundance and +he compares their numbers to the dust of the earth. Even in the open, +you cannot draw your breath without having your mouth and nostrils +filled with them; and ointments of tar and cream or of fish grease are +scarcely sufficient to protect even the case-hardened cuticle of the +Laplander from their bite. Even in their cabins, the natives cannot take +a mouthful of food or lie down to sleep unless they are fumigated almost +to suffocation." In some parts of the Northwestern and Southwestern +United States it is necessary to protect horses working in the fields by +the use of sheets or burlaps, against the ferocious attacks of these +insects. It is a surprising fact that even in the dry deserts of the +western United States they sometimes occur in enormous numbers. + +Until comparatively recent years, but few species of mosquitoes were +known and most of the statements regarding their life-history were based +upon the classic work of Reaumur (1738) on the biology of the rain +barrel mosquito, _Culex pipiens_. In 1896, Dr. Howard refers to +twenty-one species in the United States, now over fifty are known; +Giles, in 1900, gives a total of two hundred and forty-two for the world +fauna, now over seven hundred species are known. We have found eighteen +species at Ithaca, N. Y. + +All of the known species of mosquitoes are aquatic in the larval stage, +but in their life-histories and habits such great differences occur that +we now know that it is not possible to select any one species as typical +of the group. For our present purpose we shall first discuss the general +characteristics and structure of mosquitoes, and shall then give the +life-history of a common species, following this by a brief +consideration of some of the more striking departures from what have +been supposed to be the typical condition. + +The CULICIDÆ are slender, nematocerous Diptera with narrow wings, +antennæ plumose in the males, and usually with the proboscis much longer +than the head, slender, firm and adapted for piercing in the female. +The most characteristic feature is that the margins of the wings and, in +most cases, the wing veins possess a fringe of scale-like hairs. These +may also cover in part, or entirely, the head, thorax, abdomen and legs. +The females, only, suck blood. + +On account of the importance of the group in this country and the +desirability of the student being able to determine material in various +stages, we show in the accompanying figures the characters most used in +classification. + +The larvæ (fig. 73) are elongate, with the head and thorax sharply +distinct. The larval antennæ are prominent, consisting of a single +cylindrical and sometimes curved segment. The outer third is often +narrower and bears at its base a fan-shaped tuft of hairs, the +arrangement and abundance of which is of systematic importance. About +the mouth are the so-called rotary mouth brushes, dense masses of long +hairs borne by the labrum and having the function of sweeping food into +the mouth. The form and arrangement of thoracic, abdominal, and anal +tufts of hair vary in different species and present characteristics of +value. On either side of the eighth abdominal segment is a patch of +scales varying greatly in arrangement and number and of much value in +separating species. Respiration is by means of tracheæ which open at the +apex of the so-called anal siphon, when it is present. In addition, +there are also one or two pairs of tracheal gills which vary much in +appearance in different species. On the ventral side of the anal siphon +is a double row of flattened, toothed spines whose number and shape are +likewise of some value in separating species. They constitute the comb +or pecten. + +[Illustration: 73. Culex larva showing details of external structure.] + +The pupa (fig. 139, b) unlike that of most insects, is active, though it +takes no food. The head and thorax are not distinctly separated, but the +slender flexible abdomen in sharply marked off. The antennæ, +mouth-parts, legs, and wings of the future adult are now external, but +enclosed in chitinous cases. On the upper surface, near the base of the +wings are two trumpets, or breathing tubes, for the pupal spiracles are +towards the anterior end instead of at the caudal end, as in the larva. +At the tip of the abdomen is a pair of large chitinous swimming paddles. + +As illustrative of the life cycle of a mosquito we shall discuss the +development of a common house mosquito, _Culex pipiens_, often referred +to in the Northern United States as the rain barrel mosquito. Its life +cycle is often given as typical for the entire group, but, as we have +already emphasized, no one species can serve this purpose. + +The adults of _Culex pipiens_ hibernate throughout the winter in +cellars, buildings, hollow trees, or similar dark shelters. Early in the +spring they emerge and deposit their eggs in a raft-like mass. The +number of eggs in a single mass is in the neighborhood of two hundred, +recorded counts varying considerably. A single female may deposit +several masses during her life time. The duration of the egg stage is +dependent upon temperature. In the warm summer time the larvæ may emerge +within a day. The larvæ undergo four molts and under optimum conditions +may transform into pupæ in about a week's time. Under the same favorable +conditions, the pupal stage may be completed in a day's time. The total +life cycle of _Culex pipiens_, under optimum conditions, may thus be +completed in a week to ten days. This period may be considerably +extended under less favorable conditions of temperature and food supply. + +_Culex pipiens_ breeds continuously throughout the summer, developing in +rain barrels, horse troughs, tin cans, or indeed in any standing water +about houses, which lasts for a week or more. The catch basins of sewers +furnish an abundant supply of the pests under some conditions. Such +places, the tin gutters on residences, and all possible breeding places +must be considered in attempts to exterminate this species. + +Other species of mosquitoes may exhibit radical departures from _Culex +pipiens_ in life-history and habits. To control them it is essential +that the biological details be thoroughly worked out for, as Howard, +Dyar, and Knab have emphasized, "much useless labor and expense can be +avoided by an accurate knowledge of the habits of the species." For a +critical discussion of the known facts the reader is referred to their +monograph. We shall confine ourselves to a few illustrations. + +The majority of mosquitoes in temperate climates hibernate in the egg +stage, hatching in the spring or even mild winter days in water from +melting snow. It is such single-brooded species which appear in +astounding numbers in the far North. Similarly, in dry regions the eggs +may stand thorough dessication, and yet hatch out with great promptness +when submerged by the rains. "Another provision to insure the species +against destruction in such a case, exists in the fact * * * that not +all the eggs hatch, a part of them lying over until again submerged by +subsequent rains." In temperate North America, a few species pass the +winter in the larval state. An interesting illustration of this is +afforded by _Wyeomia smithii_, whose larvæ live in pitcher plants and +are to be found on the coldest winter days imbedded in the solid ice. +Late in the spring, the adults emerge and produce several broods during +the summer. + +In the United States, one of the most important facts which has been +brought out by the intensive studies of recent years is that certain +species are migratory and that they can travel long distances and become +an intolerable pest many miles from their breeding places. This was +forcibly emphasized in Dr. Smith's work in New Jersey, when he found +that migratory mosquitoes, developing in the salt marshes along the +coast, are the dominant species largely responsible for the fame of the +New Jersey mosquito. The species concerned are _Aedes sollicitans_, _A. +cantator_ and _A. tæniorhynchus_. Dr. Smith decided that the first of +these might migrate at least forty miles inland. It is obvious that +where such species are the dominant pest, local control measures are a +useless waste of time and money. Such migratory habits are rare, +however, and it is probable that the majority of mosquitoes do not fly +any great distance from their breeding places. + +While mosquitoes are thought of primarily as a pest of man, there are +many species which have never been known to feed upon human or mammalian +blood, no matter how favorable the opportunity. According to Howard, +Dyar, and Knab, this is true of _Culex territans_, one of the common +mosquitoes in the summer months in the Northern United States. There are +some species, probably many, in which the females, like the males, are +plant feeders. In experimental work, both sexes are often kept alive for +long periods by feeding them upon ripe banana, dried fig, raisins, and +the like, and in spite of sweeping assertions that mosquitoes must have +a meal of blood in order to stimulate the ovaries to development, some +of the common blood-sucking species, notably _Culex pipiens_, have been +bred repeatedly without opportunity to feed upon blood. + +The effect of the bite varies greatly with different species and depends +upon the susceptibility of the individual bitten. Some persons are +driven almost frantic by the attacks of the pests when their companions +seem almost unconscious of any inconvenience. Usually, irritation and +some degree of inflammation appear shortly following the bite. Not +infrequently a hardened wheal or even a nodule forms, and sometimes +scratching leads to secondary infection and serious results. + +The source of the poison is usually supposed to be the salivary glands +of the insect. As we have already pointed out, (p. 34), Macloskie +believed that one lobe of the gland, on each side, was specialized for +forming the poison, while a radically different view is that of +Schaudinn, who believed that the irritation is due to the expelled +contents of the Å“sophageal diverticula, which contain a gas and a +peculiar type of fungi or bacteria. In numerous attempts, Schaudinn was +unable to produce any irritation by applying the triturated salivary +glands to a wound, but obtained the typical result when he used the +isolated diverticula. + +The irritation of the bite may be relieved to some extent by using +ammonia water, a one per cent. alcoholic solution of menthol, or +preparations of cresol, or carbolic acid. Dr. Howard recommends rubbing +the bite gently with a piece of moist toilet soap. Castellani and +Chalmers recommend cleansing inflamed bites with one in forty carbolic +lotion, followed by dressing with boracic ointment. Of course, +scratching should be avoided as much as possible. + +Repellents of various kinds are used, for warding off the attacks of the +insects. We have often used a mixture of equal parts of oil of +pennyroyal and kerosene, applied to the hands and face. Oil of +citronella is much used and is less objectionable to some persons. A +recommended formula is, oil of citronella one ounce, spirits of camphor +one ounce, oil of cedar one-half ounce. A last resort would seem to be +the following mixture recommended by Howard, Dyar, and Knab for use by +hunters and fishermen in badly infested regions, against mosquitoes and +blackflies. + +Take 2¼ lbs. of mutton tallow and strain it. While still hot add ½ +lb. black tar (Canadian tar). Stir thoroughly and pour into the +receptacle in which it is to be contained. When nearly cool stir in +three ounces of oil of citronella and 1¼ oz. of pennyroyal. + +At night the surest protection is a good bed net. There are many types +of these in use, but in order to be serviceable and at the same time +comfortable it should be roomy and hung in such a way as to be stretched +tightly in every direction. We prefer one suspended from a broad, square +frame, supported by a right-angled standard which is fastened to the +head of the bed. It must be absolutely free from rents or holes and +tucked in securely under the mattress or it will serve merely as a +convenient cage to retain mosquitoes which gain an entrance. While such +nets are a convenience in any mosquito ridden community, they are +essential in regions where disease-carrying species abound. Screening of +doors, windows and porches, against the pests is so commonly practiced +in this country that its importance and convenience need hardly be +urged. + +Destruction of mosquitoes and prevention of breeding are of fundamental +importance. Such measures demand first, as we have seen, the correct +determination of the species which is to be dealt with, and a knowledge +of its life-history and habits. If it prove to be one of the migratory +forms, it is beyond mere local effort and becomes a problem demanding +careful organization and state control. An excellent illustration of the +importance and effectiveness of work along these lines is afforded by +that in New Jersey, begun by the late Dr. John B. Smith and being pushed +with vigor by his successor, Dr. Headlee. + +In any case, there is necessity for community action. Even near the +coast, where the migratory species are dominant, there are the local +species which demand attention and which cannot be reached by any +measures directed against the species of the salt marshes. The most +important of local measures consist in the destruction of breeding +places by filling or draining ponds and pools, clearing up of more +temporary breeding places, such as cans, pails, water barrels and the +like. Under conditions where complete drainage of swamps is +impracticable or undesirable, judicious dredging may result in a pool or +series of steep-sided pools deep enough to maintain a supply of fish, +which will keep down the mosquito larvæ. Where water receptacles are +needed for storage of rain water, they should be protected by careful +screening or a film of kerosene over the top of the water, renewed every +two weeks or so, so as to prevent mosquitoes from depositing their eggs. +When kerosene is used, Water drawn from the bottom of the receptacle +will not be contaminated by it to any injurious extent. Where ponds +cannot be drained much good will be accomplished by spraying kerosene +oil on the surface of the water, or by the introduction of fish which +will feed on the larvæ. + +Detailed consideration of the most efficient measures for controlling +mosquitoes is to be found in Dr. Howard's Bulletin No. 88 of the Bureau +of Entomology, "Preventive and remedial work against mosquitoes" or, in +more summarized form, in Farmers' Bulletin No. 444. One of these should +be obtained by any person interested in the problems of mosquito control +and public health. + +[Illustration: 74. Mouth parts of Simulium. After Grünberg.] + + +The Simuliidæ, or Black Flies + +The SIMULIIDÆ, or black flies, are small, dark, or black flies, with a +stout body and a hump-back appearance. The antennæ are short but +eleven-segmented, the wings broad, without scales or hairs, and with the +anterior veins stout but the others very weak. The mouth-parts (fig. 74) +are fitted for biting. + +The larvæ of the Simuliidæ (fig. 75) are aquatic and, unlike those of +mosquitoes, require a well ærated, or swiftly running water. Here they +attach to stones, logs, or vegetation and feed upon various +micro-organisms. They pupate in silken cocoons open at the top. Detailed +life-histories have not been worked out for most of the species. We +shall consider as typical that of _Simulium pictipes_, an inoffensive +species widely distributed in the Eastern United States, which has been +studied especially at Ithaca, N.Y. (Johannsen, 1903). + +[Illustration: 75. Larva of Simulium, (×8). After Garman.] + +The eggs are deposited in a compact yellowish layer on the surface of +rock, on the brinks of falls and rapids where the water is flowing +swiftly. They are elongate ellipsoidal in shape, about .4 by .18 mm. As +myriads of females deposit in the same place the egg patches may be +conspicuous coatings of a foot or much more in diameter. When first laid +they are enveloped in a yellowish white slime, which becomes darker, +until finally it becomes black just before the emerging of the larvæ. +The egg stage lasts a week. + +The larvæ (fig. 75) are black, soft skinned, somewhat cylindrical in +shape, enlarged at both ends and attenuated in the middle. The posterior +half is much stouter than the anterior part and almost club-shaped. The +head bears two large fan-shaped organs which aid in procuring food. +Respiration is accomplished by means of three so-called blood gills +which are pushed out from the dorsal part of the rectum. The larvæ occur +in enormous numbers, in moss-like patches. If removed from their natural +habitat and placed in quiet water they die within three or four hours. +Fastened to the rock by means of a disk-like sucker at the caudal end of +the body, they ordinarily assume an erect position. They move about on +the surface of the rocks, to a limited extent, with a looping gait +similar to that of a measuring worm, and a web is secreted which +prevents their being washed away by the swiftly flowing water. They feed +chiefly upon algæ and diatoms. + +The complete larval stage during the summer months occupies about four +weeks, varying somewhat with the temperature and velocity of the water. +At the end of this period they spin from cephalic glands, boot-shaped +silken cocoons within which they pupate. The cocoon when spun is firmly +attached to the rock and also to adjacent cocoons. Clustered +continuously over a large area and sometimes one above another, they +form a compact, carpet-like covering on the rocks, the reddish-brown +color of which is easily distinguishable from the jet-black appearance +of the larvæ. The pupal stage lasts about three weeks. The adult fly, +surrounded by a bubble of air, quickly rises to the surface of the water +and escapes. The adults (fig. 76) are apparently short lived and thus +the entire life cycle, from egg to egg is completed in approximately +eight weeks. + +[Illustration: 76. Simulium venustum, (×8). After Garman.] + +In the case of _Simulium pictipes_ at Ithaca, N. Y., the first brood of +adults emerges early in May and successive generations are produced +throughout the summer and early autumn. This species winters in the +larval condition. Most of the other species of _Simulium_ which have +been studied seem to be single brooded. + +While _Simulium pictipes_ does not attack man, there are a number of the +species which are blood-sucking and in some regions they are a veritable +scourge. In recent years the greatest interest in the group has been +aroused by Sambon's hypothesis that they transmit pellagra from man to +man. This has not been established, and, indeed, seems very doubtful, +but the importance of these insects as pests and the possibility that +they may carry disease make it urgent that detailed life-histories of +the hominoxious species be worked out. + +As pests a vivid account of their attacks is in Agassiz's "Lake +Superior" (p. 61), quoted by Forbes (1912). + +"Neither the love of the picturesque, however, nor the interests of +science, could tempt us into the woods, so terrible were the black +flies. This pest of flies which all the way hither had confined our +ramblings on shore pretty closely to the rocks and the beach, and had +been growing constantly worse, here reached its climax. Although +detained nearly two days, * * * we could only sit with folded hands, or +employ ourselves in arranging specimens, and such other operations as +could be pursued in camp, and under the protection of a 'smudge.' One, +whom scientific ardor tempted a little way up the river in a canoe, +after water plants, came back a frightful spectacle, with blood-red +rings round his eyes, his face bloody, and covered with punctures. The +next morning his head and neck were swollen as if from an attack of +erysipelas." + +There are even well authenticated accounts on record of death of humans +from the attacks of large swarms of these gnats. In some regions, and +especially in the Mississippi Valley in this country, certain species of +black flies have been the cause of enormous losses to farmers and +stockmen, through their attacks on poultry and domestic animals. C. V. +Riley states that in 1874 the loss occasioned in one county in Tennessee +was estimated at $500,000. + +The measures of prevention and protection against these insects have +been well summarized by Forbes (1912). They are of two kinds: "the use +of repellents intended to drive away the winged flies, and measures for +the local destruction of the aquatic larvæ. The repellents used are +either smudges, or surface applications made to keep the flies from +biting. The black-fly will not endure a dense smoke, and the well-known +mosquito smudge seems to be ordinarily sufficient for the protection of +man. In the South, leather, cloth, and other materials which will make +the densest and most stifling smoke, are often preserved for this use in +the spring. Smudges are built in pastures for the protection of stock, +and are kept burning before the doors of barns and stables. As the +black-flies do not readily enter a dark room, light is excluded from +stables as much as possible during the gnat season. If teams must be +used in the open field while gnats are abroad, they may be protected +against the attacks of the gnats by applying cotton-seed oil or axle +grease to the surface, especially to the less hairy parts of the +animals, at least twice a day. A mixture of oil and tar and, indeed, +several other preventives, are of practical use in badly infested +regions; but no definite test or exact comparison has been made with any +them in a way to give a record of the precise results." + +"It is easy to drive the flies from houses or tents by burning pyrethrum +powder inside; this either kills the flies or stupifies them so that +they do not bite for some time thereafter." * * * "Oil of tar is +commonly applied to the exposed parts of the body for the purpose of +repelling the insects, and this preparation is supplied by the Hudson +Bay Company to its employees. Minnesota fishermen frequently grease +their faces and hands with a mixture of kerosene and mutton tallow for +the same purpose." We have found a mixture of equal parts of kerosene +and oil of pennyroyal efficient. + +Under most circumstances very little can be done to destroy this insect +in its early stage, but occasionally conditions are such that a +larvicide can be used effectively. Weed (1904), and Sanderson (1910) +both report excellent results from the use of phinotas oil, a +proprietary compound. The first-mentioned also found that in some places +the larvæ could be removed by sweeping them loose in masses with stiff +stable brooms and then catching them downstream on wire netting +stretched in the water. + + +Chironomidæ or Midges + +The flies of this family, commonly known as midges, resemble mosquitoes +in form and size but are usually more delicate, and the wing-veins, +though sometimes hairy, are not fringed with scales. The venation is +simpler than in the mosquitoes and the veins are usually less distinct. + +These midges, especially in spring or autumn, are often seen in immense +swarms arising like smoke over swamps and producing a humming noise +which can be heard for a considerable distance. At these seasons they +are frequently to be found upon the windows of dwellings, where they are +often mistaken for mosquitoes. + +The larvæ are worm-like, but vary somewhat in form in the different +genera. Most of them are aquatic, but a few live in the earth, in +manure, decaying wood, under bark, or in the sap of trees, especially in +the sap which collects in wounds. + +[Illustration: 77. Culicoides guttipennis; (_a_) adult, (×15); (_b_) +head of same; (_c_) larva; (_d_) head; (_e_) pupa. After Pratt.] + +Of the many species of CHIRONOMIDÆ, (over eight hundred known), the vast +majority are inoffensive. The sub-family Ceratopogoninæ, however, forms +an exception, for some of the members of this group, known as sandflies, +or punkies, suck blood and are particularly troublesome in the +mountains, along streams, and at the seashore. Most of these have been +classed under the genus _Ceratopogon_, but the group has been broken up +into a number of genera and _Ceratopogon_, in the strict sense, is not +known to contain any species which sucks the blood of vertebrates. + +THE CERATOPOGONINÆ--The Ceratopogoninæ are among the smallest of the +Diptera, many of them being hardly a millimeter long and some not even +so large. They are Chironomidæ in which the thorax is not prolonged over +the head. The antennæ are filiform with fourteen (rarely thirteen) +segments in both sexes, those of the male being brush-like. The basal +segment is enlarged, the last segment never longer than the two +preceding combined, while the last five are sub-equal to, or longer than +the preceding segment. The legs are relatively stouter than in the other +Chironomidæ. The following three genera of this subfamily are best known +as blood suckers in this country. + +Of the genus _Culicoides_ there are many species occurring in various +parts of the world. A number are known to bite man and animals and it is +probable that all are capable of inflicting injury. In some localities +they are called punkies, in others, sand-flies, a name sometimes also +applied to the species of _Simulium_ and _Phlebotomus_. Owing to their +very small size they are known by some tribes of Indians as No-see-ums. +The larvæ are found in ponds, pools, water standing in hollow tree +stumps, and the like. Though probably living chiefly in fresh water, we +have found a species occurring in salt water. The larvæ are small, +slender, legless, worm-like creatures (fig. 77_c_) with small brown head +and twelve body segments. The pupæ (fig. 77_e_) are slender, more +swollen at the anterior end and terminating in a forked process. They +float nearly motionless in a vertical position, the respiratory tubes in +contact with the surface film. The adults are all small, rarely +exceeding 2¼ mm. in length. The wings are more or less covered with +erect setulæ or hairs and in many species variously spotted and marked +with iridescent blotches. The antennæ have fourteen segments, the palpi +usually five. The wing venation and mouth-parts are shown in figures 77 +and 78. Of the twenty or more species of this genus occurring in the +United States the following are known to bite: _C. cinctus_, _C. +guttipennis_, _C. sanguisuga_, _C. stellifer_, _C. variipennis_, _C. +unicolor_. + +[Illustration: 78. Culicoides guttipennis; mouth parts of adult. After +Pratt.] + +One of the most widely distributed and commonest species in the Eastern +States is _C. guttipennis_ (fig. 77a). It is black with brown legs, a +whitish ring before the apex of each femur and both ends of each tibia; +tarsi yellow, knobs of halteres yellow. Mesonotum opaque, brown, two +vittæ in the middle, enlarging into a large spot on the posterior half, +also a curved row of three spots in front of each wing, and the narrow +lateral margins, light gray pruinose. Wings nearly wholly covered with +brown hairs, gray, with markings as shown in the figure. Length one mm. + +_Johannseniella_ Will. is a wide-spread genus related to the foregoing. +Its mouth-parts are well adapted for piercing and it is said to be a +persistent blood sucker, particularly in Greenland. This genus is +distinguished from _Culicoides_ by its bare wings, the venation (fig. +163, c), and the longer tarsal claws. There are over twenty North +American species. + +[Illustration: 79. Chrysops univittatus, (×4). After Osborn.] + +In the Southwestern United States, _Tersesthes torrens_ Towns. occurs, a +little gnat which annoys horses, and perhaps man also, by its bite. It +is related to _Culicoides_ but differs in the number of antennal +segments and in its wing venation (fig. 163, e). The fly measures but +two mm. in length and is blackish in color. The antennæ of the female +have thirteen segments, the palpi but three, of which the second is +enlarged and swollen. + + +Tabanidæ or Horse-Flies + +The TABANIDÆ,--horse-flies, ear-flies, and deer-flies,--are well-known +pests of cattle and horses and are often extremely annoying to man. The +characteristics of the family and of the principal North American genera +are given in the keys of Chapter XII. There are over 2500 recorded +species. As in the mosquitoes, the females alone are blood suckers. The +males are flower feeders or live on plant juices. This is apparently +true also of the females of some of the genera. + +The eggs are deposited in masses on water plants or grasses and sedges +growing in marshy or wet ground. Those of a common species of _Tabanus_ +are illustrated in figure 80, _a_. They are placed in masses of several +hundred, light colored when first deposited but turning black. In a week +or so the cylindrical larvæ, tapering at both ends (fig. 80, _b_), +escape to the water, or damp earth, and lead an active, carnivorous +life, feeding mainly on insect larvæ, and worms. In the forms which have +been best studied the larval life is a long one, lasting for months or +even for more than a year. Until recently, little was known concerning +the life-histories of this group, but the studies of Hart (1895), and +Hine (1903 +) have added greatly to the knowledge concerning North +American forms. + +Many of the species attack man with avidity and are able to inflict +painful bites, which may smart for hours. In some instances the wound is +so considerable that blood will continue to flow after the fly has left. +We have seen several cases of secondary infection following such bites. + +[Illustration: 80. (_a_) Eggs of Tabanus. Photograph by J. T. Lloyd.] + +[Illustration: 80. (_b_) Larva of Tabanus. Photograph by M. V. S.] + +The horse-flies have been definitely convicted of transferring the +trypanosome of surra from diseased to healthy animals and there is good +evidence that they transfer anthrax. The possibility of their being +important agents in the conveyal of human diseases should not be +overlooked. Indeed, Leiper has recently determined that a species of +_Chrysops_ transfers the blood parasite _Filaria diurna_. + + +Leptidæ or Snipe-Flies + +The family LEPTIDÆ is made up of moderate or large sized flies, +predaceous in habit. They are sufficiently characterized in the keys of +Chapter XII. Four blood-sucking species belonging to three genera have +been reported. Of these _Symphoromyia pachyceras_ is a western species. +Dr. J. C. Bradley, from personal experience, reports it as a vicious +biter. + +[Illustration: 80. (_c_) Mouth parts of Tabanus. After Grünberg.] + + +Oestridæ or Bot-flies + +To the family OESTRIDÆ belong the bot and warble-flies so frequently +injurious to animals. The adults are large, or of medium size, heavy +bodied, rather hairy, and usually resemble bees in appearance. + +The larvæ live parasitically in various parts of the body of mammals, +such as the stomach (horse bot-fly), the subcutaneous connective tissue +(warble-fly of cattle), or the nasal passage (sheep bot-fly or head +maggot). + +There are on record many cases of the occurrence of the larvæ of +Oestridæ as occasional parasites of man. A number of these have been +collected and reviewed in a thesis by Mme. Pètrovskaia (1910). The +majority of them relate to the following species. + +_Gastrophilus hæmorrhoidalis_, the red tailed bot-fly, is one of the +species whose larvæ are most commonly found in the stomach of the horse. +Schoch (1877) cites the case of a woman who suffered from a severe case +of chronic catarrh of the stomach, and who vomited, and also passed from +the anus, larvæ which apparently belonged to this species. Such cases +are exceedingly rare but instances of subcutaneous infestation are +fairly numerous. In the latter type these larvæ are sometimes the cause +of the peculiar "creeping myasis." This is characterized at its +beginning by a very painful swelling which gradually migrates, producing +a narrow raised line four to twenty-five millimeters broad. When the +larva is mature, sometimes after several months, it becomes stationary +and a tumor is formed which opens and discharges the larva along with +pus and serum. + +_Gastrophilus equi_ is the most widespread and common of the horse +bot-flies. Portschinsky reports it as commonly causing subcutaneous +myasis of man in Russia. + +_Hypoderma bovis_ (= _Oestrus bovis_), and _Hypoderma lineata_ are the +so-called warble-flies of cattle. The latter species is the more common +in North America but Dr. C. G. Hewitt has recently shown that _H. bovis_ +also occurs. Though warbles are very common in cattle in this country, +the adult flies are very rarely seen. They are about half an inch in +length, very hairy, dark, and closely resemble common honey-bees. + +They deposit their eggs on the hairs of cattle and the animals in +licking themselves take in the young larvæ. These pass out through the +walls of the Å“sophagus and migrate through the tissues of the animal, +to finally settle down in the subcutaneous tissue of the back. The +possibility of their entering directly through the skin, especially in +case of infestation of man, is not absolutely precluded, although it is +doubtful. + +For both species of _Hypoderma_ there are numerous cases on record of +their occurrence in man. Hamilton (1893) saw a boy, six years of age, +who had been suffering for some months from the glands on one side of +his neck being swollen and from a fetid ulceration around the back teeth +of the lower jaw of the same side. Three months' treatment was of no +avail and the end seemed near; one day a white object, which was seen to +move, was observed in the ulcer at the root of the tongue, and on being +extracted was recognized as a full grown larva of _Hypoderma_. It was of +usual tawny color, about half an inch long when contracted, about one +third that thickness, and quite lively. The case resulted fatally. The +boy had been on a dairy farm the previous fall, where probably the egg +(or larva) was in some way taken into his mouth, and the larva found +between the base of the tongue and the jaw suitable tissue in which to +develop. + +Topsent (1901) reports a case of "creeping myasis" caused by _H. +lineata_ in the skin of the neck and shoulders of a girl eight years of +age. The larva travelled a distance of nearly six and a half inches. The +little patient suffered excruciating pain in the place occupied by the +larva. + +_Hypoderma diana_ infests deer, and has been known to occur in man. + +_Oestris ovis_, the sheep bot-fly, or head maggot, is widely distributed +in all parts of the world. In mid-summer the flies deposit living +maggots in the nostrils of sheep. These larvæ promptly pass up the nasal +passages into the frontal and maxillary sinuses, where they feed on the +mucous to be found there. In their migrations they cause great +irritation to their host, and when present in numbers may cause vertigo, +paroxysms, and even death. Portschinsky in an important monograph on +this species, has discussed in detail its relation to man. He shows that +it is not uncommon for the fly to attack man and that the minute living +larvæ are deposited in the eyes, nostrils, lips, or mouth. A typical +case in which the larvæ were deposited in the eye was described by a +German oculist Kayser, in 1905. A woman brought her six year old +daughter to him and said that the day before, about noontime, a flying +insect struck the eye of the child and that since then she had felt a +pain which increased towards evening. In the morning the pain ceased but +the eye was very red. She was examined at about noon, at which time she +was quiet and felt no pain. She was not sensitive to light, and the only +thing noticed was a slight congestion and accumulation of secretion in +the corner of the right eye. A careful examination of the eye disclosed +small, active, white larvæ that crawled out from the folds of the +conjunctiva and then back and disappeared. Five of these larvæ were +removed and although an uncomfortable feeling persisted for a while, the +eye became normal in about three weeks. + +Some of the other recorded cases have not resulted so favorably, for the +eyesight has been seriously affected or even lost. + +According to Edmund and Etienne Sergent (1907), myasis caused by the +larvæ of _Oestris ovis_ is very common among the shepherds in Algeria. +The natives say that the fly deposits its larvæ quickly, while on the +wing, without pause. The greatest pain is caused when these larvæ +establish themselves in the nasal cavities. They then produce severe +frontal headaches, making sleep impossible. This is accompanied by +continuous secretion from the nasal cavities and itching pains in the +sinuses. If the larvæ happen to get into the mouth, the throat becomes +inflamed, swallowing is painful, and sometimes vomiting results. The +diseased condition may last for from three to ten days or in the case of +nasal infection, longer, but recovery always follows. The natives remove +the larvæ from the eye mechanically by means of a small rag. When the +nose is infested, tobacco fumigations are applied, and in case of throat +infestation gargles of pepper, onion, or garlic extracts are used. + +_RhinÅ“strus nasalis_, the Russian gad-fly, parasitizes the +nasopharyngeal region of the horse. According to Portschinsky, it not +infrequently attacks man and then, in all the known cases deposits its +larvæ in the eye, only. This is generally done while the person is +quiet, but not during sleep. The fly strikes without stopping and +deposits its larva instantaneously. Immediately after, the victim +experiences lancinating pains which without intermission increase in +violence. There is an intense conjunctivitis and if the larvæ are not +removed promptly the envelopes of the eye are gradually destroyed and +the organ lost. + +[Illustration: 81. Larvæ of Dermatobia cyaniventris. After Blanchard.] + +[Illustration: 82. Young larva of Dermatobia cyaniventris. After +Surcouf.] + +_Dermatobia cyaniventris_--This fly (fig. 83) is widely distributed +throughout tropical America, and in its larval stage is well known as a +parasite of man. The larvæ (figs. 81 and 82) which are known as the "ver +macaque," "torcel," "ver moyocuil" or by several other local names, +enter the skin and give rise to a boil-like swelling, open at the top, +and comparable with the swelling produced by the warble fly larvæ, in +cattle. They cause itching and occasional excruciating pain. When +mature, nearly an inch in length, they voluntarily leave their host, +drop to the ground and complete their development. The adult female is +about 12 mm. in length. The face is yellow, the frons black with a +grayish bloom; antennæ yellow, the third segment four times as long as +the second, the arista pectinate. The thorax is bluish black with +grayish bloom; the abdomen depressed, brilliant metallescent blue with +violet tinge. The legs are yellowish, the squamæ and wings brownish. + +The different types of larvæ represented in figure 81 were formerly +supposed to belong to different species but Blanchard regards them as +merely various stages of the same species. It is only very recently that +the early stage and the method by which man becomes infested were made +known. + +[Illustration: 83. Dermatobia cyaniventris (×1¾). After +Manson.] + +[Illustration: 84. Mosquito carrying eggs of Dermatobia cyaniventris. +After Surcouf.] + +About 1900, Blanchard observed the presence of packets of large-sized +eggs under the abdomen of certain mosquitoes from Central America; and +in 1910, Dr. Moralès, of Costa Rica, declared that the Dermatobia +deposited its eggs directly under the abdomen of the mosquito and that +they were thus carried to vertebrates. Dr. Nunez Tovar observed the +mosquito carriers of the eggs and placing larvæ from this source on +animals, produced typical tumors and reared the adult flies. It remained +for Surcouf (1913) to work out the full details. He found that the +Dermatobia deposits its eggs in packets covered by a very viscid +substance, on leaves. These become attached to mosquitoes of the species +_Janthinosoma lutzi_ (fig. 84) which walk over the leaves. The eggs +which adhere to the abdomen, remain attached and are thus transported. +The embryo develops, but the young larva (fig. 82) remains in the egg +until it has opportunity to drop upon a vertebrate fed upon by the +mosquito. + + +Muscidæ + +The following MUSCIDÆ, characterized elsewhere, deserve special mention +under our present grouping of parasitic species. Other important species +will be considered as facultative parasites. + +_Stomoxys calcitrans_, the stable-fly, or the biting house-fly, is often +confused with _Musca domestica_ and therefore is discussed especially in +our consideration of the latter species as an accidental carrier of +disease. Its possible relation to the spread of infantile paralysis is +also considered later. + +The _tsetse flies_, belonging to the genus _Glossina_, are African +species of blood-sucking Muscidæ which have attracted much attention +because of their rôle in transmitting various trypanosome diseases of +man and animals. They are characterized in Chapter XII and are also +discussed in connection with the diseases which they convey. + +_Chrysomyia macellaria_, (= _Compsomyia_), the "screw worm"-fly is one +of the most important species of flies directly affecting man, in North +America. It is not normally parasitic, however, and hence will be +considered with other facultative parasites in Chapter IV. + +[Illustration: 85. Larva of Auchmeromyia luteola. After Graham-Smith.] + +[Illustration: 86. Auchmeromyia luteola (×4). After Graham-Smith.] + +_Auchmeromyia luteola_, the Congo floor maggot. This is a muscid of +grewsome habits, which has a wide distribution throughout Africa. The +fly (fig. 86) deposits its eggs on the ground of the huts of the +natives. The whitish larvæ (fig. 85) on hatching are slightly flattened +ventrally, and each segment bears posteriorly three foot-pads +transversely arranged. At night the larvæ find their way into the low +beds or couches of the natives and suck their blood. The adult flies do +not bite man and, as far as known, the larvæ do not play any rôle in the +transmission of sleeping sickness or other diseases. + +This habit of blood-sucking by muscid larvæ is usually referred to as +peculiar to _Auchmeromyia luteola_ but it should be noted that the larvæ +of _Protocalliphora_ frequent the nests of birds and feed upon the +young. Mr. A. F. Coutant has studied especially the life-history and +habits of _P. azurea_, whose larvæ he found attacking young crows at +Ithaca, N.Y. He was unable to induce the larvæ to feed on man. + +[Illustration: 87. Cordylobia anthropophaga (×3). After Fülleborn.] + +[Illustration: 88. Larva of Cordylobia anthropophaga. After Blanchard.] + +_Cordylobia anthropophaga_, (_Ochromyia anthropophaga_), or Tumbu-fly +(fig. 87) is an African species whose larvæ affect man much as do those +of _Dermatobia cyniventris_, of Central and South America. The larva +(fig. 88), which is known as "ver du Cayor" because it was first +observed in Cayor, in Senegambia, develops in the skin of man and of +various animals, such as dogs, cats, and monkeys. It is about 12 mm. in +length, and of the form of the larvæ of other muscids. Upon the +intermediate segments are minute, brownish recurved spines which give to +the larva its characteristic appearance. The life-history is not +satisfactorily worked out, but Fuller (1914), after reviewing the +evidence believes that, as a rule, it deposits its young in the sleeping +places of man and animals, whether such be a bed, a board, the floor, or +the bare ground. In the case of babies, the maggots may be deposited on +the scalp. The minute maggots bore their way painlessly into the skin. +As many as forty parasites have been found in one individual and one +author has reported finding more than three hundred in a spaniel puppy. +Though their attacks are at times extremely painful, it is seldom that +any serious results follow. + + +THE SIPHONAPTERA OR FLEAS + +The SIPHONAPTERA, or fleas (fig. 89) are wingless insects, with highly +chitinized and laterally compressed bodies. The mouth-parts are formed +for piercing and sucking. Compound eyes are lacking but some species +possess ocelli. The metamorphosis is complete. + +This group of parasites, concerning which little was known until +recently, has assumed a very great importance since it was learned that +fleas are the carriers of bubonic plague. Now over four hundred species +are known. Of these, several species commonly attack man. The most +common hominoxious species are _Pulex irritans_, _Xenopsylla cheopis_, +_Ctenocephalus canis_, _Ctenocephalus felis_, _Ceratophyllus fasciatus_ +and _Dermatophilus penetrans_, but many others will feed readily on +human blood if occasion arises. + +[Illustration: 89. Xenopsylla cheopis, male (×25). After Jordan and +Rothschild.] + +We shall treat in this place of the general biology and habits of the +hominoxious forms and reserve for the systematic section the discussion +of the characteristics of the different genera. + +The most common fleas infesting houses in the Eastern United States are +the cosmopolitan dog and cat fleas, _Ctenocephalus canis_ (fig. 90) and +_C. felis_. Their life cycles will serve as typical. These two species +have until recently been considered as one, under the name _Pulex +serraticeps_. See figure 92. + +[Illustration: 90. Dog flea (×15). After Howard.] + +The eggs are oval, slightly translucent or pearly white, and measure +about .5 mm. in their long diameter. They are deposited loosely in the +hairs of the host and readily drop off as the animal moves around. +Howard found that these eggs hatch in one to two days. The larvæ are +elongate, legless, white, worm-like creatures. They are exceedingly +active, and avoid the light in every way possible. They cast their first +skin in from three to seven days and their second in from three to four +days. They commenced spinning in from seven to fourteen days after +hatching and the imago appeared five days later. Thus in summer, at +Washington, the entire life cycle may be completed in about two weeks. +(cf. fig. 91, 92). + +Strickland's (1914) studies on the biology of the rat flea, +_Ceratophyllus fasciatus_, have so important a general bearing that we +shall cite them in considerable detail. + +[Illustration: 91. Larva of Xenopsylla cheopis. After Bacot and +Ridewood.] + +He found, to begin with, that there is a marked inherent range in the +rate of development. Thus, of a batch of seventy-three eggs, all laid in +the same day and kept together under the same conditions, one hatched in +ten days; four in eleven days; twenty-five in twelve days; thirty-one in +thirteen days; ten in fourteen days; one in fifteen days; and one in +sixteen days. Within these limits the duration of the egg period seems +to depend mainly on the degree of humidity. The incubation period is +never abnormally prolonged as in the case of lice, (Warburton) and +varying conditions of temperature and humidity have practically no +effect on the percentage of eggs which ultimately hatch. + +The same investigator found that the most favorable condition for the +larva is a low temperature, combined with a high degree of humidity; and +that the presence of rubbish in which the larva may bury itself is +essential to its successful development. When larvæ are placed in a +bottle containing either wood-wool soiled by excrement, or with feathers +or filter paper covered with dried blood they will thrive readily and +pupate. They seem to have no choice between dried blood and powdered rat +feces for food, and also feed readily on flea excrement. They possess +the curious habit of always devouring their molted skins. + +[Illustration: 92. Head and pronotum of (_a_) dog flea; (_b_) of cat +flea; (_c_) of hen flea. After Rothschild. (_d_) Nycteridiphilus +(Ischnopsyllus) hexactenus. After Oudemans.] + +An important part of Strickland's experiments dealt with the question of +duration of the pupal stage under the influence of temperature and with +the longevity and habits of the adult. In October, he placed a batch of +freshly formed cocoons in a small dish that was kept near a white rat in +a deep glass jar in the laboratory. Two months later one small and +feeble flea had emerged, but no more until February, four months after +the beginning of the experiment. Eight cocoons were then dissected and +seven more found to contain the imago fully formed but in a resting +state. The remainder of the batch was then placed at 70° F. for one +night, near a white rat. The next day all the cocoons were empty and the +fleas were found on the white rat. + +Thus, temperature greatly influences the duration of the pupal period, +which in _Ceratophyllus fasciatus_ averages seventeen days. Moreover, +when metamorphosis is complete a low temperature will cause the imago to +remain within the cocoon. + +Sexually mature and ovipositing fleas, he fed at intervals and kept +alive for two months, when the experiment was discontinued. In the +presence of rubbish in which they could bury themselves, unfed rat fleas +were kept alive for many months, whereas in the absence of any such +substratum they rarely lived a month. In the former case, it was found +that the length of life is influenced to some degree by the temperature +and humidity. In an experiment carried out at 70° F. and 45 per cent +humidity, the fleas did not live for more than four months, while in an +experiment at 60° F. and 70 per cent humidity they lived for at least +seventeen months. There was no indication that fleas kept under these +conditions sucked moisture from surrounding objects, and those kept in +bell jars, with an extract of flea-rubbish on filter paper, did not live +any longer than those which were not so supplied. + +Curiously enough, although the rat is the normal host of _Ceratophyllus +fasciatus_, it was found that when given the choice these fleas would +feed upon man in preference to rats. However, none of the fleas laid +eggs unless they fed on rat blood. + +The experiments of Strickland on copulation and oviposition in the rat +flea showed that fleas do not copulate until they are sexually mature +and that, at least in the case of _Ceratophyllus fasciatus_, the +reproductive organs are imperfectly developed for some time (more than a +week) after emerging from the pupa. When mature, copulation takes place +soon after the fleas have fed on their true host--the rat--but not if +they have fed on a facultative host only, such as man. Copulation is +always followed by oviposition within a very short time. + +The effect of the rat's blood on the female with regard to egg-laying, +Strickland concludes, is stimulating rather than nutritive, as fleas +that were without food for many months were observed to lay eggs +immediately after one feed. Similarly, the male requires the stimulus of +a meal of rat's blood before it displays any copulatory activity. + +Mitzmain (1910) has described in detail the act of biting on man, as +observed in the squirrel flea, _Ceratophyllus acutus_. "The flea when +permitted to walk freely on the arm selects a suitable hairy space where +it ceases abruptly in its locomotion, takes a firm hold with the tarsi, +projects its proboscis, and prepares to puncture the skin. A puncture is +drilled by the pricking epipharynx, the saw-tooth mandibles +supplementing the movement by lacerating the cavity formed. The two +organs of the rostrum work alternately, the middle piece boring, while +the two lateral elements execute a sawing movement. The mandibles, owing +to their basal attachments, are, as is expressed by the advisory +committee on plague investigations in India (_Journal of Hygiene_, vol. +6, No. 4, p. 499), 'capable of independent action, sliding up and down +but maintaining their relative positions and preserving the lumen of the +aspiratory channel.' The labium doubles back, the V-shaped groove of +this organ guiding the mandibles on either side." + +"The action of the proboscis is executed with a forward movement of the +head and a lateral and downward thrust of the entire body. As the +mouth-parts are sharply inserted, the abdomen rises simultaneously. The +hind and middle legs are elevated, resembling oars. The forelegs are +doubled under the thorax, the tibia and tarsi resting firmly on the +epidermis serve as a support for the body during the feeding. The +maxillary palpi are retracted beneath the head and thorax. The labium +continues to bend, at first acting as a sheath for the sawing mandibles, +and as these are more deeply inserted, it bends beneath the head with +the elasticity of a bow, forcing the mandibles into the wound until the +maxillæ are embedded in the skin of the victim. When the proboscis is +fully inserted, the abdomen ceases for a time its lateral swinging." + +"The acute pain of biting is first felt when the mandibles have not +quite penetrated and subsequently during each distinct movement of the +abdomen. The swinging of the abdomen gradually ceases as it becomes +filled with blood. The sting of the biting gradually becomes duller and +less sensitive as feeding progresses. The movements of the elevated +abdomen grow noticeably feebler as the downward thrusts of the springy +bow-like labium becomes less frequent." + +"As the feeding process advances one can discern through the translucent +walls of the abdomen a constant flow of blood, caudally from the +pharynx, accompanied by a peristaltic movement. The end of the meal is +signified in an abrupt manner. The flea shakes its entire body, and +gradually withdraws its proboscis by lowering the abdomen and legs and +violently twisting the head." + +"When starved for several days the feeding of the rat fleas is conducted +in a rather vigorous manner. As soon as the proboscis is buried to the +full length the abdomen is raised and there ensues a gradual lateral +swaying motion, increasing the altitude of the raised end of the abdomen +until it assumes the perpendicular. The flea is observed at this point +to gain a better foothold by advancing the fore tarsi, and then, +gradually doubling back the abdomen, it turns with extreme agility, +nearly touching with its dorsal side the skin of the hand upon which it +is feeding. Meanwhile, the hungry parasite feeds ravenously." + +"It is interesting to note the peculiar nervous action which the rodent +fleas exhibit immediately when the feeding process is completed or when +disturbed during the biting. Even while the rostrum is inserted to the +fullest the parasite shakes its head spasmodically; in a twinkling the +mouth is withdrawn and then the flea hops away." + +A habit of fleas which we shall see is of significance in considering +their agency in the spread of bubonic plague, is that of ejecting blood +from the anus as they feed. + +Fleas are famous for their jumping powers, and in control measures it is +of importance to determine their ability along this line. It is often +stated that they can jump about four inches, or, according to the Indian +Plague Commission _Xenopsylla cheopis_ cannot hop farther than five +inches. Mitzmain (1910) conducted some careful experiments in which he +found that the human flea, _Pulex irritans_, was able to jump as far as +thirteen inches on a horizontal plane. The mean average of five +specimens permitted to jump at will was seven and three-tenths inches. +The same species was observed to jump perpendicularly to a height of at +least seven and three-fourths inches. Other species were not able to +equal this record. + +The effect of the bite of fleas on man varies considerably according to +the individual susceptibility. According to Patton and Cragg, this was +borne out in a curious manner by the experiments of Chick and Martin. +"In these, eight human hosts were tried; in seven, little or no +irritation was produced, while in one quite severe inflammation was set +up around each bite." Of two individuals, equally accustomed to the +insects, going into an infested room, one may be literally tormented by +them while the other will not notice them. Indeed it is not altogether +a question of susceptibility, for fleas seem to have a special +predilection for certain individuals. The typical itching wheals +produced by the bites are sometimes followed, especially after +scratching, by inflammatory papules. + +The itching can be relieved by the use of lotions of carbolic acid (2-3 +per cent), camphor, menthol lotion, or carbolated vaseline. If forced to +sleep in an infested room, protection from attacks can be in a large +measure gained by sprinkling pyrethrum, bubach, or California insect +powder between the sheets. The use of camphor, menthol, or oil of +eucalyptus, or oil of pennyroyal is also said to afford protection to a +certain extent. + +In the Eastern United States the occurrence of fleas as household pests +is usually due to infested cats and dogs which have the run of the +house. We have seen that the eggs are not attached to the host but drop +to the floor when they are laid. Verrill, cited by Osborn, states that +on one occasion he was able to collect fully a teaspoonful of eggs from +the dress of a lady in whose lap a half-grown kitten had been held for a +short time. Patton and Cragg record seeing the inside of a hat in which +a kitten had spent the night, so covered with flea eggs that it looked +"as if it had been sprinkled with sugar from a sifter." It is no wonder +that houses in which pets live become overrun with the fleas. + +One of the first control measures, then, consists in keeping such +animals out of the house or in rigorously keeping them free from fleas. +The latter can best be accomplished by the use of strong tar soap or +Armour's "Flesope," which may be obtained from most druggists. The use +of a three per cent solution of creolin, approximately four teaspoonfuls +to a quart of warm water, has also been recommended. While this is +satisfactory in the case of dogs, it is liable to sicken cats, who will +lick their fur in an effort to dry themselves. Howard recommends +thoroughly rubbing into the fur a quantity of pyrethrum powder. This +partially stupifies the fleas which should be promptly swept up and +burned. + +He also recommends providing a rug for the dog or cat to sleep on and +giving this rug a frequent shaking and brushing, afterwards sweeping up +and burning the dust thus removed. + +Since the larvæ of fleas are very susceptible to exposure, the use of +bare floors, with few rugs, instead of carpets or matting, is to be +recommended. Thorough sweeping, so as to allow no accumulation of dust +in cracks and crevices will prove efficient. If a house is once +infested it may be necessary to thoroughly scrub the floors with hot +soapsuds, or to spray them with gasoline. If the latter method is +adopted, care must be taken to avoid the possibility of fire. + +To clear a house of fleas Skinner recommends the use of flake +naphthalene. In a badly infested house he took one room at a time, +scattering on the floor five pounds of flake naphthalene, and closed it +for twenty-four hours. It proved to be a perfect and effectual remedy +and very inexpensive, as the naphthalene could be swept up and +transferred to other rooms. Dr. Skinner adds, "so far as I am concerned, +the flea question is solved and if I have further trouble I know the +remedy. I intend to keep the dog and cat." + +The late Professor Slingerland very effectively used hydrocyanic acid +gas fumigation in exterminating fleas in houses. In one case, where +failure was reported, he found on investigation that the house had +become thoroughly reinfested from pet cats, which had been left +untreated. Fumigation with sulphur is likewise efficient. + +The fact that adult fleas are usually to be found on the floor, when not +on their hosts, was ingeniously taken advantage of by Professor S. H. +Gage in ridding an animal room at Cornell University of the pests. He +swathed the legs of a janitor with sticky fly-paper and had him walk +back and forth in the room. Large numbers of the fleas were collected in +this manner. + +In some parts of the southern United States hogs are commonly infested +and in turn infest sheds, barns and even houses. Mr. H. E. Vick informs +us that it is a common practice to turn sheep into barn-lots and sheds +in the spring of the year to collect in their wool, the fleas which +abound in these places after the hogs have been turned out. + +It is a common belief that adult fleas are attracted to fresh meat and +that advantage of this can be taken in trapping them. Various workers, +notably Mitzman (1910), have shown that there is no basis for such a +belief. + + +THE TRUE CHIGGERS--The chigoes, or true chiggers, are the most +completely parasitic of any of the fleas. Of the dozen or more known +species, one commonly attacks man. This is _Dermatophilus penetrans_, +more commonly known as _Sarcopsylla penetrans_ or _Pulex penetrans_. + +This species occurs in Mexico, the West Indies, Central and South +America. There are no authentic records of its occurrence in the United +States although, as Baker has pointed out, there is no reason why it +should not become established in Florida and Texas. It is usually +believed that Brazil was its original home. Sometime about the middle of +the nineteenth century it was introduced into West Africa and has spread +across that continent. + +The males and the immature females of _Dermatophilus penetrans_ (fig. +93) closely resemble those of other fleas. They are very active little +brown insects about 1-1.2 mm. in size, which live in the dust of native +huts and stables, and in dry, sandy soil. In such places they often +occur in enormous numbers and become a veritable plague. + +[Illustration: 93. Dermatophilus penetrans. Much enlarged. After +Karsten.] + +They attack not only man but various animals. According to Castellani +and Chalmers, "Perhaps the most noted feature is the way in which it +attacks pigs. On the Gold Coast it appeared to be largely kept in +existence by these animals. It is very easily captured in the free state +by taking a little pig with a pale abdomen, and placing it on its back +on the ground on which infected pigs are living. After watching a few +moments, a black speck will appear on the pig's abdomen, and quickly +another and another. These black specks are jiggers which can easily be +transferred to a test tube. On examination they will be found to be +males and females in about equal numbers." + +Both the males and females suck blood. That which characterizes this +species as distinguished from other fleas attacking man is that when the +impregnated female attacks she burrows into the skin and there swells +until in a few days she has the size and appearance of a small pea (fig. +94). Where they are abundant, hundreds of the pests may attack a single +individual (fig. 95). Here they lie with the apex of the abdomen +blocking the opening. According to Fülleborn (1908) they do not +penetrate beneath the epidermis. The eggs are not laid in the flesh of +the victim, as is sometimes stated, but are expelled through this +opening. The female then dies, withers and falls away or is expelled by +ulceration. According to Brumpt, she first quits the skin and then, +falling to the ground, deposits her eggs. The subsequent development in +so far as known, is like that of other fleas. + +[Illustration: 94. Dermatophilus penetrans, gravid female. After +Moniez.] + +[Illustration: 95. Chiggers in the sole of foot of man. Manson's +Tropical Diseases. Permission of Cassell and Co.] + +The chigoe usually enters between the toes, the skin about the roots of +the nails, or the soles of the feet, although it may attack other parts +of the body. Mense records the occurrence in folds of the epidermis, as +in the neighborhood of the anus. They give rise to irritation and unless +promptly and aseptically removed there often occurs pus formation and +the development of a more or less serious abscess. Gangrene and even +tetanus may ensue. + +Treatment consists in the careful removal of the insect, an operation +more easily accomplished a day or two after its entrance, than at first, +when it is unswollen. The ulcerated point should then be treated with +weak carbolic acid, or tincture of iodine, or dusted thoroughly with an +antiseptic powder. + +[Illustration: 96. Echidnophaga gallinacea.] + +[Illustration: 97. Echidnophaga gallinacea infesting head of chicken. +After Enderlein.] + +Castellani and Chalmers recommend as prophylactic measures, keeping the +house clean and keeping pigs, poultry, and cattle away therefrom. "High +boots should be used, and especial care should be taken not to go to a +ground floor bathroom with bare feet. The feet, especially the toes, and +under the nails, should be carefully examined every morning to see if +any black dots can be discovered, when the jigger should be at once +removed, and in this way suppuration will be prevented. It is +advisable, also, to sprinkle the floors with carbolic lotion, Jeyes' +fluid, or with pyrethrum powder, or with a strong infusion of native +tobacco, as recommended by Law and Castellani." + +_Echidnophaga gallinacea_ (fig. 96) is a widely distributed Hectopsyllid +attacking poultry (fig. 97). It occurs in the Southern and Southwestern +United States and has been occasionally reported as attacking man, +especially children. It is less highly specialized than _Dermatophilus +penetrans_, and does not ordinarily cause serious trouble in man. + + + + +CHAPTER IV + +ACCIDENTAL OR FACULTATIVE PARASITES + + +In addition to the many species of Arthropods which are normally +parasitic on man and animals, there is a considerable number of those +which may be classed as _accidental_ or _facultative_ parasites. + +Accidental or facultative parasites are species which are normally +free-living, but which are able to exist as parasites when accidentally +introduced into the body of man or other animal. A wide range of forms +is included under this grouping. + + +ACARINA + +A considerable number of mites have been reported as accidental or even +normal, endoparasites of man, but the authentic cases are comparatively +few. + +In considering such reports it is well to keep in mind von Siebold's +warning that in view of the universal distribution of mites one should +be on his guard. In vessels in which animal and other organic fluids and +moist substances gradually dry out, mites are very abundantly found. If +such vessels are used without very careful preliminary cleaning, for the +reception of evacuations of the sick, or for the reception of parts +removed from the body, such things may be readily contaminated by mites, +which have no other relation whatever to them. + +Nevertheless, there is no doubt but that certain mites, normally +free-living, have occurred as accidental parasites of man. Of these the +most commonly met with is _Tyroglyphus siro_, the cheese-mite. + +_Tyroglyphus siro_ is a small mite of a whitish color. The male measures +about 500µ long by 250µ wide, the female slightly larger. They live in +cheese of almost any kind, especially such as is a little decayed. "The +individuals gather together in winter in groups or heaps in the hollows +and chinks of the cheese and there remain motionless. As soon as the +temperature rises a little, they gnaw away at the cheese and reduce it +to a powder. The powder is composed of excrement having the appearance +of little grayish microscopic balls; eggs, old and new, cracked and +empty; larvæ, nymphs, and perfect mites, cast skins and fragments of +cheese, to which must be added numerous spores of microscopic +fungi."--Murray. + +_Tyroglyphus siro_, and related species, have been found many times in +human feces, under conditions which preclude the explanation that the +contamination occurred outside of the body. They have been supposed to +be the cause of dysentery, or diarrhÅ“a, and it is probable that the +_Acarus dysenteriæ_ of Linnæus, and Latreille, was this species. +However, there is little evidence that the mites cause any noteworthy +symptoms, even when taken into the body in large numbers. + +_Histiogaster spermaticus_ (fig. 152) is a Tyroglyphid mite which was +reported by Trouessart (1902) as having been found in a cyst in the +groin, adherent to the testis. When the cyst was punctured, it yielded +about two ounces of opalescent fluid containing spermatozoa and numerous +mites in all stages of development. The evidence indicated that a +fecundated female mite had been introduced into the urethra by means of +an unclean catheter. Though Trouessart reported the case as that of a +Sarcoptid, Banks places the genus _Histiogaster_ with the Tyroglyphidæ. +He states that our species feeds on the oyster-shell bark louse, +possibly only after the latter is dead, and that in England a species +feeds within decaying reeds. + +_Nephrophages sanguinarius_ is a peculiarly-shaped, angular mite which +was found by Miyake and Scriba (1893) for eight successive days in the +urine of a Japanese suffering from fibrinuria. Males, .117 mm. long by +.079 mm. wide, females .36 mm. by. 12 mm., and eggs were found both in +the spontaneously emitted urine and in that drawn by means of a +catheter. All the mites found were dead. The describers regarded this +mite as a true endoparasite, but it is more probable that it should be +classed as an accidental parasite. + + +MYRIAPODA + +There are on record a number of cases of myriapods occurring as +accidental parasites of man. The subject has been treated in detail by +Blanchard (1898 and 1902), who discussed forty cases. Since then at +least eight additions have been made to the list. + +Neveau-Lamaire (1908) lists thirteen species implicated, representing +eight different genera. Of the _Chilognatha_ there are three, _Julus +terrestris_, _J. londinensis_ and _Polydesmus complanatus_. The +remainder are _Chilopoda_, namely, _Lithobius forficatus_, _L. +malenops_, _Geophilus carpophagus_, _G. electricus_, _G. similis_, _G. +cephalicus_, _Scutigera coleoptrata_, _Himantarium gervaisi_, +_Chætechelyne vesuviana_ and _Stigmatogaster subterraneus_. + +The majority of the cases relate to infestation of the nasal fossæ, or +the frontal sinus, but intestinal infestation also occurs and there is +one recorded case of the presence of a species in _Julus_ (fig. 13) in +the auditory canal of a child. + +In the nose, the myriapods have been known to live for months and +according to some records, even for years. The symptoms caused by their +presence are inflammation, with or without increased flow of mucus, +itching, more or less intense headache, and at times general symptoms +such as vertigo, delirium, convulsions, and the like. These symptoms +disappear suddenly when the parasites are expelled. + +In the intestine of man, myriapods give rise to obscure symptoms +suggestive of infestation by parasitic worms. In a case reported by +Verdun and Bruyant (1912), a child twenty months of age had been +affected for fifteen days by digestive disturbances characterized by +loss of appetite, nausea and vomiting. The latter had been particularly +pronounced for three days, when there was discovered in the midst of the +material expelled a living myriapod of the species _Chætechelyne +vesuviana_. Anthelminthics had been administered without result. In some +of the other cases, the administration of such drugs had resulted in the +expulsion of the parasite through the anus. + +One of the extreme cases on record is that reported by Shipley (1914). +Specimens of _Geophilus gorizensis_ (= _G. subterraneus_) "were vomited +and passed by a woman of 68 years of age. Some of the centipedes emerged +through the patient's nose, and it must be mentioned that she was also +suffering from a round worm. One of her doctors was of the opinion that +the centipedes were certainly breeding inside the lady's intestines, and +as many as seven or eight, sometimes more, were daily leaving the +alimentary canal." + +"According to her attendant's statements those centipedes had left the +body in some hundreds during a period of twelve or eighteen months. +Their presence produced vomiting and some hæmatemesis, and treatment +with thymol, male-fern and turpentine had no effect in removing the +creatures." + +The clinical details, as supplied by Dr. Theodore Thompson were as +follows: + +"Examined by me July, 1912, her tongue was dry and glazed. There was +bleeding taking place from the nose and I saw a living centipede she had +just extracted from her nostril. Her heart, lungs and abdomen appeared +normal. She was not very wasted, and did not think she had lost much +flesh, nor was there any marked degree of anemia." + +Shipley gives the following reasons for believing it impossible that +these centipedes could have multiplied in the patient's intestine. "The +breeding habits of the genus _Geophilus_ are peculiar, and ill adapted +for reproducing in such a habitat. The male builds a small web or nest, +in which he places his sperm, and the female fertilizes herself from +this nest or web, and when the eggs are fertilized they are again laid +in a nest or web in which they incubate and in two or three weeks hatch +out. The young _Geophilus_ differ but very little from the adult, except +in size. It is just possible, but improbable, that a clutch of eggs had +been swallowed by the host when eating some vegetables or fruit, but +against this is the fact that the _Geophilus_ does not lay its eggs upon +vegetables or fruit, but upon dry wood or earth. The egg-shell is very +tough and if the eggs had been swallowed the egg-shells could certainly +have been detected if the dejecta were examined. The specimens of the +centipede showed very little signs of being digested, and it is almost +impossible to reconcile the story of the patient with what one knows of +the habits of the centipedes." + +In none of the observed cases have there been any clear indications as +to the manner of infestation. It is possible that the myriapods have +been taken up in uncooked fruit or vegetables. + + +LEPIDOPTEROUS LARVÆ + +SCHOLECIASIS--Hope (1837) brought together six records of infestation of +man by lepidopterous larvæ and proposed to apply the name scholeciasis +to this type of parasitism. The clearest case was that of a young boy +who had repeatedly eaten raw cabbage and who vomited larvæ of the +cabbage butterfly, _Pieris brassicæ_. Such cases are extremely rare, and +there are few reliable data relative to the subject. In this connection +it may be noted that Spuler (1906) has described a moth whose larvæ live +as ectoparasites of the sloth. + + +COLEOPTERA + +CANTHARIASIS--By this term Hope designated instances of accidental +parasitism by the larvæ or adults of beetles. Reports of such cases are +usually scouted by parasitologists but there seems no good basis for +wholly rejecting them. Cobbold refers to a half dozen cases of +accidental parasitism by the larvæ of _Blaps mortisaga_. In one of +these cases upwards of 1200 larvæ and several perfect insects were said +to have been passed _per annum_. French (1905) reports the case of a man +who for a considerable period voided adult living beetles of the species +_Nitidula bipustulata_. Most of the other cases on record relate to the +larvæ of _Dermestidæ_ (larder beetles _et al._) or _Tenebrionidæ_ (meal +infesting species). Infestation probably occurs through eating raw or +imperfectly cooked foods containing eggs or minute larvæ of these +insects. + +[Illustration: 98. Larva of Piophila casei. Caudal aspect of larva. +Posterior stigmata.] + +Brumpt cites a curious case of accidental parasitism by a coleopterous +larva belonging to the genus _Necrobia_. This larva was extracted from a +small tumor, several millimeters long, on the surface of the conjunctiva +of the eye. The larvæ of this genus ordinarily live in decomposing flesh +and cadavers. + + +DIPTEROUS LARVÆ + +[Illustration: 99. Piophila casei. After Graham-Smith.] + +MYASIS--By this term (spelled also myiasis, and myiosis), is meant +parasitism by dipterous larvæ. Such parasitism may be normal, as in the +cases already described under the heading parasitic Diptera, or it may +be facultative, due to free-living larvæ being accidentally introduced +into wounds or the body-cavities of man. Of this latter type, there is a +multitude of cases on record, relating to comparatively few species. The +literature of the subject, like that relating to facultative parasitism +in general, is unsatisfactory, for most of the determinations of species +have been very loose. Indeed, so little has been known regarding the +characteristics of the larvæ concerned that in many instances they could +not be exactly determined. Fortunately, several workers have undertaken +comparative studies along this line. The most comprehensive publication +is that of Banks (1912), entitled "The structure of certain dipterous +larvæ, with particular reference to those in human food." + +Without attempting an exhaustive list, we shall discuss here the more +important species of Diptera whose larvæ are known to cause myasis, +either external or internal. The following key will serve to determine +those most likely to be encountered. The writers would be glad to +examine specimens not readily identifiable, if accompanied by exact data +relative to occurrence. + +_a._ Body more or less flattened, depressed; broadest in the middle, + each segment with dorsal, lateral, and ventral fleshy processes, of + which the laterals, at least, are more or less spiniferous (fig. + 101). _Fannia_ (= _Homalomyia_). + + In _F. canicularis_ the dorsal processes are nearly as long as the + laterals; in _F. scalaris_ the dorsal processes are short spinose + tubercles. + +_aa._ Body cylindrical, or slender conical tapering toward the head; + without fleshy lateral processes (fig. 105). + +_b._ With the posterior stigmata at the end of shorter or longer + tubercles, or if not placed upon tubercles, then not in pit; usually + without a "marginal button" and without a chitinous ring surrounding + the three slits; the slits narrowly or broadly oval, not bent (fig. + 171 i). _Acalyptrate muscidæ_ and some species of _Anthomyiidæ_. To + this group belong the cheese skipper (_Piophila casei_, figs. 98, + 99), the pomace-fly (_Drosophila ampelophila_), the apple maggot + (_Rhagoletis pomonella_), the cherry fruit fly (_Rhagoletis + cingulata_), the small dung fly (_Sepsis violacea_, fig. 170), the + beet leaf-miner (_Pegomyia vicina_, fig. 171 i), the cabbage, bean + and onion maggots (_Phorbia_ spp.) et. al. + +_bb._ Posterior stigmata of various forms, if the slits are narrowly + oval (fig. 171) then they are surrounded by a chitin ring which may + be open ventro-mesally. + +_c._ Integument leathery and usually strongly spinulose; larvæ + hypodermatic or endoparasitic. Bot flies (fig. 171, f, g, + k).--_Oestridæ_ + +_cc._ Integument not leathery and (except in _Protocalliphora_) spinulæ + restricted to transverse patches near the incisures of the segments. + +_d._ The stigmal plates in a pit; the lip-like margin of the pit with a + number of fleshy tubercles; chitin ring of the stigma not complete; + open ventro-mesally, button absent (fig. 171 e). Flesh + flies.--_Sarcophaga_ + +_dd._ Stigmata not in a pit. + +_e._ The chitin ring open ventra-mesally; button absent (fig. 171 c). + Screw-worm fly. _Chrysomyia_ + +_ee._ The chitin ring closed. + +_f._ Slits of the posterior stigmata straight; marginal "button" present + (fig. 171 b); two distinct mouth hooks, fleshy tubercles around the + anal area. _Phormia_ (fig. 171 f), _Lucilia_ and _Calliphora_ (fig. + 172, a, b), _Protocalliphora_ (fig. 171, j), _Cynomyia_ (fig. 171, + a). Blow flies, bluebottle flies. _Calliphorinæ_ + +_ff._ Slits of the posterior stigmata sinuous or bent. Subfamily + Muscinæ. + +_g._ Slits of the posterior stigmata bent; usually two mouth hooks. + _Muscina stabulans_ (fig. 171, l), _Muscina similis_, _Myiospila + meditabunda_ (fig. 172, i), and some of the higher _Anthomyiidæ_. + +_gg._ Slits of the posterior stigmata sinuous; mouth hooks usually + consolidated into one. The house-fly (_Musca domestica_ fig. 171, + d), the stable fly (_Stomoxys calcitrans_), the horn fly (_Lyperosia + irritans_), _Pyrellia_, _Pseudopyrellia_, _Morellia_, _Mesembrina_. + _Polietes_, et. al. (fig. 172 in part). + +_Eristalis_--The larvæ of _Eristalis_ are the so-called rat-tailed +maggots, which develop in foul water. In a few instances these larvæ +have been known to pass through the human alimentary canal uninjured. +Hall and Muir (1913) report the case of a boy five years of age, who had +been ailing for ten weeks and who was under treatment for indigestion +and chronic constipation. For some time he had vomited everything he +ate. On administration of a vermifuge he voided one of the rat-tailed +maggots of _Eristalis_. He admitted having drunk water from a ditch full +of all manner of rotting matter. It was doubtless through this that he +became infested. It is worth noting that the above described symptoms +may have been due to other organisms or substances in the filthy water. + +_Piophila casei_, the cheese-fly (fig. 99), deposits its eggs not only +in old cheeses, but on ham, bacon, and other fats. The larvæ (fig. 98) +are the well-known cheese skippers, which sometimes occur in great +abundance on certain kinds of cheese. Indeed, some people have a +comfortable theory that such infested cheese is especially good. Such +being the case, it is small wonder that this species has been repeatedly +reported as causing intestinal myasis. Thebault (1901) describes the +case of a girl who, shortly after consuming a large piece of badly +infested cheese, became ill and experienced severe pains in the region +of the navel. Later these extended through the entire alimentary canal, +the excrement was mixed with blood and she suffered from vertigo and +severe headaches. During the four following days the girl felt no +change, although the excretion of the blood gradually diminished and +stopped. On the fourth day she voided two half-digested larvæ and, +later, seven or eight, of which two were alive and moving. + +That these symptoms may be directly attributed to the larvæ, or +"skippers," has been abundantly shown by experimental evidence. +Portschinsky cites the case of a dog fed on cheese containing the larvæ. +The animal suffered much pain and its excrement contained blood. On +_post mortem_ it was found that the small intestine throughout almost +its entire length was marked by bloody bruises. The papillæ on these +places were destroyed, although the walls were not entirely perforated. +In the appendix were found two or three dead larvæ. Alessandri (1910) +has likewise shown that the larvæ cause intestinal lesions. + +According to Graham-Smith, Austen (1912) has recorded a case of myasis +of the nose, attended with a profuse watery discharge of several weeks +duration and pain, due to the larvæ of _Piophila casei_. + + +ANTHYOMYIIDÆ--The characteristic larvæ of two species of _Fannia_ (= +_Homalomyia_ or _Anthomyia_, in part) (fig. 101) are the most commonly +reported of dipterous larvæ causing intestinal myasis. Hewitt (1912) has +presented a valuable study of the bionomics and of the larvæ of these +flies, a type of what is needed for all the species concerned in myasis. +We have seen two cases of their having been passed in stools, without +having caused any special symptoms. In other instances their presence in +the alimentary canal has given rise to symptoms vaguely described as +those of tapeworm infestation, or helminthiasis. More specifically, they +have been described as causing vertigo, severe headache, nausea and +vomiting, severe abdominal pains, and in some instances, bloody +diarrhÅ“a. + +[Illustration: 100. Fannia canicularis (×4). After Graham-Smith.] + +One of the most striking cases is that reported by Blankmeyer (1914), of +a woman whose illness began fourteen years previously with nausea and +vomiting. After several months of illness she began passing larvæ and +was compelled to resort to enemas. Three years previous to the report, +she noticed frequent shooting pains in the rectal region and at times +abdominal tenderness was marked. There was much mucus in the stools and +she "experienced the sensation of larvæ crawling in the intestine." +Occipital headaches were marked, with remissions, and constipation +became chronic. The appetite was variable, there was a bad taste in the +mouth, tongue furred and ridged, and red at the edges. Her complexion +was sallow, and general nervousness was marked. As treatment, there were +given doses of magnesium sulphate before breakfast and at 4 P. M., with +five grain doses of salol four times a day. The customary parasiticides +yielded no marked benefit. At the time of the report the patient passed +from four to fifty larvæ per day, and was showing some signs of +improvement. The nausea had disappeared, her nervousness was less +evident, and there was a slight gain in weight. + +The case was complicated by various other disorders, but the symptoms +given above seem to be in large part attributable to the myasis. There +is nothing in the case to justify the assumption that larvæ were +continuously present, for years. It seems more reasonable to suppose +that something in the habits of the patient favored repeated +infestation. Nevertheless, a study of the various cases of intestinal +myasis caused by these and other species of dipterous larvæ seems to +indicate that the normal life cycle may be considerably prolonged under +the unusual conditions. + +The best authenticated cases of myasis of the urinary passage have been +due to larvæ of _Fannia_. Chevril (1909) collected and described twenty +cases, of which seven seemed beyond doubt. One of these was that of a +woman of fifty-five who suffered from albuminuria, and urinated with +much difficulty, and finally passed thirty to forty larvæ of _Fannia +canicularis_. + +It is probable that infestation usually occurs through eating partially +decayed fruit or vegetables on which the flies have deposited their +eggs. Wellman points out that the flies may deposit their eggs in or +about the anus of persons using outside privies and Hewitt believes that +this latter method of infection is probably the common one in the case +of infants belonging to careless mothers. "Such infants are sometimes +left about in an exposed and not very clean condition, in consequence of +which flies are readily attracted to them and deposit their eggs." + +[Illustration: 101. Larva of Fannia scalaris.] + + +MUSCINÆ--The larvæ of the common house-fly, _Musca domestica_, are +occasionally recorded as having been passed with the feces or vomit of +man. While such cases may occur, it is probable that in most instances +similar appearing larvæ of other insects have been mistakenly +identified. + +_Muscina stabulans_ is regarded by Portschinsky (1913) as responsible +for many of the cases of intestinal myasis attributed to other species. +He records the case of a peasant who suffered from pains in the lower +part of the breast and intestines, and whose stools were mixed with +blood. From November until March he had felt particularly ill, being +troubled with nausea and vomiting in addition to the pain in his +intestines. In March, his physician prescribed injections of a +concentrated solution of tannin, which resulted in the expulsion of +fifty living larvæ of _Muscina stabulans_. Thereafter the patient felt +much better, although he suffered from intestinal catarrh in a less +severe form. + +[Illustration: 102 Muscina stabulans (×4). After Graham-Smith.] + + +CALLIPHORINÆ--Closely related to the Sarcophagidæ are the +_Calliphorinæ_, to which group belong many of the so-called "blue +bottle" flies. Their larvæ feed upon dead animals, and upon fresh and +cooked meat. Those of _Protocalliphora_, already mentioned, are +ectoparasitic on living nestling birds. Larva of _Lucilia_, we have +taken from tumors on living turtles. To this sub-family belongs also +_Aucheromyia luteola_, the Congo floor maggot. Some of these, and at +least the last mentioned, are confirmed, rather than faculative +parasites. Various species of Calliphorinæ are occasionally met with as +facultative parasites of man. + +[Illustration: 103. Lucilia cæsar, (×3). After Howard.] + +_Chrysomyia macellaria_, the screw worm fly (fig. 107), is the fly which +is responsible for the most serious cases of human myasis in the United +States. It is widely distributed in the United States but is especially +abundant in the south. While the larvæ breed in decaying matter in +general, they so commonly breed in the living flesh of animals that they +merit rank as true parasites. The females are attracted to open wounds +of all kinds on cattle and other animals and quickly deposit large +numbers of eggs. Animals which have been recently castrated, dehorned, +or branded, injured by barbed wire, or even by the attacks of ticks are +promptly attacked in the regions where the fly abounds. Even the navel +of young calves or discharges from the vulva of cows may attract the +insect. + +[Illustration: 104. Calliphora erythrocephala, (×6). After +Graham-Smith.] + +Not infrequently the fly attacks man, being attracted by an offensive +breath, a chronic catarrh, or a purulent discharge from the ears. Most +common are the cases where the eggs are deposited in the nostrils. The +larvæ, which are hatched in a day or two, are provided with strong +spines and proceed to bore into the tissues of the nose, even down into +or through the bone, into the frontal sinus, the pharynx, larynx, and +neighboring parts. + +Osborn (1896) quotes a number of detailed accounts of the attacks of the +_Chrysomyia_ on man. A vivid picture of the symptomology of rhinal +myasis caused by the larvæ of this fly is given by Castellani and +Chalmers: "Some couple of days after a person suffering from a chronic +catarrh, foul breath, or ozæna, has slept in the open or has been +attacked by a fly when riding or driving,--_i.e._, when the hands are +engaged--signs of severe catarrh appear, accompanied with inordinate +sneezing and severe pain over the root of the nose or the frontal bone. +Quickly the nose becomes swollen, and later the face also may swell, +while examination of the nose may show the presence of the larvæ. Left +untreated, the patient rapidly becomes worse, and pus and blood are +discharged from the nose, from which an offensive odor issues. Cough +appears as well as fever, and often some delirium. If the patient lives +long enough, the septum of the nose may fall in, the soft and hard +palates may be pierced, the wall of the pharynx may be destroyed. By +this time, however, the course of the disease will have become quite +evident by the larvæ dropping out of the nose, and if the patient +continues to live all the larvæ may come away naturally." + +For treatment of rhinal myasis these writers recommend douching the nose +with chloroform water or a solution of chloroform in sweet milk (10-20 +per cent), followed by douches of mild antiseptics. Surgical treatment +may be necessary. + +[Illustration: 105. Larva of a flesh fly (Sarcophaga). Caudal aspect. +Anterior stigmata. Pharyngeal skeleton.] + + +SARCOPHAGIDÆ--The larvæ (fig. 105) of flies of this family usually feed +upon meats, but have been found in cheese, oleomargerine, pickled +herring, dead and living insects, cow dung and human feces. Certain +species are parasitic in insects. Higgins (1890) reported an instance of +"hundreds" of larvæ of _Sarcophaga_ being vomited by a child eighteen +months of age. There was no doubt as to their origin for they were +voided while the physician was in the room. There are many other reports +of their occurrence in the alimentary canal. We have recorded elsewhere +(Riley, 1906) a case in which some ten or twelve larvæ of _Sarcophaga_ +were found feeding on the diseased tissues of a malignant tumor. The +tumor, a melanotic sarcoma, was about the size of a small walnut, and +located in the small of the back of an elderly lady. Although they had +irritated and caused a slight hæmorrhage, neither the patient nor others +of the family knew of their presence. Any discomfort which they had +caused had been attributed to the sarcomatous growth. The infestation +occurred in mid-summer. It is probable that the adult was attracted by +the odor of the discharges and deposited the living maggots upon the +diseased tissues. + +[Illustration: 106. A flesh fly (Sarcophaga), (×4). After Graham-Smith.] + +According to Küchenmeister, _Sarcophaga carnaria_ (fig. 106), attracted +by the odor, deposits its eggs and larvæ in the vagina of girls and +women when they lie naked in hot summer days upon dirty clothes, or when +they have a discharge from the vagina. In malignant inflammations of the +eyes the larvæ even nestle under the eyelids and in Egypt, for example, +produce a very serious addition to the effects of small-pox upon the +cornea, as according to Pruner, in such cases perforation of the cornea +usually takes place. + +[Illustration: 107. Chrysomyia macellaria, (×3).] + +_Wohlfartia magnifica_ is another Sarcophagid which commonly infests man +in the regions where it is abundant. It is found in all Europe but is +especially common in Russia, where Portschinsky has devoted much +attention to its ravages. It deposits living larvæ in wounds, the nasal +fossæ, the ears and the eyes, causing injuries even more revolting than +those described for _Chrysomyia_. + + + + +CHAPTER V + +ARTHROPODS AS SIMPLE CARRIERS OF DISEASE + + +The fact that certain arthropods are poisonous, or may affect the health +of man as direct parasites has always received attention in the medical +literature. We come now to the more modern aspect of our subject,--the +consideration of insects and other arthropods as transmitters and +disseminators of disease. + +The simplest way in which arthropods may function in this capacity is as +_simple carriers_ of pathogenic organisms. It is conceivable that any +insect which has access to, and comes in contact with such organisms and +then passes to the food, or drink, or to the body of man, may in a +wholly accidental and incidental manner convey infection. That this +occurs is abundantly proved by the work of recent years. We shall +consider as typical the case against the house-fly, which has attracted +so much attention, both popular and scientific. The excellent general +treatises of Hewitt (1910), Howard (1911), and Graham-Smith (1913), and +the flood of bulletins and popular literature render it unnecessary to +consider the topic in any great detail. + + +THE HOUSE-FLY AS A CARRIER OF DISEASE + +Up to the past decade the house-fly has usually been regarded as a mere +pest. Repeatedly, however, it had been suggested that it might +disseminate disease. We have seen that as far back as the sixteenth +century, Mercurialis suggested that it was the agent in the spread of +bubonic plague, and in 1658, Kircher reiterated this view. In 1871, +Leidy expressed the opinion that flies were probably a means of +communicating contagious diseases to a greater degree than was generally +suspected. From what he had observed regarding gangrene in hospitals, he +thought flies should be carefully excluded from wounds. In the same +year, the editor of the _London Lancet_, referring to the belief that +they play a useful rôle in purifying the air said, "Far from looking +upon them as dipterous angels dancing attendance on Hygeia, regard them +rather in the light of winged sponges spreading hither and thither to +carry out the foul behests of Contagion." + +These suggestions attracted little attention from medical men, for it is +only within very recent years that the charges have been supported by +direct evidence. Before considering this evidence, it is necessary that +we define what is meant by "house-fly" and that we then consider the +life-history of the insect. + +There are many flies which are occasionally to be found in houses, but +according to various counts, from 95 per cent to 99 per cent of these in +warm weather in the Eastern United States belong to the one species +_Musca domestica_ (fig. 108). This is the dominant house-fly the world +over and is the one which merits the name. It has been well +characterized by Schiner (1864), whose description has been freely +translated by Hewitt, as follows: + +"Frons of male occupying a fourth part of the breadth of the head. +Frontal stripe of female narrow in front, so broad behind that it +entirely fills up the width of the frons. The dorsal region of the +thorax dusty grey in color with four equally broad longitudinal stripes. +Scutellum gray with black sides. The light regions of the abdomen +yellowish, transparent, the darkest parts at least at the base of the +ventral side yellow. The last segment and a dorsal line blackish brown. +Seen from behind and against the light, the whole abdomen shimmering +yellow, and only on each side of the dorsal line on each segment a dull +transverse band. The lower part of the face silky yellow, shot with +blackish brown. Median stripe velvety black. Antennæ brown. Palpi black. +Legs blackish brown. Wings tinged with pale gray with yellowish base. +The female has a broad velvety back, often reddishly shimmering frontal +stripe, which is not broader at the anterior end than at the bases of +the antennæ, but become so very much broader above that the light +dustiness of the sides is entirely obliterated. The abdomen gradually +becoming darker. The shimmering areas on the separate segments generally +brownish. All the other parts are the same as in the male." + +The other species of flies found in houses in the Eastern United States +which are frequently mistaken for the house or typhoid fly may readily +be distinguished by the characters of the following key: + +_a._ Apical cell (R_5) of the wide wing open, i.e., the bounding veins + parallel or divergent (fig. 100). Their larvæ are flattened, the + intermediate body segments each fringed with fleshy, more or less + spinose, processes. _Fannia_ + + b. Male with the sides of the second and third abdominal segments + translucent yellowish. The larva with three pairs of nearly equal + spiniferous appendages on each segment, arranged in a + longitudinal series and in addition two pairs of series of smaller + processes (fig. 100) _F. canicularis_ + + bb. Male with blackish abdomen, middle tibia with a tubercle beyond + the middle. The larva with spiniferous appendages of which the + dorsal and ventral series are short, the lateral series long and + feathered (fig. 101) _F. scalaris_ + +aa. Apical cell (R) of the wing more or less narrowed in the margin; i. + e., the bounding veins more or less converging (fig. 108). + + b. The mouth-parts produced and pointed, fitted for piercing. + + c. Palpi much shorter than the proboscis; a brownish gray fly, its + thorax with three rather broad whitish stripes; on each border + of the middle stripe and on the mesal borders of the lateral + stripes is a blackish brown line. Abdomen yellowish brown; on + the second, third and fourth segments are three brown spots + which may be faint or even absent. The larvæ live in dung. The + stable-fly (fig. 110) _Stomoxys calcitrans_ + + cc. Palpi nearly as long as the proboscis. Smaller species than the + house-fly. The horn-fly (fig. 167) _Hæmatobia irritans_ + + bb. Mouth-parts blunt, fitted for lapping. + + c. Thorax, particularly on the sides and near the base of the wings + with soft golden yellow hairs among the bristles. This fly is + often found in the house in very early spring or even in the + winter. The cluster-fly, _Pollenia rudis_ + + cc. Thorax without golden yellow hairs among the bristles. + + d. The last segment of the vein M with an abrupt angle. (fig. + 108). The larvæ live in manure, etc. House-fly, _Musca + domestica_ + + dd. The last segment of vein M with a broad, gentle curve (fig. + 102). + + e. Eyes microscopically hairy; each abdominal segment with two + spots. Larvæ in dung. _Myiospila meditabunda_ + + ee. Eyes bare; abdomen gray and brown marbled. _Muscina_ + + f. With black legs and palpi. _M. assimilis_ + + ff. With legs more or less yellowish; palpi yellow. Larvæ in + decaying vegetable substances, dung, etc. _M. stabulans_ + +It is almost universally believed that the adults of _Musca domestica_ +hibernate, remaining dormant throughout the winter in attics, around +chimneys, and in sheltered but cold situations. This belief has been +challenged by Skinner (1913), who maintains that all the adult flies die +off during the fall and early winter and that the species is carried +over in the pupal stage, and in no other way. The cluster-fly, _Pollenia +rudis_, undoubtedly does hibernate in attics and similar situations and +is often mistaken for the house-fly. In so far as concerns _Musca +domestica_, the important question as to hibernation in the adult stage +is an open one. Many observations by one of the writers (Johannsen) tend +to confirm Dr. Skinner's conclusion, in so far as it applies to +conditions in the latitude of New York State. Opposed, is the fact that +various experimenters, notably Hewitt (1910) and Jepson (1909) wholly +failed to carry pupæ through the winter. + +[Illustration: 108. The house or typhoid fly (Musca domestica (×4)). +After Howard.] + +The house-fly breeds by preference in horse manure. Indeed, Dr. Howard, +whose extensive studies of the species especially qualify him for +expressing an opinion on the subject, has estimated that under ordinary +city and town conditions, more than ninety per cent of the flies present +in houses have come from horse stables or their vicinity. They are not +limited to such localities, by any means, for it has been found that +they would develop in almost any fermenting organic substance. Thus, +they have been bred from pig, chicken, and cow manure, dirty waste +paper, decaying vegetation, decaying meat, slaughter-house refuse, +sawdust-sweepings, and many other sources. A fact which makes them +especially dangerous as disease-carriers is that they breed readily in +human excrement. + +The eggs are pure white, elongate ovoid, somewhat broader at the +anterior end. They measure about one millimeter (1-25 inch) in length. +They are deposited in small, irregular clusters, one hundred and twenty +to one hundred and fifty from a single fly. A female may deposit as many +as four batches in her life time. The eggs hatch in from eight to +twenty-four hours. + +The newly hatched larva, or maggot (fig. 108), measures about two +millimeters (1-12 inch) in length. It is pointed at the head end and +blunt at the opposite end, where the spiracular openings are borne. It +grows rapidly, molts three times and reaches maturity in from six to +seven days, under favorable conditions. + +The pupal stage, like that of related flies, is passed in the old larval +skin which, instead of being molted, becomes contracted and heavily +chitinized, forming the so-called _puparium_ (fig. 108). The pupal stage +may be completed in from three to six days. + +Thus during the warm summer months a generation of flies may be produced +in ten to twelve days. Hewitt at Manchester, England, found the minimum +to be eight days but states that larvæ bred in the open air in horse +manure which had an average daily temperature of 22.5° C., occupied +fourteen to twenty days in their development, according to the air +temperature. + +After emergence, a period of time must elapse before the fly is capable +of depositing eggs. This period has been tuned the _preoviposition_ +period. Unfortunately we have few exact data regarding this period. +Hewitt found that the flies became sexually mature in ten to fourteen +days after their emergence from the pupal state and four days after +copulation they began to deposit their eggs; in other words the +preoviposition stage was fourteen days or longer. Griffith (1908) found +this period to be ten days. Dr. Howard believes that the time "must +surely be shorter, and perhaps much shorter, under midsummer conditions, +and in the freedom of the open air." He emphasizes that the point is of +great practical importance, since it is during this period that the +trapping and other methods of destroying the adult flies, will prove +most useful. + +Howard estimates that there may be nine generations of flies a year +under outdoor conditions in places comparable in climate to Washington. +The number may be considerably increased in warmer climates. + +The rate at which flies may increase under favorable conditions is +astounding. Various writers have given estimates of the numbers of flies +which may develop as the progeny of a single individual, providing all +the eggs and all the individual flies survived. Thus, Howard estimates +that from a single female, depositing one hundred and twenty eggs on +April 15th, there may be by September 10th, 5,598,720,000,000 adults. +Fortunately, living forms do not produce in any such mathematical manner +and the chief value of the figures is to illustrate the enormous +struggle for existence which is constantly taking place in nature. + +Flies may travel for a considerable distance to reach food and shelter, +though normally they pass to dwellings and other sources of food supply +in the immediate neighborhood of their breeding places. Copeman, Howlett +and Merriman (1911) marked flies by shaking them in a bag containing +colored chalk. Such flies were repeatedly recovered at distances of +eight to one thousand yards and even at a distance of seventeen hundred +yards, nearly a mile. + +Hindle and Merriman (1914) continued these experiments on a large scale +at Cambridge, England. They "do not think it likely that, as a rule, +flies travel more than a quarter of a mile in thickly-housed areas." In +one case a single fly was recovered at a distance of 770 yards but a +part of this distance was across open fen-land. The surprising fact was +brought out that flies tend to travel either _against_ or across the +wind. The actual direction followed may be determined either directly by +the action of the wind (positive anemotropism), or indirectly owing to +the flies being attracted by any odor that it may convey from a source +of food. They conclude that it is likely that the chief conditions +favoring the disposal of flies are fine weather and a warm temperature. +The nature of the locality is another considerable factor. Hodge (1913) +has shown that when aided by the wind they may fly to much greater +distances over the water. He reports that at Cleveland, Ohio, the cribs +of the water works, situated a mile and a quarter, five miles, and six +miles out in Lake Erie are invaded by a regular plague of flies when the +wind blows from the city. Investigation showed that there was absolutely +nothing of any kind in which flies could breed on the crib. + +The omnivorous habits of the house-fly are matters of everyday +observation. From our view point, it is sufficient to emphasize that +from feeding on excrement, on sputum, on open sores, or on putrifying +matter, the flies may pass to the food or milk upon the table or to +healthy mucous membranes, or uncontaminated wounds. There is nothing in +its appearance to tell whether the fly that comes blithely to sup with +you is merely unclean, or whether it has just finished feeding upon +dejecta teeming with typhoid bacilli. + +[Illustration: 100. Pulvillus of foot of house-fly, showing glandular +hairs.] + +The method of feeding of the house-fly has an important bearing on the +question of its ability to transmit pathogenic organisms. Graham-Smith +(1910) has shown that when feeding, flies frequently moisten soluble +substances with "vomit" which is regurgitated from the crop. This is, of +course, loaded with bacteria from previous food. When not sucked up +again these drops of liquid dry, and produce round marks with an opaque +center and rim and an intervening less opaque area. Fly-specks, then, +consist of both vomit spots and feces. Graham-Smith shows a photograph +of a cupboard window where, on an area six inches square, there were +counted eleven hundred and two vomit marks and nine fecal deposits. + +From a bacteriologist's viewpoint a discussion of the possibility of a +fly's carrying bacteria would seem superfluous. Any exposed object, +animate or inanimate, is contaminated by bacteria and will transfer them +if brought into contact with suitable culture media, whether such +substance be food, or drink, open wounds, or the sterile culture media +of the laboratory. A needle point may convey enough germs to produce +disease. Much more readily may the house-fly with its covering of hairs +and its sponge-like pulvilli (fig. 109) pick up and transfer bits of +filth and other contaminated material. + +For popular instruction this inevitable transfer of germs by the +house-fly is strikingly demonstrated by the oft copied illustration of +the tracks of a fly on a sterile culture plate. Two plates of gelatine +or, better, agar medium are prepared. Over one of these a fly (with +wings clipped) is allowed to walk, the other is kept as a check. Both +are put aside at room temperature, to be examined after twenty-four to +forty-eight hours. At the end of that time, the check plate is as clear +as ever, the one which the fly has walked is dotted with colonies of +bacteria and fungi. The value in the experiment consists in emphasizing +that by this method we merely render visible what is constantly +occurring in nature. + +A comparable experiment which we use in our elementary laboratory work +is to take three samples of _clean_ (preferably, sterile) fresh milk in +sterile bottles. One of them is plugged with a pledget of cotton, into +the second is dropped a fly from the laboratory and into the third is +dropped a fly which has been caught feeding upon garbage or other filth. +After a minute or two the flies are removed and the vials plugged as was +number one. The three are then set aside at room temperature. When +examined after twenty-four hours the milk in the first vial is either +still sweet or has a "clean" sour odor; that of the remaining two is +very different, for it has a putrid odor, which is usually more +pronounced in the case of sample number three. + +Several workers have carried out experiments to determine the number of +bacteria carried by flies under natural conditions. One of the most +extended and best known of these is the series by Esten and Mason +(1908). These workers caught flies from various sources in a sterilized +net, placed them in a sterile bottle and poured over them a known +quantity of sterilized water, in which they were shaken so as to wash +the bacteria from their bodies. They found the number of bacteria on a +single fly to range from 550 to 6,600,000. Early in the fly season the +numbers of bacteria on flies are comparatively small, while later the +numbers are comparatively very large. The place where flies live also +determines largely the numbers that they carry. The lowest number, 550, +was from a fly caught in the bacteriological laboratory, the highest +number, 6,600,000 was the average from eighteen swill-barrel flies. +Torrey (1912) made examination of "wild" flies from a tenement house +district of New York City. He found "that the surface contamination of +these 'wild' flies may vary from 570 to 4,400,000 bacteria per insect, +and the intestinal bacterial content from 16,000 to 28,000,000." + +Less well known in this country is the work of Cox, Lewis, and Glynn +(1912). They examined over four hundred and fifty naturally infected +house-flies in Liverpool during September and early October. Instead of +washing the flies they were allowed to swim on the surface of sterile +water for five, fifteen, or thirty minutes, thus giving natural +conditions, where infection occurs from vomit and dejecta of the flies, +as well as from their bodies. They found, as might be expected, that +flies from either insanitary or congested areas of the city contain far +more bacteria than those from the more sanitary, less congested, or +suburban areas. The number of aerobic bacteria from the former varied +from 800,000 to 500,000,000 per fly and from the latter from 21,000 to +100,000. The number of intestinal forms conveyed by flies from +insanitary or congested areas was from 10,000 to 333,000,000 as compared +with from 100 to 10,000 carried by flies from the more sanitary areas. + +Pathogenic bacteria and those allied to the food poisoning group were +only obtained from the congested or moderately congested areas and not +from the suburban areas, where the chances of infestation were less. + +The interesting fact was brought out that flies caught in milk shops +apparently carry and obtain more bacteria than those from other shops +with exposed food in a similar neighborhood. The writers explained this +as probably due to the fact that milk when accessible, especially during +the summer months, is suitable culture medium for bacteria, and the +flies first inoculate the milk and later reinoculate themselves, and +then more of the milk, so establishing a vicious circle. + +They conclude that in cities where food is plentiful flies rarely +migrate from the locality in which they are bred, and consequently the +number of bacteria which they carry depends upon the general standard +of cleanliness in that locality. Flies caught in a street of modern, +fairly high class, workmen's dwellings forming a sanitary oasis in the +midst of a slum area, carried far less bacteria than those caught in the +adjacent neighborhood. + +Thus, as the amount of dirt carried by flies in any particular locality, +measured in the terms of bacteria, bears a definite relation to the +habits of the people and to the state of the streets, it demonstrates +the necessity of efficient municipal and domestic cleanliness, if the +food of the inhabitants is to escape pollution, not only with harmless +but also with occasional pathogenic bacteria. + +The above cited work is of a general nature, but, especially in recent +years, many attempts have been made to determine more specifically the +ability of flies to transmit pathogenic organisms. The critical reviews +of Nuttall and Jepson (1909), Howard (1911), and Graham-Smith (1913) +should be consulted by the student of the subject. We can only cite here +a few of the more striking experiments. + +Celli (1888) fed flies on pure cultures of _Bacillus typhosus_ and +declared that he was able to recover these organisms from the intestinal +contents and excrement. + +Firth and Horrocks (1902), cited by Nuttall and Jepson, "kept _Musca +domestica_ (also bluebottles) in a large box measuring 4 × 3 × 3 feet, +with one side made of glass. They were fed on material contaminated with +cultures of _B. typhosus_. Agar plates, litmus, glucose broth and a +sheet of clean paper were at the same time exposed in the box. After a +few days the plates and broth were removed and incubated with a positive +result." Graham-Smith (1910) "carried out experiments with large numbers +of flies kept in gauze cages and fed for eight hours on emulsions of _B. +typhosus_ in syrup. After that time the infested syrup was removed and +the flies were fed on plain syrup. _B. typhosus_ was isolated up to 48 +hours (but not later) from emulsions of their feces and from plates over +which they walked." + +Several other workers, notably Hamilton (1903), Ficker (1903), +Bertarelli (1910) Faichnie (1909), and Cochrane (1912), have isolated +_B. typhosus_ from "wild" flies, naturally infected. The papers of +Faichnie and of Cochrane we have not seen, but they are quoted in +_extenso_ by Graham-Smith (1913). + +On the whole, the evidence is conclusive that typhoid germs not only may +be accidentally carried on the bodies of house-flies but may pass +through their bodies and be scattered in a viable condition in the feces +of the fly for at least two days after feeding. Similar, results have +been reached in experiments with cholera, tuberculosis and yaws, the +last-mentioned being a spirochæte disease. Darling (1913) has shown that +murrina, a trypanosome disease of horses and mules in the Canal zone is +transmitted by house-flies which feed upon excoriated patches of +diseased animals and then pass to cuts and galls of healthy animals. + +Since it is clear that flies are abundantly able to disseminate viable +pathogenic bacteria, it is important to consider whether they have +access to such organisms in nature. A consideration of the method of +spread of typhoid will serve to illustrate the way in which flies may +play an important rôle. + +Typhoid fever is a specific disease caused by _Bacillus typhosus_, and +by it alone. The causative organism is to be found in the excrement and +urine of patients suffering from the disease. More than that, it is +often present in the dejecta for days, weeks, or even months and years, +after the individual has recovered from the disease. Individuals so +infested are known as "typhoid carriers" and they, together with those +suffering from mild cases, or "walking typhoid," are a constant menace +to the health of the community in which they are found. + +Human excrement is greedily visited by flies, both for feeding and for +ovipositing. The discharges of typhoid patients, or of chronic +"carriers," when passed in the open, in box privies, or camp latrines, +or the like, serve to contaminate myriads of the insects which may then +spread the germ to human food and drink. Other intestinal diseases may +be similarly spread. There is abundant epidæmiological evidence that +infantile diarrhÅ“a, dysentery, and cholera may be so spread. + +Stiles and Keister (1913) have shown that spores of _Lamblia +intestinalis_, a flagellate protozoan living in the human intestine, may +be carried by house-flies. Though this species is not normally +pathogenic, one or more species of _EntamÅ“ba_ are the cause of a type +of a highly fatal tropical dysentery. Concerning it, and another +protozoan parasite of man, they say, "If flies can carry _Lamblia_ +spores measuring 10 to 7µ, and bacteria that are much smaller, and +particles of lime that are much larger, there is no ground to assume +that flies may not carry _EntamÅ“ba_ and _Trichomonas_ spores." + +Tuberculosis is one of the diseases which it is quite conceivable may be +carried occasionally. The sputum of tubercular patients is very +attractive to flies, and various workers, notably Graham-Smith, have +found that _Musca domestica_ may distribute the bacillus for several +days after feeding on infected material. + +A type of purulent opthalmia which is very prevalent in Egypt is often +said to be carried by flies. Nuttall and Jepson (1909) consider that the +evidence regarding the spread of this disease by flies is conclusive and +that the possibility of gonorrhÅ“al secretions being likewise conveyed +cannot be denied. + +Many studies have been published, showing a marked agreement between the +occurrence of typhoid and other intestinal diseases and the prevalence +of house-flies. The most clear-cut of these are the studies of the Army +Commission appointed to investigate the cause of epidemics of enteric +fever in the volunteer camps in the Southern United States during the +Spanish-American War. Though their findings as presented by Vaughan +(1909), have been quoted very many times, they are so germane to our +discussion that they will bear repetition: + +"Flies swarmed over infected fecal matter in the pits and fed upon the +food prepared for the soldiers in the mess tents. In some instances +where lime had recently been sprinkled over the contents of the pits, +flies with their feet whitened with lime were seen walking over the +food." Under such conditions it is no wonder that "These pests had +inflicted greater loss upon American soldiers than the arms of Spain." + +Similar conditions prevailed in South Africa during the Boer War. Seamon +believes that very much of the success of the Japanese in their fight +against Russia was due to the rigid precautions taken to prevent the +spread of disease by these insects and other means. + +Veeder has pointed out that the characteristics of a typical fly-borne +epidemic of typhoid are that it occurs in little neighborhood epidemics, +extending by short leaps from house to house, without regard to water +supply or anything else in common. It tends to follow the direction of +prevailing winds (cf. the conclusions of Hindle and Merriman). It occurs +during warm weather. Of course, when the epidemic is once well under +way, other factors enter into its spread. + +In general, flies may be said to be the chief agency in the spread of +typhoid in villages and camps. In cities with modern sewer systems they +are less important, though even under the best of such conditions, they +are important factors. Howard has emphasized that in such cities there +are still many uncared-for box privies and that, in addition, the +deposition of feces overnight in uncared-for waste lots and alleys is +common. + +Not only unicellular organisms, such as bacteria and protozoa, but also +the eggs, embryos and larvæ of parasitic worms have been found to be +transported by house-flies. Ransom (1911) has found that _Habronema +muscæ_, a nematode worm often found in adult flies, is the immature +stage of a parasite occurring in the stomach of the horse. The eggs or +embryos passing out with the feces of the horse, are taken up by fly +larvæ and carried over to the imago stage. + +Grassi (1883), Stiles (1889), Calandruccio (1906), and especially Nicoll +(1911), have been the chief investigators of the ability of house-flies +to carry the ova and embryos of human intestinal parasites. Graham-Smith +(1913) summarizes the work along this line as follows: + +"It is evident from the investigations that have been quoted that +house-flies and other species are greatly attracted to the ova of +parasitic worms contained in feces and other materials, and make great +efforts to ingest them. Unless the ova are too large they often succeed, +and the eggs are deposited uninjured in their feces, in some cases up to +the third day at least. The eggs may also be carried on their legs or +bodies. Under suitable conditions, food and fluids may be contaminated +with the eggs of various parasitic worms by flies, and in one case +infection of the human subject has been observed. Feces containing +tape-worm segments may continue to be a source of infection for as long +as a fortnight. Up to the present, however, there is no evidence to show +what part flies play in the dissemination of parasitic worms under +natural conditions." + +Enough has been said to show that the house-fly must be dealt with as a +direct menace to public health. Control measures are not merely matters +of convenience but are of vital importance. + +Under present conditions the speedy elimination of the house-fly is +impossible and the first thing to be considered is methods of protecting +food and drink from contamination. The first of these methods is the +thorough screening of doors and windows to prevent the entrance of +flies. In the case of kitchen doors, the flies, attracted by odors, are +likely to swarm onto the screen and improve the first opportunity for +gaining an entrance. This difficulty can be largely avoided by +screening-in the back porch and placing the screen door at one end +rather than directly before the door. + +The use of sticky fly paper to catch the pests that gain entrance to the +house is preferable to the various poisons often used. Of the latter, +formalin (40 per cent formaldehyde) in the proportion of two +tablespoonfuls to a pint of water is very efficient, if all other +liquids are removed or covered, so that the flies must depend on the +formalin for drink. The mixture is said to be made more attractive by +the addition of sugar or milk, though we have found the plain solution +wholly satisfactory, under proper conditions. It should be emphasized +that this formalin mixture is not perfectly harmless, as so often +stated. There are on record cases of severe and even fatal poisoning +from the accidental drinking of solutions. + +When flies are very abundant in a room they can be most readily gotten +rid of by fumigation with sulphur, or by the use of pure pyrethrum +powder either burned or puffed into the air. Herrick (1913) recommends +the following method: "At night all the doors and windows of the kitchen +should be closed; fresh powder should be sprinkled over the stove, on +the window ledges, tables, and in the air. In the morning flies will be +found lying around dead or stupified. They may then be swept up and +burned." This method has proved very efficaceous in some of the large +dining halls in Ithaca. + +The writers have had little success in fumigating with the vapors of +carbolic acid, or carbolic acid and gum camphor, although these methods +will aid in driving flies from a darkened room. + +All of these methods are but makeshifts. As Howard has so well put it, +"the truest and simplest way of attacking the fly problem is to prevent +them from breeding, by the treatment or abolition of all places in which +they can breed. To permit them to breed undisturbed and in countless +numbers, and to devote all our energy to the problem of keeping them out +of our dwellings, or to destroy them after they have once entered in +spite of all obstacles, seems the wrong way to go about it." + +We have already seen that _Musca domestica_ breeds in almost any +fermenting organic material. While it prefers horse manure, it breeds +also in human feces, cow dung and that of other animals, and in refuse +of many kinds. To efficiently combat the insect, these breeding places +must be removed or must be treated in some such way as to render them +unsuitable for the development of the larvæ. Under some conditions +individual work may prove effective, but to be truly efficient there +must be extensive and thorough coöperative efforts. + +Manure, garbage, and the like should be stored in tight receptacles and +carted away at least once a week. The manure may be carted to the fields +and spread. Even in spread manure the larvæ may continue their +development. Howard points out that "it often happens that after a lawn +has been heavily manured in early summer the occupants of the house will +be pestered with flies for a time, but finding no available breeding +place these disappear sooner or later. Another generation will not breed +in the spread manure." + +Hutchinson (1914) has emphasized that the larvæ of houseflies have +deeply engrained the habit of migrating in the prepupal stage and has +shown that this offers an important point of attack in attempts to +control the pest. He has suggested that maggot traps might be developed +into an efficient weapon in the warfare against the house-fly. Certain +it is that the habit greatly simplifies the problem of treating the +manure for the purpose of killing the larvæ. + +There have been many attempts to find some cheap chemical which would +destroy fly larvæ in horse manure without injuring the bacteria or +reducing the fertilizing values of the manure. The literature abounds in +recommendations of kerosene, lime, chloride of lime, iron sulphate, and +other substances, but none of them has met the situation. The whole +question has been gone into thoroughly by Cook, Hutchinson and Scales +(1914), who tested practically all of the substances which have been +recommended. They find that by far the most effective, economical, and +practical of the substances is borax in the commercial form in which it +is available throughout the country. + +"Borax increases the water-soluble nitrogen, ammonia and alkalinity of +manure and apparently does not permanently injure the bacterial flora. +The application of manure treated with borax at the rate of 0.62 pound +per eight bushels (10 cubic feet) to soil does not injure the plants +thus far tested, although its cumulative effect, if any, has not been +determined." + +As their results clearly show that the substances so often recommended +are inferior to borax, we shall quote in detail their directions for +treating manure so as to kill fly eggs and maggots. + +"Apply 0.62 pound borax or 0.75 pound calcined colemanite to every 10 +cubic feet (8 bushels) of manure immediately on its removal from the +barn. Apply the borax particularly around the outer edges of the pile +with a flour sifter or any fine sieve, and sprinkle two or three gallons +of water over the borax-treated manure. + +"The reason for applying the borax to the fresh manure immediately after +its removal from the stable is that the flies lay their eggs on the +fresh manure, and borax, when it comes in contact with the eggs, +prevents their hatching. As the maggots congregate at the outer edge of +the pile, most of the borax should be applied there. The treatment +should be repeated with each addition of fresh manure, but when the +manure is kept in closed boxes, less frequent applications will be +sufficient. When the calcined colemanite is available, it may be used at +the rate of 0.75 pound per 10 cubic feet of manure, and is a cheaper +means of killing the maggots. In addition to the application of borax to +horse manure to kill fly larvæ, it may be applied in the same proportion +to other manures, as well as to refuse and garbage. Borax may also be +applied to the floors and crevices in barns, stables, markets, etc., as +well as to street sweepings, and water should be added as in the +treatment of horse manure. After estimating the amount of material to be +treated and weighing the necessary amount of borax, a measure may be +used which will hold the proper amount, thus avoiding the subsequent +weighings. + +"While it can be safely stated that no injurious action will follow the +application of manure treated with borax at the rate of 0.62 pound for +eight bushels, or even larger amounts in the case of some plants, +nevertheless the borax-treated manure has not been studied in connection +with the growth of all crops, nor has its cumulative effect been +determined. It is therefore recommended that not more than 15 tons per +acre of the borax-treated manure should be applied to the field. As +truckmen use considerably more than this amount, it is suggested that +all cars containing borax-treated manure be so marked, and that +public-health officials stipulate in their directions for this treatment +that not over 0.62 pound for eight bushels of manure be used, as it has +been shown that larger amounts of borax will injure most plants. It is +also recommended that all public-health officials and others, in +recommending the borax treatment for killing fly eggs and maggots in +manure, warn the public against the injurious effects of large amounts +of borax on the growth of plants." + +"The amount of manure from a horse varies with the straw or other +bedding used, but 12 or 15 bushels per week represent the approximate +amount obtained. As borax costs from five to six cents per pound in +100-pound lots in Washington, it will make the cost of the borax +practically one cent per horse, per day. And if calcined colemanite is +purchased in large shipments the cost should be considerably less." + +Hodge (1910) has approached the problem of fly extermination from +another viewpoint. He believes that it is practical to trap flies out of +doors during the preoviposition period, when they are sexually immature, +and to destroy such numbers of them that the comparatively few which +survive will not be able to lay eggs in sufficient numbers to make the +next generation a nuisance. To the end of capturing them in enormous +numbers he has devised traps to be fitted over garbage cans, into stable +windows, and connected with the kitchen window screens. Under some +conditions this method of attack has proved very satisfactory. + +One of the most important measures for preventing the spread of disease +by flies is the abolition of the common box privy. In villages and rural +districts this is today almost the only type to be found. It is the +chief factor in the spread of typhoid and other intestinal diseases, as +well as intestinal parasites. Open and exposed to myriads of flies which +not only breed there but which feed upon the excrement, they furnish +ideal conditions for spreading contamination. Even where efforts are +made to cover the contents with dust, or ashes, or lime, flies may +continue to breed unchecked. Stiles and Gardner have shown that +house-flies buried in a screened stand-pipe forty-eight inches under +sterile sand came to the surface. Other flies of undetermined species +struggled up through seventy-two inches of sand. + +So great is the menace of the ordinary box privy that a number of +inexpensive and simple sanitary privies have been designed for use where +there are not modern sewer systems. Stiles and Lumsden (1911) have given +minute directions for the construction of one of the best types, and +their bulletin should be obtained by those interested. + +Another precaution which is of fundamental importance in preventing the +spread of typhoid, is that of disinfecting all discharges from patients +suffering with the disease. For this purpose, quick-lime is the cheapest +and is wholly satisfactory. In chamber vessels it should be used in a +quantity equal to that of the discharge to be treated. It should be +allowed to act for two hours. Air-slaked lime is of no value whatever. +Chloride of lime, carbolic acid, or formalin may be used, but are more +expensive. Other intestinal diseases demand similar precautions. + + +STOMOXYS CALCITRANS, THE STABLE-FLY--It is a popular belief that +house-flies bite more viciously just before a rain. As a matter of +fact, the true house-flies never bite, for their mouth-parts are not +fitted for piercing. The basis of the misconception is the fact that a +true biting fly, _Stomoxys calcitrans_ (fig. 110), closely resembling +the house-fly, is frequently found in houses and may be driven in in +greater numbers by muggy weather. From its usual habitat this fly is +known as the "stable-fly" or, sometimes as the "biting house-fly." + +_Stomoxys calcitrans_ may be separated from the house-fly by the use of +the key on p. 145. It may be more fully characterized as follows: + +The eyes of the male are separated by a distance equal to one-fourth of +the diameter of the head, in the female by one-third. The frontal stripe +is black, the cheeks and margins of the orbits silvery-white. The +antennæ are black, the arista feathered on the upper side only. The +proboscis is black, slender, fitted for piercing and projects forward in +front of the head. The thorax is grayish, marked by four conspicuous, +more or less complete black longitudinal stripes; the scutellum is +paler; the macrochætæ are black. The abdomen is gray, dorsally with +three brown spots on the second and third segments and a median spot on +the fourth. These spots are more pronounced in the female. The legs are +black, the pulvilli distinct. The wings are hyaline, the vein M_{1+2} +less sharply curved than in the house-fly, the apical cell being thus +more widely open (cf. fig. 110). Length 7 mm. + +[Illustration: 110. Stomoxys calcitrans; adult, larva, puparium and +details, (×5). After Howard.] + +This fly is widely distributed, being found the world over. It was +probably introduced into the United States, but has spread to all parts +of the country. Bishopp (1913) regards it as of much more importance as +a pest of domestic animals in the grain belt than elsewhere in the +United States. The life-history and habits of this species have assumed +a new significance since it has been suggested that it may transmit the +human diseases, infantile paralysis and pellagra. In this country, the +most detailed study of the fly is that of Bishopp (1913) whose data +regarding the life cycle are as follows: + +The eggs like those of the house-fly, are about one mm. in length. Under +a magnifying glass they show a distinct furrow along one side. When +placed on any moist substance they hatch in from one to three days after +being deposited. + +The larva or maggots (fig. 110) have the typical shape and actions of +most maggots of the Muscid group. They can be distinguished from those +of the house-fly as the stigma-plates are smaller, much further apart, +with the slits less sinuous. Development takes place fairly rapidly when +the proper food conditions are available and the growth is completed +within eleven to thirty or more days. + +The pupa (fig. 110), like that of related flies, undergoes its +development within the contracted and hardened last larval skin, or +puparium. This is elongate oval, slightly thicker towards the head end, +and one-sixth to one-fourth of an inch in length. The pupal stage +requires six to twenty days, or in cool weather considerably longer. + +The life-cycle of the stable-fly is therefore considerably longer than +that of _Musca domestica_. Bishopp found that complete development might +be undergone in nineteen days, but that the average period was somewhat +longer, ranging from twenty-one to twenty-five days, where conditions +are very favorable. The longest period which he observed was forty-three +days, though his finding of full grown larvæ and pupæ in straw during +the latter part of March, in Northern Texas, showed that development may +require about three months, as he considered that these stages almost +certainly developed from eggs deposited the previous December. + +The favorite breeding place, where available, seems to be straw or +manure mixed with straw. It also breeds in great numbers in +horse-manure, in company with _Musca domestica_. + +Newstead considers that in England the stable-fly hibernates in the +pupal stage. Bishopp finds that in the southern part of the United +States there is no true hibernation, as the adults have been found to +emerge at various times during the winter. He believes that in the +northern United States the winter is normally passed in the larval and +pupal stages, and that the adults which have been observed in heated +stables in the dead of winter were bred out in refuse within the warm +barns and were not hibernating adults. + +Graham-Smith (1913) states that although the stable-fly frequents stable +manure, it is probably not an important agent in distributing the +organisms of intestinal diseases. Bishopp makes the important +observation that "it has never been found breeding in human excrement +and does not frequent malodorous places, which are so attractive to the +house-fly. Hence it is much less likely to carry typhoid and other germs +which may be found in such places." + +Questions of the possible agency of _Stomoxys calcitrans_ in the +transmission of infantile paralysis and of pellagra, we shall consider +later. + + +OTHER ARTHROPODS WHICH MAY SERVE AS SIMPLE CARRIERS OF PATHOGENIC +ORGANISMS--It should be again emphasized that any insect which has +access to, and comes in contact with, pathogenic organisms and then +passes to the food, or drink, or the body of man, may serve as a simple +carrier of disease. In addition to the more obvious illustrations, an +interesting one is the previously cited case of the transfer of +_Dermatobia cyaniventris_ by a mosquito (fig. 81-84). Darling (1913) has +shown that in the tropics, the omnipresent ants may be important factors +in the spread of disease. + + + + +CHAPTER VI + +ARTHROPODS AS DIRECT INOCULATORS OF DISEASE GERMS + + +We have seen that any insect which, like the house-fly, has access to +disease germs and then comes into contact with the food or drink of man, +may serve to disseminate disease. Moreover, it has been clearly +established that a contaminated insect, alighting upon wounded or +abraded surfaces, may infect them. These are instances of mere +accidental, mechanical transfer of pathogenic organisms. + +Closely related are the instances of direct inoculation of disease germs +by insects and other arthropods. In this type, a blood-sucking species +not only takes up the germs but, passing to a healthy individual, it +inserts its contaminated mouth-parts and thus directly inoculates its +victim. In other words, the disease is transferred just as blood +poisoning may be induced by the prick of a contaminated needle, or as +the laboratory worker may inoculate an experimental animal. + +Formerly, it was supposed that this method of the transfer of disease by +arthropods was a very common one and many instances are cited in the +earlier literature of the subject. It is, however, difficult to draw a +sharp line between such cases and those in which, on the one hand, the +arthropod serves as a mere passive carrier or, on the other hand, serves +as an essential host of the pathogenic organism. More critical study of +the subject has led to the belief that the importance of the rôle of +arthropods as direct inoculators has been much overestimated. + +The principal reason for regarding this phase of the subject as +relatively unimportant, is derived from a study of the habits of the +blood-sucking species. It is found that, in general, they are +intermittent feeders, visiting their hosts at intervals and then +abstaining from feeding for a more or less extended period, while +digesting their meal. In the meantime, most species of bacteria or of +protozoan parasites with which they might have contaminated their +mouth-parts, would have perished, through inability to withstand drying. + +In spite of this, it must be recognized that this method of transfer +does occur and must be reckoned with in any consideration of the +relations of insects to disease. We shall first cite some general +illustrations and shall then discuss the rôle of fleas in the spreading +of bubonic plague, an illustration which cannot be regarded as typical, +since it involves more than mere passive carriage. + + +SOME ILLUSTRATIONS OF DIRECT INOCULATION OF DISEASE GERMS BY ARTHROPODS + +In discussing poisonous arthropods, we have already emphasized that +species which are of themselves innocuous to man, may occasionally +introduce bacteria by their bite or sting and thus cause more or less +severe secondary symptoms. That such cases should occur, is no more than +is to be expected. The mouth-parts or the sting of the insect are not +sterile and the chances of their carrying pyogenic organisms are always +present. + +More strictly falling in the category of transmission of disease germs +by direct inoculation are the instances where the insect, or related +form, feeds upon a diseased animal and passes promptly to a healthy +individual which it infects. Of such a nature are the following: + +Various species of biting flies are factors in the dissemination of +anthrax, an infectious and usually fatal disease of animals and, +occasionally, of man. That the bacteria with which the blood of diseased +animals teem shortly before death might be transmitted by such insects +has long been contended, but the evidence in support of the view has +been unsatisfactory. Recently, Mitzmain (1914) has reported a series of +experiments which show conclusively that the disease may be so conveyed +by a horse-fly, _Tabanus striatus_, and by the stable-fly, _Stomoxys +calcitrans_. + +Mitzmain's experiments were tried with an artificially infected guinea +pig, which died of the disease upon the third day. The flies were +applied two and one-half hours, to a few minutes, before the death of +the animal. With both species the infection was successfully transferred +to healthy guinea pigs by the direct method, in which the flies were +interrupted while feeding on the sick animal. The evidence at hand does +not warrant the conclusion that insect transmission is the rule in the +case of this disease. + +The nagana, or tsetse-fly disease of cattle is the most virulent disease +of domestic animals in certain parts of Africa. It is caused by a +protozoan blood parasite, _Trypanosoma brucei_, which is conveyed to +healthy animals by the bite of _Glossina morsitans_ and possibly other +species of tsetse-flies. The flies remain infective for forty-eight +hours after feeding on a diseased animal. The insect also serves as an +essential host of the parasite. + +Surra, a similar trypanosomiasis affecting especially horses and mules, +occurs in southern Asia, Malaysia, and the Philippines where the +tsetse-flies are not to be found. It is thought to be spread by various +species of blood-sucking flies belonging to the genera _Stomoxys_, +_Hæmatobia_, and _Tabanus_. Mitzmain (1913) demonstrated that in the +Philippines it is conveyed mechanically by _Tabanus striatus_. + +The sleeping sickness of man, in Africa, has also been supposed to be +directly inoculated by one, or several, species of tsetse-flies. It is +now known that the fly may convey the disease for a short time after +feeding, but that there is then a latent period of from fourteen to +twenty-one days, after which it again becomes infectious. This indicates +that in the meantime the parasite has been undergoing some phase of its +life-cycle and that the fly serves as an intermediate host. We shall +therefore consider it more fully under that grouping. + +These are a few of the cases of direct inoculation which may be cited as +of the simpler type. We shall next consider the rôle of the flea in the +dissemination of the bubonic plague, an illustration complicated by the +fact that the bacillus multiples within the insect and may be indirectly +inoculated. + + +THE RÔLE OF FLEAS IN THE TRANSMISSION OF THE PLAGUE + +The plague is a specific infectious disease caused by _Bacillus pestis_. +It occurs in several forms, of which the bubonic and the pneumonic are +the most common. According to Wyman, 80 per cent of the human cases are +of the bubonic type. It is a disease which, under the name of oriental +plague, the pest, or the black death, has ravaged almost from time +immemorial the countries of Africa, Asia, and Europe. The record of its +ravages are almost beyond belief. In 542 A. D. it caused in one day ten +thousand deaths in Constantinople. In the 14th century it was introduced +from the East and prevailed throughout Armenia, Asia Minor, Egypt and +Northern Africa and Europe. Hecker estimates that one-fourth of the +population of Europe, or twenty-five million persons, died in the +epidemic of that century. From then until the 17th century it was almost +constantly present in Europe, the great plague of London, in 1665 +killing 68,596 out of a population of 460,000. Such an epidemic would +mean for New York City a proportionate loss of over 600,000 in a single +year. It is little wonder that in the face of such an appalling disaster +suspicion and credulity were rife and the wildest demoralization ensued. + +During the 14th century the Jews were regarded as responsible for the +disease, through poisoning wells, and were subjected to the most +incredible persecution and torture. In Milan the visitation of 1630 was +credited to the so-called anointers,--men who were supposed to spread +the plague by anointing the walls with magic ointment--and the most +horrible tortures that human ingenuity could devise were imposed on +scores of victims, regardless of rank or of public service (fig. 112, +a). Manzoni's great historical novel, "The Betrothed" has well pictured +conditions in Italy during this period. + +[Illustration: 111. A contemporaneous engraving of the pest hospital in +Vienna in 1679. After Peters.] + +In modern times the plague is confined primarily to warm climates, a +condition which has been brought about largely through general +improvement in sanitary conditions. + +At present, the hotbed of the disease is India, where there were +1,040,429 deaths in 1904 and where in a period of fifteen years, ending +with January 1912, there were over 15,000,000 deaths. The reported +deaths in that country for 1913 totaled 198,875. + +During the winter of 1910-11 there occurred in Manchuria and North China +a virulent epidemic of the pneumonic plague which caused the death of +nearly 50,000 people. The question as to its origin and means of spread +will be especially referred to later. + +[Illustration: 112 a. A medieval method of combating the plague. The +persecution of the anointers in Milan in 1630. From a copy of "Il +processi originale degli untori" in the library of Cornell University.] + +Until recent years, the plague had not been known to occur in the New +World but there were outbreaks in Brazil and Hawaii in 1899, and in 1900 +there occurred the first cases in San Francisco. In California there +were 125 cases in the period 1900-04; three cases in the next three +years and then from May 1907 to March 1908, during the height of the +outbreak, 170 cases. Since that time there have been only sporadic +cases, the last case reported being in May 1914. Still more recent were +the outbreaks in the Philippine Islands, Porto Rico, and Cuba. + +On June 24, 1914, there was recognized a case of human plague in New +Orleans. The Federal Health Service immediately took charge, and +measures for the eradication of the disease were vigorously enforced. Up +to October 10, 1914 there had been reported 30 cases of the disease in +man, and 181 cases of plague in rats. + +[Illustration: 112 b. The modern method of combating the plague. A day's +catch of rats in the fight against plague in San Francisco. Courtesy of +Review of Reviews.] + +The present-day methods of combating bubonic plague are well illustrated +by the fight in San Francisco. Had it not been for the strenuous and +radical anti-plague campaign directed by the United States Marine +Hospital Service we might have had in our own country an illustration of +what the disease can accomplish. On what newly acquired knowledge was +this fight based? + +The basis was laid in 1894, when the plague bacillus was first +discovered. All through the centuries, before and during the Christian +era, down to 1894, the subject was enveloped in darkness and there had +been a helpless, almost hopeless struggle in ignorance on the part of +physicians, sanitarians, and public health officials against the ravages +of this dread disease. Now its cause, method of propagation and means to +prevent its spread are matters of scientific certainty. + +After the discovery of the causative organism, one of the first advances +was the establishment of the identity of human plague and that of +rodents. It had often been noted that epidemics of the human disease +were preceded by great epizootics among rats and mice. So well +established was this fact that with the Chinese, unusual mortality among +these rodents was regarded as foretelling a visitation of the human +disease. That there was more than an accidental connection between the +two was obvious when Yersin, the discoverer of _Bacillus pestis_, +announced that during an epidemic the rats found dead in the houses and +in the streets almost always contain the bacillus in great abundance in +their organs, and that many of them exhibit veritable buboes. + +Once it was established that the diseases were identical, the attention +of the investigators was directed to a study of the relations between +that of rats and of humans, and evidence accumulated to show that the +bubonic plague was primarily a disease of rodents and that in some +manner it was conveyed from them to man. + +There yet remained unexplained the method of transfer from rat to man. +As long ago as the 16th century, Mercuralis suggested that house-flies +were guilty of disseminating the plague but modern investigation, while +blaming the fly for much in the way of spreading disease, show that it +is an insignificant factor in this case. + +Search for blood-sucking insects which would feed on both rodents and +man, and which might therefore be implicated, indicated that the fleas +most nearly met the conditions. At first it was urged that rat fleas +would not feed upon man and that the fleas ordinarily attacking man +would not feed upon rats. More critical study of the habits of fleas +soon showed that these objections were not well-founded. Especially +important was the evidence that soon after the death of their host, rat +fleas deserted its body and might then become a pest in houses where +they had not been noticed before. + +Attention was directed to the fact that while feeding, fleas are in the +habit of squirting blood from the anus and that in the case of those +which had fed upon rats and mice dying of the plague, virulent plague +bacilli were to be found in such blood. Liston (1905) even found, and +subsequent investigations confirmed, that the plague bacilli multiply in +the stomach of the insect and that thus the blood ejected was richer in +the organisms than was that of the diseased animal. It was found that a +film of this infected blood spread out under the body of the flea and +that thus the bacilli might be inoculated by the bite of the insect and +by scratching. + +Very recently, Bacot and Martin (1914) have paid especial attention to +the question of the mechanism of the transmission of the plague bacilli +by fleas. They believe that plague infested fleas regurgitate blood +through the mouth, and that under conditions precluding the possibility +of infection by dejecta, the disease may be thus transmitted. The +evidence does not seem sufficient to establish that this is the chief +method of transmission. + +Conclusive experimental proof that fleas transmit the disease is further +available from a number of sources. The most extensive series of +experiments is that of the English Plague Commission in India, which +reported in 1906 that: + +On thirty occasions a healthy rat contracted plague in sequence of +living in the neighborhood of a plague infected rat under circumstances +which prevented the healthy rat coming in contact with either the body +or excreta of the diseased animal. + +In twenty-one experiments out of thirty-eight, healthy rats living in +flea-proof cages contracted plague when exposed to rat fleas +(_Xenopsylla cheopis_), collected from rats dead or dying of septicæmic +plague. + +Close contact of plague-infected with healthy animals, if fleas are +excluded, does not give rise to an epizootic among the latter. As the +huts were never cleaned out, close contact included contact with feces +and urine of infected animals, and contact with, and eating of food +contaminated with feces and urine of infected animals, as well as pus +from open plague ulcers. Close contact of young, even when suckled by +plague-infected mothers, did not give the disease to the former. + +If fleas are present, then the epizootic, once started, spreads from +animal to animal, the rate of progress being in direct proportion to the +number of fleas. + +Aerial infection was excluded. Thus guinea-pigs suspended in a cage two +feet above the ground did not contract the disease, while in the same +hut those animals allowed to run about and those placed two inches above +the floor became infected. It had previously been found that a rat flea +could not hop farther than about five inches. + +Guinea pigs and monkeys were placed in plague houses in pairs, both +protected from soil contact infection and both equally exposed to aerial +infection, but one surrounded with a layer of tangle-foot paper and the +other surrounded with a layer of sand. The following observations were +made: + +(_a_) Many fleas were caught in the tangle-foot, a certain proportion of +which were found on dissection to contain in their stomachs abundant +bacilli microscopically identical with plague bacilli. Out of +eighty-five human fleas dissected only one contained these bacilli, +while out of seventy-seven rat fleas twenty-three were found thus +infected. + +(_b_) The animals surrounded with tangle-foot in no instance developed +plague, while several (24 per cent) of the non-protected animals died of +the disease. + +Thus, the experimental evidence that fleas transmit the plague from rat +to rat, from rats to guinea pigs, and from rats to monkeys is +indisputable. There is lacking direct experimental proof of its transfer +from rodents to man but the whole chain of indirect evidence is so +complete that there can be no doubt that such a transfer does occur so +commonly that in the case of bubonic plague it must be regarded as the +normal method. + +Rats are not the only animals naturally attacked by the plague but as +already suggested, it occurs in various other rodents. In California the +disease has spread from rats to ground squirrels (_Otospermophilus +beecheyi_), a condition readily arising from the frequency of +association of rats with the squirrels in the neighborhood of towns, and +from the fact that the two species of fleas found on them are also found +on rats. While the danger of the disease being conveyed from squirrels +to man is comparatively slight, the menace in the situation is that the +squirrels may become a more or less permanent reservoir of the disease +and infect rats, which may come into more frequent contact with man. + +The tarbagan (_Arctomys bobac_), is a rodent found in North Manchuria, +which is much prized for its fur. It is claimed that this animal is +extremely susceptible to the plague and there is evidence to indicate +that it was the primary source of the great outbreak of pneumonic plague +which occurred in Manchuria and North China during the winter of +1910-11. + +Of fleas, any species which attacks both rodents and man may be an agent +in the transmission of the plague. We have seen that in India the +species most commonly implicated is the rat flea, _Xenopsylla cheopis_, +(= _LÅ“mopsylla_ or _Pulex cheopis_) (fig. 89). This species has also +been found commonly on rats in San Francisco. The cat flea, +_Ctenocephalus felis_, the dog flea, _Ctenocephalus canis_, the human +flea, _Pulex irritans_, the rat fleas, _Ceratophyllus fasciatus_ and +_Ctenopsyllus musculi_ have all been shown to meet the conditions. + +But, however clear the evidence that fleas are the most important agent +in the transfer of plague, it is a mistake fraught with danger to assume +that they are the only factor in the spread of the disease. The +causative organism is a bacillus and is not dependent upon any insect +for the completion of its development. + +Therefore, any blood-sucking insect which feeds upon a plague infected +man or animal and then passes to a healthy individual, conceivably might +transfer the bacilli. Verjbitski (1908) has shown experimentally that +bed-bugs may thus convey the disease. Hertzog found the bacilli in a +head-louse, _Pediculus humanus_, taken from a child which had died from +the plague, and McCoy found them in a louse taken from a plague-infected +squirrel. On account of their stationary habits, the latter insects +could be of little significance in spreading the disease. + +Contaminated food may also be a source of danger. While this source, +formerly supposed to be the principal one, is now regarded as +unimportant, there is abundant experimental evidence to show that it +cannot be disregarded. It is believed that infection in this way can +occur only when there is some lesion in the alimentary canal. + +Still more important is the proof that in pneumonic plague the patient +is directly infective and that the disease is spread from man to man +without any intermediary. Especially conclusive is the evidence obtained +by Drs. Strong and Teague during the Manchurian epidemic of 1910-11. +They found that during coughing, in pneumonic plague cases, even when +sputum visible to the naked eye is not expelled, plague bacilli in large +numbers may become widely disseminated into the surrounding air. By +exposing sterile plates before patients who coughed a single time, very +numerous colonies of the bacillus were obtained. + +But the great advance which has been made rests on the discovery that +bubonic plague is in the vast majority of cases transmitted by the flea. +The pneumonic type forms a very small percentage of the human cases and +even with it, the evidence indicates that the original infection is +derived from a rodent through the intermediary of the insect. + +So modern prophylactic measures are directed primarily against the rat +and fleas. Ships coming from infected ports are no longer disinfected +for the purpose of killing the plague germs, but are fumigated to +destroy the rats and the fleas which they might harbor. When anchored at +infected ports, ships must observe strenuous precautions to prevent the +ingress of rats. Cargo must be inspected just before being brought on +board, in order to insure its freedom from rats. Even lines and hawsers +must be protected by large metal discs or funnels, for rats readily run +along a rope to reach the ship. Once infested, the ship must be +thoroughly fumigated, not only to avoid carrying the disease to other +ports but to obviate an outbreak on board. + +When an epidemic begins, rats must be destroyed by trapping and +poisoning. Various so-called biological poisons have not proved +practicable. Sources of food supply should be cut off by thorough +cleaning up, by use of rat-proof garbage cans and similar measures. Hand +in hand with these, must go the destruction of breeding places, and the +rat-proofing of dwellings, stables, markets, warehouses, docks and +sewers. All these measures are expensive, and a few years ago would have +been thought wholly impossible to put into practice but now they are +being enforced on a large scale in every fight against the disease. + +Rats and other rodents are regularly caught in the danger zone and +examined for evidence of infection, for the sequence of the epizootic +and of the human disease is now understood. In London, rats are +regularly trapped and poisoned in the vicinity of the principal docks, +to guard against the introduction of infected animals in shipping. +During the past six years infected rats have been found yearly, thirteen +having been found in 1912. In Seattle, Washington, seven infected rats +were found along the water front in October, 1913, and infected ground +squirrels are still being found in connection with the anti-plague +measures in California. + +The procedure during an outbreak of the human plague was well +illustrated by the fight in San Francisco. The city was districted, and +captured rats, after being dipped in some fluid to destroy the fleas, +were carefully tagged to indicate their source, and were sent to the +laboratory for examination. If an infected rat was found, the officers +in charge of the work in the district involved were immediately notified +by telephone, and the infected building was subjected to a thorough +fumigation. In addition, special attention was given to all the +territory in the four contiguous blocks. + +By measures such as these, this dread scourge of the human race is being +brought under control. Incidentally, the enormous losses due to the +direct ravages of rats are being obviated and this alone would justify +the expenditure many times over of the money and labor involved in the +anti-rat measures. + + + + +CHAPTER VII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC ORGANISMS + + +We now have to consider the cases in which the arthropod acts as the +essential host of a pathogenic organism. In other words, cases in which +the organism, instead of being passively carried or merely accidentally +inoculated by the bite of its carrier, or vector, is taken up and +undergoes an essential part of its development within the arthropod. + +[Illustration: 113. Dipylidium caninum. The double pored tapeworm of the +dog.] + +In some cases, the sexual cycle of the parasite is undergone in the +arthropod, which then serves as the _definitive_ or _primary host_. In +other cases, it is the asexual stage of the parasite which is undergone, +and the arthropod then acts as the _intermediate host_. This distinction +is often overlooked and all the cases incorrectly referred to as those +in which the insect or other arthropod acts as intermediate host. + +We have already emphasized that this is the most important way in which +insects may transmit disease, for without them the particular organisms +concerned could never complete their development. Exterminate the +arthropod host and the life cycle of the parasite is broken, the disease +is exterminated. + +As the phenomenon of alternation of generations, as exhibited by many of +the parasitic protozoa, is a complicated one and usually new to the +student, we shall first take up some of the grosser cases illustrated by +certain parasitic worms. There is the additional reason that these were +the first cases known of arthropod transmission of pathogenic organisms. + + +INSECTS AS INTERMEDIATE HOSTS OF TAPEWORMS + +A number of tapeworms are known to undergo their sexual stage in an +insect or other arthropod. Of these at least two are occasional +parasites of man. + +_Dipylidium caninum_ (figs. 113 and 114), more generally known as +_Taenia cucumerina_ or _T. elliptica_, is the commonest intestinal +parasite of pet dogs and cats. It is occasionally found as a human +parasite, 70 per cent of the cases reported being in young children. + +In 1869, Melnikoff found in a dog louse, _Trichodectes canis_, some +peculiar bodies which Leuckart identified as the larval form of this +tapeworm. The worm is, however, much more common in dogs and cats than +is the skin parasite, and hence it appears that the _Trichodectes_ could +not be the only intermediate host. In 1888, Grassi found that it could +also develop in the cat and dog fleas, _Ctenocephalus felis_ and _C. +canis_, and in the human flea, _Pulex irritans_. + +[Illustration: 114. Dipylidium caninum. Rostrum evaginated and +invaginated. After Blanchard.] + +[Illustration: 115. Dipylidium caninum. Immature cysticercoid. After +Grassi and Rovelli.] + +The eggs, scattered among the hairs of the dog or cat, are ingested by +the insect host and in its body cavity they develop into pyriform +bodies, about 300µ in length, almost entirely destitute of a bladder, +but in the immature stage provided with a caudal appendage (fig. 115). +Within the pear-shaped body (fig. 116) are the invaginated head and +suckers of the future tapeworm. This larval form is known as a +cysticercoid, in contradistinction to the bladder-like cysticercus of +many other cestodes. It is often referred to in literature as +_Cryptocystis trichodectis_ Villot. + +As many as fifty of the cysticercoids have been found in the body cavity +of a single flea. When the dog takes up an infested flea or louse, by +biting itself, or when the cat licks them up, the larvæ quickly develop +into tapeworms, reaching sexual maturity in about twenty days in the +intestine of their host. Puppies and kittens are quickly infested when +suckling a flea-infested mother, the developing worms having been found +in the intestines of puppies not more than five or six days old. + +[Illustration: 116. Dipylidium caninum. Cysticercoid. After Villet.] + +Infestation of human beings occurs only through accidental ingestion of +an infested flea. It is natural that such cases should occur largely in +children, where they may come about in some such way as illustrated in +the accompanying figures 117 and 118. + +_Hymenolepis diminuta_, very commonly living in the intestine of mice +and rats, is also known to occur in man. Its cysticercoid develops in +the body cavity of a surprising range of meal-infesting insects. Grassi +and Rovelli (abstract in Ransom, 1904) found it in the larvæ and adult +of a moth, _Asopia farinalis_, in the earwig, _Anisolabis annulipes_, +the Tenebrionid beetles _Akis spinosa_ and _Scaurus striatus_. Grassi +considers that the lepidopter is the normal intermediate host. The +insect takes up the eggs scattered by rats and mice. It has been +experimentally demonstrated that man may develop the tapeworm by +swallowing infested insects. Natural infection probably occurs by +ingesting such insects with cereals, or imperfectly cooked foods. + +[Illustration: 117. One way in which Dipylidium infection in children +may occur. After Blanchard.] + +_Hymenolepis lanceolata_, a parasite of geese and ducks, has been +reported once for man. The supposed cysticercoid occurs in various small +crustaceans of the family Cyclopidæ. + +[Illustration: 118. The probable method by which Dipylidium infection +usually occurs.] + +Several other cestode parasites of domestic animals are believed to +develop their intermediate stage in certain arthropods. Among these may +be mentioned: + +_Choanotænia infundibulformis_, of chickens, developing in the house-fly +(Grassi and Rovelli); + +_Davainea cesticillus_, of chickens, in some lepidopter or coleopter +(Grassi and Rovelli); + +_Hymenolepis anatina_, _H. gracilis_, _H. sinuosa_, _H. coronula_ and +_Fimbriaria fasciolaris_, all occurring in ducks, have been reported as +developing in small aquatic crustaceans. In these cases, cysticercoids +have been found which, on account of superficial characters, have been +regarded as belonging to the several species, but direct experimental +evidence is scant. + + +ARTHROPODS AS INTERMEDIATE HOSTS OF NEMATODE WORMS + + +FILARIASIS AND MOSQUITOES--A number of species of Nematode worms +belonging to the genus _Filaria_, infest man and other vertebrates and +in the larval condition are to be found in the blood. Such infestation +is known as _filariasis_. The sexually mature worms are to be found in +the blood, the lymphatics, the mesentery and subcutaneous connective +tissue. In the cases best studied it has been found that the larval +forms are taken up by mosquitoes and undergo a transformation before +they can attain maturity in man. + +The larvæ circulating in the blood are conveniently designated as +microfilariæ. In this stage they are harmless and only one species, +_Filaria bancrofti_, appears to be of any great pathological +significance at any stage. + +_Filaria bancrofti_ in its adult state, lives in the lymphatics of man. +Though often causing no injury it has been clearly established that they +and their eggs may cause various disorders due to stoppage of the +lymphatic trunks (fig. 119). Manson lists among other effects, abscess, +varicose groin glands, lymph scrotum, chyluria, and elephantiasis. + +The geographical distribution of this parasite is usually given as +coextensive with that of elephantiasis, but it is by no means certain +that it is the only cause of this disease and so actual findings of the +parasites are necessary. Manson reports that it is "an indigenous +parasite in almost every country throughout the tropical and subtropical +world, as far north as Spain in Europe and Charlestown in the United +States, and as far south as Brisbane in Australia." In some sections, +fully 50 per cent of the natives are infested. Labredo (1910) found +17.82 per cent infestation in Havana. + +[Illustration: 119. Elephantiasis in Man. From "New Sydenham Society's +Atlas."] + +The larval forms of _Filaria bancrofti_ were first discovered in 1863, +by Demarquay, in a case of chylous dropsy. They were subsequently noted +under similar conditions, by several workers, and by Wücherer in the +urine of twenty-eight cases of tropical chyluria, but in 1872 Lewis +found that the blood of man was the normal habitat, and gave them the +name _Filaria sanguinis hominis_. The adult worm was found in 1876 by +Bancroft, and in 1877, Cobbold gave it the name _Filaria bancrofti_. It +has since been found repeatedly in various parts of the lymphatic +system, and its life-history has been the subject of detailed studies by +Manson (1884), Bancroft (1899), Low (1900), Grassi and Noé (1900), Noé +(1901) and Fülleborn (1910). + +The larvæ as they exist in the circulating blood, exhibit a very active +wriggling movement, without material progression. They may exist in +enormous numbers, as many as five or six hundred swarming in a single +drop of blood. This is the more surprising when we consider that they +measure about 300µ × 8µ, that is, their width is equal to the diameter +of the red blood corpuscle of their host and their length over +thirty-seven times as great. + +Their organs are very immature and the structure obscure. When they have +quieted down somewhat in a preparation it may be seen that at the head +end there is a six-lipped and very delicate prepuce, enclosing a short +"fang" which may be suddenly exserted and retracted. Completely +enclosing the larva is a delicate sheath, which is considerably longer +than the worm itself. To enter into further details of anatomy is beyond +the scope of this discussion and readers interested are referred to the +work of Manson and of Fülleborn. + +One of the most surprising features of the habits of these larvæ is the +periodicity which they exhibit in their occurrence in the peripheral +blood. If a preparation be made during the day time there may be no +evidence whatever of filarial infestation, whereas a preparation from +the same patient taken late in the evening or during the night may be +literally swarming with the parasites. Manson quotes Mackenzie as having +brought out the further interesting fact that should a "filarial subject +be made to sleep during the day and remain awake at night, the +periodicity is reversed; that is to say, the parasites come into the +blood during the day and disappear from it during the night." There have +been numerous attempts to explain this peculiar phenomenon of +periodicity but in spite of objections which have been raised, the most +plausible remains that of Manson, who believes that it is an adaptation +correlated with the life-habits of the liberating agent of the parasite, +the mosquito. + +The next stages in the development of _Filaria nocturna_ occur in +mosquitoes, a fact suggested almost simultaneously by Bancroft and +Manson in 1877, and first demonstrated by the latter very soon +thereafter. The experiments were first carried out with _Culex +quinquefasciatus_ (= _fatigans_) as a host, but it is now known that a +number of species of mosquitoes, both anopheline and culicine, may serve +equally well. + +When the blood of an infested individual is sucked up and reaches the +stomach of such a mosquito, the larvæ, by very active movements, escape +from their sheaths and within a very few hours actively migrate to the +body cavity of their new host and settle down primarily in the thoracic +muscles. There in the course of sixteen to twenty days they undergo a +metamorphosis of which the more conspicuous features are the formation +of a mouth, an alimentary canal and a trilobed tail. At the same time +there is an enormous increase in size, the larvæ which measured .3 mm. +in the blood becoming 1.5 mm. in length. This developmental period may +be somewhat shortened in some cases and on the other hand may be +considerably extended. The controlling factor seems to be the one of +temperature. + +The transformed larvæ then reenter the body cavity and finally the +majority of them reach the interior of the labium (fig. 120). A few +enter the legs and antennæ, and the abdomen, but these are wanderers +which, it is possible, may likewise ultimately reach the labium, where +they await the opportunity to enter their human host. + +It was formerly supposed that when the infested mosquito punctured the +skin of man, the mature larvæ were injected into the circulation. The +manner in which this occurred was not obvious, for when the insect feeds +it inserts only the stylets, the labium itself remaining on the surface +of the skin. Fülleborn has cleared up the question by showing that at +this time the filariæ escape and, like the hookworm, actively bore into +the skin of their new host. + +[Illustration: 120. Filaria in the muscles and labium of Culex. After +Blanchard.] + +Once entered, they migrate to the lymphatics and there quickly become +sexually mature. The full grown females measure 85-90 mm. in length by +.24-.28 mm. in diameter, while the males are less than half this size, +being about 40 mm. by .1 mm. Fecundation occurs and the females will be +found filled with eggs in various stages of development, for they are +normally viviparous. + +_Filaria philippinensis_ is reported by Ashburn and Craig (1907) as a +common blood filaria in the Philippine Islands. As they describe it, it +differs from _Filaria bancrofti_ primarily in that it does not exhibit +periodicity. Its development has been found to occur in _Culex +quinquefasciatus_, where it undergoes metamorphosis in about fourteen or +fifteen days. There is doubt as to the species being distinct from +_bancrofti_. + +Several other species occur in man and are thought to be transferred by +various insects, among which have been mentioned Tabanidæ and +tsetse-flies, but there is no experimental proof in support of such +conjectures. + +_Filaria immitis_ is a dangerous parasite of the dog, the adult worm +living in the heart and veins of this animal. It is one of the species +which has been clearly shown to undergo its development in the mosquito, +particularly in _Anopheles maculipennis_ and _Aedes calopus_ (= +Stegomyia). The larval form occurs in the peripheral blood, especially +at night. When taken up by mosquitoes they differ from _Filaria +bancrofti_ in that they undergo their development in the Malpighian +tubules rather than in the thoracic muscles. In about twelve days they +have completed their growth in the tubules, pierce the distal end, and +pass to the labium. This species occurs primarily in China and Japan, +but is also found in Europe and in the United States. It is an +especially favorable species for studying the transformations in the +mosquito. + +[Illustration: 121. Dracunculus medinensis; female; mouth; embryo. After +Bastian and Leuckart.] + +_Filariæ_ are also commonly found in birds, and in this country this is +the most available source of laboratory material. We have found them +locally (Ithaca, N. Y.) in the blood of over sixty per cent of all the +crows examined, at any season of the year, and have also found them in +English sparrows. + +In the crows, they often occur in enormous numbers, as many as two +thousand having been found in a single drop of the blood of the most +heavily infested specimen examined. For study, a small drop of blood +should be mounted on a clean slide and the coverglass rung with vaseline +or oil to prevent evaporation. In this way they can be kept for hours. + +Permanent preparations may be made by spreading out the blood in a film +on a perfectly clean slide and staining. This is easiest done by +touching the fresh drop of blood with the end of a second slide which is +then held at an angle of about 45° to the first slide and drawn over it +without pressure. Allow the smear to dry in the air and stain in the +usual way with hæmatoxylin. + + +OTHER NEMATODE PARASITES OF MAN AND ANIMALS DEVELOPING IN ARTHROPODS + +_Dracunculus medinensis_ (fig. 121), the so-called guinea-worm, is a +nematode parasite of man which is widely distributed in tropical Africa, +Asia, certain parts of Brazil and is occasionally imported into North +America. + +The female worm is excessively long and slender, measuring nearly three +feet in length and not more than one-fifteenth of an inch in diameter. +It is found in the subcutaneous connective tissue and when mature +usually migrates to some part of the leg. Here it pierces the skin and +there is formed a small superficial ulcer through which the larvæ reach +the exterior after bursting the body of the mother. + +[Illustration: 122. Cyclops, the intermediate host of Dracunculus.] + +Fedtschenko (1879) found that when these larvæ reach the water they +penetrate the carapace of the little crustacean, _Cyclops_ (fig. 122). +Here they molt several times and undergo a metamorphosis. Fedtschenko, +in Turkestan, found that these stages required about five weeks, while +Manson who confirmed these general results, found that eight or nine +weeks were required in the cooler climate of England. + +Infection of the vertebrate host probably occurs through swallowing +infested cyclops in drinking water. Fedtschenko was unable to +demonstrate this experimentally and objection has been raised against +the theory, but Leiper (1907), and Strassen (1907) succeeded in +infesting monkeys by feeding them on cyclops containing the larvæ. + +_Habronema muscæ_ is a worm which has long been known in its larval +stage, as a parasite of the house-fly. Carter found them in 33 per cent +of the house-flies examined in Bombay during July, 1860, and since that +time they have been shown to be very widely distributed. Italian workers +reported them in 12 per cent to 30 per cent of the flies examined. +Hewitt reported finding it rarely in England. In this country it was +first reported by Leidy who found it in about 20 per cent of the flies +examined at Philadelphia, Pa. Since then it has been reported by several +American workers. We have found it at Ithaca, N. Y., but have not made +sufficient examinations to justify stating percentage. Ransom (1913) +reports it in thirty-nine out of one hundred and thirty-seven flies, or +28 per cent. + +[Illustration: 123. An Echinorhynchid, showing the spinose retractile +proboscis.] + +[Illustration: 124. June beetle (Lachnosterna).] + +[Illustration: Larva] + +Until very recently the life-history of this parasite was unknown but +the thorough work of Ransom (1911, 1913) has shown clearly that the +adult stage occurs in the stomach of horses. The embryos, produced by +the parent worms in the stomach of the horse, pass out with the feces +and enter the bodies of fly larvæ which are developing in the manure. In +these they reach their final stage of larval development at about the +time the adult flies emerge from the pupal stage. In the adult fly they +are commonly found in the head, frequently in the proboscis, but they +occur also in the thorax and abdomen. Infested flies are accidentally +swallowed by horses and the parasite completes its development to +maturity in the stomach of its definitive host. + +_Gigantorhynchus hirudinaceus_ (= _Echinorhynchus gigas_) is a common +parasite of the pig and has been reported as occurring in man. The adult +female is 20-35 cm. long and 4-9 mm. in diameter. It lacks an alimentary +canal and is provided with a strongly spined protractile rostrum, by +means of which it attaches to the intestinal mucosa of its host. + +The eggs are scattered with the feces of the host and are taken up by +certain beetle larvæ. In Europe the usual intermediate hosts are the +larvæ of the cockchafer, _Melolontha vulgaris_, or of the flower beetle, +_Cetonia aurata_. Stiles has shown that in the United States the +intermediate host is the larva of the June bug, _Lachnosterna_ (fig. +124). It is probable that several of the native species serve in this +capacity. + +A number of other nematode parasites of birds and mammals have been +reported as developing in arthropods but here, as in the case of the +cestodes, experimental proof is scant. The cases above cited are the +better established and will serve as illustrations. + + + + +CHAPTER VIII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA + + +MOSQUITOES AND MALARIA + +Under the name of malaria is included a group of morbid symptoms +formerly supposed to be due to a miasm or bad air, but now known to be +caused by protozoan parasites of the genus _Plasmodium_, which attack +the red blood corpuscles. It occurs in paroxysms, each marked by a +chill, followed by high fever and sweating. The fever is either +intermittent or remittent. + +There are three principal types of the disease, due to different species +of the parasite. They are: + +1. The benign-tertian, caused by _Plasmodium vivax_, which undergoes its +schizogony or asexual cycle in the blood in forty-eight hours or even +less. This type of the disease,--characterized by fever every two days, +is the most wide-spread and common. + +2. The quartan fever is due to the presence of _Plasmodium malariæ_, +which has an asexual cycle of seventy-two hours, and therefore the fever +recurs every three days. This type is more prevalent in temperate and +sub-tropical regions, but appears to be rare everywhere. + +3. The sub-tertian "æstivo-autumnal," or "pernicious" fever is caused by +_Plasmodium falciparum_. Schizogony usually occurs in the internal +organs, particularly in the spleen, instead of in the peripheral +circulation, as is the case of the tertian and quartan forms. The fever +produced is of an irregular type and the period of schizogony has not +been definitely determined. It is claimed by some that the variations +are due to different species of malignant parasites. + +It is one of the most wide-spread of human diseases, occurring in almost +all parts of the world, except in the polar regions and in waterless +deserts. It is most prevalent in marshy regions. + +So commonplace is malaria that it causes little of the dread inspired by +most of the epidemic diseases, and yet, as Ross says, it is perhaps the +most important of human diseases. Figures regarding its ravages are +astounding. Celli estimated that in Italy it caused an average annual +mortality of fifteen thousand, representing about two million cases. In +India alone, according to Ross (1910) "it has been officially estimated +to cause a mean annual death-rate of five per thousand; that is, to kill +every year, on the average, one million one hundred and thirty +thousand." In the United States it is widespread and though being +restricted as the country develops, it still causes enormous losses. +During the year 1911, "in Alabama alone there were seventy thousand +cases and seven hundred and seventy deaths." The weakening effects of +the disease, the invasion of other diseases due to the attacks of +malaria, are among the very serious results, but they cannot be +estimated. + +Not only is there direct effect on man, but the disease has been one of +the greatest factors in retarding the development of certain regions. +Everywhere pioneers have had to face it, and the most fertile regions +have, in many instances been those most fully dominated by it. Herrick +(1903) has presented an interesting study of its effects on the +development of the southern United States and has shown that some parts, +which are among the most fertile in the world, are rendered practically +uninhabitable by the ravages of malaria. Howard (1909) estimates that +the annual money loss from the disease in the United States is not less +than $100,000,000. + +It was formerly supposed that the disease was due to a miasm, to a +noxious effluvia, or infectious matter rising in the air from swamps. In +other words its cause was, as the name indicated "mal aria," and the +deep seated fear of night air is based largely on the belief that this +miasm was given off at night. Its production was thought to be favored +by stirring of the soil, dredging operations and the like. + +The idea of some intimate connection between malaria and mosquitoes is +not a new one. According to Manson, Lancisi noted that in some parts of +Italy the peasants for centuries have believed that malaria is produced +by the bite of mosquitoes. Celli states that one not rarely hears from +such peasants the statement that "In such a place, there is much fever, +because it is full of mosquitoes." Koch points out that in German East +Africa the natives call malaria and the mosquito by the same name, +_Mbù_. The opinion was not lacking support from medical men. Celli +quotes passages from the writings of the Italian physician, Lancisi, +which indicate that he favored the view in 1717. + +Dr. Josiah Nott is almost universally credited with having supported the +theory, in 1848, but as we have already pointed out his work has been +misinterpreted. The statements of Beauperthuy, (1853) were more +explicit. + +The clearest early presentation of the circumstantial evidence in favor +of the theory of mosquito transmission was that of A. F. A. King, an +American physician, in 1883. He presented a series of epidemiological +data and showed "how they may be explicable by the supposition that the +mosquito is the real source of the disease, rather than the inhalation +or cutaneous absorption of a marsh vapor." We may well give the space to +summarizing his argument here for it has been so remarkably +substantiated by subsequent work: + +1. Malaria, like mosquitoes, affects by preference low and moist +localities, such as swamps, fens, jungles, marshes, etc. + +2. Malaria is hardly ever developed at a lower temperature than 60° +Fahr., and such a temperature is necessary for the development of the +mosquito. + +3. Mosquitoes, like malaria, may both accumulate in and be obstructed by +forests lying in the course of winds blowing from malarious localities. + +4. By atmospheric currents malaria and mosquitoes are alike capable of +being transported for considerable distances. + +5. Malaria may be developed in previously healthy places by turning up +the soil, as in making excavations for the foundation of houses, tracks +for railroads, and beds for canals, because these operations afford +breeding places for mosquitoes. + +6. In proportion as countries, previously malarious, are cleared up and +thickly settled, periodical fevers disappear, because swamps and pools +are drained so that the mosquito cannot readily find a place suitable to +deposit her eggs. + +7. Malaria is most dangerous when the sun is down and the danger of +exposure after sunset is greatly increased by the person exposed +sleeping in the night air. Both facts are readily explicable by the +mosquito malaria theory. + +8. In malarial districts the use of fire, both indoors and to those who +sleep out, affords a comparative security against malaria, because of +the destruction of mosquitoes. + +9. It is claimed that the air of cities in some way renders the poison +innocuous, for, though a malarial disease may be raging outside, it does +not penetrate far into the interior. We may easily conceive that +mosquitoes, while invading cities during their nocturnal pilgrimages +will be so far arrested by walls and houses, as well as attracted by +lights in the suburbs, that many of them will in this way be prevented +from penetrating "far into the interior." + +10. Malarial diseases and likewise mosquitoes are most prevalent toward +the latter part of summer and in the autumn. + +11. Various writers have maintained that malaria is arrested by canvas +curtains, gauze veils and mosquito nets and have recommended the rise of +mosquito curtains, "through which malaria can seldom or never pass." It +can hardly be conceived that these intercept marsh-air but they +certainly do protect from mosquitoes. + +12. Malaria spares no age, but it affects infants much less frequently +than adults, because young infants are usually carefully housed and +protected from mosquito inoculation. + +Correlated with the miasmatic theory was the belief that some animal or +vegetable organism which lived in marshes, produced malaria, and +frequent searches were made for it. Salisbury (1862) thought this +causative organism to be an alga, of the genus _Palmella_; others +attributed it to certain fungi or bacteria. + +In 1880, the French physician, Laveran, working in Algeria, discovered +an amÅ“boid organism in the blood of malarial patients and definitely +established the parasitic nature of this disease. Pigmented granules had +been noted by Meckel as long ago as 1847, in the spleen and blood of a +patient who had died of malaria, and his observations had been +repeatedly verified, but the granules had been regarded as degeneration +products, and the fact that they occurred in the body of a foreign +organism had been overlooked. + +Soon after the discovery of the parasites in the blood, Gerhardt (1884) +succeeded in transferring the disease to healthy individuals by +inoculation of malarious blood, and thus proved that it is a true +infection. This was verified by numerous experimenters and it was found +that inoculation with a very minute quantity of the diseased blood would +not only produce malaria but the particular type of disease. + +Laveran traced out the life cycle of the malarial parasite as it occurs +in man. The details as we now know them and as they are illustrated by +the accompanying figure 125, are as follows: + +The infecting organism or _sporozoite_, is introduced into the +circulation, penetrates a red blood corpuscle, and forms the amÅ“boid +_schizont_. This lives at the expense of the corpuscle and as it +develops there are deposited in its body scattered black or reddish +black particles. These are generally called melanin granules, but are +much better referred to as hæmozoin, as they are not related to +melanin. The hæmozoin is the most conspicuous part of the parasite, a +feature of advantage in diagnosing from unstained preparations. + +[Illustration: 125. Life cycle of the malaria parasite. Adapted from +Leuckart's chart, by Miss Anna Stryke.] + +As the schizont matures, its nucleus breaks up into a number of daughter +nuclei, each with a rounded mass of protoplasm about it, and finally the +corpuscles are broken down and these rounded bodies are liberated in the +plasma as _merozoites_. These merozoites infect new corpuscles and thus +the asexual cycle is continued. The malarial paroxysm is coincident with +sporulation. + +As early as Laveran's time it was known that under conditions not yet +determined there are to be found in the blood of malarious patients +another phase of the parasite, differing in form according to the type +of the disease. In the pernicious type these appear as large, +crescent-shaped organisms which have commonly been called "crescents." +We now know that these are sexual forms. + +When the parasite became known there immediately arose speculations as +to the way in which it was transferred from man to man. It was thought +by some that in nature it occurred as a free-living amÅ“ba, and that +it gained access to man through being taken up with impure water. +However, numerous attempts to infect healthy persons by having them +drink or inhale marsh water, or by injecting it into their circulation +resulted in failure, and influenced by Leuckart's and Melnikoff's work +on _Dipylidium_, that of Fedtschenko on _Dracunculus_, and more +especially by that of Manson on _Filaria_, search was made for some +insect which might transfer the parasite. + +Laveran had early suggested that the rôle of carrier might be played by +the mosquito, but Manson first clearly formulated the hypothesis, and it +was largely due to his suggestions that Ross in India, undertook to +solve the problem. With no knowledge of the form or of the appearance in +this stage, or of the species of mosquito concerned, Ross spent almost +two and a half years of the most arduous work in the search and finally +in August, 1897, seventeen years after the discovery of the parasite in +man, he obtained his first definite clue. In dissecting a +"dappled-winged mosquito," "every cell was searched and to my intense +disappointment nothing whatever was found, until I came to the insect's +stomach. Here, however, just as I was about to abandon the examination, +I saw a very delicate circular cell, apparently lying amongst the +ordinary cells of the organ and scarcely distinguishable from them. On +looking further, another and another similar object presented itself. I +now focused the lens carefully on one of these, and found that it +contained a few minute granules of some black substance, exactly like +the pigment of the parasite of malaria. I counted altogether twelve of +these cells in the insect." + +Further search showed that "the contents of the mature pigment cells did +not consist of clear fluid but of a multitude of delicate, thread-like +bodies which on the rupture of the parent cell, were poured into the +body cavity of the insect. They were evidently spores." + +With these facts established, confirmation and extension of Ross's +results quickly followed, from many different sources. We cannot trace +this work in detail but will only point out that much of the credit is +due to the Italian workers, Grassi, Bignami, and Bastianelli, and to +Koch and Daniels. + +It had already been found that when fresh blood was mounted and properly +protected against evaporation, a peculiar change occurred in these +crescents after about half an hour's time. From certain of them there +were pushed out long whip-like processes which moved with a very active, +lashing movement. The parasite at this stage is known as the +"flagellated body." Others, differing somewhat in details of structure, +become rounded but do not give off "flagella." + +The American worker, MacCallum (1897), in studying bird malaria as found +in crows, first recognized the true nature of these bodies. He regarded +them as sexual forms and believed that the so-called flagella played the +part of spermatozoa. Thus, the "flagellated body" is in reality a +_microgametoblast_, producing _microgametes_, or the male sexual +element, while the others constitute the _macrogametes_, or female +elements. + +It was found that when blood containing these sexual forms was sucked up +by an Anopheline mosquito and taken into its stomach, a microgamete +penetrated and fertilized a macrogamete in a way analogous to what takes +place in the fertilization of the egg in higher forms. The resultant, +mobile organism is known as the _migratory ookinete_. In this stage the +parasite bores through the epithelial lining of the "stomach" +(mid-intestine) of the mosquito and becomes encysted under the muscle +layers. Here the _oocyst_, as it is now known, matures and breaks up +into the body cavity and finally its products come to lie in the +salivary glands of the mosquito. Ten to twelve days are required for +these changes, after which the mosquito is infective, capable of +introducing the parasite with its saliva, when feeding upon a healthy +person. + +Thus the malarial parasite is known to have a double cycle, an +alternation of generations, of which the asexual stage is undergone in +man, the sexual in certain species of mosquitoes. The mosquito is +therefore the definitive host rather than the _intermediate_, as usually +stated. + +The complicated cycle may be made clearer by the diagram of Miss Stryke +(1912) which, by means of a double-headed mosquito (fig. 126) endeavors +to show how infection takes place through the biting of the human +victim, (at A), in whom asexual multiplication then takes place, and how +the sexual stages, taken up at B in the diagram, are passed in the body +of the mosquito. + +[Illustration: 126. Life cycle of the malarial parasite. After Miss Anna +Stryke.] + +The experimental proof that mosquitoes of the Anopheline group are +necessary agents in the transmission of malaria was afforded in 1900 +when two English physicians, Drs. Sambon and Low lived for the three +most malarial months in the midst of the Roman Campagna, a region +famous for centuries as a hot-bed of malaria. The two experimenters +moved about freely throughout the day, exposed themselves to rains and +all kinds of weather, drank marsh water, slept exposed to the marsh air, +and, in short, did everything which was supposed to cause malaria, +except that they protected themselves thoroughly from mosquito bites, +retiring at sunset to a mosquito-proof hut. Though they took no quinine +and all of their neighbors suffered from malaria, they were absolutely +free from the disease. + +To complete the proof, mosquitoes which had fed in Rome on malarious +patients were sent to England and allowed to bite two volunteers, one of +them Dr. Manson's own son, who had not been otherwise exposed to the +disease. Both of these gentlemen contracted typical cases of malaria and +the parasites were to be found in abundance in their blood. + +[Illustration: 127. Eggs of Anopheles. After Howard.] + +Since that time there have been many practical demonstrations of the +fact that malaria is transmitted exclusively by the bite of mosquitoes +and that the destruction of the mosquitoes means the elimination of the +disease. + +We have said that the malarial parasite is able to undergo its +development only in certain species of mosquitoes belonging to the +Anopheline group. It is by no means certain that all of this group even, +are capable of acting as the definitive host of the parasites, and much +careful experiment work is still needed along this line. In the United +States, several species have been found to be implicated, _Anopheles +quadrimaculatus_ and _Anopheles crucians_ being the most common. The +characteristics of these species and the distinctions between them and +other mosquitoes will be discussed in Chapter XII. + +In antimalarial work it is desirable to distinguish the anopheline +mosquitoes from the culicine species in all stages. The following +tabulation presents the more striking distinctions between the groups as +represented in the United States. + +_Anopheles_ _Culex, Aedes, etc_. + +_Eggs_: Laid singly in small Deposited in clumps in the +numbers upon the surface of the form of a raft (Culex group) or +water. Eggs lie upon their sides deposited singly in the water or +and float by means of lateral on the ground in places which +expansions (fig. 127). may later be submerged. + +_Larva_: When at rest floats in When at rest (with few exceptions) +a horizontal position beneath the floats suspended in an +surface film. No respiratory oblique or vertical position, or +tube but instead a flattened more rarely nearly horizontal, +area on the eighth abdominal with the respiratory tube in +segment into which the two contact with the surface film +spiracles open (fig. 128). (fig. 128). + +_Adults_: Palpi in both sexes Palpi short in the female, in +nearly or quite as long as the the male usually elongate. +proboscis. Proboscis projecting Proboscis projects forward at an +forward nearly on line with the angle with the axis of the body. +axis of the body. When at rest When at rest on a vertical wall +on a vertical wall the body is the body is usually held parallel +usually held at an angle with the or the tip of the abdomen inclined +vertical (fig. 128). Wings frequently towards the wall (fig. 128). +spotted (fig. 130). Wings usually not spotted. + +[Illustration: 128. (_a_) Normal position of the larvæ of Culex and +Anopheles in the water. Culex, left; Anopheles, middle; Culex pupa, +right hand figure.] + +These malarial-bearing species are essentially domesticated mosquitoes. +They develop in any accumulation of water which stands for a week or +more. Ponds, puddles, rain barrels, horse troughs, cess-pools, cans, +even the foot-prints of animals in marshy ground may afford them +breeding places. + +[Illustration: 128. (_b_) Normal position of Culex and Anopheles on the +wall.] + +It is clear from what has been said regarding the life cycle of the +malarial parasite that the mosquito is harmless if not itself diseased. +Hence malarial-bearing species may abound in the neighborhood where +there is no malaria, the disease being absent simply because the +mosquitoes are uninfected. Such a locality is potentially malarious and +needs only the introduction of a malarial patient who is exposed to the +mosquitoes. It is found that such patients may harbor the parasites in +their blood long after they are apparently well and thus may serve as a +menace, just as do the so-called typhoid carriers. In some malarious +regions as high as 80-90 per cent of the natives are such +malaria-carriers and must be reckoned with in antimalaria measures. + +Based upon our present day knowledge of the life cycle of the malarial +parasite the fight against the disease becomes primarily a problem in +economic entomology,--it is a question of insect control, in its +broadest interpretation. + +[Illustration: 129. Larva of Anopheles. After Howard.] + +The lines of defence and offence against the disease as outlined by +Boyce (1909) are: + + 1. Measures to avoid the reservoir (man): + + Segregation. + + Screening of patients. + + 2. Measures to avoid Anopheles: + + Choice of suitable locality, when possible. + + Screening of houses and porches. + + Sleeping under mosquito nets. + + 3. Measures to exterminate the Anopheles: + + Use of natural enemies. + + Use of culicides, oiling ponds, etc. + + Drainage and scavenging to destroy breeding places. + + Enforcement of penalties for harboring larvæ or keeping stagnant + water. + + Educational methods. + + 4. Systematic treatment with quinine to exterminate the parasites. + + +MOSQUITOES AND YELLOW FEVER + +Yellow fever was until recently one of the most dreaded of epidemic +diseases. It is an acute, specific and infectious disease, +non-contagious in character but occurring in epidemics, or endemics, +within a peculiarly limited geographical area. It is highly fatal, but +those who recover are generally immune from subsequent attacks. + +It is generally regarded as an American disease, having been found by +Cortez, in Mexico, and being confined principally to the American +continents and islands. It also occurs in Africa and attempts have been +made to show that it was originally an African disease but there is not +sufficient evidence to establish this view. + +There have been many noted outbreaks in the United States. Boston +suffered from it in 1691 and again in 1693; New York in 1668 and as late +as 1856; Baltimore in 1819. In 1793 occurred the great epidemic in +Philadelphia, with a death rate of one in ten of the population. In the +past century it was present almost every year in some locality of our +Southern States, New Orleans being the greatest sufferer. In the latter +city there were 7848 deaths from the disease in 1853, 4854 in 1858, and +4046 in 1878. The last notable outbreak was in 1905. Reed and Carroll +(1901) estimated that during the period from 1793 to 1900 there had not +been less than 500,000 cases in the United States. + +[Illustration: 130. Anopheles quadrimaculatus, male and female, +(×3½). After Howard.] + +As in the case of the plague, the most stringent methods of control +proved ineffective and helplessness, almost hopelessness marked the +great epidemics. A vivid picture of conditions is that given by Mathew +Cary, 1793 (quoted by Kelly, 1906) in "A Short Account of the Malignant +Fever Lately Prevalent in Philadelphia." + +[Illustration: 131. Anopheles punctipennis. Female, (×4). After Howard.] + +"The consternation of the people of Philadelphia at this period was +carried beyond all bounds. Dismay and affright were visible in the +countenance of almost every person. Of those who remained, many shut +themselves in their houses and were afraid to walk the streets. * * * +The corpses of the most respectable citizens, even those who did not die +of the epidemic, were carried to the grave on the shafts of a chair +(chaise), the horse driven by a negro, unattended by friends or +relative, and without any sort of ceremony. People hastily shifted their +course at the sight of a hearse coming toward them. Many never walked on +the footpath, but went into the middle of the streets to avoid being +infected by passing by houses wherein people had died. Acquaintances and +friends avoided each other in the streets and only signified their +regard by a cold nod. The old custom of shaking hands fell into such +disuse that many shrunk back with affright at even the offer of the +hand. A person with a crape, or any appearance of mourning was shunned +like a viper. And many valued themselves highly on the skill and address +with which they got to the windward of every person they met. Indeed, it +is not probable that London, at the last stage of the plague, exhibited +stronger marks of terror than were to be seen in Philadelphia from the +24th or 25th of August until pretty late in September." + +[Illustration: 132. Anopheles crucians. Female (×4). After Howard.] + +Such was the condition in Philadelphia in 1793 and, as far as methods of +control of the disease were concerned, there was practically no advance +during the last century. The dominant theory was that yellow fever was +spread by _fomites_, that is, exposed bedding, clothing, baggage, and +the like. As late as 1898 a bulletin of the United States Marine +Hospital Service stated: + +"While yellow fever is a communicable disease, it is not contagious in +the ordinary acceptance of the term, but is spread by the infection of +places and articles of bedding, clothing, and furniture." + +Based upon this theory, houses, baggage, freight, even mail, were +disinfected, and the most rigid quarantine regulations were enforced. +The hardships to which people of the stricken regions were subjected and +the financial losses are incalculable. And withal, the only efficient +check upon the disease seemed to be the heavy frosts. It was found that +for some reason, the epidemic abated with cold weather,--a measure +beyond human control. + +[Illustration: 133. Culex sollicitans. Female (×4). After Howard.] + +It is not strange that among the multitude of theories advanced to +explain the cause and method of dissemination of the disease there +should be suggestions that yellow fever was transmitted by the mosquito. +We have seen that Beauperthuy (1855) clearly urged this theory. + +More detailed, and of the greatest influence in the final solution of +the problem were the arguments of Dr. Cárlos Finlay, of Havana. In 1881, +in a paper presented before the "Real Academia de Ciencias Médicas, +FÃsicas y Naturales de la Habana," he said: + +"I feel convinced that any theory which attributes the origin and the +propagation of yellow fever to atmospheric influences, to miasmatic or +meteorological conditions, to filth, or to the neglect of general +hygienic precautions, must be considered as utterly indefensible." + +He postulated the existence of a material transportable substance +causing yellow fever,--"something tangible which requires to be conveyed +from the sick to the healthy before the disease can be propagated" and +after discussing the peculiarities of the spread of the disease and the +influence of meteorological conditions, he decides that the carriers of +the disease must be sought among insects. He continues: + +"On the other hand, the fact of yellow fever being characterized both +clinically and (according to recent findings) histologically, by lesions +of the blood vessels and by alterations of the physical and chemical +conditions of the blood, suggested that the insect which should convey +the infectious particles from the patient to the healthy should be +looked for among those which drive their sting into blood vessels in +order to suck human blood. Finally, by reason of other considerations +which need not be stated here, I came to think that the mosquito might +be the transmitter of yellow fever." + +"Assimilating the disease to small-pox and to vaccination, it occurred +to me that in order to inoculate yellow fever it would be necessary to +pick out the inoculable material from within the blood vessels of a +yellow fever patient and to carry it likewise into the interior of a +blood vessel of a person who was to be inoculated. All of which +conditions the mosquito satisfies most admirably through its bite." + +In the course of his study of the problem, Finlay made detailed studies +of the life history and habits of the common mosquitoes at Havana, and +arrived at the conclusion that the carrier of the yellow fever was the +_Culex mosquito_ or _Aëdes calopus_, as it is now known. With this +species he undertook direct experimental tests, and believed that he +succeeded in transmitting the disease by the bite of infected mosquitoes +in three cases. Unfortunately, possibility of other exposure was not +absolutely excluded, and the experiments attracted little attention. + +Throughout the next twenty years Finlay continued his work on yellow +fever, modifying his original theory somewhat as time went on. Among his +later suggestions was that in the light of Smith's work on Texas fever, +his theory must be "somewhat modified so as to include the important +circumstance that the faculty of transmitting the yellow fever germ need +not be limited to the parent insect, directly contaminated by stinging a +yellow fever patient (or perhaps by contact with or feeding from his +discharges), but may be likewise inherited by the next generation of +mosquitoes issued from the contaminated parent." He believed that the +bite of a single mosquito produced a light attack of the disease and was +thus effective in immunizing the patient. Throughout the period, many +apparently successful attempts to transmit the disease by mosquitoes +were made. In the light of present day knowledge we must regard these as +defective not only because possibility of other infection was not +absolutely excluded but because no account was taken of the incubation +period within the body of the mosquito. + +In 1900, while the American army was stationed in Cuba there occurred an +epidemic of yellow fever and an army medical board was appointed for +"the purpose of pursuing scientific investigations with reference to the +acute infectious diseases prevalent on the island." This was headed by +Walter Reed and associated with him were James Carroll, Jesse W. Lazear +and Aristides Agramonte, the latter a Cuban immune. For a detailed +summary of this work the lay reader cannot do better than read Dr. +Kelly's fascinating biography "Walter Reed and Yellow Fever." + +Arriving at the army barracks near Havana the Commission first took up +the study of _Bacillus icteroides_, the organism which Sanarelli, an +Italian physician, had declared the causative agent in yellow fever. +They were unable to isolate this bacillus either from the blood during +life or from the blood and organs of cadavers and therefore turned their +attention to Finlay's theory of the propagation of yellow fever by means +of the mosquito. In this work they had the unselfish and enthusiastic +support of Dr. Finlay himself, who not only consulted with them and +placed his publications at their disposal, but furnished eggs from which +their experimental mosquitoes were obtained. Inoculations of eleven +non-immunes through the bite of infected mosquitoes were made, and of +these, two gave positive results. The first of the two was Dr. Carroll +who allowed himself to be bitten by a mosquito which had been caused to +feed upon four cases of yellow fever, two of them severe and two mild. +The first patient had been bitten twelve days before. + +Three days after being bitten, Dr. Carroll came down with a typical case +of yellow fever. So severe was the attack that for three days his life +hung in the balance. During his convalescence an incident occurred which +showed how the theory of mosquito transmission of the disease was +generally regarded. We quote from Dr. Kelly: "One of his nurses who came +from Tennessee had had considerable experience with yellow fever, having +indeed, lost her husband and several children from it. One day early in +his illness Dr. Carroll mentioned to her that he had contracted the +disease through the bite of a mosquito, and noticed that she looked +surprised. Some time later, when well enough to look over the daily +records of his condition, he found this entry: 'Says he got his illness +through the bite of a mosquito,--delirious'." + +The second case was that of an American who was bitten by four +mosquitoes, two of which had bitten severe (fatal) cases of yellow fever +twelve days previously, one of which had bitten a severe case (second +day) sixteen days before and one which had bitten a severe case eight +days before. Five days later, the subject developed a well pronounced +but mild case of the disease. + +In the meantime, another member of the Commission, Dr. Lazear, was +accidentally bitten by a mosquito while collecting blood from yellow +fever patients. Five days later he contracted a typical case which +resulted fatally. + +So clear was the evidence from these preliminary experiments that the +commission felt warranted in announcing, October 27, 1900, that, "The +mosquito serves as the intermediate host for the parasite of yellow +fever, and it is highly probable that the disease is only propagated +through the bite of this insect." + +In order to extend the experimental evidence under conditions which +could leave no possibility of infection from other sources, a special +experimental sanitary station, named in honor of the deceased member of +the Commission, was established in an open field near the town of +Quemados, Cuba. Here there were constructed two small buildings known +respectively as the "infected clothing building" and the "infected +mosquito building." + +The infected clothing building, 14 × 20 feet in size, was purposely so +constructed as to exclude anything like efficient ventilation, but was +thoroughly screened to prevent the entrance of mosquitoes. Into this +building were brought sheets, pillow-slips, blankets, etc., contaminated +by contact with cases of yellow fever and their discharges,--many of +them purposely soiled with a liberal quantity of black vomit, urine, and +fecal matter from patients sick with yellow fever. Nothing could better +serve as the fomites which were supposed to convey the dread disease. + +Three non-immunes unpacked these articles, giving each a thorough +handling and shaking in order to disseminate through the air of the room +the specific agent of the disease. They were then used in making up the +beds which the volunteers occupied each night for a period of twenty +days. The experiment was repeated three times, volunteers even sleeping +in the soiled garments of yellow fever victims but in not a single case +was there the slightest symptom of disease. The theory of the spread of +yellow fever by fomites was completely demolished. + +The infected mosquito building, equal in size to its companion, was the +antithesis as far as other features were concerned. It was so +constructed as to give the best possible ventilation, and bedding which +was brought into it was thoroughly sterilized. Like the infected +clothing building it was carefully screened, but in this case it was in +order to keep mosquitoes in it as well as to prevent entrance of others. +Through the middle of the room ran a mosquito-proof screen. + +On December 5, 1900, a non-immune volunteer who had been in the +quarantine camp for fifteen days and had had no other possible exposure, +allowed himself to be bitten by five mosquitoes which had fed on yellow +fever patients fifteen or more days previously. The results were fully +confirmatory of the earlier experiments of the Commission--at the end of +three days, nine and a half hours, the subject came down with a well +marked case of yellow fever. + +In all, ten cases of experimental yellow fever, caused by the bite of +infected mosquitoes were developed in Camp Lazear. Throughout the period +of the disease, other non-immunes slept in the little building, +separated from the patient only by the mosquito-proof screen, but in no +circumstances did they suffer any ill effects. + +It was found that a yellow fever patient was capable of infecting +mosquitoes only during the first three or four days after coming down +with the disease. Moreover, after the mosquito has bitten such a +patient, a period of at least twelve days must elapse before the insect +is capable of transmitting the disease. + +Once the organism has undergone its twelve day development, the mosquito +may remain infective for weeks. In experiments of the Commission, two of +the mosquitoes transmitted the disease to a volunteer fifty-seven days +after their contamination. No other volunteers presenting themselves, +one of these mosquitoes died the sixty-ninth and one the seventy-first +day after their original contamination, without it being determined +whether they were still capable of transmitting the disease. + +So carefully carried out was this work and so conclusive were the +results that Dr. Reed was justified in writing: + +"Six months ago, when we landed on this island, absolutely nothing was +known concerning the propagation and spread of yellow fever--it was all +an unfathomable mystery--but today the curtain has been drawn--its mode +of propagation is established and we know that a case minus mosquitoes +is no more dangerous than one of chills and fever." + +The conclusions of the Commission were fully substantiated by numerous +workers, notably Dr. Guiteras of the Havana Board of Health, who had +taken a lively interest in the work and whose results were made known in +1901, and by the Brazilian and French Commission at Sao Paulo, Brazil, +in 1903. + +Throughout the work of the Army Commission and down to the present time +many fruitless efforts have been made to discover the specific organism +of yellow fever. It was clearly established that the claims of Sanarelli +for _Bacillus icteroides_ were without foundation. It was found, too, +that whatever the infective agent might be it was capable of passing +through a Berkefeld filter and thus belongs to the puzzling group of +"filterable viruses." It was further found that the virus was destroyed +by heating up to 55° C for ten minutes. It is generally believed that +the organism is a Protozoan. + +The question of the hereditary transmission of the yellow fever organism +within the mosquito was left unsettled by the Army Commission, though, +as we have seen, it was raised by Finlay. Marchoux and Simond, of the +French Commission devoted much attention to this phase of the problem +and basing their conclusions on one apparently positive case, they +decided that the disease could be transmitted through the egg of an +infected _Aëdes calopus_ to the second generation and thence to man. The +conclusion, which is of very great importance in the control of yellow +fever, has not been verified by other workers. + +Once clearly established that yellow fever was transmitted solely by +mosquitoes, the question of the characteristics, habits, and +geographical distribution of the insect carrier became of vital +importance. + +_Aëdes calopus_, more commonly known as _Stegomyia fasciata_ or +_Stegomyia calopus_ (fig. 134) is a moderate sized, rather strikingly +marked mosquito. The general color is dark-brown or reddish-brown, but +the thorax has a conspicuous broad, silvery-white curved line on each +side, with two parallel median silvery lines. Between the latter there +is a slender, broken line. The whole gives a lyre-shaped pattern to the +thorax. The abdomen is dark with silvery-white basal bands and silvery +white spots on each side of the abdominal segments. Legs black with +rings of pure white at the base of the segments. + +Size of the female 3.3 to 5 mm.; male 3 to 4.5 mm. + +[Illustration: 134. The yellow fever mosquito (Aëdes calopus). (×7). +After Howard.] + +It is preeminently a domesticated species, being found almost +exclusively about the habitation of man. "Its long association with man +is shown by many of its habits. It approaches stealthily from behind. It +retreats upon the slightest alarm. The ankles and, when one is sitting +at a table or desk, the underside of the hands and wrists are favorable +points of attack. It attacks silently, whereas other mosquitoes have a +piping or humming note. The warning sound has doubtless been suppressed +in the evolutionary process of its adaptation to man. It is extremely +wary. It hides whenever it can, concealing itself in garments, working +into the pockets, and under the lapels of coats, and crawling up under +the clothes to bite the legs. In houses, it will hide in dark corners, +under picture moldings and behind the heads of old-fashioned bedsteads. +It will enter closets and hide in the folds of garments."--Howard. + +It was claimed by the French Commission, and subsequently often stated +in discussions of the relation of the mosquito to yellow fever that the +mature _Aëdes calopus_ will bite only at night. If this were true it +would be of the greatest importance in measures to avoid the disease. +Unfortunately, the claim was illy founded and numerous workers have +clearly established that the exact converse is more nearly true, this +mosquito being pre-eminently a day species, feeding most actively in +early morning, about sunrise, and late in the afternoon. On cloudy days +it attacks at any time during the day. Thus there is peril in the +doctrine that infected regions may be visited with perfect safety during +the daytime and that measures to avoid the mosquito attack need be taken +only at night. + +[Illustration: 135_a_. Aëdes calopus. Pupa. After Howard.] + +Dr. Finlay maintained that the adult, even when starved, would not bite +when the temperature was below 23° C, but subsequent studies have shown +that this statement needs modification. The French Commission, working +at Rio de Janeiro, found that _Aëdes calopus_ would bite regularly at +temperatures between 22° and 25° and that the optimum temperature was +between 27° and 30° C, but their experiments led them to believe that it +would bite in nature at a temperature as low as 17° C. + +The yellow fever mosquito breeds in cisterns, water barrels, pitchers +and in the various water receptacles about the house. In our own +Southern States it very commonly breeds in the above-ground cisterns +which are in general use. Often the larvæ (fig. 135b) are found in +flower vases, or even in the little cups of water which are placed under +the legs of tables to prevent their being overrun by ants. They have +been repeatedly found breeding in the holy water font in churches. In +short, they breed in any collection of water in close proximity to the +dwellings or gathering places of man. + +The life cycle under favorable conditions is completed in from twelve to +fifteen days. These figures are of course very dependent upon the +temperature. The Army Commission in Cuba found that the cycle might be +completed in as brief a period as nine and a half days. Under less +favorable conditions it may be greatly lengthened. + +The adults are long lived. We have seen that during the experimental +work in Cuba specimens were kept in captivity for sixty-nine and +seventy-one days, respectively, and that they were proved to retain +their infectivity for at least fifty-seven days. Dr. Guiteras +subsequently kept an infected adult for one hundred and fifty-four days. + +Low temperatures have a very great effect not only on development, but +on the activity and even life of the adults. Long before the method of +transmission of yellow fever was discovered it was well known that the +epidemics were brought to a close by heavy frosts, and it is now known +that this is due to the killing of the mosquitoes which alone could +spread the disease. + +[Illustration: 135_b_. Aëdes calopus; larva. (×7). After Howard.] + +_Aëdes calopus_ has a very wide distribution since, as Howard says, +being a domestic mosquito, having a fairly long life in the adult stage, +and having the custom of hiding itself in the most ingenious ways, it is +particularly subject to carriage for long distances on board vessels, in +railway trains, even packed in baggage. In general, its permanent +distribution is from 40 degrees north latitude to 40 degrees south +latitude (Brumpt), in a belt extending around the world. In the United +States it breeds in most of our Southern States. + +Thus, as in the case of malaria, there are many places where the insect +carrier is abundant but where yellow fever does not occur. Such, for +instance, are Hawaii, Australia and Asia. An outbreak may occur at any +time that a patient suffering from the disease is allowed to enter and +become a source of infection for the mosquitoes. In this connection +various writers have called attention to the menace from the Panama +Canal. When it is completed, it will allow of direct passage from +regions where yellow fever is endemic and this will greatly increase the +possibility of its introduction into these places where it is now +unknown. The result, with a wholly non-immune population, would be +appalling. + +On the other hand, there are places wholly outside of the normal range +of _Aëdes calopus_ where the disease has raged. Such are New York, +Boston, and even Philadelphia, which have suffered notable epidemics. +These outbreaks have been due to the introduction of infected mosquitoes +during the heat of summer, when they have not only conveyed the disease +but have found conditions favorable for their multiplication. Or, +uninfected mosquitoes have been thus accidentally brought in and +developed in large numbers, needing then only the accidental +introduction of cases of the disease to start an epidemic. + +Methods of control of various diseases have been revolutionized by the +discovery that they were insect-borne, but in no other case has the +change been as radical or the results as spectacular as in the case of +yellow fever. The "shot-gun quarantine," the sufferings and horrors, the +hopelessness of fighting absolutely blindly have given way to an +efficient, clear-cut method of control, based upon the knowledge that +the disease is carried from man to man solely by the mosquito, _Aëdes +calopus_. The lines of defense and offense are essentially as follows: + +In the first place, when a case of yellow fever occurs, stringent +precautions must be adopted to prevent the infection of mosquitoes and +the escape of any already infected. This means that the patient must be +removed to a mosquito-proof room, or ward beyond reach of the insects, +and that the infected room must be thoroughly fumigated at once, to kill +the mosquitoes hiding within it. All cracks and openings should be +closed with strips of paper and fumigation with burning sulphur or +pyrethrum carefully carried out. + +It should be remembered that if the first case noted is that of a +resident rather than imported, it means that the mosquito carriers +became infected more than two weeks before the case was diagnosed, for +as we have seen, the germ must undergo a twelve-day period of +development within its insect host. Therefore a careful search must be +made for mild cases which, though unrecognized, may serve as foci for +the spread of the disease. + +In face of a threatened epidemic one of the most essential measures is +to educate the citizens and to gain their complete coöperation in the +fight along modern lines. This may be done through the schools, the +pulpit, places of amusement, newspapers and even bulletin boards. + +Emphasis should be placed on the necessity of both non-immunes and +immunes using mosquito curtains, and in all possible ways avoiding +exposure to the mosquitoes. + +Then the backbone of the fight must be the anti-mosquito measures. In +general, these involve screening and fumigating against adults, and +control of water supply, oiling, and drainage against the larvæ. The +region involved must be districted and a thorough survey undertaken to +locate breeding places, which must, if possible, be eradicated. If they +are necessary for water supplies, such as casks, or cisterns, they +should be carefully screened to prevent access of egg-laying adults. + +The practical results of anti-mosquito measures in the fight against +yellow fever are well illustrated by the classic examples of the work in +Havana, immediately following the discoveries of the Army Commission and +by the stamping out of the New Orleans epidemic in 1905. + +The opportunities for an immediate practical application of the theories +of the Army Commission in Havana were ideal. The city had always been a +hotbed of yellow fever and was the principal source from which the +disease was introduced year after year into our Southern States. It was +under martial law and with a military governor who was himself a +physician and thoroughly in sympathy with the views of the Commission, +the rigid enforcement of the necessary regulations was possible. The +story of the first campaign has been often told, but nowhere more +clearly than in Dr. Reed's own account, published in the _Journal of +Hygiene_ for 1902. + +Closer home was the demonstration of the efficacy of these measures in +controlling the yellow fever outbreak in New Orleans in 1905. During the +spring and early summer of the year the disease had, unperceived, gained +a firm foothold in that city and when, in early July the local Board of +Health took cognizance of its existence, it was estimated that there had +been in the neighborhood of one hundred cases. + +Conditions were not as favorable as they had been under martial law in +Havana for carrying on a rigid fight along anti-mosquito lines. The +densely populated city was unprepared, the public had to be educated, +and an efficient organization built up. The local authorities actively +began a general fight against the mosquito but in spite of their best +efforts the disease continued to spread. It was recognized that more +rigid organization was needed and on August 12th the United States +Public Health and Marine Hospital Service was put in absolute charge of +the fight. Up to this time there had been one hundred and forty-two +deaths from a total of nine hundred and thirteen cases and all of the +conditions seemed to threaten an outbreak to exceed the memorable one of +1878 when, as we have seen there were four thousand and forty-six +deaths. + +With the hearty coöperation of the citizens,--physicians and laymen +alike,--the fight was waged and long before frost or any near approach +thereto the disease was stamped out,--a thing unheard of in previous +epidemics. The total loss of life was four hundred and sixty--about 11 +per cent as great as that from the comparable epidemic of 1878. If the +disease had been promptly recognized and combated with the energy which +marked the fight later in the summer, the outbreak would have made +little headway and the great proportion of these lives would have been +saved. + + + + +CHAPTER IX + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA (Continued) + + +INSECTS AND TRYPANOSOMIASES + +By trypanosomiasis is meant a condition of animal parasitism, common to +man and the lower animals, in which trypanosomes, peculiar flagellate +protozoa, infest the blood. Depending upon the species, they may be +harmless, producing no appreciable ill-effect, or pathogenic, giving +rise to conditions of disease. A number of these are known to be +transferred by insects. + +In order that we may consider more fully the developmental stage of +these parasites within their insect host, it is necessary that we +describe briefly the structure of the blood-inhabiting stage. + +[Illustration: 136. Trypanosoma brucei. After Bruce.] + +The trypanosomes are elongated, usually pointed, flagellated protozoa +(fig. 136) in which the single flagellum, bent under the body, forms the +outer limit of a delicate undulating membrane. It arises near one end of +the organism from a minute centrosome-like body which is known as the +blepheroplast, and at the opposite end extends for a greater or less +distance as a free flagellum. Enclosing, or close beside the +blepheroplast is the small kinetonucleus. The principal nucleus, round +or oval in form, is situated near the center of the body. Asexual +reproductions occurs in this stage, by longitudinal fission, the nucleus +and the blepheroplast dividing independently of one another. From the +blepheroplast of one of the daughter cells a new flagellum is formed. + +Among the pathogenic species are to be found the causative organisms of +some of the most serious diseases of domestic animals and even of man. +It is probable that these pathogenic species secrete a specific poison. +The majority of them are tropical in distribution. + +Though we are concerned especially with the species which infest man, we +shall first consider two of the trypanosomes of lower animals, known +long before any of those of man had been found. + + +FLEAS AND LICE AS CARRIERS OF TRYPANOSOMA LEWISI.--_Trypanosoma lewisi_, +the first mammalian trypanosome known, is to be found in the blood of +wild rats. Like its host, it appears to be cosmopolitan in distribution, +having been reported from several localities in the United States, +Brazil, Argentine, England, Germany, France, Italy, Russia, Asia and +Africa. + +This species is usually regarded as non-pathogenic, but in experimental +work, especially with white rats, heavy infestations often result +fatally, and naturally infested specimens sometimes show evidence of +injury. Rats which have been infested exhibit at least temporary +immunity against new infection. + +_Trypanosoma lewisi_ is transmitted from rat to rat by fleas and by +lice. Rabinowitsch and Kempner (1899) first found that healthy rats +which were kept with infested rats, showed trypanosomes in their blood +after about two weeks. They found the trypanosomes in the alimentary +canal of fleas which had fed on the diseased rats. On teasing such fleas +in physiological salt solution and inoculating them into fresh rats they +were able to produce the infection. Finally, they showed that the fleas +which had fed upon infested rats were able to carry the parasites to +healthy rats. Corresponding experiments with lice were not successful. +Prowazek (1905) found in the rat louse (_Hæmatopinus spinulosus_) +organisms which he regarded as developmental stages of the _Trypanosoma +lewisi_. He believed that the sexual cycle was undergone in this insect. + +Nuttall (1908) readily transmitted the trypanosomes through the agency +of fleas, (_Ceratophyllus fasciatus_ and _Ctenopthalmus agyrtes_). He +believes that these insects are probably the chief transmitters of the +parasite. He was also able to transmit it from diseased to healthy rats +through the agency of the rat louse. He was unable to trace any +developmental stages in the louse and inclined to the opinion that +Prowazek was deceived by the presence of extraneous flagellates such as +are known to exist in a number of blood-sucking arthropods. + +Nuttall concludes that since three distinct kinds of blood-sucking +insects are capable of transmitting _Trypanosoma lewisi_ it appears +doubtful that this flagellate is a parasite of the invertebrate "host" +in the sense claimed by Prowazek and other investigators. + + +TSETSE-FLIES AND NAGANA--One of the greatest factors in retarding the +development of certain regions of Africa has been the presence of a +small fly, little larger than the common house-fly. This is the +tsetse-fly, _Glossina morsitans_ (fig. 165) renowned on account of the +supposed virulence of its bite for cattle, horses and other domestic +mammals. + +The technical characteristics of the tsetse-flies, or Glossinas, and +their several species, will be found in a later chapter. We need +emphasize only that they are blood-sucking Muscidæ and that, unlike the +mosquitoes, the sexes resemble each other closely in structure of the +mouth-parts, and in feeding habits. + +In 1894, Colonel David Bruce discovered that the fly was not in itself +poisonous but that the deadly effect of its bite was due to the fact +that it transmitted a highly pathogenic blood parasite, _Trypanosoma +brucei_. This trypanosome Bruce had discovered in the blood of South +African cattle suffering from a highly fatal disease known as "nagana". +On inoculating the blood of infected cattle into horses and dogs he +produced the disease and found the blood teeming with the causative +organism. In the course of his work he established beyond question that +the "nagana" and the tsetse-fly disease were identical. + +Tsetse-flies of the species _Glossina morsitans_, which fed upon +diseased animals, were found capable of giving rise to the disease in +healthy animals up to forty-eight hours after feeding. Wild tsetse-flies +taken from an infected region to a region where they did not normally +occur were able to transmit the disease to healthy animals. It was found +that many of the wild animals in the tsetse-fly regions harbored +_Trypanosoma brucei_ in their blood, though they showed no evidence of +disease. As in the case of natives of malarial districts, these animals +acted as reservoirs of the parasite. Non-immune animals subjected to the +attacks of the insect carrier, quickly succumbed to the disease. + +A question of prime importance is as to whether the insect serves as an +essential host of the pathogenic protozoan or whether it is a mere +mechanical carrier. Bruce inclined to the latter view. He was unable to +find living trypanosomes in the intestines or excrements of the fly or +to produce the disease on the many occasions when he injected the +excrement into healthy animals. Moreover, he had found that the +experimental flies were infective only during the first forty-eight +hours and that if wild flies were taken from the infected region, "kept +without food for three days and then fed on a healthy dog, they never +gave rise to the disease." + +Koch had early described what he regarded as sexual forms from the +intestine of the fly but it remained for Kleine (1909) to experimentally +demonstrate that a part of the life cycle of the parasite was undergone +in the fly. Working with _Glossina palpalis_, he found that for a period +of ten days or longer after feeding on an animal suffering from nagana +it was non-infective, but that then it became infective and was able to +transmit the disease for weeks thereafter. He discovered and described +developmental stages of the parasite within the intestine of the insect. +In other words, the tsetse-fly (in nature, _Glossina morsitans_), serves +as an essential host, within which an important part of the life cycle +of the parasite is undergone. These conclusions were quickly verified by +Bruce and numerous other workers and are no longer open to question. +Klein and Taute are even inclined to think that mechanical transmission +plays practically no rôle in nature, unless the fly is interrupted while +feeding and passes immediately to a new animal. + + +TSETSE-FLIES AND SLEEPING SICKNESS OF MAN--About the beginning of the +present century a hitherto little known disease of man began to attract +great attention on account of its ravages in Uganda and the region of +Victoria Nyanza in South Africa. It was slow, insiduous and absolutely +fatal, characterized in its later stages by dullness, apathy, and +finally absolute lethargy all day long, symptoms which gave it the name +of "sleeping sickness." + +It was soon found that the disease was not a new one but that it had +been known for over a hundred years on the west coast of Africa. Its +introduction into Central and East Africa and its rapid spread have been +attributed primarily to the development of the country, the formation of +new trade routes and the free mingling of native tribes formerly +isolated. It is estimated that in the first ten years of the present +century there were approximately two hundred thousand deaths from the +disease in the Uganda protectorate. In the British province Bugosa, on +the Victoria Nyanza there were thirty thousand deaths in the period from +1902-1905. + +While the disease is peculiarly African there are a number of instances +of its accidental introduction into temperate regions. Slaves suffering +from it were occasionally brought to America in the early part of the +last century and cases have sometimes been imported into England. In +none of the cases did the disease gain a foothold or spread at all to +other individuals. + +In 1902 Dutton described a trypanosome, _T. gambiense_, which he and +Forde had found the year before in the blood of a patient suffering from +a peculiar type of fever in Gambia. In 1902-1903 Castellani found the +same parasite in the cerebro-spinal fluid of sleeping-sickness patients +and definitely reported it as the causative organism of the disease. His +work soon found abundant confirmation, and it was discovered that the +sleeping sickness was but the ultimate phase of the fever discovered by +Dutton and Forde. + +When Castellani made known his discovery of the trypanosome of sleeping +sickness, Brumpt, in France, and Sambon, in England, independently +advanced the theory that the disease was transmitted by the tsetse-fly, +_Glossina palpalis_. This theory was based upon the geographical +distribution and epidemiology of the disease. Since then it has been +abundantly verified by experimental evidence. + +Fortunately for the elucidation of problems relating to the methods of +transfer of sleeping sickness, _Trypanosoma gambiense_ is pathogenic for +many species of animals. In monkeys it produces symptoms very similar to +those caused in man. Bruce early showed that _Glossina palpalis_ "fed on +healthy monkeys eight, twelve, twenty-four and forty-eight hours after +having fed on a native suffering from trypanosomiasis, invariably +transmitted the disease. After three days the flies failed to transmit +it." In his summary in Osler's Modern Medicine, he continues "But this +is not the only proof that these flies can carry the infective agent. On +the lake shore there was a large native population among whom we had +found about one-third to be harboring trypanosomes in their blood. The +tsetse-flies caught on this lake shore, brought to the laboratory in +cages, and placed straightway on healthy monkeys, gave them the disease +in every instance, and furnished a startling proof of the danger of +loitering along the lake shore among those infected flies." + +As in the case of nagana, Bruce and most of the earlier investigators +supposed the transmission of the sleeping sickness trypanosome by +_Glossina palpalis_ to be purely mechanical. The work of Kleine (1909) +clearly showed that for _Trypanosoma gambiense_ as well as for +_Trypanosoma brucei_ the fly served as an essential host. Indeed, Kleine +and many subsequent investigators are inclined to think that there is +practically no mechanical transmission of trypanosomes from animal to +animal by _Glossina_ in nature, and that the many successful experiments +of the earlier investigators were due to the fact that they used wild +flies which already harbored the transformed parasite rather than +directly inoculated it from the blood of the diseased experimental +animals. While the criticism is applicable to some of the work, this +extreme view is not fully justified by the evidence at hand. + +Kleine states (1912) that _Glossina palpalis_ can no longer be regarded +as the sole transmitter of sleeping sickness. Taute (1911) had shown +that under experimental conditions _Glossina morsitans_ was capable of +transferring the disease and Kleine calls attention to the fact that in +German East Africa, in the district of the Rovuma River, at least a +dozen cases of the disease have occurred recently, though only _Glossina +morsitans_ exists in the district. It appears, however, that these cases +are due to a different parasite, _Trypanosoma rhodesiense_. This +species, found especially in north-east Rhodesia and in Nyassaland, is +transferred by _Glossina morsitans_. + +Other workers maintain that the disease may be transmitted by various +blood-sucking flies, or even bugs and lice which attack man. Fülleborn +and Mayer (1907) have shown by conclusive experiments that _Aedes +(Stegomyia) calopus_ may transmit it from one animal to another if the +two bites immediately succeed each other. + +It is not possible that insects other than the tsetse-flies (and only +certain species of these), play an important rôle in the transmission of +the disease, else it would be much more wide-spread. Sambon (1908) +pointed out that the hypothesis that is spread by _Aedes calopus_ is +opposed by the fact that the disease never spread in the Antilles, +though frequently imported there by West African slaves. The same +observation would apply also to conditions in our own Southern States in +the early part of the past century. + +Since _Glossina palpalis_ acts as an essential host of the parasite and +the chief, if not the only, transmitter, the fight against sleeping +sickness, like that against malaria and yellow fever, becomes primarily +a problem in economic entomology. The minutest detail of the +life-history, biology, and habits of the fly, and of its parasites and +other natural enemies becomes of importance in attempts to eradicate the +disease. Here we can consider only the general features of the subject. + +_Glossina palpalis_ lives in limited areas, where the forest and +undergrowth is dense, along the lake shore or river banks. According to +Hodges, the natural range from shore is under thirty yards, though the +distance to which the flies may follow man greatly exceed this. + +It is a day feeder, a fact which may be taken advantage of in avoiding +exposure to its attacks. The young are brought forth alive and +full-grown, one every nine or ten days. Without feeding, they enter the +ground and under favorable conditions, complete their development in a +month or more. + +[Illustration: 137. Sleeping sickness concentration camp in German East +Africa. Report of German Commission.] + +Methods of control of the disease must look to the prevention of +infection of the flies, and to their avoidance and destruction. Along +the first line, much was hoped from temporary segregation of the sick in +regions where the fly was not found. On the assumption that the flies +acted as carriers only during the first two or three days, it was +supposed that even the "fly belts" would become safe within a few days +after the sick were removed. The problem was found to be a much more +difficult one when it was learned that after a given brief period the +fly again became infective and remained so for an indeterminate period. +Nevertheless, isolation of the sick is one of the most important +measures in preventing the spread of the disease into new districts. +Much, too, is being accomplished by moving native villages from the fly +belts. (c.f. fig. 137.) + +All measures to avoid the flies should be adopted. This means locating +and avoiding the fly belts as far as possible, careful screening of +houses, and protection of the body against bites. + +Clearing the jungle along the water courses for some yards beyond the +natural range of the fly has proved a very important measure. Castellani +recommends that the area be one hundred yards and around a village three +hundred yards at least. + +Detailed studies of the parasites and the natural enemies of the +tsetse-fly are being undertaken and may ultimately yield valuable +results. + + +SOUTH AMERICAN TRYPANOSOMIASIS--The tsetse-flies are distinctively +African in distribution and until recently there were no trypanosomes +known to infest man in America. In 1909 Dr. Chagas, of Rio de Janeiro +described a new species, _Trypanosoma cruzi_, pathogenic to man. + +_Trypanosoma cruzi_ is the causative organism of a disease common in +some regions of Brazil, where it is known as "opilacao." It is +especially to be met with in children and is characterized by extreme +anemia, wasting, and stunted development associated with fever, and +enlargement of the thyroid glands. The disease is transmitted by the +bites of several species of assassin-bugs, or Reduviidæ, notably by +_Conorhinus megistus_. The evolution of the parasite within the bug has +been studied especially by Chagas and by Brumpt. From the latter's text +we take the following summary. + +The adult trypanosomes, ingested by a _Conorhinus megistus_, of any +stage, first change into Crithidia-like forms and then those which +remain in the stomach become ovoid and non-motile. Brumpt found these +forms in immense numbers, in a _Conorhinus_ which had been infested +fourteen months before. The forms which pass into the intestine quickly +assume the _Crithidia_ form and continue to develop rapidly under this +form. Some weeks later they evolve into the trypanosome forms, +pathogenic for man. They then pass out with the excrement of the bug and +infect the vertebrate host as soon as they come in contact with any +mucous layer (buccal, ocular or rectal). More rarely they enter through +the epidermis. + +Brumpt showed that the development could take place in three species; +bed-bugs (_Cimex lectularius_, _C. hemipterus_) and in the tick +_Ornithodoros moubata_. The evolution proceeds in the first two species +of bed-bugs as rapidly as in Conorhinus, or even more rapidly, but they +remain infective for a much shorter time and hence Brumpt considers that +they play a much less important rôle in the spread of the disease. + +_Conorhinus megistus_, like related forms in our Southern States, very +commonly frequents houses and attacks man with avidity. Chagas states +that the bites are painless and do not leave any traces. They are +usually inflicted on the lips, or the cheeks and thus the buccal mucosa +of a sleeper may be soiled by the dejections of the insect and the bite +serving as a port of entry of the virus, remain unnoticed. + +The possibility of some of our North American Reduviidæ playing a +similar rôle in the transmission of disease should not be overlooked. + + +LEISHMANIOSES AND INSECTS--Closely related to the trypanosomes is a +group of intracellular parasites which have recently been grouped by +Ross under the genus _Leishmania_. Five species are known to affect man. +Three of these produce local skin infestations, but two of them, +_Leishmania donovani_ and _L. infantum_, produce serious and often fatal +systemic diseases. + +The first of these, that produced by _L. donovani_, is an exceedingly +virulent disease common in certain regions of India and China. It is +commonly known as "Kala-azar," or "dum-dum" fever, and more technically +as tropical leishmaniasis. Patton (1907) believes that the parasite is +transmitted by the bed-bug _Cimex hemipterus_, and has described a +developmental cycle similar to that which can be found in artificial +cultures. On the other hand, Donovan was unable to confirm Patton's work +and believes that the true intermediate host is a Reduviid bug, +_Conorhinus rubrofasciatus_. + +_Leishmania infantum_ is the cause of the so-called infantile splenic +leishmaniasis, occurring in northern Africa, Spain, Portugal, Italy, and +possibly other parts of Europe. The parasite occurs habitually in the +dog and is only accidentally transferred to children. Alvares and da +Silva, in Portugal (according to Brumpt, 1913) have found that the +excrement of a flea from a diseased dog contains flagellates, and they +suggest that the infection may be transmitted by the accidental +inoculation of this excrement by means of the proboscis of the flea, as +has been thought to occur in the case of the plague. To this Brumpt +objects that they and other workers who thought to trace the development +of _Leishmania infantum_ were apparently misled by the presence of a +harmless _Herpetomonas_ which infests dog fleas in all countries, even +where the leishmaniasis is unknown. + +Basile (1910 and 1911) however, carried on numerous experiments +indicating that the disease was transferred from children to dogs and +from dog to dog by the dog flea, and was able to find in the tissues of +the insects forms perfectly identical with those found in children and +in dogs suffering from leishmaniasis. He also found that _Pulex +irritans_ was capable of acting as the carrier. + +Of the cutaneous type of leishmaniasis, the best known is the so-called +"Oriental sore," an ulcerative disease of the skin which is epidemic in +many tropical and subtropical regions. The causative organism is +_Leishmania tropica_, which occurs in the diseased tissues as bodies +very similar to those found in the spleen in cases of Kala-azar. The +disease is readily inoculable and there is no doubt that it may be +transferred from the open sores to abraded surfaces of a healthy +individual by house-flies. It is also believed by a number of +investigators that it may be transferred and directly inoculated by +various blood-sucking insects. + + +TICKS AND DISEASES OF MAN AND ANIMALS + +We have seen that the way to the discoveries of the relations of +arthropods to disease was pointed out by the work of Leuckart and +Melnikoff on the life cycle of _Dipylidium_, and of Fedtschenko and +Manson on that of _Filaria_. They dealt with grosser forms, belonging to +well-recognized parasitic groups. + +This was long before the rôle of any insect as a carrier of pathogenic +micro-organisms had been established, and before the Protozoa were +generally regarded as of importance in the causation of disease. The +next important step was taken in 1889 when Smith and Kilbourne +conclusively showed that the so-called Texas fever of cattle, in the +United States, is due to an intracorpuscular blood parasite transmitted +exclusively by a tick. This discovery, antedating by eight years the +work on the relation of the mosquito to malaria, had a very great +influence on subsequent studies along these lines. + +While much of the recent work has dealt with the true insects, or +hexapods, it is now known that several of the most serious diseases of +animals, and at least two important diseases of man are tick borne. +These belong to the types known collectively as _babesioses_ (or +"_piroplasmoses_"), and _spirochætoses_. + +The term _babesiosis_ is applied to a disease of man or animals which is +caused by minute protozoan parasites of the genus _Babesia_, living in +the red blood corpuscles. These parasites have usually been given the +generic name _Piroplasma_ and hence the type of disease which they cause +is often referred to as "_piroplasmosis_." The best known illustration +is the disease known in this country as Texas fever of cattle. + + +CATTLE TICKS AND TEXAS FEVER--The cattle disease, which in the United +States is known as Texas fever, is a widely distributed, exceedingly +acute disease. In Australia it is known as _redwater fever_ and in +Europe as hæmoglobinuria, due to the fact that the urine of the diseased +animals is discolored by the breaking down of the red blood corpuscles +infested by the parasite. + +In their historical discussion, Smith and Kilbourne, point out that as +far back as 1796 it was noted that Southern cattle, in a state of +apparent health, might spread a fatal disease among Northern herds. As +observations accumulated, it was learned that this infection was carried +only during the warm season of the year and in the depth of winter +Southern cattle were harmless. Moreover, Southern cattle after remaining +for a short time in the North lost their power to transmit the disease, +and the same was true of cattle which had been driven for a considerable +distance. + +Very significant was the fact that the infection was not communicated +directly from the Southern to Northern cattle but that the ground over +which the former passed was infected by them, and that the infection was +transmitted thence to susceptible cattle _after a period of not less +than thirty days had elapsed_. + +Of course a disease as striking as this, and which caused such enormous +losses of cattle in the region invaded was fruitful in theories +concerning its causation. The most widespread was the belief that +pastures were infected by the saliva, urine, or manure of Southern +cattle. There were not wanting keen observers who suggested that the +disease was caused by ticks, but little weight was given to their view. + +Various workers had described bacteria which they had isolated from the +organs of the diseased animals, but their findings could not be +verified. In 1889, Smith and Kilbourne discovered a minute, pear-shaped +organism (fig. 138) in the red blood corpuscles of a cow which had +succumbed to Texas fever. On account of their shape they were given the +generic name _Pyrososma_ and because they were usually found two in a +corpuscle, the specific name, _bigeminum_. It is now generally accepted +that the parasite is the same which Babes had observed the year before +in Roumanian cattle suffering from hæmoglobinuria, and should be known +as _Babesia bovis_ (Babes). + +[Illustration: 138. Babesia bovis in blood corpuscles. After Calli.] + +[Illustration: 139. The cattle tick (Boophilus annulatus). (_a_) Female; +(_b_) male. After Comstock.] + +By a series of perfectly conclusive experiments carried on near +Washington, D.C., Smith and Kilbourne showed that this organism was +carried from Southern cattle to non-immune animals by the so-called +Southern cattle tick, _Boophilus annulatus_ (= _Margaropus annulatus_) +(fig. 139). + +Of fourteen head of native cattle placed in a field with tick-infested +Northern cattle all but two contracted the disease. This experiment was +repeated with similar results. Four head of native cattle kept in a plot +with three North Carolina cattle which had been carefully freed from +ticks remained healthy. A second experiment the same year gave similar +results. + +Still more conclusive was the experiment showing that fields which had +not been entered by Southern cattle but which had been infected by +mature ticks taken from such animals would produce Texas fever in native +cattle. On September 13, 1889, several thousand ticks collected from +cattle in North Carolina three and four days before, were scattered in a +small field near Washington. Three out of four native animals placed in +this field contracted the disease. The fourth animal was not examined as +to its blood but it showed no external symptoms of the disease. + +[Illustration: 140. Hyalomma ægypticum. After Nuttall and Warburton.] + +In these earlier experiments it was believed that the cattle tick acted +as a carrier of the disease between the Southern cattle and the _soil_ +of the Northern pastures. "It was believed that the tick obtained the +parasite from the blood of its host and in its dissolution on the +pasture a certain resistant spore form was set free which produced the +disease when taken in with the food." The feeding of one animal for some +time with grass from the most abundantly infected field, without any +appearance of the disease, made this hypothesis untenable. + +In the experimental work in 1890 the astonishing fact was brought out +that the disease was conveyed neither by infected ticks disintegrating +nor by their directly transferring the parasite, but that it was +conveyed by the young hatched from eggs of infected ticks. In other +words, the disease was hereditarily transferred to ticks of the second +generation and they alone were capable of conveying it. + +Thus was explained the fact that Texas fever did not appear immediately +along the route of Southern cattle being driven to Northern markets but +that after a certain definite period it manifested itself. It was +conveyed by the progeny of ticks which had dropped from the Southern +cattle and deposited their eggs on the ground. + +These results have been fully confirmed by workers in different parts of +the world,--notably by Koch, in Africa, and by Pound, in Australia. + +The disease is apparently transmitted by _Boophilus annulatus_ alone, in +the United States, but it, or an almost identical disease, is conveyed +by _Ixodes hexagonus_ in Norway, _Ixodes ricinus_ in Finland and France +and by the three species, _Boophilus decoloratus_, _Hyalomma ægypticum_ +(fig. 140 and 141), and _Hæmaphysalis punctata_ in Africa. + +[Illustration: 141. Hyalomma ægypticum. Capitulum of female; (_a_) +dorsal, (_b_) ventral aspect.] + +In spite of the detailed study which it has received, the life cycle of +_Babesia bovis_ has not been satisfactorily worked out. The asexual +reproduction in the blood of the vertebrate host has been described but +the cycle in the tick is practically unknown. + +More successful attempts have been made to work out the life cycle of a +related species, _Babesia canis_, which causes malignant jaundice in +dogs in Africa and parts of Southern Europe. In this instance, also, the +disease is transmitted by heredity to the ticks of the second +generation. Yet the larval, or "seed ticks," from an infected female are +not capable of conveying the disease, but only the nymphs and adults. +Still more complicated is the condition in the case of _Babesia ovis_ of +sheep, which Motas has shown can be conveyed solely by the adult, +sexually mature ticks of the second generation. + +In _Babesia canis_, Christopher (1907) observed developmental stages in +the tick. He found in the stomach of adult ticks, large motile +club-shaped bodies which he considered as oökinetes. These bodies pass +to the ovaries of the tick and enter the eggs where they become globular +in form and probably represent an oocyst. This breaks up into a number +of sporoblasts which enter the tissues of the developing tick and give +rise to numerous sporozoites, which collect in the salivary glands and +thence are transferred to the vertebrate host. A number of other species +of _Babesia_ are known to infest vertebrates and in all the cases where +the method has been worked out it has been found that the conveyal was +by ticks. We shall not consider the cases more fully here, as we are +concerned especially with the method of transfer of human diseases. + + +TICKS AND ROCKY MOUNTAIN SPOTTED FEVER OF MAN--Ever since 1873 there has +been known in Montana and Idaho a peculiar febrile disease of man, which +has gained the name of "Rocky Mountain spotted fever." Its onset is +marked by chills and fever which rapidly become acute. In about four to +seven days there appears a characteristic eruption on the wrists, ankles +or back, which quickly covers the body. + +McClintic (1912) states that the disease has now been reported from +practically all of the Rocky Mountain States, including Arizona, +California, Colorado, Idaho, Montana, Nevada, Oregon, Utah, Washington, +and Wyoming. "Although the disease is far more prevalent in Montana and +Idaho than in any of the other States, its spread has assumed such +proportions in the last decade as to call for the gravest consideration +on the part of both the state and national health authorities. In fact, +the disease has so spread from state to state that it has undoubtedly +become a very serious interstate problem demanding the institution of +measures for its control and suppression." + +A peculiar feature of the Rocky Mountain spotted fever is a marked +variation in its severity in different localities. In Montana, and +especially in the famous Bitter Root Valley, from 33 per cent to 75 per +cent of the cases result fatally. On the other hand, the fatality does +not exceed four per cent in Idaho. + +In 1902, Wilson and Chowning reported the causative organism of spotted +fever to be a blood parasite akin to the _Babesia_ of Texas fever, and +made the suggestion that the disease was tick-borne. The careful studies +of Stiles (1905) failed to confirm the supposed discovery of the +organism, and the disease is now generally classed as due to an +invisible virus. On the other hand, the accumulated evidence has fully +substantiated the hypothesis that it is tick-borne. + +According to Ricketts (1907) the experimental evidence in support of +this hypothesis was first afforded by Dr. L. P. McCalla and Dr. H. A. +Brereton, in 1905. These investigators transmitted the disease from man +to man in two experiments. "The tick was obtained 'from the chest of a +man very ill with spotted fever' and 'applied to the arm of a man who +had been in the hospital for two months and a half, and had lost both +feet from gangrene due to freezing.' On the eighth day the patient +became very ill and passed through a mild course of spotted fever, +leaving a characteristic eruption. The experiment was repeated by +placing the tick on a woman's leg and she likewise was infected with +spotted fever." + +The most detailed studies were those of the late Dr. H. T. Ricketts, and +it was he who clearly established the tick hypothesis. In the summer of +1906 he found that guinea pigs and monkeys are very susceptible to +spotted fever and can readily be infected by inoculation of blood from +patients suffering from the disease. This opened the way to experimental +work on tick transmission. A female tick was fed upon an infected guinea +pig for two days, removed and isolated for two days and then placed upon +a healthy guinea pig. After an incubation period of three and a half +days the experimental animal contracted a well-marked case of the +disease. + +A similar result was obtained at the same time by King, and later in the +season Ricketts proved that the male tick was also capable of +transmitting the disease. He found that there was a very intimate +relation of the virus to the tick and that the transmission must be +regarded as biological throughout. Ticks remained infective as long as +they lived and would feed for a period of several months. If they +acquired the disease in the larval or nymphal stage they retained it +during molting and were infective in the subsequent stages. In a few +cases the larvæ from an infected female were infective. + +The evidence indicated that the tick suffers from a relatively harmless, +generalized infection and the virus proliferates in its body. The +disease probably is transferred through the salivary secretion of the +tick since inoculation experiments show that the salivary glands of the +infected adult contain the virus. + +It is probable that in nature the reservoir of the virus of spotted +fever is some one or more of the native small animals. Infected ticks +have been found in nature, and as various wild animals are susceptible +to the disease, it is obvious that it may exist among them unnoticed. +Wilson and Chowning suggested that the ground squirrel plays the +principal rôle. + +Unfortunately, much confusion exists regarding the correct name of the +tick which normally conveys the disease. In the medical literature it is +usually referred to as _Dermacentor occidentalis_, but students of the +group now agree that it is specifically distinct. Banks has designated +it as _Dermacentor venustus_ and this name is used in the publications +of the Bureau of Entomology. On the other hand, Stiles maintains that +the common tick of the Bitter Root Valley, and the form which has been +collected by the authors who have worked on Rocky Mountain spotted fever +in that region, is separable from _D. venustus_, and he has described it +under the name of _Dermacentor andersoni_. + +Mayer (1911) has shown experimentally that spotted fever may be +transmitted by several different species of ticks, notably _Dermacentor +marginatus_, _Dermacentor variabilis_ and _Amblyomma americanum_. This +being the case, the question of the exact systematic status of the +species experimented upon in the Bitter Root Valley becomes less +important, for since _Dermacentor occidentalis_, _Dermacentor venustus_ +and _Dermacentor andersoni_ all readily attack man, it is probable that +either species would readily disseminate the disease if it should spread +into their range. + +Hunter and Bishop (1911) have emphasized the fact that in the eastern +and southern United States there occur several species which attack man, +and any one of which might transmit the disease from animal to animal +and from animal to man. The following species, they state, would +probably be of principal importance in the Southern and Eastern States: +the lone star tick (_Amblyomma americanum_); the American dog tick +(_Dermacentor variabilis_); and the gulf-coast tick (_Amblyomma +maculatum_). In the extreme southern portions of Texas, _Amblyomma +cajennense_, is a common pest of man. + +Since the evidence all indicates that Rocky Mountain spotted fever is +transmitted solely by the tick, and that some of the wild animals serve +as reservoirs of the virus, it is obvious that personal prophylaxis +consists in avoiding the ticks as fully as possible, and in quickly +removing those which do attack. General measures along the line of tick +eradication must be carried out if the disease is to be controlled. That +such measures are feasible has been shown by the work which has been +done in controlling the tick-borne Texas fever of cattle, and by such +work as has already been done against the spotted fever tick, which +occurs on both wild and domestic animals. Detailed consideration of +these measures is to be found in the publications of the Public Health +and Marine Hospital Service, and the Bureau of Entomology. Hunter and +Bishopp give the following summarized recommendations for control or +eradication measures in the Bitter Root Valley. + +(1) A campaign of education, whereby all the residents of the valley +will be made thoroughly familiar with the feasibility of the plan of +eradication, and with what it will mean in the development of the +valley. + +(2) The obtaining of legislation to make it possible to dip or oil all +live stock in the Bitter Root Valley. + +(3) The obtaining of an accurate census of the horses, cattle, sheep, +mules, and dogs in the valley. + +(4) The construction of ten or more dipping vats. + +(5) The providing of materials to be used in the dipping mixture. + +(6) The organization of a corps of workers to carry on the operations. + +(7) The systematic dipping of the horses, cattle, sheep, and dogs of the +valley on a definite weekly schedule from approximately March 10 to June +9. + +(8) The treatment by hand of the animals in localities remote from vats, +on the same schedule. + +They estimate that after three seasons' operations a very small annual +expenditure would provide against reinfestation of the valley by the +incoming of cattle from other places. + +Supplementary measures consist in the killing of wild mammals which may +harbor the tick; systematic burning of the brush and debris on the +mountain side; and in clearing, since the tick is seldom found on land +under cultivation. + + + + +CHAPTER X + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA [_Continued_] + + +ARTHROPODS AND SPIROCHÆTOSES OF MAN AND ANIMALS + +The term spirochætoses is applied to diseases of man or animals which +are due to protistan parasites belonging to the group of slender, spiral +organisms known as spirochætes. + +There has been much discussion concerning the relationship Of the +spirochætes. Formerly, they were regarded as bacteria closely related to +the forms grouped in the genus _Spirillum_. The results of the detailed +study which the spirochætes have received in recent years, have led most +of the workers to consider them as belonging to the protozoa. The merits +of the discussion we are not concerned with here, but rather with the +fact that a number of diseases caused by spirochætes are +arthropod-borne. The better known of these we shall discuss. + + +AFRICAN RELAPSING FEVER OF MAN--It has long been known to the natives of +Africa and to travelers in that country, that the bite of a certain +tick, _Ornithodoros moubata_, may be followed by severe or even fatal +fever of the relapsing type. Until recent years, it was supposed that +the effect was due to some special virulence of the tick, just as nagana +of cattle was attributed to the direct effect of the bite of the +tsetse-fly. The disease is commonly known as "tick-fever" or by the +various native names of the tick. + +In 1904, Ross and Milne, in Uganda, and Dutton and Todd on the Congo, +discovered that the cause of the disease is a spirochæte which is +transmitted by the tick. This organism has been designated by Novy and +Knapp as _Spirochæta duttoni_. + +_Ornithodoros moubata_ (fig. 142), the carrier of African relapsing +fever, or "tick-fever," is widely distributed in tropical Africa, and +occurs in great numbers in the huts of natives, in the dust, cracks and +crevices of the dirt floors, or the walls. It feeds voraciously on man +as well as upon birds and mammals. Like others of the _Argasidæ_, it +resembles the bed-bug in its habit of feeding primarily at night. Dutton +and Todd observed that the larval stage is undergone in the egg and that +the first free stage is that of the octopod nymph. + +[Illustration: 142. Ornithodoros moubata. (_a_) Anterior part of venter; +(_b_) second stage nymph; (_c_) capitulum; (_d_) dorsal and (_e_) +ventral aspect of female; (_f_) ventral aspect of nymph; (_g_) capitulum +of nymph. After Nuttall and Warburton.] + +The evidence that the fever is transmitted by this tick is conclusive. +Koch found that from five per cent to fifteen per cent, and in some +places, fifty per cent of the ticks captured, harbored the spirochæte. +The disease is readily transmitted to monkeys, rats, mice and other +animals and the earlier experiments along these lines have been +confirmed by many workers. + +Not only are the ticks which have fed on infected individuals capable of +conveying the disease to healthy animals but they transmit the causative +organism to their progeny. Thus Möllers (1907), working in Berlin, +repeatedly infected monkeys through the bites of nymphs which had been +bred in the laboratory from infected ticks. Still more astonishing was +his discovery that ticks of the third generation were infective. In +other words, if the progeny of infected ticks were fed throughout life +on healthy animals, and on maturity deposited eggs, the nymphs which +hatched from these eggs would still be capable of carrying the +infection. + +The developmental cycle of the spirochæte within the tick has not been +fully worked out, though the general conclusions of Leishman (1910) have +been supported by the recent works of Balfour (1911 and 1912), and +Hindle (1912), on the life cycle of spirochætes affecting fowls. + +_Spirochæta duttoni_ ingested by _Ornithodoros moubata_ apparently +disappear within a few days, but Leishman believed that in reality they +break up into minute granules which are to be found in the alimentary +canal, the salivary glands and the Malpighian tubes of the tick. These +granules, or "coccoid bodies," as Hindle calls them, are supposed to be +the form in which the spirochætes infect the new host. We shall see +later that Marchoux and Couvy (1913) dissent wholly from this +interpretation. + +According to Leishman, and Hindle, the coccoid bodies are not injected +into the vertebrate host with the saliva of the tick, as are the +sporozoites of malaria with that of the mosquito. Instead, they pass out +with the excrement and secondarily gain access to the wound inflicted by +the tick. + +Nuttall (1912) calls attention to the fact that the geographical +distribution of _Ornithodoros moubata_ is far wider than our present +records show for the distribution of the relapsing fever in man and that +there is every reason to fear the extension of the disease. Huts where +the ticks occur should be avoided and it should be remembered that in +infected localities there is special danger in sleeping on the ground. + + +EUROPEAN RELAPSING FEVER--There is widely distributed in Europe a type +of relapsing fever which is caused by _Spirochæta recurrentis_. It has +long been supposed that this disease is spread by the bed-bug and there +is some experimental evidence to show that it may be conveyed by these +insects. + +In 1897, Tictin found that he could infect monkeys by inoculating the +contents of bed-bugs which had fed upon a patient within forty-eight +hours. Nuttall, in 1907, in one experiment succeeded in transmitting +_Spirochæta recurrentis_ from mouse to mouse by bites of bed-bugs. The +bugs, thirty-five in number, were transferred at short intervals from +one mouse to another, not being allowed to take a full meal on the +first, or infected mouse. + +On the other hand, there is much clinical evidence to show that the +European relapsing fever like various other types of the disease is +transmitted from man to man by head and body lice (_Pediculus humanus_ +and _Pediculus corporis_). + +Interesting supplementary evidence is that of Bayon's observations +(1912), in Moscow. "Having visited the big municipal night hospitals at +Moscow I soon noticed that they were kept with such scrupulous +cleanliness, disinfected so lavishly, the beds of iron, the floor +cemented, that it was not possible for bed-bugs to thrive to any extent +on the premises. The people sleeping there were allowed, however, to +sleep in their own clothes. The introduction of these model homes had +not had any effect on the incidence of relapsing fever, for the places +were still hot-beds of the fever during winter. On the other hand, +though I changed my rooms several times, I found bugs in every +successive lodging, and I was told in Moscow, this can hardly be +avoided. Yet no foreigner, or Russian of the better class, ever catches +relapsing fever. To this may be added the fact that when I asked for +clothes-lice and promised to pay a kopec for two, the attendants from +the night hostel brought me next morning a small ounce bottle crammed +with _Pediculus capitis_ (= _P. humanus_), and _Pediculus vestimentorum_ +(= _P. corporis_) collected off the sleepers. If relapsing fever were +transmitted by bed-bugs, it would be much more disseminated than it is +at present in Moscow." + +Direct experimental evidence of the agency of lice in transmitting +relapsing fever is especially clear in the case of a type of the disease +prevalent in parts of North Africa. We shall consider this evidence +later. + + +OTHER TYPES OF RELAPSING FEVER OF MAN--In addition to the three types of +human relapsing fever already referred to, several others have been +distinguished and have been attributed to distinct species of +spirochætes. The various spirochætoses of man are: + +African, caused by _S. duttoni_; European, caused by _S. recurrentis_; +North African, caused by _S. berbera_; East African, caused by _S. +rossi_; East Indian, caused by _S. carteri_; North American, caused by +_S. novyi_; South American, caused by _S. duttoni_ (?). + +Nuttall (1912) in his valuable résumé of the subject, has emphasized +that "in view of the morphological similarity of the supposedly +different species of spirochætes and their individual variations in +virulence, we may well doubt if any of the 'species' are valid. As I +pointed out four years ago, the various specific names given to the +spirochætes causing relapsing fever in man may be used merely for +convenience _to distinguish strains or races_ of different origin. They +cannot be regarded as valid names, in the sense of scientific +nomenclature, for virulence and immunity reactions are not adequate +tests of specificity." + + +NORTH AFRICAN RELAPSING FEVER OF MAN--The type of human relapsing fever +to be met with in Algeria, Tunis, and Tripoli, is due to a _Spirochæta_ +to which does not differ morphologically from _Spirochæta duttoni_, but +which has been separated on biological grounds as _Spirochæta berberi_. + +Experimenting with this type of disease in Algeria, Sergent and Foly +(1910), twice succeeded in transmitting it from man to monkeys by +inoculation of crushed body lice and in two cases obtained infection of +human subjects who had received infected lice under their clothing and +who slept under coverings harboring many of the lice which had fed upon +a patient. Their results were negative with _Argas persicus_, _Cimex +lectularius_, _Musca domestica_, _Hæmatopinus spinulosus_ and +_Ceratophyllus fasciatus_. They found body lice associated with every +case of relapsing fever which they found in Algeria. + +Nicolle, Blaizot, and Conseil (1912) showed that the louse did not +transmit the parasite by its bite. Two or three hours after it has fed +on a patient, the spirochætes begin to break up and finally they +disappear, so that after a day, repeated examinations fail to reveal +them. They persist, nevertheless, in some unknown form, for if the +observations are continued they reappear in eight to twelve days. These +new forms are virulent, for a monkey was infected by inoculating a +single crushed louse which had fed on infected blood fifteen days +before. + +Natural infection is indirect. Those attacked by the insect scratch, and +in this act they excoriate the skin, crush the lice and contaminate +their fingers. The least abrasion of the skin serves for the entrance of +the spirochætes. Even the contact of the soiled fingers on the various +mucosa, such as the conjunctive of the eye, is sufficient. + +As in the case of _Spirochæta duttoni_, the organism is transmitted +hereditarily in the arthropod vector. The progeny of lice which have fed +on infected blood may themselves be infective. + + +SPIROCHÆTOSIS OF FOWLS--One of the best known of the spirochætes +transmitted by arthropods is _Spirochæta gallinarum_, the cause of a +very fatal disease of domestic fowls in widely separated regions of the +world. According to Nuttall, it occurs in Southeastern Europe, Asia, +Africa, South America and Australia. + +In 1903, Marchoux and Salimbeni, working in Brazil, made the first +detailed study of the disease, and showed that the causative organism is +transmitted from fowl to fowl by the tick _Argas persicus_. They found +that the ticks remained infective for at least five months. Specimens +which had fed upon diseased birds in Brazil were sent to Nuttall and he +promptly confirmed the experiments. Since that date many investigators, +notably Balfour and Hindle, have contributed to the elucidating of the +life-cycle of the parasite. Since it has been worked out more fully than +has that of any of the human spirochætes, we present Hindle's diagram +(fig. 143) and quote the brief summary from his preliminary paper +(1911_b_). + +"Commencing with the ordinary parasite in the blood of the fowl, the +spirochæte grows until it reaches a certain length (16-19µ) and then +divides by transverse division. This process is repeated, and is +probably the only method of multiplication of the parasite within the +blood. When the spirochætes disappear from the circulation, some of them +break up into the coccoid bodies which, however, do not usually develop +in the fowl. When the spirochætes are ingested by _Argas persicus_, some +of them pass through the gut wall into the cÅ“lomic fluid. From this +medium they bore their way into the cells of the various organs of the +tick and there break up into a number of coccoid bodies. These +intracellular forms multiply by ordinary fission in the cells of the +Malpighian tubules and gonads. Some of the coccoid bodies are formed in +the lumen of the gut and Malpighian tubules. The result is that some of +the coccoid bodies may be present in the Malpighian secretion and +excrement of an infected tick and when mixed with the coxal fluid may +gain entry into another fowl by the open wound caused by the tick's +bite. They then elongate and redevelop into ordinary spirochætes in the +blood of the fowl, and the cycle may be repeated." + +[Illustration: 143. Spirochæta gallinarum. After Hindle.] + +Hindle's account is clear cut and circumstantial, and is quite in line +with the work of Balfour, and of Leishman. Radically different is the +interpretation of Marchoux and Couvy (1913). These investigators +maintain that the granules localized in the Malpighian tubules in the +larvæ and, in the adult, also in the ovules and the genital ducts of the +male and female, are not derived from spirochætes but that they exist +normally in many acariens. They interpret the supposed disassociation +of the spirochæte into granules as simply the first phase, not of a +process of multiplication, but of a degeneration ending in the death of +the parasite. The fragmented chromatin has lost its affinity for stains, +remaining always paler than that of the normal spirochætes. On the other +hand, the granules of Leishman stain energetically with all the basic +stains. + +Further, according to Marchoux and Couvy, infection takes place without +the emission of the coxal fluid and indeed, soiling of the host by the +coxal fluid diluting the excrement is exceptional. All of the organs of +the Argasid are invaded by the parasites, but they pass from the +cÅ“lom into the acini of the salivary glands and collect in its +efferent canal. The saliva serves as the vehicle of infection. + +Thus, the question of the life cycle of _Spirochæta gallinarum_, and of +spirochætes in general, is an open one. + +It should be noted that _Argas persicus_, the carrier of _Spirochæta +gallinarum_, is a common pest of poultry in the southwestern United +States. Though the disease has not been reported from this country, +conditions are such that if accidentally introduced, it might do great +damage. + + +OTHER SPIROCHÆTE DISEASES OF ANIMALS--About a score of other blood +inhabiting spirochætes have been reported as occurring in mammals, but +little is known concerning their life-histories. One of the most +important is _Spirochæta theileri_ which produces a spirochætosis of +cattle in the Transvaal. Theiler has determined that it is transmitted +by an Ixodid tick, _Margaropus decoloratus_. + + +TYPHUS FEVER AND PEDICULIDÆ + +Typhus is an acute, and continued fever, formerly epidemically prevalent +in camps, hospitals, jails, and similar places where persons were +crowded together under insanitary conditions. It is accompanied by a +characteristic rash, which gives the disease the common name of +"spotted" or "lenticular" fever. The causative organism is unknown. + +Typhus fever has not generally been supposed to occur in the United +States, but there have been a few outbreaks and sporadic cases +recognized. According to Anderson and Goldberger (1912_a_), it has been +a subject of speculation among health authorities why, in spite of the +arrival of occasional cases in this country and of many persons from +endemic foci of the disease, typhus fever apparently does not gain a +foothold in the United States. These same workers showed that the +so-called Brill's disease, studied especially in New York City, is +identical with the typhus fever of Mexico and of Europe. + +The conditions under which the disease occurs and under which it spreads +most rapidly are such as to suggest that it is carried by some parasitic +insect. On epidemiological grounds the insects most open to suspicion +are the lice, bed-bugs and fleas. + +In 1909, Nicolle, Comte and Conseil, succeeded in transmitting typhus +fever from infected to healthy monkeys by means of the body louse +(_Pediculus corporis_). Independently of this work, Anderson and +Goldberger had undertaken work along this line in Mexico, and in 1910 +reported two attempts to transmit the disease to monkeys by means of +body lice. The first experiment resulted negatively, but the second +resulted in a slight rise in temperature, and in view of later results +it seems that this was due to infection with typhus. + +Shortly after, Ricketts and Wilder (1910) succeeded in transmitting the +disease to the monkey by the bite of body lice in two experiments, the +lice in one instance deriving their infection from a man and in another +from the monkey. Another monkey was infected by typhus through the +introduction of the feces and abdominal contents of infested lice into +small incisions. Experiments with fleas and bed-bugs resulted +negatively. + +Subsequently, Goldberger and Anderson (1912_b_) indicated that the head +louse (_Pediculus humanus_) as well, may become infected with typhus. In +an attempt to transmit typhus fever (Mexican virus) from man to monkey +by subcutaneous injection of a saline suspension of crushed head lice, +the monkeys developed a typical febrile reaction with subsequent +resistance to an inoculation of virulent typhus (Mexican) blood. In one +of the three experiments to transmit the disease from man to monkey by +means of the bite of the head louse, the animal bitten by the presumably +infected head lice proved resistant to two successive immunity tests +with virulent typhus blood. + +In 1910, Ricketts and Wilder reported an experiment undertaken with a +view to determining whether the young of infected lice were themselves +infected. Young lice were reared to maturity on the bodies of typhus +patients, so that if the eggs were susceptible to infection at any stage +of their development, they would have every opportunity of being +infected within the ovary. The eggs of these infected lice were +obtained, they were incubated, and the young lice of the second +generation were placed on a normal rhesus monkey. The experimenters were +unable to keep the monkey under very close observation during the +following three or four weeks, but from the fact that he proved +resistant to a subsequent immunity test they concluded that he probably +owed this immunity to infection by these lice of the second generation. + +Anderson and Goldberger (1912_b_) object that due consideration was not +given to the possibility of a variable susceptibility of the monkey to +typhus. Their similar experiment was "frankly negative." + +Prophylaxis against typhus fever is, therefore, primarily a question of +vermin extermination. A brief article by Dr. Goldberger (1914) so +clearly shows the practical application of his work and that of the +other investigators of the subject, that we abstract from it the +following account: + +"In general terms it may be stated that association with a case of +typhus fever in the absence of the transmitting insect is no more +dangerous than is association with a case of yellow fever in the absence +of the yellow fever mosquito. Danger threatens only when the insect +appears on the scene." + +"We may say, therefore, that to prevent infection of the individual it +is necessary for him only to avoid being bitten by the louse. In theory +this may readily be done, for we know that the body louse infests and +attaches itself almost entirely to the body linen, and that boiling +kills this insect and its eggs. Individual prophylaxis is based +essentially, therefore, on the avoidance of contact with individuals +likely to harbor lice. Practically, however, this is not always as easy +as it may seem, especially under the conditions of such intimate +association as is imposed by urban life. Particularly is this the case +in places such as some of the large Mexican cities, where a large +proportion of the population harbors this vermin. Under such +circumstances it is well to avoid crowds or crowded places, such as +public markets, crowded streets, or public assemblies at which the +'peon' gathers." + +"Community prophylaxis efficiently and intelligently carried out is, +from a certain point of view, probably easier and more effective in +protecting the individual than is the individual's own effort to guard +himself. Typhus emphasizes, perhaps better than any other disease, the +fact that fundamentally, sanitation and health are economic problems. In +proportion as the economic condition of the masses has improved--that +is, in proportion as they could afford to keep clean--the notorious +filth disease has decreased or disappeared. In localities where it still +prevails, its further reduction or complete eradication waits on a +further improvement in, or extension of, the improved economic status of +those afflicted. Economic evolution is very slow process, and, while +doing what we can to hasten it, we must take such precautions as +existing conditions permit, looking to a reduction in or complete +eradication of the disease." + +"When possible, public bath houses and public wash houses, where the +poor may bathe and do their washings at a minimum or without cost, +should be provided. Similar provision should be made in military and +construction camps. Troops in the field should be given the opportunity +as frequently as possible to wash and _scald_ or _boil_ their body +linen." + +"Lodging houses, cheap boarding houses, night shelters, hospitals, jails +and prisons, are important factors in the spread and frequently +constitute foci of the disease. They should receive rigid sanitary +supervision, including the enforcement of measures to free all inmates +of such institutions of lice on admission." + +"So far as individual foci of the disease are concerned these should be +dealt with by segregating and keeping under observation all exposed +individuals for 14 days--the period of incubation--from the last +exposure, by disinfecting (boiling or steaming) the suspected bedding, +body linen, and clothes, for the destruction of any possible vermin that +they may harbor, and by fumigating (with sulphur) the quarters that they +may have occupied." + +"It will be noted that nothing has been said as to the disposition of +the patient. So far as the patient is concerned, he should be removed to +'clean' surroundings, making sure that he does not take with him any +vermin. This may be done by bathing, treating the hair with an +insecticide (coal oil, tincture of larkspur), and a complete change of +body linen. Aside from this, the patient may be treated or cared for in +a general hospital ward or in a private house, provided the sanitary +officer is satisfied that the new surroundings to which the patient has +been removed are 'clean,' that is, free from vermin. Indeed, it is +reasonably safe to permit a 'clean' patient to remain in his own home if +this is 'clean,' for, as has already been emphasized, there can be no +spread in the absence of lice. This is a common experience in native +families of the better class and of Europeans in Mexico City." + +"Similarly the sulphur fumigation above prescribed may be dispensed with +as unnecessary in this class of cases." + + + + +CHAPTER XI + +SOME POSSIBLE, BUT IMPERFECTLY ESTABLISHED CASES OF ARTHROPOD +TRANSMISSION OF DISEASE + + +INFANTILE PARALYSIS OR ACUTE ANTERIOR POLIOMYELITIS + +The disease usually known in this country as infantile paralysis or, +more technically, as acute anterior poliomyelitis, is one which has +aroused much attention in recent years. + +The causative organism of infantile paralysis is unknown, but it has +been demonstrated that it belongs to the group of filterable viruses. It +gives rise to a general infection, producing characteristic lesions in +the central nervous system. The result of the injury to the motor nerves +is a more or less complete paralysis of the corresponding muscle. This +usually manifests itself in the legs and arms. The fatal cases are +usually the result of paralysis of the muscles of respiration. Of the +non-fatal cases about 60 per cent remain permanently crippled in varying +degrees. + +Though long known, it was not until about 1890 that it was emphasized +that the disease occurs in epidemic form. At this time Medin reported +his observations on an epidemic of forty-three cases which occurred in +and around Stockholm in 1887. Since then, according to Frost (1911), +epidemics have been observed with increasing frequency in various parts +of the world. The largest recorded epidemics have been those in Vermont, +1894, 126 cases; Norway and Sweden, 1905, about 1,500 cases; New York +City, 1907, about 2,500 cases. Since 1907 many epidemics have been +reported in the United States, and especially in the Northern States +east of the Dakotas. In 1912 there were over 300 cases of the disease in +Buffalo, N. Y., with a mortality of somewhat over 11 per cent. + +In view of the sudden prominence and the alarming spread of infantile +paralysis, there have been many attempts to determine the cause, and the +manner in which the disease spreads and develops in epidemic form. In +the course of these studies, the question of possible transmission by +insects was naturally suggested. + +C. W. Howard and Clark (1912) presented the results of studies in this +phase of the subject. They dealt especially with the house-fly, bedbug, +head, and body lice, and mosquitoes. It was found that the house-fly +(_Musca domestica_) can carry the virus of poliomyelitis in an active +state for several days upon the surface of the body and for several +hours within the gastro-intestinal tract. Mosquitoes and lice were found +not to take up or maintain the virus. On the other hand, the bedbug +(_Cimex lectularius_) was found to take the virus from the infected +monkeys and to maintain it in a living state within the body for a +period of seven days. This was demonstrated by grinding up in salt +solution, insects which had fed on poliomyelitic animals and injecting +the filtrate into a healthy monkey. The experimenters doubted that the +bedbug is a carrier of the virus in nature. + +Earlier in the same year, Brues and Sheppard published the results of an +intensive epidemiological study of the outbreak of 1911, in +Massachusetts. Special attention had been paid to the possibility of +insect transfer and the following conclusion was reached: + +"Field work during the past summer together with a consideration of the +epidemiology of the disease so far as known, points strongly toward +biting flies as possible carriers of the virus. It seems probable that +the common stable-fly (_Stomoxys calcitrans_ L.) may be responsible to a +certain extent for the spread of acute epidemic poliomyelitis, possibly +aided by other biting flies, such as _Tabanus lineola_. No facts which +disprove such a hypothesis have as yet been adduced, and experiments +based upon it are now in progress." + +As stated by Brues (1913), especial suspicion fell upon the stable-fly +because: + +1. The blood-sucking habits of the adult fly suit it for the transfer of +virus present in the blood. + +2. The seasonal abundance of the fly is very closely correlated with the +incidence of the disease, rising rapidly during the summer and reaching +a maximum in July and August, then slowly declining in September and +October. + +3. The geographical distribution of the fly is, so far as can be +ascertained, wider, or at least co-extensive with that of poliomyelitis. + +4. _Stomoxys_ is distinctly more abundant under rural conditions, than +in cities and thickly populated areas. + +5. While the disease spreads over districts quickly and in a rather +erratic way, it often appears to follow along lines of travel, and it is +known that _Stomoxys_ flies will often follow horses for long distances +along highways. + +6. In a surprisingly large number of cases, it appeared probable that +the children affected had been in the habit of frequenting places where +_Stomoxys_ is particularly abundant, i.e., about stables, barnyards, +etc. + +The experiments referred to were carried on during the summer of 1912 +and in September Dr. Rosenau announced that the disease was transferred +by the bite of the stable-fly. + +A monkey infected by inoculation was exposed to the bites of upwards of +a thousand of the _Stomoxys_ flies daily, by stretching it at full +length and rolling it in a piece of chicken wire, and then placing it on +the floor of the cage in which the flies were confined. The flies fed +freely from the first, as well as later, after paralysis had set in. +Alternating with the inoculated monkey, healthy monkeys were similarly +introduced into the cage at intervals. New monkeys were inoculated to +keep a supply of such infected animals and additional healthy ones were +exposed to the flies, which fed willingly and in considerable numbers on +each occasion. "Thus the flies were given every opportunity to obtain +infection from the monkeys, since the animals were bitten during +practically every stage of the disease from the time of the inoculation +of the virus till their death following the appearance of paralysis. By +the same arrangement the healthy monkeys were likely to be bitten by +flies that had previously fed during the various stages of the disease +on the infected monkeys. The flies had meanwhile enjoyed the opportunity +of incubating the virus for periods varying from the day or two which +usually elapses between consecutive feedings, to the two or three-week +period for which at least some (although a very small percentage) of the +flies lived in the cage." + +"In all, twelve apparently healthy monkeys of a small Japan species were +exposed to the flies in the manner described for the infected monkeys. +Some were placed in the cage only once or twice and others a number of +times after varying intervals. These exposures usually lasted for about +half an hour, but were sometimes more protracted. No results were +apparent until two or three weeks after the experiment was well under +way, and then in rather rapid succession six of the animals developed +symptoms of poliomyelitis. In three, the disease appeared in a virulent +form, resulting in death, while the other three experienced transient +tremblings, diarrhÅ“a, partial paralysis and recovery."--Brues, 1913. + +Very soon after the announcement of the results of experiments by +Rosenau and Brues, they were apparently conclusively confirmed by +Anderson and Frost (1912), who repeated the experiments, at Washington. +They announced that through the bites of the _Stomoxys_ flies that had +previously fed on infected monkeys, they had succeeded in experimentally +infecting three healthy monkeys. + +The results of these experiments gained much publicity and in spite of +the conservative manner in which they had been announced, it was widely +proclaimed that infantile paralysis was conveyed in nature by the +stable-fly and by it alone. + +Serious doubt was cast on this theory by the results of further +experiments by Anderson and Frost, reported in May of 1913. Contrary to +the expectations justified by their first experience, the results of all +the later, and more extended, experiments were wholly negative. Not once +were these investigators again able to transmit the infection of +poliomyelitis through _Stomoxys_. They concluded that it was extremely +doubtful that the insect was an important factor in the natural +transmission of the disease, not only because of their series of +negative results, "but also because recent experiments have afforded +additional evidence of the direct transmissibility or contagiousness of +poliomyelitis, and because epidemiological studies appear to us to +indicate that the disease is more likely transmitted largely through +passive human virus carriers." + +Soon after this, Kling and Levaditi (1913) published their detailed +studies on acute anterior poliomyelitis. They considered that the +experiments of Flexner and Clark (and Howard and Clark), who fed +house-flies on emulsion of infected spinal cord, were under conditions +so different from what could occur in nature that one could not draw +precise conclusions from them regarding the epidemiology of the disease. +They cited the experiments of Josefson (1912), as being under more +reasonable conditions. He sought to determine whether the inoculation of +monkeys with flies caught in the wards of the Hospital for Contagious +Diseases at Stockholm, where they had been in contact with cases of +poliomyelitis, would produce the disease. The results were completely +negative. + +Kling and Lavaditi made four attempts of this kind. The flies were +collected in places where poliomyelitics had dwelt, three, four and +twenty-four after the beginning of the disease in the family and one, +three, and fifteen days after the patient had left the house. These +insects were for the greater part living and had certainly been in +contact with the infected person. In addition, flies were used which had +been caught in the wards of the Hospital for Contagious Diseases at +Söderkoping, when numbers of poliomyelitics were confined there. +Finally, to make the conditions as favorable as possible, the emulsions +prepared from these flies were injected without previous filtering, +since filtration often causes a weakening of the virus. In spite of +these precautions, all their results were negative, none of the +inoculated animals having contracted poliomyelitis. They also +experimented with bedbugs which had fed upon infected patients at +various stages of the disease, but the results in these cases also were +wholly negative. + +Kling and Levaditi considered at length the possibility of transmission +of the disease by _Stomoxys_. As a result of their epidemiological +studies, they found that infantile paralysis continued to spread in +epidemic form in the dead of winter, when these flies were very rare and +torpid, or were even completely absent. Numerous cases developed in the +northern part of Sweden late in October and November, long after snow +had fallen. On account of the rarity of the Stomoxys flies during the +period of their investigations they were unable to conduct satisfactory +experiments. In one instance, during a severe epidemic, they found a +number of the flies in a stable near a house inhabited by an infected +family, though none was found in the house itself. These flies were used +in preparing an emulsion which, after filtering, was injected into the +peritoneal cavity of a monkey. The result was wholly negative. + +As for the earlier experiments, Kling and Levaditi believe if the flies +were responsible for the transmission of the disease in the cases +reported by Rosenau and Brues, and the first experiments of Anderson and +Frost, it was because the virus of infantile paralysis is eliminated +with the nasal secretions of paralyzed monkeys and the flies, becoming +contaminated, had merely acted as accidental carriers. + +Still further evidence against the hypothesis of the transmission of +acute anterior poliomyelitis by _Stomoxys calcitrans_ was brought +forward by Sawyer and Herms (1913). Special precautions were used to +prevent the transference of saliva or other possibly infectious material +from the surface of one monkey to that of another, and to avoid the +possibility of complicating the experiments by introducing other +pathogenic organisms from wild flies, only laboratory-bred flies were +used. In a series of seven carefully performed experiments, in which the +conditions were varied, Sawyer and Herms were unable to transmit +poliomyelitis from monkey to monkey through the agency of _Stomoxys_, or +to obtain any indication that the fly is the usual agent for spreading +the disease in nature. + +The evidence at hand to date indicates that acute anterior +poliomyelitis, or infantile paralysis, is transmitted by contact with +infected persons. Under certain conditions insects may be agents in +spreading the disease, but their rôle is a subordinate one. + + +Pellagra + +PELLAGRA is an endemic and epidemic disease characterized by a peculiar +eruption or erythema of the skin (figs 144 and 145), digestive +disturbances and nervous trouble. Insanity is a common result, rather +than a precursor of the disease. The manifestations of pellagra are +periodic and its duration indeterminate. + +[Illustration: 144. Pellagrous eruption on the face. After Watson.] + +The disease is one the very name of which was almost unknown in the +United States until within the past decade. It has usually been regarded +as tropical, though it occurs commonly in Italy and in various parts of +Europe. Now it is known that it not only occurs quite generally in the +United States but that it is spreading. Lavinder (1911) says that "There +are certainly many thousand cases of the disease in this country, and +the present situation must be looked upon with grave concern." + +It is not within the scope of this book to undertake a general +discussion of pellagra. The subject is of such importance to every +medical man that we cannot do better than refer to Lavinder's valuable +précis. We can only touch briefly upon the entomological phases of the +problems presented. + +The most commonly accepted theories regarding the etiology of the +disease have attributed it to the use of Indian corn as an article of +diet. This supposed relationship was explained either on the basis of, +(a) insufficiency of nutriment and inappropriateness of corn as a prime +article of food; (b) toxicity of corn or, (c) parasitism of certain +organisms--fungi or bacteria--ingested with either sound or deteriorated +corn. + +In 1905, Sambon proposed the theory of the protozoal origin of pellagra +and in 1910 he marshalled an imposing array of objections to the theory +that there existed any relationship between corn and the disease. He +presented clear evidence that pellagra existed in Europe before the +introduction of Indian corn from America, as an article of diet, and +that its spread was not _pari passu_ with that of the use of corn. Cases +were found in which the patients had apparently never used corn, though +that is obviously difficult to establish. He showed that preventive +measures based on the theory had been a failure. Finally, he believed +that the recurrence of symptoms of the disease for successive springs, +in patients who abstained absolutely from the use of corn, militated +against the theory. + +[Illustration: 145. Pellagrous eruption on the hand. After Watson.] + +On the other hand, Sambon believed that the periodicity of the symptoms, +peculiarities of distribution and seasonal incidence, and analogies of +the symptoms to those of other parasitic diseases indicated that +pellagra was of protozoal origin, and that it was insect-borne. + +The insect carriers, he believed to be one or more species of Simuliidæ, +or black-flies. In support of this he stated that _Simulium_ appears to +effect the same topographical conditions as pellagra, that in its imago +stage it seems to present the same seasonal incidence, that it has a +wide geographical distribution which seems to cover that of pellagra, +and that species of the genus are known to cause severe epizootics. +Concluding from his studies in Italy, that pellagra was limited almost +wholly to agricultural laborers, he pointed out that the Simulium flies +are found only in rural districts, and as a rule do not enter towns, +villages, or houses. + +[Illustration: 146. A favorite breeding place of Simulium. Ithaca, N. +Y.] + +When Sambon's detailed report was published in 1910, his theory was +seized upon everywhere by workers who were anxious to test it and who, +in most cases, were favorably disposed towards it because of the +wonderful progress which had been made in the understanding of other +insect-borne diseases. In this country, the entomological aspects of the +subject have been dealt with especially by Forbes (1912), and by King +and Jennings, under the direction of W. D. Hunter, of the Bureau of +Entomology, and in coöperation with the Thompson-McFadden Pellagra +Commission of the Department of Tropical Medicine of the New York +Post-Graduate Medical School. An important series of experiments with +monkeys has been undertaken by S. J. Hunter, of Kansas, but +unfortunately we have as yet no satisfactory evidence that these animals +are susceptible to the disease--a fact which renders the whole problem +difficult. + +The accumulated evidence is increasingly opposed to Sambon's hypothesis +of the transmission of pellagra by _Simulium_. This has been so clearly +manifested in the work of the Thompson-McFadden Commission that we quote +here from the report by Jennings (1914): + +"Our studies in 1912 convinced us that there was little evidence to +support the incrimination of any species of _Simulium_ in South Carolina +in the transmission of pellagra. Reviewing the group as a whole, we find +that its species are essentially "wild" and lack those habits of +intimate association with man which would be expected in the vector of +such a disease as pellagra. Although these flies are excessively +abundant in some parts of their range and are moderately so in +Spartanburg County, man is merely an incidental host, and no disposition +whatever to seek him out or to invade his domicile seems to be +manifested. Critically considered, it is nearer the fact that usually +man is attacked only when he invades their habitat." + +"As our knowledge of pellagra accumulates, it is more and more evident +that its origin is in some way closely associated with the domicile. The +possibility that an insect whose association with man and his immediate +environment is, at the best, casual and desultory, can be active in the +causation of the disease becomes increasingly remote." + +"Our knowledge of the biting habits of _Simulium_ is not complete, but +it is evident, as regards American species at least, that these are +sometimes not constant for the same species in different localities. +Certain species will bite man freely when opportunity offers, while +others have never been known to attack him. To assume that the proximity +of a _Simulium_-breeding stream necessarily implies that persons in its +vicinity must be attacked and bitten is highly fallacious. In +Spartanburg County attacks by _Simulium_ seems to be confined to the +immediate vicinity of the breeding-places. Our records and observations, +exceedingly few in number, refer almost exclusively to such locations. +Statements regarding such attacks, secured with much care and +discrimination from a large number of persons, including many +pellagrins, indicate conclusively that these insects are seldom a pest +of man in this county. A certain number of the persons questioned were +familiar with the gnats in other localities, but the majority were +seemingly ignorant of the existence of such flies with biting habits. +This is especially striking, in view of the fact that the average +distance of streams from the homes of the pellagra cases studied was +about 200 yards, many being at a distance of less than 200 yards, and +that 78 per cent of these streams were found to be infested by larval +_Simulium_. Such ignorance in a large number of persons cannot be +overlooked and indicates strongly that our belief in the negligible +character of local attacks by _Simulium_ is well founded." + +"In localities infested by 'sand-flies,' mosquitoes, etc., these pests +are always well known and the ignorance described above is very +significant." + +"Such positive reports as we received nearly always referred to bites +received in the open, along streams, etc., and observations made of +their attack were of those on field laborers in similar situations. +Males engaged in agricultural pursuits are almost exempt from pellagra +in Spartanburg County. During the season of 1913, in some two or three +instances, observations were made of the biting of _Simulium_ and some +additional and entirely creditable reports were received. These +observations and reports were under conditions identical with those +referred to in the reports of 1912 and confirm the conclusions based on +the observations of that year. I would repeat with emphasis that it is +inconceivable that a fly of the appearance and habits of the prevalent +species of _Simulium_ could be present in such a region, especially +about the haunts of man and attack him with sufficient frequency and +regularity to satisfactorily account for so active and prevalent a +disease as pellagra without being a well-known and recognized pest." + +"In connection with the conditions in the Piedmont region of South +Carolina, it may be well to cite the results of a study of those in the +arid region of western Texas." + +"In May, 1913, in company with Capt. J. F. Siler of the +Thompson-McFadden Pellagra Commission, I visited the region of which +Midland in Midland County is the center. This region is very dry and +totally devoid of running water for a long distance in every direction. +The only natural source of water-supply, a few water holes and ponds, +were visited and found to be of such a nature that the survival of +_Simulium_, far less its propagation in them, is absolutely impossible. +The nearest stream affording possibilities as a source of _Simulium_ is +60 miles away, while the average distance of such possibility is not +less than 100 miles." + +"Artificial sources of water-supply were also investigated carefully and +were found to offer no opportunity for the breeding of _Simulium_." + +"At Midland the histories of five cases of pellagra were obtained, which +gave clear evidence that this place or its immediate vicinity was the +point of origin. Persons of long residence in the country were +questioned as to the occurrence of such flies as _Simulium_ and returned +negative answers. These included a retired cattle owner, who is a man of +education and a keen observer, an expert veterinarian stationed in the +country who has the cattle of the country under constant observation, +and a practical cattle man, manager of a ranch and of wide experience. +The latter had had experience with 'Buffalo gnats' in other localities +(in the East) and is well acquainted with them. His close personal +supervision of the cattle under his charge, makes it practically certain +that he would have discovered these gnats had they been present in the +country." + +"At the time the study was made, _Simulium_ was breeding and active in +the adult state in the vicinity of Dallas, Texas, in the eastern part of +the state. We have here a region in which cases of pellagra have +originated, yet in which _Simulium_ does not and cannot breed." + +Other possible insect vectors of pellagra have been studied in great +detail and the available evidence indicates that if _any_ insect plays a +rôle in the spread of the disease, _Stomoxys calcitrans_ most nearly +fills the conditions. This conclusion was announced by Jennings and King +in 1912, and has been supported by their subsequent work. + +Yet, after all the studies of the past decade, the old belief that +pellagra is essentially of dietary origin is gaining ground. Goldberger, +Waring and Willets (1914) of the United States Public Health Service +summarize their conclusions in the statement, (1) that it is dependent +on some yet undetermined fault in a diet in which the animal or +leguminous protein component is disproportionately large and (2) that no +pellagra develops in those who _consume_ a mixed, well-balanced, and +varied diet, such, for example, as that furnished by the Government to +the enlisted men of the Army, Navy, and Marine Corps. + + +Leprosy + +LEPROSY is a specific, infectious disease due to _Bacillus lepræ_, and +characterized by the formation of tubercular nodules, ulcerations, and +disturbances of sensation. In spite of the long time that the disease +has been known and the dread with which it is regarded, little is known +concerning the method of transfer of the causative organism or the means +by which it gains access to the human body. + +It is known that the bacilli are to be found in the tubercles, the scurf +of the skin, nasal secretions, the sputum and, in fact in practically +all the discharges of the leper. Under such conditions it is quite +conceivable that they may be transferred in some instances from diseased +to healthy individuals through the agency of insects and other +arthropods. Many attempts have been made to demonstrate this method of +spread of the disease, but with little success. + +Of the suggested insect carriers none seem to meet the conditions better +than mosquitoes, and there are many suggestions in literature that these +insects play an important rôle in the transmission of leprosy. The +literature has been reviewed and important experimental evidence +presented by Currie (1910). He found that mosquitoes feeding, under +natural conditions, upon cases of nodular leprosy so rarely, if ever, +imbibe the lepra bacillus that they cannot be regarded as one of the +ordinary means of transference of this bacillus from lepers to the skin +of healthy persons. He believes that the reason that mosquitoes that +have fed on lepers do not contain the lepra bacillus is that when these +insects feed they insert their proboscis directly into a blood vessel +and thus obtain bacilli-free blood, unmixed with lymph. + +The same worker undertook to determine whether flies are able to +transmit leprosy. He experimented with five species found in +Honolulu,--_Musca domestica_, _Sarcophaga pallinervis_, _Sarcophaga +barbata_, _Volucella obesa_ and an undetermined species of _Lucilia_. +The experiments with _Musca domestica_ were the most detailed. From +these experiments he concluded, first, that all of the above-named +flies, when given an opportunity to feed upon leprous fluids, will +contain the bacilli in their intestinal tracts and feces for several +days after such feeding. Second, that considering the habits of these +flies, and especially those of _Musca domestica_, it is certain that, +given an exposed leprous ulcer, these insects will frequently convey +immense numbers of lepra bacilli, directly or indirectly, to the skins, +nasal mucosa, and digestive tracts of healthy persons. Additional +evidence along this line has recently been brought forward by Honeij and +Parker (1914), who incriminate both _Musca domestica_ and _Stomoxys +calcitrans_. Whether or not such insect-borne bacilli are capable of +infecting persons whose skin and mucosa are thus contaminated, Currie +was unwilling to maintain, but he concludes that until we have more +accurate knowledge on this point, we are justified in regarding these +insects with grave suspicion of being one of the means of disseminating +leprous infection. + +Various students of the subject have suggested that bed-bugs may be the +carriers of leprosy and have determined the presence of acid-fast +bacilli in the intestines of bed-bugs which had fed on leprous patients. +Opposed to this, the careful experiments of Thompson (1913) and of +Skelton and Parkham (1913) have been wholly negative. + +Borrel has recently suggested that _Demodex_, may play a rôle in +spreading the infection in families. Many other insects and acariens +have been suggested as possible vectors, but the experimental data are +few and in no wise conclusive. The most that can be said is that it is +quite possible that under favorable conditions the infection might be +spread by any of the several blood-sucking forms or by house-flies. + + +Verruga peruviana + +VERRUGA PERUVIANA is defined by Castellani and Chalmers as "a chronic, +endemic, specific, general disorder of unknown origin, not contagious, +but apparently inoculable, and characterized by an irregular fever +associated with rheumatoid pains, anemia, followed by granulomatous +swellings in the skin, mucous membranes, and organs of the body." It has +been generally believed by medical men interested that the comparatively +benign eruptive verruga is identical with the so-called Oroya, or +Carrion's fever, a malignant type. This view is not supported by the +work of Strong, Tyzzer and Brues, (1913). + +The disease is confined to South America and to definitely limited areas +of those countries in which it does occur. It is especially prevalent in +some parts of Peru. + +The causative organism and the method of transfer of verruga are +unknown. Castellani and Chalmers pointed out in 1910 that the study of +the distribution of the disease in Peru would impress one with the +similarity to the distribution of the Rocky Mountain fever and would +lead to the conclusion that the ætiological cause must in some way be +associated with some blood-sucking animal, perhaps an arachnid, and that +this is supported by the fact that the persons most prone to the +infection are those who work in the fields. + +More recently, Townsend (1913), in a series of papers, has maintained +that verruga and Carrion's disease are identical, and that they are +transmitted to man by the bites of the Psychodid fly, _Phlebotomus +verrucarum_. He succeeded in producing the eruptive type of the disease +in experimental animals by injecting a physiological salt trituration of +wild Phlebotomus flies. A cebus monkey was exposed from October so to +November 6, by chaining him to a tree in the verruga zone, next to a +stone wall from which the flies emerged in large numbers every night. +Miliar eruption began to appear on the orbits November 13 and by +November 21, there were a number of typical eruptions, with exudation on +various parts of the body exactly like miliar eruptive sores commonly +seen on legs of human cases. + +An assistant in the verruga work, George E. Nicholson, contracted the +eruptive type of the disease, apparently as a result of being bitten by +the Phlebotomus flies. He had slept in a verruga zone, under a tight +net. During the night he evidently put his hands in contact with the +net, for in the morning there were fifty-five unmistakable Phlebotomus +bites on the backs of his hands and wrists. + +Townsend believes that in nature, lizards constitute the reservoir of +the disease and that it is from them that the Phlebotomus flies receive +the infection. + + +Cancer + +There are not wanting suggestions that this dread disease is carried, or +even caused, by arthropods. Borrel (1909) stated that he had found mites +of the genus _Demodex_ in carcinoma of the face and of the mammæ. He +believed that they acted as carriers of the virus. + +Saul (1910) and Dahl (1910) go much further, since they attribute the +production of the malignant growth to the presence of mites which Saul +had found in cancers. These Dahl described as belonging to a new +species, which he designated _Tarsonemus hominis_. These findings have +since been confirmed by several workers. Nevertheless, the presence of +the mite is so rare that it cannot be regarded as an important factor in +the causation of the disease. The theory that cancer is caused by an +external parasite is given little credence by investigators in this +field. + + +IN CONCLUSION, it should be noted that the medical and entomological +literature of the past few years abounds in suggestions, and in +unsupported direct statements that various other diseases are +insect-borne. Knab (1912) has well said "Since the discovery that +certain blood-sucking insects are the secondary hosts of pathogenic +parasites, nearly every insect that sucks blood, whether habitually or +occasionally, has been suspected or considered a possible transmitter of +disease. No thought seems to have been given to the conditions and the +characteristics of the individual species of blood-sucking insects, +which make disease transmission possible." + +He points out that "in order to be a potential transmitter of human +blood-parasites, an insect must be closely associated with man and +normally have opportunity to suck his blood repeatedly. It is not +sufficient that occasional specimens bite man, as, for example, is the +case with forest mosquitoes. Although a person may be bitten by a large +number of such mosquitoes, the chances that any of these mosquitoes +survive to develop the parasites in question, (assuming such development +to be possible), and then find opportunity to bite and infect another +person, are altogether too remote. Applying this criterion, not only the +majority of mosquitoes but many other blood-sucking insects, such as +Tabanidæ and Simuliidæ, may be confidently eliminated. Moreover, these +insects are mostly in evidence only during a brief season, so that we +have an additional difficulty of a very long interval during which there +could be no propagation of the disease in question." He makes an +exception of tick-borne diseases, where the parasites are directly +transmitted from the tick host to its offspring and where, for this +reason, the insect remains a potential transmitter for a very long +period. He also cites the trypanosome diseases as possible exceptions, +since the causative organisms apparently thrive in a number of different +vertebrate hosts and may be transmitted from cattle, or wild animals, to +man. + +Knab's article should serve a valuable end in checking irresponsible +theorizing on the subject of insect transmission of disease. +Nevertheless, the principles which he laid down cannot be applied to the +cases of accidental carriage of bacterial diseases, or to those of +direct inoculation of pyogenic organisms, or of blood parasites such as +the bacillus of anthrax, or of bubonic plague. Accumulated evidence has +justified the conclusion that certain trypanosomes pathogenic to man are +harbored by wild mammals, and so form an exception. Townsend believes +that lizards constitute the natural reservoir of verruga; and it seems +probable that field mice harbor the organism of tsutsugamushi disease. +Such instances are likely to accumulate as our knowledge of the relation +of arthropods to disease broadens. + + + + +CHAPTER XII + +HOMINOXIOUS ARTHROPODS + + +The following synoptic tables are presented in the hope that they may be +of service in giving the reader a perspective of the relationships of +the Arthropoda in general and enabling him to identify the more +important species which have been found noxious to man. Though +applicable chiefly to the arthropods found in the United States, exotic +genera and species which are concerned in the transmission of disease +are also included. For this reason the keys to the genera of the Muscids +of the world are given. As will be seen, the tables embrace a number of +groups of species which are not injurious. This was found necessary in +order that the student might not be lead to an erroneous determination +which would result were he to attempt to identify a species which +heretofore had not been considered noxious, by means of a key containing +only the noxious forms. The names printed in BOLD FACED TYPE indicate +the hominoxious arthropods which have been most commonly mentioned in +literature. + + +CRUSTACEA + +Arthropods having two pairs of antennæ which are sometimes modified for +grasping, and usually with more than five pairs of legs. With but few +exceptions they are aquatic creatures. Representatives are: Crabs, +lobsters, shrimps, crayfish, water-fleas, and woodlice. To this class +belongs the CYCLOPS (fig. 122) a genus of minute aquatic crustaceans of +which at least one species harbors _Dracunculus medinensis_, the Guinea +worm (fig. 121). + + +MYRIAPODA + +Elongate, usually vermiform, wingless, terrestrial creatures having one +pair of antennæ, legs attached to each of the many intermediate body +segments. This group is divided into two sections, now usually given +class rank: the DIPLOPODA or millipedes (fig. 13), commonly known as +thousand legs, characterized by having two pairs of legs attached to +each intermediate body segment, and the CHILOPODA or centipedes (fig. +14) having only one pair of legs to each body segment. + + +ARACHNIDA + +In this class the antennæ are apparently wanting, wings are never +present, and the adults are usually provided with four pairs of legs. +Scorpions, harvest-men, spiders, mites, etc. + + +HEXAPODA (Insects) + +True insects have a single pair of antennæ, which is rarely vestigial, +and usually one or two pairs of wings in the adult stage. Familiar +examples are cockroaches, crickets, grasshoppers, bugs, dragon-flies, +butterflies, moths, mosquitoes, flies, beetles, ants, bees and wasps. + + +ORDERS OF THE ARACHNIDA + +a. Abdomen distinctly segmented. A group of orders including scorpions, + (fig. 11), whip-scorpions (fig. 10), pseudo-scorpions, solpugids + (fig. 12) harvest-men (daddy-long-legs or harvestmen), etc. + ARTHROGASTRA + +aa. Abdomen unsegmented, though sometimes with numerous annulations + SPHÆROGASTRA + + b. A constriction between cephalothorax and abdomen (fig. 7). True + Spiders ARANEIDA + + bb. No deep constriction between these parts. + + c. Legs usually well developed, body more or less depressed (fig. + 49). Mites ACARINA + + cc. Legs stumpy or absent, body more or less elongate or vermiform, + or if shorter, the species is aquatic or semi-aquatic in habit. + + d. Four pairs of short legs; species inhabiting moss or water. + Water-bears. TARDIGRADA + + dd. Two pairs of clasping organs near the mouth, instead of legs, + in the adult; worm-like creatures parasitic within the nasal + passages, lungs, etc. of mammals and reptiles (fig. 148). + Tongue worms. LINGUATULINA + +[Illustration: 148. Linguatula. (_a_) larva; (enlarged). (_b_) adult; +(natural size).] + + +ACARINA[E] + +a. Abdomen annulate, elongate; very minute forms, often with but four + legs (fig. 62). DEMODICOIDEA + + b. With but four legs of five segments each. Living on plants, often + forming galls. ERIOPHYIDÆ + + bb. With eight legs, of three segments each. Living in the skin of + mammals. DEMODICIDÆ + + To this family belongs the genus DEMODEX found in the sebaceous + glands and hair follicles of various mammals, including man. _D. + phylloides_ Csokor has been found in Canada on swine, causing + white tubercles on the skin. _D. bovis_ Stiles has been reported + from the United States on cattle, upon the skin of which they + form swellings. D. FOLLICULORUM Simon is the species found on + man. See page 78. + +aa. Abdomen not annulate nor prolonged behind; eight legs in the adult + stage. + + b. With a distinct spiracle upon a stigmal plate on each side of the + body (usually ventral) above the third or fourth coxæ or a little + behind (fig. 50); palpi free; skin often coriaceous or leathery; + tarsi often with a sucker. + + c. Hypostome large (fig. 50), furnished below with many recurved + teeth; venter with furrows, skin leathery; large forms, usually + parasitic. IXODOIDEA + + d. Without scutum but covered by a more or less uniform leathery + integument; festoons absent; coxæ unarmed, tarsi without + ventral spurs; pulvilli absent or vestigial in the adults; + palpi cylindrical; sexual dimorphism slight. ARGASIDÆ + + e. Body flattened, oval or rounded, with a distinct flattened + margin differing in structure from the general integument; + this margin gives the body a sharp edge which is not + entirely obliterated even when the tick is full fed. + Capitulum (in adults and nymphs) entirely invisible + dorsally, distant in the adult by about its own length from + the anterior border. Eyes absent. ARGUS Latr. + + f. Body oblong; margin with quadrangular cells; anterior tibiæ + and metatarsi each about three times as long as broad. On + poultry, southwest United States. A. PERSICUS MINIATUS + + _A. brevipes_ Banks, a species with proportionately shorter + legs has been recorded from Arizona. + + ff. With another combination of characters. About six other + species of _Argas_ from various parts of the world, + parasitic on birds and mammals. + + ee. Body flattened when unfed, but usually becoming very convex + on distention; anterior end more or less pointed and + hoodlike; margin thick and not clearly defined, similar in + structure to the rest of the integument and generally + disappearing on distention; capitulum subterminal, its + anterior portions often visible dorsally in the adult; eyes + present in some species. + + f. Integument pitted, without rounded tubercles; body provided + with many short stiff bristles; eyes absent. On horses, + cattle and man (fig. 48). OTIOBIUS Banks. + + O. MEGNINI, a widely distributed species, is the type of + this genus. + + ff. Integument with rounded tubercles or granules; body + without stiff bristles. ORNITHODOROS Koch. + + g. Two pairs of eyes; tarsi IV with a prominent subterminal + spur above; leg I strongly roughened. On cattle and man. + O. CORIACEUS + + gg. No eyes; no such spur on the hind tarsi. + + h. Tarsi I without humps above. _O. talaje._ + + hh. Tarsi I with humps above. + + i. Tarsi IV without distinct humps above. On hogs, + cattle and man. O. TURICATA + + ii. Tarsi IV with humps nearly equidistant (fig. 142). + Africa. O. MOUBATA + +[Illustration: 149. Hæmaphysalis wellingtoni. Note short palpi. After +Nuttall and Warburton.] + + dd. With scutum or shield (fig. 50); festoons usually present; + coxæ usually armed with spurs, tarsi generally with one or two + ventral spurs; pulvilli present in the adults; sexual + dimorphism pronounced. IXODIDÆ + + e. With anal grooves surrounding anus in front; inornate; + without eyes; no posterior marginal festoons; venter of the + male with non-salient plates. Numerous species, 14 from the + United States, among them I. RICINUS (fig. 49 and 50), + SCAPULARIS, COOKEI, _hexagonus_, _bicornis_. IXODES Latr. + (including Ceratixodes). + + ee. With anal groove contouring anus behind, or groove faint or + obsolete. + + f. With short palpi (fig. 149). + + g. Without eyes, inornate, with posterior marginal festoons; + male without ventral plates. Numerous species. _H. + chordeilis_ and _leporis-palustris_ from the United + States. _Hæmaphysalis_ Koch. + +[Illustration: 150. Stigmal plate of Dermacentor andersoni; (_a_) of +male, (_b_) of female. After Stiles. (_c_) Dermacentor variabilis, male; +(_d_) Glyciphagus obesus; (_e_) Otodectes cynotis; (_f_) Tyroglyphus +lintneri; (_g_) Tarsonemus pallidus; (_h_) anal plate and mandible of +Liponyssus; (_c_) to (_h_) after Banks.] + + gg. With eyes. + + h. Anal groove distinct; posterior marginal festoons + present. + + i. Base of the capitulum (fig. 150c) rectangular + dorsally; usually ornate. DERMACENTOR Koch. + + j. Adults with four longitudinal rows of large + denticles on each half of hypostome; stigmal plate + nearly circular, without dorso-lateral + prolongation, goblets very large, attaining 43µ to + 115µ in diameter; not over 40 per plate, each + plate surrounded by an elevated row of regularly + arranged supporting cells; white rust wanting; + base of capitulum distinctly broader than long, + its postero-lateral angles prolonged slightly, if + at all; coxæ T with short spurs; trochanter I with + small dorso-terminal blade. Texas, Arizona, etc. + _D. nitens_ + +[Illustration: 151. Rhipicephalus bursa, male. After Nuttall and +Warburton.] + + jj. Adults with three longitudinal rows of large + denticles on each half of hypostome; goblet cells + always more than 40 per plate; whitish rust + usually present. + + k. Dorso-lateral prolongation of stigmal plate small + or absent; plates of the adults distinctly + longer than broad; goblet cells large, usually + 30µ to 85µ in diameter, appearing as very coarse + punctations on untreated specimens, but on + specimens treated with caustic potash they + appear very distinct in outline; base of + capitulum distinctly (usually about twice) + broader than long, the postero-lateral angles + distinctly produced caudad; spurs of coxæ I + long, lateral spur slightly longer than median; + trochanter I with dorso-terminal spur. _D. + albipictus_, (= _variegatus_), _salmoni_, + _nigrolineatus_. + + kk. Dorso-lateral prolongation of stigmal plate + distinct. + + l. Body of plate distinctly longer than broad; + goblet cells of medium size, usually 17.5µ to + 35µ or 40µ in diameter, appearing as medium + sized punctuations on untreated specimens, but + on the specimens treated with caustic potash + they appear very distinct in outline, which is + not circular; base of capitulum usually less + than twice as broad as long, the + postero-lateral angles always distinctly + prolonged caudad. + + m. Trochanter I with distinct dorso-subterminal + retrograde sharp, digitate spur; + postero-lateral angles of capitulum + pronouncedly prolonged caudal, 112µ to 160µ + long; goblet cells attain 13µ to 40µ in + diameter; type locality California. D. + OCCIDENTALIS + + mm. Trochanter I with dorso-terminal blade; + postero-lateral angles of capitulum with + rather short prolongations. + + n. Stigmal plate small, goblet cells not + exceeding 45 in the male or 100 in the + female; scutum with little rust, coxa I + with short spurs, the inner distinctly + shorter than the outer. _D. + parumapertus-marginatus_ + + nn. Stigmal plate larger; goblet cells over 70 + in the male and over 100 in the female; + coxa I with longer spurs, inner slightly + shorter than the outer; scutum with + considerable rust. D. VENUSTUS[F] + + ll. Goblet cells small, rarely exceeding 17.6µ, + occasionally reaching 19µ in diameter; on + untreated specimens they appear as very fine + granulations, and on specimens treated with + caustic potash they may be difficult to see, + but their large number can be determined from + the prominent stems of the goblets; surface of + outline of the goblets distinctly circular; + base of the capitulum usually less than twice + as broad as long, the postero-lateral angle + distinctly prolonged caudad; spurs of coxæ I + long. _D. reticulatus_ and _electus_ (= + _variabilis_?) + + ii. Base of the capitulum (fig. 151) usually hexagonal + (except in the male of _puchellus_); and usually + inornate. + + j. No ventral plate or shield in either sex (fig. + 153). R. BICORNIS from the United States. + RHIPICENTOR Nuttall + + jj. Males with a pair of adanal shields, and usually a + pair of accessory adanal shields. Numerous + species, among them _R. sanguineus_ (fig. 154) and + _texanus_, the latter from the United States. + _Rhipicephalus_ Koch + + hh. Anal grooves faint or obsolete; no marginal festoons. + + i. Short palpi; highly chitinized; unfed adults of large + size; coxæ conical; male with a median plate + prolonged in two long spines projecting caudad; + segments of leg pair IV greatly swollen (fig. 155, + 156). _M. winthemi_ _Margaropus_ Karsch + +[Illustration: 152. Monieziella (Histiogaster) emtomophaga-spermatica, +ventral aspect, male and female. After Trouessart.] + + ii. Very short palpi, ridged dorsally and laterally; + slightly chitinized; unfed adults of smaller size; + coxæ I bifid; male with adanal and accessory adanal + shields (fig. 139). B. ANNULATUS. BOOPHILUS Curtis + + ff. Palpi longer than broad (fig. 157). + + g. Male with pair of adanal shields, and two posterior + abdominal protrusions capped by chitinized points; + festoons present or absent. Several species, among them + H. ÆGYPTICUM (fig. 140) from the old world. HYALOMMA + Koch + + gg. Male without adanal shields but small ventral plaques + are occasionally present close to the festoons. Many + species, a few from the United States (fig. 157). + AMBLYOMMA Koch + + h. Coxa I with but one spine; metatarsi (except I) with + two thickened spurs at tips. _A. maculatum_ + + hh. Coxa I with two spines; metatarsi without stout spurs + at tips, only slender hairs. + + i. Projections of coxa I blunt and short. Large species + on the gopher tortoise in Florida. _A. tuberculatum_ + + ii. Projections of coxa I longer, and at least one of + them sharp pointed; second segment of palpus twice + as long as the third; coxa IV of the male with a + long spine. + + j. Porose areas nearly circular; shield of both sexes + pale yellowish, with some silvery streaks and + marks, and some reddish spots; shield of female as + broad as long. A. CAJENNENSE (= MIXTUM). + + jj. Porose areas elongate, shield brown, in the female + with an apical silvery mark, in the male with two + small and two or four other silvery spots; shield + of the female longer than broad (fig 158 e). A. + AMERICANUM. + +[Illustration: 153. Rhipicentor bicornis, ventral aspect, male. After +Nuttall and Warburton.] + + cc. Hypostome small, without teeth, venter without furrows; body + often with coriaceous shields, posterior margin of the body + never crenulate (i.e. without festoons); no eyes. GAMASOIDEA. + + d. Parasitic on vertebrates; mandibles fitted for piercing; body + sometimes constricted. DERMANYSSIDÆ. + + e. Anal plate present. DERMANYSSINÆ. + + f. Body short; legs stout, hind pair reaching much beyond the + tip of the body. On bats. _Pteroptus_ Dufour. + + ff. Body long; hind legs not reaching beyond the tip of the + body. + + g. Peritreme on the dorsum, very short; body distinctly + constricted. _Ptilonyssus_ Berl. + + gg. Peritreme on the venter, longer; body not distinctly + constricted. + + h. Mandibles in both sexes chelate. Parasitic on bats, + mice and birds (fig. 150, h). LIPONYSSUS Kol. + + The species L. (= LEIOGNATHUS) SYLVIARUM frequents the + nests of warblers. An instance is on record of + these mites attacking man, causing a pruritis. + + hh. Mandibles in the male chelate, in the female long, + styliform (fig. 158 j). Parasitic on birds. + DERMANYSSUS Dug. Two species of importance may be + noted, _D. hirundinus_ and D. GALLINÆ. The latter + (fig. 51) is a serious pest of poultry, sometimes + attacking man, causing itching and soreness. + + ee. Anal plate absent. In lungs and air passages of some + mammals. HALARACHNINÆ. + + dd. Free or attached to insects, rarely on vertebrates. + + e. First pair of legs inserted within the same body opening as + the oral tube; genital apertures surrounded by the sternum. + On insects. UROPODIDÆ. + +[Illustration: 154. Rhicephalus sanguineus, male. After Nuttall and +Warburton.] + + ee. First pair of legs inserted at one side of the mouth + opening; male genital aperture usually on the anterior + margin of the sternal plate. GAMASIDÆ. + + This family contains a number of genera, some of which are + found upon mammals, though the majority affect only other + arthropods. One species, LÆLAPS STABULARIS, frequents the + bedding in stables, and in one instance at least, has + occasioned irritation and itching, in man. + + bb. No distinct spiracle in the stigmal plate on each side of the + body. + + c. Body usually coriaceous, with few hairs, with a specialized seta + arising from a pore near each posterior corner of the + cephalothorax; no eyes; mouth parts and palpi very small; + ventral openings of the abdomen large; tarsi without sucker. Not + parasitic. ORIBATOIDEA. + + cc. Body softer; without such specialized seta. + + d. Aquatic species. HYDRACHNOIDEA. + + dd. Not aquatic. + + e. Palpi small, three segmented, adhering for some distance to + the lip; ventral suckers at genital opening or near anal + opening usually present; no eyes; tarsi often end in + suckers; beneath the skin on the venter are seen rod-like + epimera that support the legs; body often entire. Adults + frequently parasitic. SARCOPTOIDEA. + + f. With tracheæ; no ventral suckers; legs ending in claws; + body divided into cephalothorax and abdomen; the female + with a clavate hair between legs I and II. Usually not + parasitic on birds and mammals. TARSONEMIDÆ + + g. Hind legs of female ending in claw and sucker as in the + other pairs. PEDICULOIDINÆ + + To this sub-family belongs the genus PEDICULOIDES + + P. ventricosus is described on page 69. + +[Illustration: 155. Margaropus winthemi, male. After Nuttall and +Warburton.] + +[Illustration: 156. Margaropus winthemi, capitulum and scutum. +After Nuttall and Warburton.] + + gg. Hind legs of the female end in long hairs. TARSONEMINÆ + + TARSONEMUS INTECTUS Karpelles, normally found upon grain, + is said to attack man in Hungary and Russia. Other + species of the genus affect various plants (c.f. fig. + 150, g). + + ff. Without tracheæ; no such clavate hair. + + g. Genital suckers usually present; integument usually + without fine parallel lines. + + h. Legs short, without clavate hair on tarsi I and II. On + insects. CANESTRINIDÆ. + + hh. Legs longer, with a clavate hair on tarsi I and II. + Not normally parasitic except on bees. TYROGLYPHIDÆ + + i. Dorsal integument more or less granulate; claws very + weak, almost invisible; some hairs of the body + plainly feathered; ventral apertures large. + GLYCIPHAGUS Her. + + This genus occurs in the United States. In Europe the + mites have been found feeding on all sorts of + substances. They are known as sugar mites and + cause the disease known as grocer's itch. G. + DOMESTICUS and G. PRUNORUM are old world species + (fig. 150, d). + + ii. Dorsal integument not granulate; claws distinct; no + prominent feathered hairs; ventral aperture small. + + j. Mandibles not chelate; elongate, and toothed below; + body without long hairs; palpi enlarged at tip and + provided with two divergent bristles. Species feed + on decaying substances. _Histiostoma_ Kram. + + jj. Mandibles chelate; palpi not enlarged at the tip, + nor with two bristles. + + k. No clavate hair on the base of tarsi I and II; no + suture between cephalothorax and abdomen. Live + on bees or in their nests. _Trichotarsus_ Can. + + kk. A clavate or thickened hair at the base of tarsi + I and II. + + l. The bristle on the penultimate segment of the + legs arises from near the middle; no suture + between the cephalothorax and abdomen. The + species, some of which occur in the United + States, feed on dried fruit, etc. + _Carpoglyphus_ Robin. + + ll. The bristle on the penultimate segment of the + legs arise from near the tip; a suture between + cephalothorax and abdomen. + + m. Cephalothorax with four distinct and long + bristles in a transverse row; tarsi I and II + about twice as long as the preceding segment + (fig. 150 f). TYROGLYPHUS Latr. + + n. Some bristles on tarsi I and II near + middle, distinctly spine-like; the sense + hair about its length from the base of the + segment. Several species in the United + States belong to this group. + + nn. No spine-like bristles near the middle of + the tarsi; sense hair not its length from + the base of the segment. + + o. Of the terminal abdominal bristles, only + two are about as long as the abdomen; + leg I of the male greatly thickened and + with a spine at apex of the femur below. + T. FARINÆ. + + oo. Of the terminal abdominal bristles at + least six or more are very long, nearly + as long as the body. + + p. Bristles of the body distinctly plumose + or pectinate; tarsi very long. T. + LONGIOR. + + pp. Bristles of the body not pectinate. + + q. In mills, stored foods, grains, etc. + Third and fourth joints of hind legs + scarcely twice as long as broad; + abdominal bristles not unusually + long; legs I and II of the male not + unusually stout. T. AMERICANUS. + + qq. With other characters and habits. + _T. lintneri_ (fig. 150 f) the + mushroom mite, and several other + species. + + mm. Cephalothorax with but two long distinct + bristles (besides the frontal pair), but + sometimes a very minute intermediate pair; + tarsi I and II unusually short and not twice + as long as the preceding segment. + + n. Tarsi with some stout spines. RHIZOGLYPHUS + Clap. + + The species of this genus are vegetable + feeders. Several occur in the United + States. R. PARASITICUS and R. + SPINITARSUS have been recorded from the + old world, attacking human beings who + handle affected plants. + + nn. Tarsi with only fine hairs. MONIEZIELLA + Berl. + + The species of this genus, as far as known, + are predaceous or feed on recently + killed animal matter. Several species + occur in the United States. M. (= + HISTIOGASTER) ENTOMOPHAGA (fig. 152) + from the old world has been recorded as + injurious to man. + + gg. Genital suckers absent; integument with fine parallel + lines. Parasitic on birds and mammals. + + h. Possessing a specially developed apparatus for clinging + to hairs of mammals. LISTROPHORIDÆ. + + hh. Without such apparatus. + + i. Living on the plumage of birds. ANALGESIDÆ. + + ii. In the living tissues of birds and mammals. + + j. Vulva longitudinal. In the skin and cellular + tissues of birds. CYTOLEICHIDÆ. + + This family contains two species, both occurring in + the United States on the common fowl. + _Laminosioptes cysticola_ occurs on the skin and + also bores into the subcutaneous tissue where it + gives rise to a calcareous cyst. _Cytoleichus + nudus_ is most commonly found in the air + passages and air cells. + + jj. Vulva transverse. In the skin of mammals and + birds. SARCOPTIDÆ + + k. Anal opening on the dorsum. + + l. Third pair of legs in the male without apical + suckers. On cats and rabbits. NOTÅ’DRES + Rail. + + The itch mite of the cat, N. CATI (fig. 61) has + been recorded on man. + + ll. Third leg in the male with suckers. On bats + _Prosopodectes_ Can. + + kk. Anal opening below. + + l. Pedicel of the suckers jointed; mandibles + styliform and serrate near the tip. PSOROPTES + Gerv. P. COMMUNIS OVIS is the cause of sheep + scab. + + ll. Pedicel of the suckers not jointed; mandibles + chelate. + + m. No suckers on the legs of the females; + parasitic on birds, including chickens. _C. + mutans_ is itch mite of chickens. + _Cnemidocoptes_ Fürst. + + mm. Suckers at least on legs I and II; parasitic + on mammals. + + n. Legs very short; in the male the hind pairs + equal in size; body usually short. + SARCOPTES Latr. + + S. SCABIEI is the itch mite of man (fig. + 56). + +[Illustration: 157. Amblyomma, female. After Nuttall and Warburton.] + + nn. Legs more slender; in the male the third + pair is much larger than the fourth; body + more elongate. + + o. Female with suckers on the fourth pair of + legs. Species do not burrow in the skin, + but produce a scab similar to sheep + scab. They occur in the ox, horse, sheep + and goat. CHORIOPTES Gerv. C. SYMBIOTES + BOVIS of the ox has been recorded a few + times on man. + + oo. Female without suckers to the fourth + legs. + + p. Hind part of the male abdomen with two + lobes. On a few wild animals. + _Caparinia_ Can. + + pp. Hind part of the male abdomen without + lobes. Live in ears of dogs and cats. + _Otodectes_ Canestr. + + _O. cynotis_ Hering (fig. 150 e) has + been taken in the United States. + + ee. Palpi usually of four or five segments, free; rarely with + ventral suckers near genital or anal openings; eyes often + present; tarsi never end in suckers; body usually divided + into cephalothorax and abdomen; rod-like epimera rarely + visible; adults rarely parasitic. + + f. Last segment of the palpi never forms a thumb to the + preceding segment; palpi simple, or rarely formed to hold + prey; body with but few hairs. EUPODOIDEA. + + g. Palpi often geniculate, or else fitted for grasping prey; + mandibles large and snout like; cephalothorax with four + long bristles above, two in front, two behind; last + segment of leg I longer than the preceding segment, + often twice as long. BDELLIDÆ. + + gg. Palpi never geniculate (fig. 158a), nor fitted for + grasping prey: beak small; cephalothorax with bristles + in different arrangement; last segment of leg I shorter + or but little longer than the preceding joint; eyes when + present near posterior border. EUPODIDÆ + + Moniez has described a species from Belgium (TYDEUS + MOLESTUS) which attacks man. It is rose colored; + eyeless; its legs are scarcely as long as its body, + the hind femur is not thickened; the mandibles are + small and the anal opening is on the venter. The + female attains a length of about 0.3 mm. + + ff. Last segment of the palpus forms a thumb to the preceding, + which ends in a claw (with few exceptions); body often + with many hairs (fig. 158 k). TROMBIDOIDEA. + + g. Legs I and II with processes bearing spines; skin with + several shields; coxæ contiguous. CÆCULIDÆ. + + gg. Legs I and II without such processes; few if any + shields. + + h. Palpi much thickened on the base, moving laterally, + last joint often with two pectinate bristles; no eyes; + legs I ending in several long hairs; adult sometimes + parasitic. CHEYLETIDÆ + + CHEYLETUS ERUDITUS, which frequents old books, has once + been found in pus discharged from the ear of man. + + hh. Palpi less thickened, moving vertically; eyes usually + present; leg I not ending in long hairs. + + i. Coxæ contiguous, radiate; legs slender, bristly; body + with few hairs; no dorsal groove; tarsi not swollen. + ERYTHRÆIDÆ. + + ii. Coxæ more or less in two groups; legs less bristly. + +[Illustration: 158. (_a_) Tydeus, beak and leg from below; (_b_) +Cheyletus pyriformis, beak and palpus; (_c_) beak and claw of +Pediculoides; (_d_) leg of Sarcoptes; (_e_) scutum of female of +Amblyomma americana; (_f_) leg I and tip of mandible of Histiostoma +americana; (_g_) Histiogaster malus, mandible and venter; (_h_) +Aleurobius farinæ, palpus and leg I of male; (_i_) Otodectes cynotis, tip +of abdomen of male, (_j_) beak and anal plate of Dermanyssus gallinæ; +(_k_) palpus of Allothrombium. (_a_) to (_j_) after Banks.] + + j. Body with fewer, longer hairs; often spinning + threads; no dorsal groove; tarsi never swollen; + mandibles styliform (for piercing). TETRANYCHIDÆ + + The genus TETRANYCHUS may be distinguished from the + other genera occurring in the United States by + the following characters: No scale-like + projections on the front of the cephalothorax; + legs I as long or longer than the body; palp + ends in a distinct thumb; the body is about 1.5 + times as long as broad. T. MOLESTISSIMUS Weyenb. + from South America, and T. TELARIUS from Europe + and America ordinarily infesting plants, are + said also to molest man. + + jj. Body with many fine hairs or short spines; not + spinning threads; often with dorsal groove; tarsi + often swollen. + + k. Mandibles styliform for piercing. RHYCHOLOPHIDÆ. + + kk. Mandibles chelate, for biting. TROMBIDIDÆ + + The genus TROMBIDIUM has recently been subdivided + by Berlese into a number of smaller ones, of + which some five or six occur in the United + States. The mature mite is not parasitic but + the larvæ which are very numerous in certain + localities will cause intense itching, + soreness, and even more serious complications. + They burrow beneath the skin and produce + inflamed spots. They have received the + popular name of "RED BUG." The names LEPTUS + AMERICANUS and L. IRRITANS have been applied + to them, although they are now known to be + immature stages. (Fig. 44.) + + +HEXAPODA (Insecta) + +The Thysanura (springtails and bristletails), the Neuropteroids +(may-flies, stone-flies, dragon-flies, caddis-flies, etc.), Mallophaga +(bird lice), Physopoda (thrips), Orthoptera (grasshoppers, crickets, +roaches), are of no special interest from our viewpoint. The remaining +orders are briefly characterized below. + + +SIPHUNCULATA (page 275) + +Mouth parts suctorial; beak fleshy, not jointed; insect wingless; +parasitic upon mammals. Metamorphosis incomplete. Lice. + + +HEMIPTERA (page 275) + +Mouth parts suctorial; beak or the sheath of the beak jointed; in the +mature state usually with four wings. In external appearance the +immature insect resembles the adult except that the immature form (i.e. +nymph) never has wings, the successive instars during the process of +growth, therefore, are quite similar; and the metamorphosis is thus +incomplete. To this order belong the true bugs, the plant lice, leaf +hoppers, frog hoppers, cicadas, etc. + + +LEPIDOPTERA + +The adult insect has the body covered with scales and (with the rare +exception of the females of a few species) with four wings also covered +with scales. Proboscis, when present, coiled, not segmented, adapted for +sucking. Metamorphosis complete, i.e. the young which hatches from the +egg is quite unlike the adult, and after undergoing several molts +transforms into a quiescent pupa which is frequently enclosed in a +cocoon from which the adult later emerges. The larvæ are known as +caterpillars. Butterflies and moths. + + +DIPTERA (page 285) + +The adult insect is provided with two, usually transparent, wings, the +second pair of wings of other insects being replaced by a pair of +halteres or balancers. In a few rare species the wings, or halteres, or +both, are wanting. The mouth parts, which are not segmented, are adapted +for sucking. The tarsi are five-segmented. Metamorphosis complete. The +larvæ, which are never provided with jointed legs, are variously known +as maggots, or grubs, or wrigglers. Flies, midges, mosquitoes. + + +SIPHONAPTERA (page 316) + +Mouth parts adapted for sucking; body naked or with bristles and spines; +prothorax well developed; body compressed; tarsi with five segments; +wings absent. Metamorphosis complete. The larva is a wormlike creature. +Fleas. + + +COLEOPTERA + +Adult with four wings (rarely wanting), the first pair horny or +leathery, veinless, forming wing covers which meet in a line along the +middle of the back. Mouth parts of both immature stages and adults +adapted for biting and chewing. Metamorphosis complete. The larvæ of +many species are known as grubs. Beetles. + + +HYMENOPTERA + +Adult insect with four, usually transparent, wings, wanting in some +species. Mouth parts adapted for biting and sucking; palpi small; tarsi +four or five-segmented. Metamorphosis complete. Parasitic four-winged +flies, ants, bees, and wasps. + + +SIPHUNCULATA AND HEMIPTERA + +a. Legs with claws fitted for clinging to hairs; wings wanting; + spiracles of the abdomen on the dorsal surface. (= ANOPLURA = + PARASITICA) SIPHUNCULATA. + + b. Legs not modified into clinging hooks; tibia and tarsus very long + and slender; tibia without thumb-like process; antennæ + five-segmented HÆMATOMYZIDÆ Endr. + + _Hæmatomyzus elephantis_ on the elephant. + + bb. Legs modified into clinging hooks; tibia and tarsus usually short + and stout; tibia with a thumb-like process; head not anteriorly + prolonged, tube-like. + + c. Body depressed; a pair of stigmata on the mesothorax, and + abdominal segments three to eight; antennæ three to + five-segmented. + + d. Eyes large, projecting, distinctly pigmented; pharynx short and + broad; fulturæ (inner skeleton of head) very strong and broad, + with broad arms; proboscis short, scarcely attaining the + thorax. PEDICULIDÆ + + e. Antennæ three-segmented. A few species occurring upon old + world monkeys. _Pedicinis_ Gerv. + + ee. Antennæ five-segmented. + + f. All legs stout; thumb-like process of the tibia very long + and slender, beset with strong spines, fore legs stouter + than the others; abdomen elongate, segments without + lateral processes; the divided telson with a conical + process posteriorly upon the ventral side. PEDICULUS L. + + g. Upon man. + + h. Each abdominal segment dorsally with from one to three + more or less regular transverse rows of small setæ; + antenna about as long as the width of the head. Head + louse (fig. 65). P. HUMANUS. + + hh. "No transverse rows of abdominal setæ; antenna longer + than the width of the head; species larger." Piaget. + Body louse of man. P. CORPORIS. + + gg. Upon apes and other mammals. _P. pusitatus_ (?). + + ff. Fore legs delicate, with very long and slender claws; + other legs very stout with short and stout claws; + thumb-like process of the tibia short and stout; abdomen + very short and broad; segment one to five closely crowded, + thus the stigmata of segments three to five apparently + lying in one segment; segments five to eight with lateral + processes; telson without lateral conical appendages (fig. + 69). Crab louse of man. PHTHIRUS PUBIS. + + dd. Eyes indistinct or wanting; pharynx long and slender, fulturæ + very slender and closely applied to the pharynx; proboscis + very long. Several genera found upon various mammals. + HÆMATOPINIDÆ. + + cc. Body swollen; meso- and metathorax, and abdominal segments two + to eight each with a pair of stigmata; eyes wanting; antennæ + four or five-segmented; body covered with stout spines. Three + genera found upon marine mammals. ECHINOPHTHIRIIDÆ + +aa. Legs fitted for walking or jumping; spiracles of abdomen usually + ventral; beak segmented. + + b. Apex of head usually directed anteriorly; beak arising from its + apex; sides of the face remote from the front coxæ; first pair of + wings when present thickened at base, with thinner margins. + HETEROPTERA + +[Illustration: 159. Taxonomic details of Hemiptera-Heteroptera. (_a_) +Dorsal aspect; (_b_) seta from bedbug; (_c_) wing of Heteropteron; (_d_) +leg; (_e_) wing of Sinea.] + + c. Front tarsi of one segment, spade-form (palæformes); beak short, + at most two-segmented; intermediate legs long, slender; + posterior pair adapted for swimming. CORIXIDÆ + + cc. Front tarsi rarely one-segmented, never spade-form; beak free, + at least three-segmented. + + d. Pulvilli wanting. + + e. Hemelytra usually with a distinct clavus (fig. 159), clavus + always ends behind the apex of the scutellum, forming the + commissure. (Species having the wings much reduced or + wanting should be sought for in both sections.) + + f. Antennæ very short; meso- and metasternum composite; eyes + always present. + + g. Ocelli present; beak four-segmented. OCHTERIDÆ and + NERTHRIDÆ. + + gg. Ocelli wanting; antennæ more or less hidden in a groove. + + h. Anterior coxæ inserted at or near anterior margin of + the prosternum; front legs raptorial; beak + three-segmented BELOSTOMIDÆ (with swimming legs), + NEPIDÆ, NAUCORIDÆ. + + i. Metasternum without a median longitudinal keel; + antennæ always four-segmented. + + j. Beak short, robust, conical; the hairy fleck on the + corium elongate, large, lying in the middle + between the inner angle of the membrane and the + outer vein parallel to the membrane margin; + membrane margin S-shaped. + + k. The thick fore femur with a relatively deep + longitudinal furrow to receive the tibia. + Several American species (fig. 19f.). BELOSTOMA + (= Lethocerus Mayer) + + kk. The less thickened fore femur without such a + furrow. B. GRISEUS. BENACUS Stäl. + + jj. Beak slender, cylindrical; the hairy spot on the + corium rounded lying next to the inner angle of + the membrane. + + k. Membrane large, furrow of the embolium broadened. + _Z. aurantiacum_, _fluminea_, etc. _Zaitha_ + + kk. Membrane very short; furrow of embolium not + broadened. Western genus. _Pedinocoris_ + + ii. Metasternum with a long median longitudinal keel. + Southwestern forms. _Abedus ovatus_ and _Deniostoma + dilatato_ + + hh. Anterior coxæ inserted at the posterior margin of the + prosternum; legs natatorial. Back swimmers (fig. 19 b.). + NOTONECTIDÆ + + i. Apices of the hemelytra entire; the three pairs of legs + similar in shape; beak three-segmented; abdomen not + keeled or hairy. _Plea_ Leach + + ii. Apices of hemelytra notched; legs dissimilar; beak + four-segmented; abdomen keeled and hairy. + + j. Hemelytra usually much longer than the abdomen; + fourth segment of the antenna longer than the third + segment; hind tarsi with claws. _Bueno_ Kirk. + + jj. Hemelytra but little longer than the abdomen; fourth + segment of the antenna shorter than the third + segment; hind tarsi without claws (fig. 19b). + NOTONECTA L. + + ff. Antennæ longer than the head; or if shorter, then the eyes + and ocelli absent. + + g. Eyes, ocelli, and scutellum wanting; beak + three-segmented; head short; hemelytra always short; + membrane wanting. Insects parasitic on bats. POLYCTENIDÆ + + gg. Eyes present. + + h. First two antennal segments very short, last two long, + pilose, third thickened at the base; ocelli present, + veins of the hemelytra forming cells. DIPSOCORIDÆ (= + CERATOCOMBIDÆ) including SCHIZOPTERIDÆ. + + hh. Third segment of the antenna not thickened at the + base, second as long or longer than the third, rarely + shorter. + + i. Posterior coxæ hinged (cardinate), if rarely + rotating, the cuneus is severed, the membrane is one + or two-celled, and the meso- and metasternum are + composite. + + j. Ocelli absent, clypeus dilated toward the apex; + hemelytra always short, membrane wanting. Species + parasitic. Bed bugs, etc. CIMICIDÆ + + k. Beak short, reaching to about the anterior coxæ; + scutellum acuminate at the apex; lateral margin + of the elytra but little reflexed, apical margin + more or less rounded; intermediate and posterior + coxæ very remote. + + l. Body covered with short hairs, only the sides + of the pronotum and the hemelytra fringed with + longer hairs; antennæ with the third and + fourth segments very much more slender than + the first and second; pronotum with the + anterior margin very _deeply sinuate_. CIMEX + L. + + m. Sides of the pronotum widely dilated, broader + than the breadth of one eye, and densely + fringed with backward curved hairs; apical + margin of the hemelytra nearly straight, + rounded toward the interior or exterior + angles. + + n. Body covered with very short hairs; second + segment of the antenna shorter than the + third; sides of the pronotum feebly + reflexed, fringed with shorter hairs than + the breadth of one eye; hemelytra with the + commissural (inner) margin rounded and + shorter than the scutellum, apical margin + rounded towards the interior angle. The + common bed bug (fig. 19h). C. LECTULARIUS + Linn + + nn. Body covered with longer hairs; second and + third segments of the antenna of equal + length; side of the pronotum narrowly, but + distinctly, reflexed, fringed with longer + hairs than the breadth of one eye; + hemelytra with the commissural margin + straight and longer than the scutellum, + apical margin rounded towards the exterior + angle. Species found on bats in various + parts of the United States. _C. + pillosellus_ Hov. + + mm. Sides of the pronotum neither dilated, nor + reflexed, fringed with less dense and nearly + straight hairs; hemelytra with the apical + margin distinctly rounded. Parasitic on man, + birds and bats. Occurs in the old world, + Brazil and the West Indies. C. HEMIPTERUS + Fabr. (= rotundatus) + + ll. Body clothed with rather longer silky hairs; + third and fourth segments of the antenna + somewhat more slender than the first and + second; anterior margin of the pronotum _very + slightly sinuate_ or nearly straight in the + middle, produced at the lateral angles. This + is the species which in American collections + is known as _C. hirundinis_, the latter being + an old world form. It is found in swallows + nests. O. VICARIUS. OECIACUS Stäl + + kk. Beak long, reaching to the posterior coxæ; + scutellum rounded at the apex; lateral margins + of the elytra strongly reflexed, apical margin + slightly sinuate toward the middle; intermediate + and posterior coxæ sub-contiguous. This species + infests poultry in southwest United States and + in Mexico. H. INODORUS. HÆMATOSIPHON Champ. + +[Illustration: 160. Pselliopsis (Milyas) cinctus (×2). After C. V. +Riley.] + + jj. Ocelli present, if rarely absent in the female, + then the tarsus has two segments; or if with three + tarsal segments, the wing membrane with one or two + cells. + + k. Beak four-segmented, or with two-segmented tarsi. + ISOMETOPIDÆ, MICROPHYSIDÆ, and some CAPSIDÆ. + + kk. Beak three-segmented. + + l. Hemelytra with embolium; head horizontal, more + or less conical; membrane with one to four + veins, rarely wanting. ANTHOCORIDÆ + + Several species of this family affecting man + have been noted, ANTHOCORIS KINGI and + CONGOLENSE, from Africa and LYCTOCORIS + CAMPESTRIS from various parts of the world. + LYCTOCORIS FITCHII Reuter (fig. 19 j), later + considered by Reuter as a variety of L. + CAMPESTRIS, occurs in the United States. + + ll. Hemelytra without embolium. Superfamily + ACANTHIOIDEA (= SALDÆ Fieber and LEPTOPODÆ + Fieber) + + ii. Posterior coxæ rotating. + + j. Claws preapical; aquatic forms. GERRIDÆ and VELIADÆ + + jj. Claws apical. + + k. Prosternum without stridulatory sulcus (notch for + beak). + + l. Tarsus with three segments; membrane with two + or three longitudinal cells from which veins + radiate; rarely with free longitudinal veins + (Arachnocoris) or veins nearly obsolete + (Arbela); clavus and corium coriaceous; ocelli + rarely absent. NABIDÆ + + REDUVIOLUS (= CORISCUS) SUBCOLEOPTRATUS (fig. 19 + g), a species belonging to this family, + occurring in the United States, has been + accused of biting man. This insect is flat, + of a jet black color, bordered with yellow + on the sides of the abdomen, and with + yellowish legs. It is predaceous, feeding on + other insects. + + ll. With other combinations of characters. + HYDROMETRIDÆ, HENICOCEPHALIDÆ, NÆOGEIDÆ, + MESOVELIADÆ, JOPPEICIDÆ + + kk. Prosternum with stridulatory sulcus (notch for + beak); with three segments, short, strong. + + l. Antennæ filiform or sometimes more slender + apically, geniculate; wing membrane with two + or three large basal cells; scutellum small or + moderate REDUVIIDÆ + + For a key to the genera and species see next + page. + + ll. Last antennal segment clavate or fusiform; win + membrane with the veins often forked and + anastomosing; scutellum large; tarsi each with + two segments; fore legs strong. (= PHYMATIDÆ) + MACROCEPHALIDÆ + + ee. Clavus noticeably narrowed towards the apex, never extending + beyond the scutellum, the two not meeting to form a + commissure; head horizontal, much prolonged between the + antennæ, on each side with an antennal tubercle, sometimes + acute; ocelli absent; meso- and metasternum simple; tarsi + each with two segments; body flattened (fig. 19c). ARADIDÆ, + including DYSODIIDÆ. + + dd. Pulvilli present (absent in one Australian family + THAUMATOCORIDÆ in which case there is a membranous appendage + at the tip of the tibia). CAPSIDÆ (= MIRIDÆ),[G] _Eotrechus_ + (in family GERRIDÆ), NÆOGAIDÆ, TINGITIDÆ, PIESMIDÆ, + MYODOCHIDÆ, CORIZIDÆ, COREIDÆ, ALYDIDÆ, PENTATOMIDÆ, + SCUTELLERIDÆ, etc. + + bb. Apex of head directed ventrally, beak arising from the hinder part + of the lower side of the head; sides of face contiguous to the + front coxæ; first pair of wings, when present, of uniform + thickness. Cicadas, scale insects, plant lice (Aphids), + spittle-insects, leaf hoppers, etc. HOMOPTERA + + +REDUVIIDÆ OF THE UNITED STATES + +(Adapted from a key given by Fracker). + +a. Ocelli none; wings and hemelytra always present in the adults; no + discoidal areole in the corium near the apex of the clavus. + _Orthometrops decorata_, _Oncerotrachelus acuminatus_, etc., + Pennsylvania and south. _Sarcinæ_ + +aa. Ocelli present in the winged individuals; anterior coxæ not as long + as the femora. + + b. Hemelytra without a quadrangular or discoidal areole in the corium + near the apex of the clavus. + + c. Ocelli not farther cephalad than the caudal margins of the eyes; + segment two of the antenna single. + + d. Thorax usually constricted caudad of the middle; anterior coxæ + externally flat or concave. PIRATINÆ + + e. Middle tibiæ without spongy fossa, head long, no lateral + tubercle on neck. _S. stria_, Carolina, Ill., Cal. + _Sirthenia_ Spinola + + ee. Middle tibiæ with spongy fossa; fore tibiæ convex above; + neck with a small tubercle on each side. + + f. Apical portion of anterior tibiæ angularly dilated beneath, + the spongy fossa being preceded by a small prominence. + MELANOLESTES Stäl + + g. Black, with piceous legs and antennæ. N. E. States (fig. + 19a) M. PICIPES + + gg. Sides, and sometimes the whole dorsal surface of the + abdomen red. Ill., and southward. M. ABDOMINALIS + + ff. Tibiæ not dilated as in "f"; spongy fossa elongate; + metapleural sulci close to the margin. R. BIGUTTATUS (fig. + 22). South RASAHUS A. and S. + + dd. Thorax constricted in the middle or cephalad of the middle; + anterior tarsi each three-segmented. + + e. Apex of the scutellum narrow, without spines or with a single + spine REDUVIINÆ + + f. Antennæ inserted in the lateral or dorso-lateral margins of + the head; antenniferous tubercles slightly projecting from + the sides of the head; head produced strongly cephalad; + ocelli at least as far apart as the eyes. + + g. Antennæ inserted very near the apex of the head; segments + one and three of the beak short, segment two nearly four + times as long as segment one. R. PROLIXUS. W. I. + RHODNIUS Stäl + + gg. Antennæ inserted remote from the vertex of the head. + + h. Body slightly hairy; pronotum distinctly constricted; + angles distinct; anterior lobe four-tuberculate, with + the middle tubercles large and conical. _M. + phyllosoma_, large species from the southwest. + _Meccus_ Stäl + + hh. Body smooth, margin of the pronotum sinuous, scarcely + constricted; anterior lobe lined with little + tubercles. CONORHINUS Lap. + + i. Surface of the pronotum and prosternum more or less + granular. + + j. Eyes small, head black; body very narrow, a fifth + as wide as long; beak reaches the middle of the + prosternum. California. C. PROTRACTUS + + jj. Eyes large, head fuscous; body at least a fourth + as wide as long. Southern species. _C. + rubrofasciatus_ + + ii. Pronotum and prosternum destitute of granules. + + j. Border of abdomen entirely black except for a + narrow yellowish spot at the apex of one segment. + Texas. _C. gerstaeckeri_ + + jj. Border of abdomen otherwise marked. + + k. Beak slender, joints one and two slightly pilose, + two more than twice as long as one; tubercles at + the apical angles of the pronotum slightly + acute, conical. Md. to Ill. and south. The + masked bed bug hunter (fig. 71). C. SANGUISUGUS + + kk. Beak entirely pilose, joint two a third longer + than joint one; joint one much longer than + three; tubercles at the apical angles of + pronotum slightly elevated, obtuse. Ga., Ill., + Tex., Cal. _C. variegatus_ + + ff. Antenna inserted on top of the head between margins, close + to the eyes; antenniferous tubercles not projecting from + the side of the head. + + g. Anterior lobe of the pronotum with a bispinous or + bituberculate disc; femora unarmed. _S. arizonica_, _S. + bicolor_. Southwestern species. _Spiniger_ Burm. + + gg. Disc of pronotum unarmed; apex of scutellum produced + into a spine; ocelli close to the eyes; eyes large and + close together. REDUVIUS Lamarck + + h. Color piceous. Widely distributed in the United States. + (Fig. 20). R. PERSONATUS + + hh. More or less testaceous in color. Southwestern states. + R. SENILIS + + ee. Apex of scutellum broad, with two or three spines. + ECTRICHODIINÆ + + f. First segment of the antenna about as long as the head. _E. + cruciata_ Pa. and south; _E. cinctiventris_, Tex. and Mex. + _Ectrichodia_ L. et S. + + ff. First segment of the antennæ short. _P. æneo-nitens_. + South. _Pothea_ A. et S. + + cc. Ocelli cephalad of the hind margins of the eyes; first segment + of the antennæ stout, second segment divided into many smaller + segments. South and west. _Homalocoris maculicollis_, and + _Hammatocerus purcis_. HAMMATOCERINÆ + + bb. Hemelytra with a quadrangular or discoidal areole in the corium + near the apex of the clavus (fig. 159e). + + c. Anal areole of the membrane not extending as far proximad as the + costal areole; basal segment of the antenna thickened, porrect; + the other segments slender, folding back beneath the head and + the first segment STENOPODINÆ + + d. Head armed with a ramous or furcate spine below each side, + caudad of the eyes. + + e. First segment of the antenna thickened, apex produced in a + spine beyond the insertion of the second segment. Species + from Va., Ill. and south. _Pnirontis_ Stäl. + + ee. First segment of the antenna not produced beyond the + insertion of the second segment. _Pygolampis_, N. E. states + and south; _Gnathobleda_, S. W. and Mex. + + dd. Head unarmed below or armed with a simple spine; rarely with a + subfurcate spine at the side of the base. Carolina, Missouri + and south. _Stenopoda_, _Schumannia_, _Diaditus_, _Narvesus_, + _Oncocephalus_ + + cc. Anal areole of membrane extending farther proximad than the + costal areole. + + d. Ocelli farther apart than the eyes. _A. crassipes_, widely + distributed in the United States; other species occur in the + southwest. _Apiomerus_ Hahn. + + dd. Ocelli not so far apart as the eyes. ZELINÆ + + e. Sides of mesosternum without a tubercle or fold in front. + + f. Fore femur as long as or longer than the hind femur; first + segment of the beak much shorter than the second. _Z. + audax_, in the north eastern states; other species south + and west. _Zelus_ Fabr. + + ff. Fore femur shorter than the hind femur, rarely of equal + length, in this case the first segment of the beak as long + or longer than the second. + + g. First segment of the beak shorter than the second; fore + femur a little shorter than the hind femur; the first + segment of the beak distinctly longer than the head + before the eyes. _P. cinctus_ a widely distributed + species (fig. 160). _P. punctipes_, _P. spinicollis_, + Cal., Mex. (= _Milyas_) _Pselliopus_ Berg. + + gg. First segment of the beak as long or longer than the + second. + + h. Pronotum armed with spines on the disc. + + i. Juga distinctly prominent at the apex and often acute + or subacute; fore femur distinctly thickened; + hemelytra usually not reaching the apex of the + abdomen. _Fitchia aptera_, N. Y., south and west; + _F. spinosula_, South; _Rocconata annulicornis_, + Texas, etc. + + ii. Juga when prominent, obtuse at apex; eyes full width + of the head; fore femur not thickened; pronotum with + four spines on posterior lobe. _R. taurus_, Pa., + south and west. _Repipta_ Stäl. + + hh. Pronotum unarmed on the disc. + + i. Spines on each apical angle of the penultimate + abdominal segment. _A. cinereus_, Pa., and south. + _Atrachelus_ A. et S. + + ii. Apical angle of the penultimate abdominal segment + unarmed. _Fitchia_ (in part); _Castolus ferox_, + Arizona. + + ee. Sides of the mesosternum with a tubercle or fold in front at + the hind angles of the prosternum; first segment of the beak + longer than the part of the head cephalad of the eyes. + + f. Fore femur thickened, densely granulated; hind femur + unarmed. + +[Illustration: 161. Taxonomic details of Diptera. (_a_) Ventral aspect +of abdomen of Cynomyia; (_b_) antenna of Tabanus; (_c_) ventral aspect +of abdomen of Chortophila; (_d_) ventral aspect of abdomen of Stomoxys; +(_e_) claw of Aedes (Culex) sylvestris, male; (_f_) claw of Hippoboscid; +(_g_) foot of dipterous insect showing empodium developed pulvilliform; +(_h_) hind tarsal segment of Simulium vittatum, female; (_i_) foot of +dipterous insect showing bristle-like empodium.] + + g. Fore tibiæ each with three long spines on the ventral + side. _S. diadema_ (fig. 159e), a widely distributed + species; and several southwestern species. _Sinea_ A. et + S. + + gg. Fore tibiæ unarmed. _A. multispinosa_, widely + distributed; _A. tabida_, Cal. _Acholla_ Stäl. + + ff. Fore femur unarmed, rarely a little thickened, a little + granulated. + + g. Pronotum produced caudad over the scutellum, with a high + mesal tuberculate ridge (fig. 19e). A. CRISTATUS. N. Y. + to Cal. and south. ARILUS Hahn. + + gg. Caudal lobe of the pronotum six sided, neither elevated + nor produced caudad. _H. americanus_, Southwest; also + several W. I. and Mexican genera. HARPACTOR Lap. + + +DIPTERA (Mosquitoes, Midges, Flies) + +a. Integument leathery, abdominal segments indistinct; wings often + wanting; parasitic forms. PUPIPARA + + b. Head folding back on the dorsum of the thorax; wingless flies + parasitic on bats. Genus _Nycteribia_. NYCTERIBIIDÆ + + bb. Head not folding back upon the dorsum of the thorax; flies either + winged or wingless; parasitic on birds and on bats and other + mammals. + + c. Antennæ reduced, wings when present, with distinct parallel veins + and outer crossveins; claws simple; palpi leaf-like, projecting + in front of the head. Flies chiefly found on bats. Several + genera occur in North America. STREBLIDÆ + +[Illustration: 162. Hippobosca equina, ×4. After Osborn.] + + cc. Antennæ more elongate, segments more or less distinctly + separated; head sunk into an emargination of the thorax; wings + when present with the veins crowded toward the anterior margin; + palpi not leaf-like. HIPPOBOSCIDÆ + + d. Wings absent or reduced and not adapted for flight. + + e. Wings and halteres (balancers) absent. _M. ovinus_, the sheep + tick. _Melophagus_ Latr. + + ee. Wing reduced (or cast off), halteres present. + + f. Claw bidentate; ocelli present. On deer after the wings are + cast off. _L. depressa_. _Lipoptena_ Nitsch + + ff. Claw tridentate (fig. 161 f). On _Macropis_. _B. + femorata._ _Brachypteromyia_ Will. + + dd. Wings present and adapted for flight. + + e. Claws bidentate. + + f. Ocelli present; head flat; wings frequently cast off. On + birds before casting of the wing. _Lipoptena_ Nitsch. + + ff. Ocelli absent; head round; wings present. The horse tick + H. EQUINA may attack man (fig. 162). HIPPOBOSCA L. + + ee. Claws tridentate (fig. 161 f.). + + f. Anal cell closed at apical margin by the anal crossvein. + + g. Ocelli absent. _Stilbometopa_ Coq. + + gg. Ocelli present. + + h. R_{4+5} does not form an angle at the crossvein. On + birds. There is a record of one species of this genus + attacking man. ORNITHOMYIA Latr. + + hh. R_{4+5} makes an angle at the crossvein. _O. + confluens_. _Ornithoica_ Rdi. + + ff. Anal cell not closed by an anal crossvein. _Lynchia_, + _Pseudolfersia_, and _Olfersia_ are chiefly bird + parasites. The first mentioned genus is said to be the + intermediate host of _Hæmoproteus columbæ_. + +aa. Abdominal segments chitinous; not parasitic in the adult stage. + + b. Antennæ with six or more segments and empodium not developed + pulvilliform; palpi often with four segments. + + c. Ocelli present. BLEPHAROCERIDÆ, RHYPHIDÆ, BIBIONIDÆ, + MYCETOPHILIDÆ, besides some isolated genera of other families. + + cc. Ocelli absent. + + d. Dorsum of the thorax with a V-shaped suture; wings usually with + numerous veins; legs often very long and slender. Crane flies. + TIPULIDÆ + + dd. Dorsum of the thorax without a V-shaped suture. + + e. Not more than four longitudinal veins ending in the wing + margin; wing usually hairy: antennæ slender; coxæ not long; + tibiæ: without spurs, legs long and slender. Small, delicate + flies often called Gall gnats. CECIDOMYIIDÆ. + + ee. More than four longitudinal veins ending in the wing margin. + + f. The costal vein is not produced beyond the tip of the wing; + radius with not more than three branches. + + g. Antennæ short, composed of ten or eleven closely united + segments; legs stout; body stout; abdomen oval; anterior + veins stout, posterior ones weak (fig. 163 b); eyes of + the male contiguous over the antennæ. Black flies, + buffalo flies, turkey gnats. Many North American + species, several of them notorious for their blood + sucking propensities. SIMULIIDÆ + + h. Second joint of the hind tarsus with basal scale-like + process and dorsal excision (fig. 161 h); radial + sector not forked; no small cell at the base of the + wing. _S. forbesi_, _jenningsi_, _johannseni_, + _meridionale_, _piscicidium_, VENUSTUM, VITTATUM, etc. + Widely distributed species (= EUSIMULIUM) SIMULIUM + Latr. + + hh. No basal scale-like process on the second joint of the + hind tarsus; radial sector usually forked (fig. 163 + b). + + i. Face broad, small basal cell of the wing present. _P. + fulvum_, HIRTIPES, _mutatum_, PECUARUM, _pleurale_. + PROSIMULIUM Roub. + + ii. Face linear; small basal cell of the wing absent. + One species, _P. furcatum_, from California. + _Parasimulium_ Malloch + + gg. Flies of a different structure. + + h. Antennæ composed of apparently two segments and a + terminal arista formed of a number of closely united + segments. Rare flies with aquatic larvæ. + ORPHNEPHILIDÆ + + hh. Antennæ of six to fifteen segments, those of the male + usually plumose; legs frequently slender and wings + narrow CHIRONOMIDÆ + + i. Media forked (except in the European genus + _Brachypogon_); thorax without longitudinal fissure + and not produced over the head (except in four + exotic genera); antennæ usually fourteen-jointed in + both sexes; fore tibia with a simple comb of setulæ, + hind tibia with two unequal combs, middle tibia + without comb. CERATOPOGONINÆ + + j. Thorax produced cap-like over the head, wing narrow + and very long. _Jenkinsia_, _Macroptilum_ and + _Calyptopogon_, eastern hemisphere; + _Paryphoconus_, Brazil. + + jj. Thorax not produced over the head. + + k. Eyes pubescent, empodium well developed, or if + short then R_{2+3} distinct and crossvein-like + or the branches of R coalescent; r-m crossvein + present; fore femora not thickened; wing either + with appressed hairs or with microscopic erect + setulæ _Dasyhelea_ Kieff. + + kk. Eyes bare, or otherwise differing from the + foregoing. + + l. Empodium well developed, nearly as long as the + claws and with long hairs at the base; femora + and fifth tarsal segments unarmed, i.e. + without spines or stout setæ; fourth tarsal + segment cylindrical. + + m. Wing with erect and microscopic setulæ. + Widely distributed. (= Atrichopogon) + _Ceratopogon_ Meig. + + mm. Wing with long and depressed hairs. Widely + distributed. _Forcipomyia_ + + n. Hind metatarsus shorter or not longer than + the following (i.e. the second tarsal) + segment. Subgenus _Prohelea_ Kieff + + nn. Hind metatarsus longer than the following + segment. Subgenus _Forcipomyia_ Meig. + + ll. Empodium short, scarcely reaching the middle + of the claws, or vestigial. + + m. R-m crossvein wanting. + + n. Palpi four segmented; inferior fork of the + media obliterated at the base. Australia. + _Leptoconops_ Skuse + + nn. Palpi three-segmented. + + o. Legs spinulose, tarsal claws of the + female with a basal tooth or strong + bristle, those of the male unequal, the + anterior with a long sinuous tooth, the + posterior with a short arcuate tooth. + Italy. MYCTEROTYPUS Noé + + oo. Legs unarmed; no crossvein between the + branches of the radius (fig. 163e). New + Mexico. TERSESTHES Townsend + + mm. R-m crossvein present. + + n. Fore femora very much swollen, armed with + spines below, fore tibia arcuate and + applied closely to the inferior margin of + the femur. + + o. R_{2+3} present, therefore cell R_1 and + R_2 both present; wing usually fasciate. + United States _Heteromyia_ Say. + + oo. R_{2+3} not distinct from R_{4+5}, hence + cell R_3 obliterated. South America + _Pachyleptus_ Arrib. (Walker) + + nn. Fore femur not distinctly swollen. + + o. R_{2+3} present therefore cells R_1 and + R_3 both present, or if not, then the + branches of the radius more or less + coalescent, obliterating the cells. + + p. At least the tip of the wing with erect + setulæ; tip of R_{4+5} scarcely + attaining the middle of the wing, + empodium rather indistinct, not + reaching the middle of the claws, the + claws not toothed, equal, with long + basal bristle; legs without stout + setæ. Widely distributed. CULICOIDES + Latr. + + HÆMATOMYIDIUM and OECACTA are probable + synonyms of this. + + pp. Wings bare, if rarely with hair, then + the radius reaches beyond two-thirds + the length of the wing, or the femur + or fifth tarsal segment with stout + black spines. + + q. Media unbranched. Europe. + _Brachypogon_ Kieff + + qq. Media branched. + + r. Hind femur much swollen and spined. + America and Europe. _Serromyia_ + Meg. + + rr. Hind femur not distinctly swollen. + + s. Cell R_1 not longer than high; + fork of the media distad of the + crossvein; wing with microscopic + setulæ _Stilobezzia_ Kieff + + ss. Cell R_1 elongate. + + t. Femora unarmed. Widely + distributed. (= Sphaeromias + Kieff. 1913 not Curtis?). + JOHANNSENIELLA Will. + + tt. Femora, at least in part, with + strong black spines. Widely + distributed. _Palpomyia_ + Megerle + + oo. R_{2+3} coalescent with R_{4+5} hence + cell R_3 is obliterated. + + p. In the female the lower branch of the + media with an elbow near its base + projecting proximad, the petiole of + the media coalescent with the basal + section of the radius, wing long and + narrow, radial sector ending near the + tip of the wing; venation of the male + as in _Bezzia_; front concave. United + States. _Stenoxenus_ Coq. + + pp. Venation otherwise, front not concave. + + q. Subcosta and R_1 more or less + coalescent with the costa; wing + pointed at the apex, much longer + than the body; antennæ fourteen + segmented, not plumose. India. + _Haasiella_ Kieff. + + qq. Subcosta and radius distinct from + the costa. + + r. Abdomen petiolate. _Dibezzia_ + Kieff. + + rr. Abdomen not petiolate. + + s. Head semi-globose; hind tarsi + unusually elongate in the + female; antennæ of the male not + plumose. Europe. _Macropeza_ + Meigen. + + ss. Head not globose, more or less + flattened in front; antennæ of + the male plumose. Widely + distributed. _Bezzia_ Kieff. + + t. Fore femora, at least, armed + with stout spines below. + Subgenus _Bezzia_ Kieff. + + tt. Femora unarmed. Subgenus + _Probezzia_ Kieff. + + ii. Media of the wing simple, and otherwise not as in + "i". To this group belong numerous Chironomid + genera, none of which are known to be noxious to + man. + + ff. The costal vein apparently is continued around the hind + margin of the wing; radius with at least four branches. + + g. Wing ovate pointed, with numerous veins; crossveins, if + evident, before the basal third of the wing; veins very + hairy; very small moth-like flies. PSYCHODIDÆ + + h. With elongate biting proboscis; the petiole of the + anterior forked cell of the wing (R_2) arises at or + beyond the middle of the wing (fig. 163d). PHLEBOTOMUS + Rdi. + +[Illustration: 163. Wings of Diptera. (_a_) Anopheles; (_b_) +Prosimulium; (_c_) Johannseniella; (_d_) Phlebotomus (After Doerr and +Russ); (_e_) Tersesthes (after Townsend); (_f_) Tabanus; (_g_) +Symphoromyia; (_h_) Aphiochæta; (_i_) Eristalis; (_j_) Gastrophilus; +(_k_) Fannia; (_l_) Musca.] + + hh. With shorter proboscis; the petiole of the anterior + forked cell arises near the base of the wing. + _Psychoda_, _Pericoma_, etc. + + gg. The r-m crossvein placed at or beyond the center of the + wing; wings not folded roof-like over the abdomen. + + h. Proboscis short, not adapted for piercing; wings bare + (DIXIDÆ); or wings scaled (CULICIDÆ, Subf. + CORETHRINÆ). + + hh. Proboscis elongate, adapted for piercing; wings + scaled, fringed on the hind margin; antennæ of the + male bushy plumose. Mosquitoes. CULICIDÆ (exclusive of + CORETHRINÆ) + + i. Metanotum without setæ. + + j. Proboscis strongly decurved; body with broad, + appressed, metalescent scales; cell R_2 less than + half as long as its petiole; claws of female + simple, some of the claws of the male toothed. + Several large southern species believed to feed + only on nectar of flowers. _Megarhinus_ R. D. + + jj. Proboscis straight or nearly so, or otherwise + different. + + k. Scutellum evenly rounded, not lobed; claws simple + in both sexes. ANOPHELES Meig. + + l. Abdomen with clusters of broad outstanding + scales along the sides; outstanding scales on + the veins of the wing rather narrow, + lanceolate; upper side of the thorax and + scutellum bearing many appressed lanceolate + scales. Florida and southward (CELLIA). + + m. Hind feet from the middle of the second + segment largely or wholly snow white. + + n. With a black band at the base of the last + segment of each hind foot. A. + ALBIMANUS*[H] and TARSIMACULATA* + + nn. Without such a band A. ARGYRITARSIS* + + mm. Hind feet black, mottled with whitish and + with bands of the same color at the sutures + of the segments. W. I. A. MACULIPES + + ll. Abdomen without such a cluster of scales; + outstanding scales of the wing veins rather + narrow, lanceolate; tarsi wholly black. + + m. Deep black, thorax obscurely lined with + violaceous, especially posteriorly; head, + abdomen and legs black; no markings on the + pleura; abdomen without trace of lighter + bindings; wing scales outstanding, uniform, + not forming spots, though little thicker at + the usual points indicating the spottings. + Florida. A. ATROPUS + + mm. Otherwise marked when the wings are + unspotted. + + n. Wings unspotted. + + o. Petiole of the first forked cell (R_2) + more than a third the length of the + cell. Mississippi valley. A. WALKERI + + oo. Petiole of the first forked cell a third + the length of the cell. Md. A. BARBERI + + nn. Wings spotted. + + o. Front margin of the wings with a patch of + whitish and yellow scales at a point + about two-thirds or three-fourths of the + way from base to apex of wing. + + p. Veins of the wing with many broad + obovate outstanding scales; thorax + with a black dot near the middle of + each side. W. I. A. GRABHAMI* + + pp. The outstanding scales of the wings + rather narrow, lanceolate. + + q. Scales of the last vein of the wings + white, those at each end black; + R_{4+5} black scaled, the extreme + apex white scaled. Widely + distributed north and south (fig. + 131). A. PUNCTIPENNIS + + A dark variety from Pennsylvania has + been named A. PERPLEXENS. + + qq. Scales of the last vein of the wing + white, those at its apex black; + R_{4+5} white scaled and with two + patches of black scales. South and + the tropics. A. FRANCISCANUS and + PSEUDOPUNCTIPENNIS* + + oo. Front margin of the wings wholly black + scaled. + + p. Last (anal) vein of the wings white + scaled with three patches of black + scales (fig. 132). New Jersey to + Texas. A. CRUCIANS* + + pp. Last vein of the wings wholly black + scaled. + + q. Widely distributed north and south + (fig. 130), (= MACULIPENNIS). A. + QUADRIMACULATUS* + + qq. Distributed from Rocky Mountains + westward. A. OCCIDENTALIS + + kk. Scutellum distinctly trilobed. + + l. Cell R_2 less than half as long as its petiole; + thorax with metallic blue scales; median lobe + of the scutellum not tuberculate; few small + species which are not common. URANOTÆNIA + Arrib. + + ll. Cell R_2 nearly or quite as long as its + petiole, or otherwise distinct. + + m. Femora with erect outstanding scales; occiput + broad and exposed. Large species. P. + CILIATA. P. HOWARDI PSOROPHORA R. D. + + mm. Femora without erect scales. + + n. Clypeus bearing several scales or hairs, + scutellum with broad scales only; back of + head with broad scales; scales along the + sides of the mesonotum narrow; some or the + claws toothed; thorax marked with a pair + of silvery scaled curved stripes; legs + black with white bands at the bases of + some of the segments (fig. 134). Yellow + Fever mosquito. AEDES (= STEGOMYIA) + CALOPUS. + + nn. With another combination of characters. + Numerous species of mosquitoes belonging + to several closely related genera, widely + distributed over the country. (_Culex_, + _Aedes_, _Ochlerotatus_, etc.). CULEX in + the wide sense. + + ii. Metanotum with setæ. _Wyeomyia_ (found in the United + States); and related tropic genera. + + bb. Antennæ composed of three segments with a differentiated style or + bristle; third segment sometimes complex or annulate, in which + case the empodium is usually developed like the pulvilli, i.e., + pad-like (fig. 161 g). + + c. Empodium developed pad-like (pulvilliform) i.e., three nearly + equal membranous appendages on the underside of the claw (fig. + 161g). + + d. Squamæ, head, and eyes large; occiput flattened or concave; + third segment of the antennæ with four to eight annuli or + segments, proboscis adapted for piercing; body with fine + hairs, never with bristles; middle tibia with two spurs; wing + venation as figured (fig. 163f); marginal vein encompasses the + entire wing. Horse flies, greenheads, deer flies, gad flies + TABANIDÆ[I] + + e. Hind tibia with spurs at tip; ocelli usually present + (PANGONINÆ) + + f. Third joint of the antennæ with seven or eight segments; + proboscis usually prolonged. + + g. Each section the third antennal segment branched. Central + American species, _P. festæ_. _Pityocera_ G. T. + + gg. Sections of the third antennal segment not branched. + + h. Upper corner of the eyes in the female terminating in + an acute angle; wings of both sexes dark anteriorly. + _G. chrysocoma_, a species from the eastern states. + _Goniops_ Ald. + + hh. Upper corner of the eye in the female not so + terminating; wings nearly uniform in color, or + hyaline. + + i. Proboscis scarcely extending beyond the palpi; front + of the female wide; much wider below than above. S. + W. States. _Apatolestes_ Will. + + ii. Proboscis extending beyond the palpi. + + j. Wing with cell M_3 closed. Tropic America. (= + _Diclisa_) _Scione_ Wlk. + + jj. Cell M_3 open; ocelli present or absent. Two or + three eastern species; many south and west + PANGONIA Rdi. + + ff. Third segment of the antenna with five divisions; ocelli + present. + + g. First and second segments of the antenna short, the + second only half as long as the first, three western + species. SILVIUS Rdi. + + gg. First and second segments of the antenna long, the + second distinctly over half as long as the first. Deer + flies. Many species, widely distributed. CHRYSOPS Meig. + + ee. Hind tibia without spurs; ocelli absent. + + f. Third segment of antenna with four divisions, no tooth or + angulation; wings marked with rings and circles of darker + coloring; front of the female very wide. Widely + distributed. _H. americana_, _H. punctulata_. HÆMATOPOTA + Meig. + + ff. Third segment of the antenna with five divisions (fig. + 161b). + + g. Third segment of the antenna not furnished with a tooth + or distinct angular projection. + + h. Body covered with metallic scales; front of female of + normal width; front and middle tibiæ greatly dilated. + _L. lepidota_. _Lepidoselaga_ Macq. + + hh. Body without metallic scales; antennæ not very long, + the third segment not cylindrical, not situated on a + projecting tubercle; front of the female narrow. + South. _D. ferrugatus._ (= _Diabasis_) _Diachlorus_ O. + S. + + gg. Third segment of the antenna furnished with a tooth or a + distinct angular projection. + + h. Hind tibiæ ciliate with long hairs. S. W. and tropics. + _Snowiella_ and _Stibasoma_. + + hh. Hind tibiæ not ciliate. + + i. Species of slender build, usually with a banded + thorax and abdomen; third segment of the antenna + slender, the basal prominence long; wings mostly + with brownish markings. Tropic America. + _Dichelacera_ Macq. + + ii. Species of a stouter build; third segment of the + antenna stout, its basal process short (fig. 161b). + Many species, widely distributed TABANUS L. + + dd. With another group of characters. + + e. Squamæ small, antennæ variable, thinly pilose or nearly bare + species, without distinct bristles; wing veins not crowded + anteriorly, R_4 and R_5 both present, basal cells large; + middle tibiæ at least with spurs. LEPTIDÆ + + f. Flagellum of the antenna more or less elongated, composed + of numerous more or less distinct divisions. XYLOPHAGINÆ + and ARTHROCERATINÆ. + + ff. Antennæ short, third segment simple, with arista or style; + face small, proboscis short LEPTINÆ + + g. Front tibiæ each with one or two spurs, or if absent, + then no discal cell. _Triptotricha_, _Pheneus_, + _Dialysis_, _Hilarimorpha_. + + gg. Front tibiæ without terminal spurs, discal cell present. + + h. Hind tibiæ each with a single spur. + + i. Anal cell open (fig. 163g); third antennal segment + kidney-shaped with dorsal or subdorsal arista; first + antennal segment elongate and thickened. About a + dozen species have been described from the United + States, of which at least one (S. PACHYCERAS) is + known to be a vicious blood sucker. SYMPHOROMYIA + Frauenf. + + ii. Anal cell closed; third antennal segment not + kidney-shaped. _Chrysopila_, _Ptiolina_, _Spania_. + + hh. Hind tibiæ each with two spurs. + + i. Third segment kidney-shaped, the arista subdorsal; + anal cell closed. _Atherix_ Meig. + + ii. Third segment of the antenna short and with terminal + arista; anal cell open _Leptis_ Fabr. Two European + species of this genus have been accused of blood + sucking habits, but the record seems to have been + based upon error in observation. + + ee. With another combination of characters. STRATIOMYIIDÆ, + CYRTIDÆ, etc. + + cc. Empodium bristlelike or absent. + + d. Antennæ apparently two segmented, with three-segmented arista, + wings (rarely wanting) with several stout veins anteriorly, + the weaker ones running obliquely across the wing (fig. 163h); + small, quick running, bristly, humpbacked flies. Several + genera; APHIOCHÆTA, PHORA, TRINEURA, etc. PHORIDÆ + + dd. Flies with other characters. + + e. No frontal lunule above the base of the antennæ; both R_4 and + R_5 often present; third segment of the antenna often with a + terminal bristle. ASILIDÆ, MYDAIDÆ, APIOCERIDÆ, THEREVIDÆ, + SCENOPINIDÆ, BOMBYLIIDÆ, EMPIDIDÆ, DOLICHOPODIDÆ, + LONCHOPTERIDÆ. + + ee. A frontal lunule above the base of the antennæ; third + segment of the antenna always simple, i.e., not ringed, + usually with a dorsal arista; R_4 and R_5 coalesced into a + simple vein. + + f. A spurious vein or fold between the radius and the media, + rarely absent; the cell R_{4+5} closed at the apex by vein + M_1; few or no bristles on the body, none on the head; + flies frequently with yellow markings. ERISTALIS (fig. + 163i), HELOPHILUS, and many other genera. SYRPHIDÆ + + ff. No spurious vein present. + + g. Body without bristles; proboscis elongate and slender, + often folding; front of both male and female broad. + CONOPIDÆ + + gg. Bristles almost always present on head, thorax, abdomen + and legs. + + h. Arista terminal; hind metatarsus enlarged, sometimes + ornamented, hind tarsus more or less flattened beneath + PLATYPEZIDÆ + + hh. Flies having a different combination of characters. + + i. Head large, eyes occupying nearly the entire head; + cell R_{4+5} narrowed in the margin; small flies. + PIPUNCULIDÆ + + ii. Head and eyes not unusually large. + + j. Squamæ (tegulæ, or calyptræ, or alulæ) not large, + often quite small, the lower one lacking, or at + most barely projecting from below the upper one + (antisquama); front of both male and female broad, + the eyes therefore widely separated; posthumeral + and intraalar macrochæta not simultaneously + present; thorax usually without a complete + transverse suture; postalar callus usually absent; + the connectiva adjoining the ventral sclerites + always visible; hypopleural macrochætæ absent; + last section of R_{4+5} and M_{1+2} with but few + exceptions nearly parallel; subcostal vein often + wanting or vestigial or closely approximated to + R_1; the latter often short, basal cells small, + the posterior ones often indistinct or wanting; + vibrissæ present or absent. ACALYPTRATE MUSCOIDEA + + k. Subcosta present, distinctly separated from R_1 + at the tip; R_1 usually ends distad of the + middle of the wing; the small basal cells of the + wing distinct. + + l. A bristle (vibrissa) on each side of the face + near the margin of the mouth. CORDYLURIDÆ, + SEPSIDÆ, PHYCODROMIDÆ, HETERONEURIDÆ, + HELOMYZIDÆ. + + ll. No vibrissæ present. + + m. Head nearly spherical, cheeks broad and + retreating; proboscis short; the cell R_5 + closed or narrowed in the margin; legs very + long; tarsi shorter than the tibiæ. CALOBATA + and other genera. MICROPEZIDÆ + + mm. Flies with another combination of + characters. RHOPALOMERIDÆ, TRYPETIDÆ, + ORTALIDÆ, SCIOMYZIDÆ. + + kk. Subcosta absent or vestigial, or if present, + then apparently ending in the costa at the point + where R_1 joins it; R_1 usually ends in the + costa at or before the middle of the wing. + + l. Arista long plumose, or pectinate above; oral + vibrissæ present; anal cell complete; costa + broken at the apex of R_1. DROSOPHILA, + PHORTICA, and other genera. DROSOPHILIDÆ + + ll. With another combination of characters. + + m. The cell M and first M_2 not separated by a + crossvein; anal cell absent; front bare or + only bristly above; usually light colored + flies. HIPPELATES, OSCINUS, and other + genera. (See also m m m.) OSCINIDÆ + + mm. Cell M and cell first M_2 often separated by + a crossvein; anal cell present, complete, + though frequently small; scutellum without + spines or protuberances; oral vibrissæ + present; arista bare or short plumose; front + bristly at vertex only; small dark flies. + PIOPHILA (fig. 99), SEPSIS and other genera. + SEPSIDÆ + + mmm. The GEOMYZIDÆ, AGROMYZIDÆ, PSILIDÆ, + TRYPETIDÆ, RHOPALOMERIDÆ, BORBORIDÆ and + DIOPSIDÆ differ in various particulars from + either the OSCINIDÆ and the SEPSIDÆ noted + above. + + jj. Squamæ well developed, usually large, the lower + one frequently projecting from below the upper + one; both posthumeral and intraalar macrochætæ + present; thorax with a complete transverse suture; + postalar callus present and separated by a + distinct suture from the dorsum of the thorax; + front of the female broad, of the male frequently + narrow, the eyes then nearly or quite contiguous; + the connectiva adjoining the ventral sclerites + either visible or not; hypopleural macrochætæ + present or absent; subcosta always distinct in its + whole course, R_1 never short. CALYPTRATE + MUSCOIDEA[J] + + k. Oral opening small, mouth parts usually much + reduced or vestigial. This family is undoubtedly + of polyphyletic origin but for convenience it is + here considered as a single family. OESTRIDÆ. + + l. The costal vein ends at the tip of R_{4+5}, + M_{1+2} straight, not reaching the wing + margin, hence cell R_5 wide open (fig. 163j); + squamæ small; arista bare; ovipositor of the + female elongate. Larvæ in the alimentary canal + of horses, etc. GASTROPHILUS + + m. Posterior crossvein (m-cu) wanting; wings + smoky or with clouds. Europe. G. PECORUM + + mm. Posterior crossvein (m-cu) present, at least + in part. + + n. Wing hyaline with smoky median cross band, + and two or three spots; posterior + trochanters with hook in the male and a + prominence in the female. World wide + distribution. G. EQUI. + + nn. Wings without spots. + + o. Posterior crossvein (m-cu) distad of the + anterior crossvein (r-m); legs, + particularly the femora, blackish brown. + Europe and North America. G. + HÆMORRHOIDALIS + + oo. Posterior crossvein opposite or proximad + of the anterior crossvein. Europe and + North America. G. NASALIS + + ll. The costal vein ends at the tip of M_{1+2}, + M_{1+2} with a bend, the cell R_5 hence much + narrowed in the margin, or closed. + + m. Proboscis geniculate, inserted in a deep + slit; female without extricate ovipositor; + arista either bare or plumose; squamæ large; + facial grooves approximated below. + + n. Arista bare, short. Larvæ in rodents. + Tropic America. _B. princeps_. _Bogeria_ + Austen + + nn. Arista pectinate above. + + o. Tarsi broadened and flattened, hairy, + anal lobe of the wing large. Larvæ in + rodents. A number of American species. + _Cuterebra_. + + oo. Tarsi slender, not hairy; anal lobe of + the wing moderate. Larvæ in man and + other mammals. Tropic America. D. + CYANIVENTRIS. DERMATOBIA Br. + + mm. Mouth parts very small, vestigial; arista + bare. + + n. Facial grooves approximated below, leaving + a narrow median depression or groove. + + o. Cell R_5 closed and petiolate, body + nearly bare. Larvæ in the nasal cavities + of the smaller Ungulates. The sheep bot + fly. O. OVIS. Widely distributed OESTRUS + L. + + oo. Cell R_5 narrowly open, body hairy. + Larvæ parasitic on deer. Europe and + America. _Cephenomyia_ Latr. + + nn. Facial grooves far apart, enclosing + between them a broad shield-shaped + surface; squamæ large; female with + elongate ovipositor. Larvæ hypodermatic on + Ungulates HYPODERMA Clark + + o. Palpi wanting; tibiæ thickened in the + middle. + + p. Hair at apex of the abdomen yellow; + legs including femora yellowish brown. + H. DIANA + + pp. Hair at the apex of the abdomen + reddish yellow. Europe and America. + + q. Tibiæ and tarsi yellow; femora black. + H. LINEATA + + qq. Legs black with black hair; tips of + hind tibia and tarsi yellowish + brown. H. BOVIS + + oo. Palpi small, globular; tibiæ + cylindrical, straight. On reindeer. _O. + tarandi_ _Oedemagena_ Latr. + + kk. Oral opening of the usual size; mouth parts not + vestigial. + + l. Hypopleurals wanting; if three sternopleurals + are present the arrangement is 1:2; + conjunctiva (fig. 161c) of the venter usually + present; if the terminal section of M_{1+2} is + bent it has neither fold nor appendage + (ANTHOMYIIDÆ of Girschner). + + m. Sternopleurals wanting; M_{1+2} straight + toward the apex, costa ends at or slightly + beyond the tip of R_{4+5}; mouth parts + vestigial. GASTROPHILINÆ. See OESTRIDÆ + + mm. Sternopleurals present, if rarely absent + then differing in other characters. + + n. Caudal margin of the fifth ventral + abdominal sclerite of the male deeply + notched on the median line usually to + beyond the middle; abdomen often + cylindrical or linear; abdomen often with + four to eight spots; eyes of the male + usually widely separated; sternopleurals + three, arranged in an equilateral + triangle; subapical seta of the hind tibia + placed very low; M_{1+2} straight; anal + vein abbreviated; wings not rilled. + _Cænosia_, _Caricea_, _Dexiopsis_, + _Hoplogaster_, _SchÅ“nomyia_, etc. + (CÅ’NOSINÆ)[K]. ANTHOMYIIDÆ in part + + nn. Caudal margin of the fifth ventral + abdominal sclerite of the male incurved, + rarely deeply cleft, rarely entire, in a + few genera deeply two or three notched; + M_{1+2} straight or curved; abdomen + usually short or elongate oval; + sternopleurals, if three are present, + arranged in the order 1:2 in a right + triangle. (MUSCINÆ-ANTHOMYIINÆ of + Girschner) + + o. M_{1+2} straight, hence cell R_5 not + narrowed in the margin. ANTHOMYIIDÆ in + part + + p. Underside of the scutellum more or less + sparsely covered with fine hairs; anal + vein nearly always reaches the hind + margin of the wing; extensor surface + of the hind tibiæ with a number of + stout setæ; squamæ often small and + equal. ANTHOMYIA, _Chortophila_, + _Eustalomyia_, _Hammomyia_, + _Hylemyia_, _Prosalpia_, _Pegomyia_, + etc. HYLEMYINÆ-PEGOMYINÆ + + pp. Underside of the scutellum bare; anal + vein does not reach the wing margin. + + q. First anal vein short, second anal + suddenly flexed upwards; hind tibiæ + each with one or two strong setæ on + the extensor surface. FANNIA (= + HOMALOMYIA), _CÅ“lomyia_, + _Choristoma_, _Euryomma_, _Azelia_, + etc. FANNINÆ-AZELINÆ + + qq. Anal veins parallel or divergent. + + r. Setæ on the exterior surface of the + hind tibiæ wanting (except in + _Limnaricia_ and _CÅ“nosites_), + lower squama not broadened to the + margin of the scutellum. + _Leucomelina_, _Limnophora_, + _Limnospila_, _Lispa_, _Mydæa_, + _Spilogaster_, etc. + MYDÆINÆ-LIMNOPHORINÆ + + rr. One (rarely more) seta on the + extensor surface of the hind + tibia; squamæ usually large and + unequal. HYDROTAEA, _Aricia_, + _Drymeia_, _Ophyra_, _Phaonia_ (= + _Hyetodesia_), _Pogonomyia_, + _Trichophthicus_, etc. ARICINÆ + + oo. M_{1+2} curved or bent, hence the cell + R_5 more or less narrowed in the margin. + (MUSCINÆ). MUSCIDÆ in part. See page 303 + for generic synopsis. + + ll. Hypopleurals present; when three + sternopleurals are present the arrangement is + 2:1 or 1:1:1. (TACHINIDÆ of Girschner) + + m. Conjunctiva of the ventral sclerites of the + abdomen present, frequently well developed, + surrounding the sclerites. + + n. Mouth parts vestigial. OESTRIDÆ. See page + 297 for generic synopsis. + + nn. Mouth parts well developed. + + o. M_{1+2} straight, hence cell R_5 wide + open in the margin; costa ending at the + tip of R_5; three sternopleurals + present; antennal arista plumose. + _Syllegoptera_. Europe. + (SYLLEGOPTERINÆ). DEXIIDÆ in part + + oo. M_{1+2} bent, hence cell R_5 narrowed in + the margin; sternopleurals rarely + wanting, usually 1:1 or 0:1; facial + plate strongly produced below vibrissal + angle like the bridge of the nose; + antennal arista bare. Parasitic on + Hemiptera and Coleoptera. _Allophora_, + _Cistogaster_, _Clytia_, _Phasia_, etc. + (PHASIINÆ) TACHINIDÆ in part. + + mm. Conjunctiva of the ventral sclerites + invisible (fig. 161a). + + n. Second ventral sclerite of the abdomen + lying with its edges either upon or in + contact with the ventral edges of the + corresponding dorsal sclerite. + + o. Outermost posthumeral almost always lower + (more ventrad) in position than the + presutural macrochæta; fifth ventral + abdominal sclerite of the male cleft + beyond the middle, often strongly + developed; body color very frequently + metallic green or blue, or yellow; + arista plumose. (CALLIPHORINÆ) MUSCIDÆ + in part. + + See page 303 for generic synopsis. + + oo. Outermost posthumeral macrochæta on + level or higher (more dorsad) than the + presutural macrochæta; arista bare, + pubescent, or plumose only on the basal + two-thirds; body coloring usually + grayish (fig. 106). SARCOPHAGIDÆ + + p. Fifth ventral sclerite of the male + either wanting or with the caudal + margin straight; presutural intraalar + rarely present. (SARCOPHAGINÆ) + + q. Fifth ventral abdominal sclerite of + the male much reduced, the remaining + segments with straight posterior + margin, overlapping scale-like; in + the female only segment one and two + scale-like, the others wholly or in + part covered; sternopleurals usually + three or more. SARCOPHAGA and + related genera. + + qq. Fifth ventral sclerite of the male + plainly visible; sternopleurals + usually two. SARCOPHILA, + WOHLFAHRTIA, _Brachycoma_, + _Hilarella_, _Miltogramma_, + _Metopia_, _Macronychia_, _Nyctia_, + _Paramacronychia_, _Pachyphthalmus_, + etc. + + pp. Fifth ventral abdominal sclerite of + the male cleft to beyond the middle; + ventral sclerites usually visible, + shield-like. _Rhinophora_, _Phyto_, + _Melanophora_ RHINOPHORINÆ + +[Illustration: 164. Glossina palpalis. (×4.) After Austen.] + + nn. Second ventral abdominal sclerite as well + as the others more or less covered, + sometimes wholly, by the edges of the + dorsal sclerite. + + o. The presutural intraalar wanting; ventral + sclerites two to five nearly or quite + covered by the edges of the + corresponding dorsal sclerites; base of + the antennæ usually at or below the + middle of the eye; arista usually + plumose; legs usually elongate; + abdominal segments with marginal and + often discal macrochætæ. DEXIIDÆ + + oo. Presutural intraalar present, if absent, + then the ventral sclerites broadly + exposed or the fifth ventral sclerite + vestigial; base of the antennæ usually + above the middle of the eye; arista + bare; at least two posthumerals and + three posterior intraalars present. + Parasitic on caterpillars, etc. + TACHINIDÆ + + +SYNOPSIS OF THE PRINCIPAL GENERA OF THE MUSCIDÆ OF THE WORLD + +a. Proboscis long, directed forward, adapted for piercing, or oral + margin much produced, snout-like. + + b. Oral margin produced snout-like; vibrissa placed high above the + oral margin; antennal arista either pectinate or more or less + plumose. + + c. Antennal arista short or long-plumose; neither sex with distinct + orbital bristles. + + d. No facial carina between the antennæ. RHYNCHOMYIINÆ + + e. Arista short-plumose. _R. speciosa._ Europe. _Rhynchomyia_ R. + D. + + ee. Arista long-plumose. _I. phasina_. Europe and Egypt. + _Idiopsis_. B. B. + + dd. With flattened carina, the bases of the antennæ separated; no + abdominal macrochætæ. COSMININÆ _C. fuscipennis_. South + Africa. _Cosmina_ + + cc. Antennal arista pectinate; bases of the antennæ separated by a + flattened carina. RHINIINÆ R. D. + + d. Cell R_5 open, or closed at the margin. + + e. Third segment of the antenna twice as long as the second; + claws of both sexes short; cell R_5 open. _I. lunata._ + Eastern Hemisphere. _Idia_ Meigen + + ee. Third segment of the antenna three times as long as the + second; cell R_5 open or closed; claws of the male long and + slender, of the female shorter than the last tarsal joint. + _I. mandarina_, China. _Idiella_ B. B. + + dd. Cell R_5 petiolate. _Rhinia_; and _Beccarimyia_ Rdi. + + bb. Proboscis long, directed forward, adapted for piercing. STOMOXINÆ + + c. Arista flat, pectinate above with plumose rays; sternopleurals + 1:2; bases of the veins R_1 and R_{4+5} without setæ; base of + the media bowed down; apical cell opens before the apex of the + wing. African species. GLOSSINA Wied. + + d. Species measuring over twelve mm. in length. _G. longipennis_ + and FUSCA. + + dd. Species less than twelve mm. in length. + + e. All segments of the hind tarsi black. + + f. The fourth and fifth segments of the fore tarsi black; + antennæ black (fig. 164). G. PALPALIS R. D. + + ff. Otherwise marked. _G. bocagei_, _tachinoides_, _pallicera_. + + ee. First three segments of the hind tarsi are yellow, the + fourth and fifth segments are black. + + f. Fourth and fifth segments of the first and second pair of + tarsi are black. + + g. The yellow bands of the abdominal segments occupy a third + of the segment (fig. 165). G. MORSITANS Westw. + + gg. The yellow band on each segment of the abdomen occupies + a sixth of the segment. G. LONGIPALPIS Wied. + + ff. Tarsi of the first and second pairs of legs wholly yellow. + _G. pallidipes_ Austen + + cc. Rays of the arista not plumose; only one or two sternopleurals; + base of the media not strongly bowed down; apical cell opens at + or very near the apex of the wing. + + d. Vein R_{4+5} without setæ at the base; palpi about as long as + the proboscis. + + e. Arista pectinate (i.e. rays on one side only), the rays often + undulate; two yellow sternopleurals often difficult to + detect; vein M_{1+2} only slightly bent, the apical cell + hence wide open. The horn fly, H. IRRITANS (= _Lyperosia + serrata_) and related species. Widely distributed (figs. + 167, 168). HÆMATOBIA R. D. not B. B. + +[Illustration: 165. Glossina morsitans. (×4.) After Austen.] + + ee. Arista also with rays below; vein M_{1+2} more strongly + bent, the apical cell hence less widely open. + + f. Palpi strongly spatulate at the tips, lower rays of the + arista about six in number, B. SANGUINOLENTUS. South Asia. + BDELLOLARYNX Austen + + ff. Palpi feebly spatulate; apical cell of the wing narrowly + open slightly before the tip; sternopleurals black, + anterior bristle sometimes absent. H. ATRIPALPIS. Europe. + HÆMATOBOSCA Bezzi + + dd. Vein R_{4+5} with setæ at the base.[L] + + e. Veins R_1 and R_{4+5} with setæ at the base; two equally + prominent sternopleural macrochætæ; arista with rays both + above and below; palpi as long as the proboscis; apical cell + of the wing wide open. L. TIBIALIS. (_Hæmatobia_ B. B. not + R. D.). LYPEROSIOPS Town. + + ee. Only vein R_{4+5} with basal setæ; anterior sternopleural + macrochæta wanting; arista pectinate. + + f. Palpi as long as the proboscis, the latter stout, with + fleshy terminal labellæ; apical cell narrowly open; + sternopleural macrochætæ black. S. MACULOSA from Africa + and related species from Asia. STYGEROMYIA Austen + + ff. Palpi much shorter than the proboscis, the latter pointed + at the apex, without fleshy labellæ; apical cell of the + wing wide open. S. CALCITRANS, the stable fly and related + species. Widely distributed in both hemispheres (fig. + 110). STOMOXYS Geof. + +aa. Proboscis neither slender nor elongate, the labellæ fleshy and not + adapted for piercing. + + b. Hypopleuræ without a vertical row of macrochætæ. MUSCINÆ + + c. Arista bare; distal portion of M_{1+2} broadly curved at the end; + hypopleuræ with a sparse cluster of fine hairs. _S. + braziliana_, Southern States and southward. _Synthesiomyia_ B. + B. + + cc. Arista pectinate or plumose. + + d. Arista pectinate. _H. vittigera_, with the posterior half of + the abdomen metallic blue. Mexico. _Hemichlora_ V. d. W. + + dd. Arista plumose. + + e. Middle tibia with one or more prominent setæ on the inner + (flexor) surface beyond the middle, or inner surface very + hairy. + + f. R_1 ends distad of the m-cu crossvein; M_{1+2} with a broad + curve near its apical end. (= _Neomesembrina_ Schnabl. = + _Metamesembrina_ Town). _M. meridiana._ Europe. + _Mesembrina_ Meigen + + ff. R_1 ends proximad of the m-cu crossvein. + + g. Eyes pilose, sometimes sparsely in the female. + + h. Female with two or three stout orbital setæ; the hind + metatarsus of the male thickened below at the base and + penicillate. _D. pratorum._ Europe. _Dasyphora_ R. + D.[M] + + hh. Neither sex with orbital setæ. + + i. Abdomen without macrochætæ; arista plumose. _C. + asiatica._ Eastern Hemisphere. _Cryptolucilia_ B. B. + + ii. Abdomen with strong macrochætæ; arista very + short-plumose, nearly bare. _B. tachinina._ Brazil. + _Reinwardtia_ B. B. + + gg. Eyes bare. + + h. Body densely pilose; thoracic macrochætæ wanting; + middle tibiæ much elongate and bent; last section of + M_{1+2} with a gentle curve. H. (_Mesembrina_) + _mystacea, et al._, Europe and _H. solitaria_, N. + America. _Hypodermodes_ Town. + + hh. Body not densely pilose. + + i. Dorsocentrals six; last section of M_{1+2} with a + gentle curve. + + j. Inner dorsocentrals ("acrostichals") wanting; + sternopleurals arranged 1:3. _P. cyanicolor_, + _cadaverina_, etc. Europe and America _Pyrellia_ + R. D. + + jj. Inner dorsocentrals ("acrostichals") present; + sternopleurals arranged 1:2. E. _latreillii._ + North America. _Eumesembrina_ Town. + + ii. Dorsocentrals five; inner dorsocentrals present; + last section of M_{1+2} with a rounded angle; + sternopleurals arranged 1:2. _P. cornicina_ Europe + and America. (_Pseudopyrellia_ Girsch.) _Orthellia_ + R. D. + + ee. Middle tibia without a prominent bristle on the inner + surface beyond the middle. + +[Illustration: 166. Pycnosoma marginale. (×4.) After Graham-Smith.] + + f. Squamula thoracalis broadened mesad and caudad as far as + the scutellum. + + g. Sternopleural macrochætæ arranged in an equilateral + triangle; front of both sexes broad; genæ bare; + dorsocentrals six, small; wing not rilled. (To + COENOSINÆ). _Atherigona_ Rdi. + + gg. Sternopleural macrochætæ when three are present, + arranged in a right triangle. + + h. Last section of M_{1+2} with a more or less rounded + angle (fig. 163l). + + i. Eyes of the male pilose or pubescent, of the female + nearly bare; m-cu crossvein usually at or proximad + of the mid-distance between the r-m crossvein and + the bend of M_{1+2}. P. (= _Placomyia_ R. D.) + _vitripennis_. _Plaxemyia_ R. D. + + ii. Eyes bare; the m-cu crossvein always nearer to the + bend of M_{1+2} than to the r-m crossvein. + + j. Apex of the proboscis when extended reveals a + circlet of stout chitinous teeth. P. INSIGNIS + Austen, of India, bites both man and animals. (= + _Pristirhynchomyia_.) PHILÆMATOMYIA Austen + + jj. Apex of the proboscis without black teeth. + + k. Eyes of male separated by a distance equal to a + fourth the width of the head. House or typhoid + fly. M. DOMESTICA L. Widely distributed. MUSCA + L. + + kk. Eyes of the male contiguous. E. CORVINA. Europe. + EUMUSCA Town + + hh. Last section of M_{1+2} with a gentle curve (fig. + 102). + + i. Eyes pilose. + + j. Claws in the male somewhat elongated; no orbitals + in either sex; antennæ separated at the base by a + flat carina; abdomen marked with red or yellow. + _G. maculata._ Europe and America. _Graphomyia_ R. + D. + + jj. Claws short and equal in the two sexes; two or + three stout orbital macrochætæ in the female; R_1 + scarcely produced beyond the r-m crossvein; eyes + contiguous in the male. _P. obsoleta._ Brazil. + _Phasiophana_ Br. + + ii. Eyes bare; fronto-orbital macrochætæ in a double + row, antennæ contiguous at the base. + + j. One or more pairs of well developed anterior inner + dorsocentral (acrostichal) macrochætæ; seta on + extensor surface of hind tibia. M. ASSIMILIS, + STABULANS, etc. Europe and America. MUSCINA R. D. + + jj. Anterior inner dorsocentrals and the setæ; on the + extensor surface of the hind tibia wanting. _M. + micans_, etc. Europe and North America. _Morellia_ + R. D. + + ff. Squamula thoracalis not broadened mesad and caudad, not + reaching the margin of the scutellum; macrochætæ on + extensor surface of the hind tibia wanting. + + g. Eyes pubescent. _M. meditabunda._ Europe and America. + _Myiospila_ Rdi. + + gg. Eyes bare; R_1 ends before the middle of the wing. A + number of species from the tropics of both hemispheres. + _Clinopera_ V. d. W. + + bb. Hypopleuræ with a vertical row of macrochætæ. + + c. Eyes pubescent. + + d. R_1 ends about opposite the r-m crossvein; basal section of + R_{4+5} bristly nearly to the crossvein; _S. enigmatica_. + Africa. _Somalia_ Hough + + dd. R_1 ends distad of the r-m crossvein. + + e. Eastern hemisphere. Australasia. _N. ochracea_, _dasypthalma_. + _Neocalliphora_ Br. + + ee. Western Hemisphere. _T. muscinum._ Mexico. _Tyreomma_ V. d. + W. + + cc. Eyes bare. + + d. The vibrissal angle situated at a noticeable distance above the + level of the margin of the mouth. + + e. Sternopleural macrochætæ arranged in the order 1:1. + + f. Genæ with microchætæ. + + g. Body grayish, with depressed yellow woolly hair among the + macrochætæ; wings folded longitudinally over the body + when at rest. Cluster flies. _P. rudis_ and related + species, widely distributed. _Pollenia_ R. D.[N] + + gg. Body metallic blue or green. Eastern Hemisphere. + + h. Vibrissal angle placed very high above the oral margin; + a carina between the antennæ; outer posthumeral + wanting; anterior intraalar present. _T. viridaurea_. + Java. _Thelychæta_ Br. + +[Illustration: 167. Horn fly. (_a_) egg; (_b_) larva; (_c_) puparium; +(_d_) adult. (×4). Bureau of Entomology] + + hh. Vibrissal angle moderately high above the oral margin; + carina small or wanting; no post humeral macrochæta; + lower squamæ hairy above. (= _Paracompsomyia_ Hough) + (fig. 166). _Pycnosoma_ Br. + + ff. Genæ bare. _S. terminata_. Eastern Hemisphere + _Strongyloneura_ Bigot + + ee. Sternopleurals arranged 2:1. + + f. Body metallic green or blue, with gray stripes; genæ hairy + to the lower margin; post humerals often wanting; lower + squamæ bare above. (= _Compsomyia_ Rdi.). CHRYSOMYIA R. D. + + g. With one or two orbitals; height of bucca less than half + the height of the eye. South and east U. S. (fig. 107). + C. MARCELLARIA + + gg. No orbitals; height of bucca about a third less than + height of eye. West U. S. _C. wheeleri_ Hough + + ff. Body black or sordidly metallic greenish gray, usually + yellow pollinose or variegate; genæ at most hairy above. + _N. stygia_. Eastern Hemisphere. _Neopollenia_ Br. + + dd. Vibrissal angle situated nearly on a level of the oral margin. + + e. Species wholly blackish, bluish, or greenish metallic in + color. + + f. First section of R_{4+5} with at most three or four small + bristles at the immediate base. + + g. The bend of M_{1+2} a gentle curve; costal spine present; + cell R_5 closed, ending before the apex of the wing. _S. + cuprina._ Java. _Synamphoneura_ Bigot + + gg. Bend of M_{1+2} angular; or the insect differs in other + characters; dorsal surface of the squamula thoracalis + hairy (except in _Melinda_); arista plumose only on the + basal two-thirds (except usually in _Calliphora_ and + _Eucalliphora_). + +[Illustration: 168. Head of horn-fly (Lyperosia irritans); (_a_) female; +(_b_) male; (_c_) lateral aspect of female. Bureau of Entomology.] + + h. Arista plumose only on the basal two-thirds. + + i. Base of the antennæ ventrad of the middle of the eye; + eyes of the male nearly contiguous; genæ hairy; + second abdominal segment with median marginal + macrochætæ; two, rarely three, postsutural intraalar + macrochætæ. + + j. Squamula thoracalis dorsally with long black hairs; + male hypopygium two-segmented, large, projecting; + claws and pulvilli of the male elongate; three + strong sternopleural macrochætæ; genæ at least + half the width of the eye; buccæ (cheeks) half the + height of the eyes; oviviparous. _O. sepulcralis._ + Europe. _Onesia_ R. D. + + jj. Dorsal surface of the squamula thoracalis bare; + male hypopygium small, scarcely projecting below; + claws and pulvilli not elongate; two stout + sternopleural macrochætæ, sometimes with a + delicate one below the anterior; genæ nearly + linear in the male; buccæ about a third of the eye + height; oviparous. _M. cærulea_. Europe. + _Melinda_. R. D. + +[Illustration: 169. Lateral and dorsal aspects of the thorax, and +frontal aspect of the head of a muscoidean fly, with designations of the +parts commonly used in taxonomic work.] + + ii. Base of the antennæ dorsad of the middle of the eye; + eyes of both sexes distinctly separated; dorsal + surface of the squamula thoracalis with black hairs; + two postsutural intraalar macrochætæ. + + j. Hypopygium of the male large, with a pair of + slightly curved forceps whose ends are concealed + in a longitudinal slit in the fifth ventral + sclerite; third posterior inner dorso-central + (acrostichal) macrochætæ absent; anterior + intraalar rarely present; abdomen usually not + pollinose; the second segment without median + marginal macrochætæ; face yellow. _C. mortuorum_, + _cadaverina_, and related species. Both + hemispheres. _Cynomyia_ R. D.[O] + +[Illustration: 170. Sepsis violacea; puparium and adult. (See page 297.) +After Howard.] + + jj. Three pairs of posterior inner dorsocentrals + (acrostichals) present; second abdominal segment + with a row of marginal macrochætæ; genæ hairy, at + least above. + + k. Hypopygium of the male with a projecting style. + _S. stylifera_. Europe. _Steringomyia_ Pok. + + + kk. Hypopygium of the male without style. _A. + stelviana_ B. B. _Acrophaga_ B. B. + + hh. Arista usually plumose nearly to the tip; posterior + dorsocentrals and inner dorsocentrals (acrostichals) + well developed; dorsal surface of the squamula + thoracalis hairy; abdomen metallic and usually + pollinose; genæ hairy. + + i. With one pair of ocellar macrochætæ. C. VOMITORIA, + ERYTHROCEPHALA, VIRIDESCENS, and related species. + Both hemispheres. CALLIPHORA R. D. + + ii. With two strong pairs of ocellar macrochætæ. _E. + latifrons._ Pacific slope of the U. S. + _Eucalliphora_ Town. + + ff. First section of R_{4+5} bristly near or quite half way to + the small crossvein; dorsal surface of the squamula + thoracalis is bare; the hypopygium of the male is + inconspicuous. + + g. Genæ bare; posterior inner and outer dorsocentrals + distinct and well developed. _L. cæsar_, _sericata_, + _sylvarum_, and related species. Widely distributed in + both hemispheres (fig. 103). LUCILIA R. D. + + gg. Genæ with microchætæ, at least down to the level of the + base of the arista. + + h. Mesonotum flattened behind the transverse suture. + + i. Posterior dorsocentrals inconstant and unequally + developed; one pair of posterior inner + dorsocentrals. _P. terrænovæ._ North America. + _Protophormia_ Town. + + ii. Posterior dorsocentrals well developed, the inner + dorsocentrals (acrostichals) unequally developed. + _P. azurea_, _chrysorrhÅ“a_, etc. Europe and America. + _Protocalliphora_ Hough + + hh. Mesonotum not flattened behind the transverse suture; + posterior inner and outer dorsocentrals inconstant and + unequally developed. _P. regina._ Europe and America. + _Phormia_ R. D. + + ee. Species more or less rufous or yellow in color. + + f. Anterior dorsocentrals wanting; first section of the + R_{4+5} at most only bristly at the base, bend near apex + of M_{1+2} rectangular, R_1 ends over the crossvein; + fronto-orbital macrochæta absent; eyes of the male + contiguous. _C. semiviridis._ Mexico. _Chloroprocta_ + V. d. W + + ff. With another combination of characters. + + g. Body robust, of large size, abdomen elongate, not round; + genæ with several ranges of microchætæ; vibrissal ridges + strongly convergent; abdomen with well developed + macrochætæ; costal spine usually absent; eyes of the + male widely separated. + +[Illustration: 171. Stigmata of the larvæ of Muscoidea. Third instar. +(_a_) Cynomyia cadaverina; (_b_) Phormia regina; (_c_) Chrysomyia +macellaria; (_d_) Musca domestica; (_e_) Sarcophaga sp.; (_f_) Oestris +ovis; (_g_) Gastrophilus equi; (_h_) Sarcophaga sp.; (_i_) Pegomyia +vicina; (_j_) Protocalliphora azurea; (_k_) Hypoderma lineata; (_l_) +Muscina stabulans. Magnification for f, g, and k, ×25; all others, × +50.] + + h. Peristome broad, pteropleural macrochætæ distinct; one + or two sternopleurals; in the female a single orbital + macrochæta; last abdominal segment without discal + macrochætæ; hypopygial processes of the male with a + long stylet; second abdominal segment of the female + sometimes much elongate. A. LUTEOLA (fig. 86). + Africa. The sub-genus _ChÅ“romyia_ Roub. is included + here. AUCHMEROMYIA B. B. + + hh. Peristome narrow; no pteropleurals, two + sternopleurals; two orbitals in the female; second + segment not elongate; the fourth with two well + developed discal macrochætæ. B. DEPRESSA. Africa. + BENGALIA R. D + + gg. With another combination of characters. + + h. Costal spine present; body in part black; antennæ + noticeably shorter than the epistome, inserted above + the middle of the eye and separated from each other by + a carina; abdominal segments with marginal macrochætæ; + sternopleurals 2:1 or 1:1. _Paratricyclea_ Villen. + + hh. Costal spine not distinct, or if present, insect + otherwise different. + + i. Genæ with several ranges of microchætæ; vibrissal + ridges strongly converging; peristome broad; arista + moderately plumose; sternopleurals usually 1:1; + color entirely testaceous. C. ANTHROPOPHAGA (fig. + 87) and GRUNBERGI. Africa. CORDYLOBIA Grünb. + + ii. Genæ bare or with but one range of setæ; vibrissal + ridges less converging; peristome narrow; arista + long plumose. + + j. Genæ with a single row of microchætæ. + + k. Sternopleurals 2:1; color entirely testaceous. + _Ochromyia_ Macq.[P] + + kk. Sternopleurals 1:1. _P. varia_ Hough. Africa. + _Parochromyia_ Hough + + jj. Genæ bare. + + k. Basal section of R_{4+5} bristly only at the + immediate base, distally M_{1+2} with a broad + curve; distal portion of the abdomen metallic; + sternopleurals usually 1:1, rarely 2:1. _M. + æneiventris_ Wd. Tropic America. + _Mesembrinella._ G. T. + + kk. R_{4+5} bristly at least nearly half way to the + small crossvein; sternopleurals 1:1. + + l. Macrochætæ of the abdomen marginal; neither sex + with orbitals; no carina between the base of + the antennæ; three pairs of presutural inner + dorsocentrals. Eastern hemisphere. _T. + ferruginea._ _Tricyclea V. d. W_. (= + _Zonochroa_ B. B. according to Villeneuve + 1914). + + ll. Abdomen without macrochætæ; wing usually with + a marginal streak and gray markings. Brazil. + Hemilucilia B. B. + +[Illustration: 172. Left hand stigmata of the larvæ of muscoidea. Third +instar. (_a_) Lucilia cæsar; (_b_) Calliphora vomitoria; (_c_) Stomoxys +calcitrans; (_d_) Orthellia cornicina; (_e_) Pyrellia cadaverina; +(_f_) Hæmatobia irritans; (_g_) Mesembrina mystacea; (_h_) Mesembrina +meridiana; (_i_) Myospila meditabunda; (_j_) Mydæa urbana; (_k_) +Polietes albolineata; (_l_) Polietes lardaria; (_m_) Morellia hortorum; +(_n_) Hydrotæa dentipes; (_o_) Hebecnema umbratica; (_p_) H. vespertina; +(_q_) Limnophora septemnotata; (_r_) Muscina stabulans. (_a_ and _b_) +after MacGregor; (_d_) after Banks; all others after Portchinsky. +Magnification varies. The relative distance to the median line is +indicated in each figure.] + + +SIPHONAPTERA. Fleas + +Adapted from a table published by Oudemans. + +a. Elongated fleas, with jointed (articulated) head, with combs + (ctenidia) on head and thorax; with long, oval, free-jointed + flagellum of the antenna (fig. 92d). Suborder FRACTICIPATA + + b. With ctenidia in front of the antennæ and on the genæ (cheeks); + maxillæ with acute apices; labial palpi five-segmented, + symmetrical; eyes poorly developed or wanting. On rodents. + HYSTRICHOPSYLLIDÆ + + c. Abdominal segments without ctenidia. + + d. Post-tibial spines in pairs and not in a very close set row; + head with ctenidia. _Ctenophthalmus_ Kol. + + dd. Post-tibial spines mostly single and in a close set row. + _Ctenopsyllus_ and _Leptopsyllus_. The last genus has recently + been erected for _L. musculi_, a widely distributed species + occurring on rats and mice. + + cc. Abdominal segments with one or more ctenidia; post-tibial spines + in numerous, short, close-set transverse rows on posterior + border with about four spines in each row. _H. americana._ + _Hystrichopsylla_ Taschenb. + + bb. With only two pairs of subfrontal ctenidia; labial palpi + five-segmented, symmetrical; eyes vestigial or wanting. On bats. + (= ISCHNOPSYLLIDÆ). NYCTERIDIPSYLLIDÆ + + With more or less blunt maxilla; all tibiæ with notch; a single + antepygidial bristle; metepimeron without ctenidium. _N. + crosbyi_ from Missouri was found on bats. Rothschild suggests + that this is probably the same as _N. insignis_. (= + _Ischnopsyllus_ = _Ceratopsyllus_), _Nycteridiphilus_ + +aa. Head not jointed, i.e. the segments coalescent, traces of the + segmentation still being visible in the presence of the vertex + tubercle, the falx (sickle-shaped process), and a suture. Suborder + INTEGRICIPITA + + b. Flagellum of the antennæ long and oval. + + c. Usually elongate fleas, with a free-segmented flagellum of the + antenna; thorax not shorter than the head, longer than the first + tergite. + + d. Genæ of the head and the pronotum with ctenidia. NEOPSYLLIDÆ + + e. Labial palpi four or five-segmented; symmetrical; hind coxæ + with patch of spines inside; row of six spatulate spines on + each side in front of the antennæ. _C. ornate_ found on a + California mole. _Corypsylla_ + + ee. Labial palpi two-segmented, transparent, membranous. On + hares. _Spilopsyllus_ Baker + + dd. No ctenidium on the head. + + e. Pronotum with ctenidium. DOLICHOPSYLLIDÆ + + f. Labial palpi five-segmented, symmetrical. + + g. Antepygidial bristles one to three; eyes present. + + h. Inner side of hind coxæ distally with a comb of minute + teeth; falx present. On rodents and carnivores. + _Odontopsyllus_ Baker + + hh. Inner side of hind coxæ without comb or teeth. Many + North American species on rodents. CERATOPHYLLUS + Curtis + + gg. Antepygidial bristles five on each side; eyes absent; + suture white. _D. stylosus_ on rodents. _Dolichopsyllus_ + Baker + + ff. Labial palpi four or five-segmented; asymmetrical + (membranous behind), apex acute. _Hoplopsyllus anomalus_ + found on Spermophiles in Colorado. HOPLOPSYLLIDÆ + + ee. Pronotum without ctenidium. _Anomiopsyllus californicus_ and + _nudatus_ on rodents. ANOMIOPSYLLIDÆ + + cc. Very short fleas; flagellum of the antenna with pseudo-segments + coalescent; thorax much shorter than the head and than the first + tergite. HECTOPSYLLIDÆ + + Flagellum of the antenna with six coalescent pseudo-segments; + maxilla blunt. The chigger on man (fig. 93). D. PENETRANS. (= + RHYNCHOPRION = SARCOPSYLLA) DERMATOPHILUS Guérin + + bb. Flagellum short, round, free portion of the first segment shaped + like a mandolin. + + c. Thorax not shorter than the head, longer than the first tergite; + flagellum either with free segments or in part with the segments + coalescent. + + d. Head and pronotum with ctenidium; labial palpi asymmetrical. + ARCHÆOPSYLLIDÆ + + With four subfrontal, four genal, and one angular ctenidia. + Widely distributed. CTENOCEPHALUS Kol. + + e. Head rounded in front (fig. 92a). Dog flea. C. CANIS + + ee. Head long and flat (fig. 92b). Cat flea. C. FELIS + + dd. Neither head nor pronotum with ctenidium. Labial palpi + asymmetrical, membranous behind. PULICIDÆ + + e. Mesosternite narrow, without internal rod-like thickening + from the insertion of the coxæ upwards. Human flea, etc. + PULEX L. + + ee. Mesosternite broad with a rod-like internal thickening from + the insertion of the coxæ upwards (fig. 89). X. + (LÅ’MOPSYLLA) CHEOPIS, plague or rat flea. XENOPSYLLA + + cc. Thorax much shorter than the head and than the first tergite. + ECHIDNOPHAGIDÆ. E. GALLINACEA, the hen flea also attacks man + (fig. 96). (= ARGOPSYLLA = XESTOPSYLLA) ECHIDNOPHAGA Olliff. + + +FOOTNOTES: + +[E] Adapted from Banks, Nuttall, Warburton, Stiles, _et al._ + +[F] Dr. C. W. Stiles considers the species which is responsible for +spotted fever distinct from the _venustus_ of Banks, separating it as +follows: + +Goblet cells about 75 in the male or 105 in the female. Texas. _D. +venustus._ + +Goblet cells 157 in the male, or 120 in the female; stigmal plate shaped +as shown in the figure (figs. 150 a, b). Montana, etc. D. ANDERSONI. + +[G] Professor C. R. Crosby who has been working upon certain capsids +states that he and his assistant have been bitten by LYGUS PRATENSIS, +the tarnished plant bug, by CHLAMYDATUS ASSOCIATUS and by ORTHOTYLUS +FLAVOSPARSUS, though without serious results. + +[H] Species marked with an * are known to transmit malaria. Species +found only in tropical North America and not known to carry malaria have +been omitted from this table, but all found in the United States are +included. + +[I] This table to the North American genera of the Tabanidæ is adapted +from one given by Miss Ricardo. + +[J] The classification of the Muscoidea as set forth by Schiner and +other earlier writers has long been followed, although it is not +satisfactory, being admittedly more or less artificial. Within the last +two or three decades several schemes have been advanced, that of Brauer +and Bergenstamm and of Girschner, with the modifications of Schnabl and +Dziedzicki having obtained most favor in Europe. Townsend, in 1908, +proposed a system which differs from Girschner's in some respects, but +unfortunately it has not yet been published in sufficient detail to +permit us to adopt it. From considerations of expediency we use here the +arrangement given in Aldrich's Catalogue of North American Diptera, +though we have drawn very freely upon Girschner's most excellent paper +for taxonomic characters to separate the various groups. + +It may sometimes be found that a species does not agree in all the +characters with the synopsis; in this case it must be placed in the +group with which it has the most characters in common. + +[K] There are several genera of flies of the family _Cordyluridæ_; (i.e. +_Acalyptratæ_) which might be placed with the _Anthomyiidæ_ (i.e. +_Calyptratæ_), owing to the relatively large size of their squamæ. As +there is no single character which will satisfactorily separate all +doubtful genera of these two groups we must arbitrarily fix the limits. +In general those forms on the border line having a costal spine, or +lower squama larger than the upper, or the lower surface of the +scutellum more or less pubescent, or the eyes of the male nearly or +quite contiguous, or the eyes hairy, or the frontal setæ decussate in +the female; or any combination of these characters may at once be placed +with the _Anthomyiidæ_. Those forms which lack these characteristics and +have at least six abdominal segments (the first and second segments +usually being more or less coalescent) are placed with the Acalyptrates. +There are other acalyptrates with squamæ of moderate size which have +either no vibrissæ, or have the subcosta either wholly lacking or +coalescent in large part with R_1 or have spotted wings; they, therefore +will not be confused with the calyptrates. + +[L] _Pachymyia_ Macq. is closely related to _Stomoxys_. It differs in +having the arista rayed both above and below. _P. vexans_, Brazil. + +[M] The genus _Eudasyphora_ Town. has recently been erected to contain +_D. lasiophthalma_. + +[N] _Nitellia_, usually included in this genus has the apical cell +petiolate. _Apollenia_ Bezzi, has recently been separated from +_Pollenia_ to contain the species _P. nudiuscula_. Both genera belong to +the Eastern hemisphere. + +[O] The following three genera are not sufficiently well defined to +place in this synopsis. In color and structural characters they are +closely related to _Cynomyia_ from which they may be distinguished as +follows. _Catapicephala_ Macq., represented by the species _C. +splendens_ from Java, has the setæ on the facial ridges rising to the +base of the antennæ and has median marginal macrochætæ on the abdominal +segments two to four: _Blepharicnema_ Macq., represented by _B. +splendens_ from Venezuela has bare genæ, oral setæ not ascending; tibiæ +villose; claws short in both sexes; _Sarconesia_ Bigot with the species +_S. chlorogaster_ from Chile, setose genæ; legs slender, not villose; +claws of the male elongate. + +[P] _Plinthomyia_ Rdi. and _Hemigymnochæta_ Corti are related to +_Ochromyia_, though too briefly described to place in the key. + + + + +APPENDIX + +HYDROCYANIC ACID GAS AGAINST HOUSEHOLD INSECTS + + +The following directions for fumigating with hydrocyanic acid +gas are taken from Professor Herrick's circular published by the +Cornell Reading Course: + +Hydrocyanic acid gas has been used successfully against household +insects and will probably be used more and more in the future. +It is particularly effective against bed-bugs, and cockroaches, but +because _it is such a deadly poison it must be used very carefully_. + +The gas is generated from the salt potassium cyanid, by treating +it with sulfuric acid diluted with water. Potassium cyanid is a +most poisonous substance and the gas emanating from it is also +deadly to most, if not all, forms of animal life. The greatest care +must always be exercised in fumigating houses or rooms in buildings +that are occupied. Before fumigation a house should be vacated. +It is not necessary to move furniture or belongings except brass or +nickel objects, which may be somewhat tarnished, and butter, milk, +and other larder supplies that are likely to absorb gas. If the nickel +and brass fixtures or objects are carefully covered with blankets +they will usually be sufficiently protected. + +There may be danger in fumigating one house in a solid row of +houses if there is a crack in the walls through which the gas may find +its way. It also follows that the fumigation of one room in a house +may endanger the occupants of an adjoining room if the walls between +the two rooms are not perfectly tight. It is necessary to keep +all these points in mind and to do the work deliberately and thoughtfully. +The writer has fumigated a large college dormitory of 253 +rooms, once a year for several years, without the slightest accident +of any kind. In order to fumigate this building about 340 pounds +of cyanid and the same amount of sulfuric acid were used each time. +In addition to this, the writer has fumigated single rooms and smaller +houses with the gas. In one instance the generating jars were too +small; the liquid boiled over and injured the floors and the rugs. +Such an accident should be avoided by the use of large jars and by +placing old rugs or a quantity of newspapers beneath the jars. + + +THE PROPORTIONS OF INGREDIENTS + +Experiments and experience have shown that the potassium +cyanid should be ninety-eight per cent pure in order to give satisfactory +results. The purchaser should insist on the cyanid being of +at least that purity, and it should be procurable at not more than +forty cents per pound. The crude form of sulfuric acid may be used. +It is a thickish, brown liquid and should not cost more than four or +five cents a pound. If a room is made tight, one ounce of cyanid for +every one hundred cubic feet of space has been shown to be sufficient. +It is combined with the acid and water in the following proportions: + + Potassium cyanid 1 ounce + Commercial sulfuric acid 1 fluid ounce + Water 3 fluid ounces + + +A SINGLE ROOM AS AN EXAMPLE + +Suppose a room to be 12 by 15 by 8 feet. It will contain 12 × 15 × 8, or +1440 cubic feet. For convenience the writer always works on the basis of +complete hundreds; in this case he would work on the basis of 1500 cubic +feet, and thus be sure to have enough. The foregoing room, then, would +require 15 ounces of cyanid, 15 ounces of sulfuric acid, and 45 ounces +of water. The room should be made as tight as possible by stopping all +the larger openings, such as fireplaces and chimney flues, with old rags +or blankets. Cracks about windows or in other places should be sealed +with narrow strips of newspaper well soaked in water. Strips of +newspaper two or three inches wide that have been thoroughly soaked in +water may be applied quickly and effectively over the cracks around the +window sash and elsewhere. Such strips will stick closely for several +hours and may be easily removed at the conclusion of the work. + +While the room is being made tight, the ingredients should be measured +according to the formula already given. The water should be measured and +_poured first_ into a stone jar for holding at least two gallons. The +jar should be placed in the middle of the room, with an old rug or +several newspapers under it in order to protect the floor. + +The required amount of sulfuric acid should then be poured rather slowly +into the water. _This process must never be reversed; that is, the acid +must never be poured into the jar first._ The cyanid should be weighed +and put into a paper bag beside the jar. All hats, coats, or other +articles that will be needed before the work is over should be removed +from the room. When everything is ready the operator should drop the bag +of cyanid gently into the jar, holding his breath, and should walk +quickly out of the room. The steam-like gas does not rise immediately +under these conditions, and ample time is given for the operator to walk +out and shut the door. If preferred, however, the paper bag may be +suspended by a string passing through a screw eye in the ceiling and +then through the keyhole of the door. In this case the bag may be +lowered from the outside after the operator has left the room and closed +the door. + +The writer has most often started the fumigation toward evening and left +it going all night, opening the doors in the morning. The work can be +done, however, at any time during the day and should extend over a +period of five or six hours at least. It is said that better results +will be obtained in a temperature of 70° F., or above, than at a lower +degree. + +At the close of the operation the windows and doors may be opened from +the outside. In the course of two or three hours the gas should be +dissipated enough to allow a person to enter the room without danger. +The odor of the gas is like that of peach kernels and is easily +recognized. The room should not be occupied until the odor has +disappeared. + + +FUMIGATING A LARGE HOUSE + +The fumigation of a large house is merely a repetition, in each room and +hall, of the operations already described for a single room. All the +rooms should be made tight, and the proper quantities of water and +sulfuric acid should be measured and poured into jars placed in each +room with the cyanid in bags besides the jars. When all is ready, the +operator should _go to the top floor and work downward_ because the gas +is lighter than air and tends to rise. + + +PRECAUTIONS + +The cyanid should be broken up into small pieces not larger than small +eggs. This can best be done on a cement or brick pavement. It would be +advantageous to wear gloves in order to protect the hands, although the +writer has broken many pounds of cyanid without any protection on the +hands. Wash the hands thoroughly at frequent intervals in order to +remove the cyanid. + +The operations of the work must be carried out according to directions. + +The work should be done by a calm, thoughtful and careful person--best +by one who has had some experience. + +Conspicuous notices of what has been done should be placed on the doors, +and the doors should be locked so that no one can stray into the rooms. + +The gas is lighter than air, therefore one should always begin in the +rooms at the top of the house and work down. + +After fumigation is over the contents of the jar should be emptied into +the sewer or some other safe place. The jars should be washed thoroughly +before they are used again. + +_It must be remembered that cyanid is a deadly poison_; but it is very +efficient against household insects, if carefully used, and is not +particularly dangerous when properly handled. + + +LESIONS PRODUCED BY THE BITE OF THE BLACK-FLY + +While this text was in press there came to hand an important paper +presenting a phase of the subject of black fly injury so different from +others heretofore given that we deem it expedient to reproduce here the +author's summary. The paper was published in _The Journal of Cutaneous +Diseases_, for November and December, 1914, under the title of "A +Clinical, Pathological and Experimental Study of the Lesions Produced by +the Bite of the Black Fly (_Simulium venustum_)," by Dr. John Hinchman +Stokes, of the University of Michigan. + + +RESUME AND DISCUSSION OF EXPERIMENTAL FINDINGS + +The principal positive result of the work has been the experimental +reproduction of the lesion produced by the black-fly in characteristic +histological detail by the use of preserved flies. The experimental +lesions not only reproduced the pathological pictures, but followed a +clinical course, which in local symptomatology especially, tallied +closely with that of the bite. This the writer interprets as +satisfactory evidence that the lesion is not produced by any living +infective agent. The experiments performed do not identify the nature of +the toxic agent. Tentatively they seem to bring out, however, the +following characteristics. + +1. The product of alcoholic extraction of flies do not contain the toxic +agent. + +2. The toxic agent is not inactivated by alcohol. + +3. The toxic agent is not destroyed by drying fixed flies. + +4. The toxic agent is not affected by glycerin, but is, if anything, +more active in pastes made from the ground fly and glycerin, than in the +ground flies as such. + +5. The toxic agent is rendered inactive or destroyed by hydrochloric +acid in a concentration of 0.25%. + +6. The toxic agent is most abundant in the region of the anatomical +structures connected with the biting and salivary apparatus (head and +thorax). + +7. The toxic agent is not affected by a 0.5% solution of sodium +bicarbonate. + +8. The toxic agent is not affected by exposure to dry heat at 100° C. +for two hours. + +9. The toxic agent is destroyed or rendered inactive in alkaline +solution by a typical hydrolytic ferment, pancreatin. + +10. Incomplete experimental evidence suggests that the activity of the +toxic agent may be heightened by a possible lytic action of the blood +serum of a sensitive individual, and that the sensitive serum itself may +contain the toxic agent in solution. + +These results, as far as they go (omitting No. 10), accord with Langer's +except on the point of alcoholic solubility and the effect of acids. The +actual nature of the toxic agent in the black-fly is left a matter of +speculation. + +The following working theories have suggested themselves to the writer. +First, the toxin may be, as Langer believes in the case of the bee, an +alkaloidal base, toxic as such, and neutralized after injection by +antibodies produced for the occasion by the body. In such a case the +view that a partial local fixation of the toxin occurs, which prevents +its immediate diffusion, is acceptable. Through chemotactic action, +special cells capable of breaking up the toxin into harmless elements +are attracted to the scene. Their function may be, on the other hand, to +neutralize directly, not by lysis. This would explain the rôle of the +eosinophiles in the black-fly lesion. If their activities be essential +to the destruction or neutralization of the toxin, one would expect them +to be most numerous where there was least reaction. This would be at the +site of a bite in an immune individual. A point of special interest for +further investigation, would be the study of such a lesion. + +Second, it is conceivable that the injected saliva of the fly does not +contain an agent toxic as such. It is possible, that like many foreign +proteins, it only becomes toxic when broken down. The completeness and +rapidity of the breaking down depends on the number of eosinophiles +present. In such a case immunity should again be marked by intense +eosinophilia. + +[Illustration: 173. Fifth day mature lesion. Lower power drawing showing +papillary Å“dema and infiltrate in the region of the puncture. After Dr. +J. H. Stokes] + +Third, lytic agents in the blood serum may play the chief rôle in the +liberation of the toxic agent from its non-toxic combination. An immune +individual would then be one whose immunity was not the positive one of +antibody formation, but the negative immunity of failure to metabolize. +An immune lesion in such a case might be conceived as presenting no +eosinophilia, since no toxin is liberated. If the liberation of the +toxin is dependent upon lytic agents present in the serum rather than in +any cellular elements, a rational explanation would be available for the +apparent results (subject to confirmation) of the experiment with +sensitive and immune sera. In this experiment it will be recalled that +the sensitive serum seemed to bring out the toxicity of the ground +flies, and the serum itself seemed even to contain some of the dissolved +or liberated toxin. The slowness with which a lesion develops in the +case of the black-fly bite supports the view of the initial lack of +toxicity of the injected material. The entire absence of early +subjective symptoms, such as pain, burning, etc., is further evidence +for this view. It would appear as if no reaction occurred until lysis of +an originally non-toxic substance had begun. Regarding the toxin itself +as the chemotactic agent which attracts eosinophiles, its liberation in +the lytic process and diffusion through the blood stream attracts the +cells in question to the point at which it is being liberated. Arriving +upon the scene, these cells assist in its neutralization. + +The last view presented is the one to which the author inclines as the +one which most adequately explains the phenomena. + +A fourth view is that the initial injection of a foreign protein by the +fly (i.e., with the first bite) sensitizes the body to that protein. Its +subsequent injection at any point in the skin gives rise to a local +expression of systematic sensitization. Such local sensitization +reactions have been described by Arthus and Breton, by Hamburger and +Pollack and by Cowie. The description of such a lesion given by the +first named authors, in the rabbit, however, does not suggest, +histopathologically at least, a strong resemblance to that of the +black-fly. Such an explanation of many insect urticariæ deserves further +investigation, however, and may align them under cutaneous expressions +of anaphylaxis to a foreign protein injected by the insect. Depending on +the chemical nature of the protein injected, a specific chemotactic +reaction like eosinophilia may or may not occur. Viewed in this light +the development of immunity to insect bites assumes a place in the +larger problem of anaphylaxis. + +[Illustration: 174. Experimental lesion produced from alcohol-fixed +flies, dried and ground into a paste with glycerin. After Dr. J. H. +Stokes] + + +SUMMARY + +In order to bring the results of the foregoing studies together, the +author appends the following résumé of the clinical data presented in +the first paper. + +The black-fly, _Simulium venustum_, inflicts a painless bite, with +ecchymosis and hæmorrhage at the site of puncture. A papulo-vesicular +lesion upon an urticarial base slowly develops, the full course of the +lesion occupying several days to several weeks. Marked differences in +individual reaction occur, but the typical course involves four stages. +These are, in chronological order, the papular stage, the vesicular or +pseudovesicular, the mature vesico-papular or weeping papular stage and +the stage of involution terminating in a scar. The papule develops in +from 3 to 24 hours. The early pseudovesicle develops in 24 to 48 hours. +The mature vesico-papular lesion develops by the third to fifth day and +may last from a few days to three weeks. Involution is marked by +cessation of oozing, subsidence of the papule and scar-like changes at +the site of the lesion. The symptoms accompanying this cycle consist of +severe localized or diffused pruritus, with some heat and burning in the +earlier stages if the Å“dema is marked. The pruritus appears with the +pseudovesicular stage and exhibits extraordinary persistence and a +marked tendency to periodic spontaneous exacerbation. The flies tend to +group their bites and confluence of the developing lesions in such cases +may result in extensive Å“dema with the formation of oozing and +crusted plaques. A special tendency on the part of the flies to attack +the skin about the cheeks, eyes and the neck along the hair line and +behind the ears, is noted. In these sites inflammation and Å“dema may +be extreme. + +A distinctive satellite adenopathy of the cervical glands develops in +the majority of susceptible persons within 48 hours after being bitten +in the typical sites. This adenopathy is marked, discrete and painful, +the glands often exquisitely tender on pressure. It subsides without +suppuration. + +Immunity may be developed to all except the earliest manifestations, by +repeated exposures. Such an immunity in natives of an infested locality +is usually highly developed. There are also apparently seasonal +variations in the virulence of the fly and variations in the reaction of +the same individual to different bites. + +Constitutional effects were not observed but have been reported. + + + + +BIBLIOGRAPHY + + +ALDRICH, J. M. 1905. A catalogue of North American Diptera. Washington, + D. C. 1-680. + +ALESSANDRI, G. 1910. Studii ed esperienze sulle larve della Piophila + casei. Arch. Parasit. xiii, p. 337-387. + +ANDERSON, J. F. and FROST, W. H. 1912. Transmission of poliomyelitis by + means of the stable-fly (Stomoxys calcitrans). Public Health Reports. + Washington. xxvii, p. 1733-1735. + +---- 1913. Further attempts to transmit the disease through the agency + of the stable-fly (Stomoxys calcitrans). Public Health Repts., + Washington. xxviii, p. 833-837. + +ANDERSON, J. F. and GOLDBERGER, J. 1910. 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The migratory habit of housefly larvæ as + indicating a favorable remedial measure. An account of progress. U. + S. Dept. Agric., Bul. 14, p. 1-11. + +JENNINGS, A. H. 1914. Summary of two years' study of insects in relation + to pellagra. Journ. of Parasitology, i, p. 10-21. + +JENNINGS, A. H. and KING, W. V. 1913. One of the possible factors in the + causation of pellagra. Journ. Amer. Med. Assoc., lx, p. 271-274. + +JEPSON, F. P. 1909. Notes on colouring flies for purposes of + identification. Rep't to the Local Gov't Board on Publ. Health, n. s. + 16, p. 4-9. + +JOHANNSEN, O. A. 1903. Aquatic Nematocerous Diptera. N. Y. State Mus. + Bul., 68, p. 328-448, pls. 32-50. + +---- 1905. Aquatic Nematocerous Diptera II. (Chironomidæ). ibid. 86, p. + 76-330, pls. 16-37. + +---- 1908. North America Chironomidæ. ibid., 124, p. 264-285. + +---- 1911. The typhoid fly and its allies. Maine Agric. Exp. Sta. Bul., + 401, p. 1-7. + +---- 1911. Simulium and pellagra. Insect Notes for 1910. Maine Agr. + Exper. Station. Bul. 187, p. 4. + +KELLOGG, V. L. 1915. Spider poison. Jour. of Parasitology, i, p. 107+ + +KELLY, H. A. 1907. Walter Reed and yellow fever. 8vo. New York, McClure, + Phillips & Co. (xix + 310 p.). + +KEPHART, CORNELIA F. 1914. The poison glands of the larva of the + browntail moth (Euproctis chrysorrhoea Linn.). Journ. Parasit., i, + p. + +KIEFFER, J. J. 1906. Chironomidæ. Genera Insectorum. Fasc. 42, p. 1-78. + +---- 1913. Nouv. étude sur les Chironomides de l'Indien Museum de + Calcutta. Records of the Indian Mus., ix, p. 119-197. + +KING, A. F. A. 1883. Insects and disease--mosquitoes and malaria. Pop. + Sci. Mo. xxiii, p. 644-658. + +KIRKLAND, A. H. 1907. Second annual report of the Superintendent for + suppressing the gypsy and browntail moths. 8vo. Boston. 170 p. + +KLEINE, E. 1909. Postive Infektionsversuche mit _Trypanosoma brucei_ + durch _Glossina palpalis._ Deutsche med. Wochenschr., xxxv, p. + 469-470. + + Weitere wissenschaftliche Beobachtungen über die Entwicklung von + Trypanosomen in Glossinen. ibid. p. 924-925. + + Weitere Untersuchungen über die Ætiologie der Schlafkrankheit. ibid., + p. 1257-1260. + + Weitere Beobachtungen über Tsetsefliegen und Trypanosomen. ibid., p. + 1956-1958. + +KLING, C. and LEVADITI, C. 1913. Études sur la poliomyélite aiguë + épidémique. Ann. Inst. Pasteur, xxvii, p. 718-749, 739-855. + +KNAB, F. 1912. Unconsidered factors in disease-transmission by + blood-sucking insects. Journ. Econ. Ent., v, p. 196-200. + +---- 1913 a. The species of Anopheles that transmit human malaria. Amer. + Journ. Trop. Dis. and Preventive Med., i, p. 24-43. + +---- 1913 b. Anopheles and malaria. ibid., i, p. 217. + +---- 1913 c. The life history of _Dermatobia hominis_. ibid., i, p. + 464-468. + +KNAB, F. See Howard, Dyar, and Knab. + +KOBERT, R. 1893. Lehrbuch der Intoxikationen. 4^o. Stuttgart, Enke. (xxii + + 816 p.). 2d ed. in 2 vols., 1906. + +---- 1901. Beiträge zur Kenntniss der Giftspinnen. 8^o. Stuttgart, Enke. + (viii + 191 p.). + +KOLBE, H. J. 1894. Der Pfeilgiftkäfer der Kalahari-Wüste, _Diamphidia + simplex_. Stett. Ent. Zeitg., iv, p. 79-86. + +KRAUSE, M. 1907. Untersuchungen über Pfeilgifte aus unseren + africanischen Kolonien. Verhand. deutsche Kolonien kong. 1905. p. + 264-288. + +LALLIER, P. 1897. Étude sur la myase du tube digestif chez l'homme. + Thesis, Paris, 8^o. 120 p. + +LANGER, J. 1897. Ueber das Gift unserer Honigbiene. Archiv. exper. Path. + und Pharm., xxxviii, p. 381-396. + +LAVINDER, C. H. 1911. Pellagra: a précis. U. S. Publ. Health Service + Bul. 48, 37 p. + +LEIDY, J. 1847. History and anatomy of the hemipterous genus + _Belostoma_. Journ. Acad. Philad. (2), i, p. 57-67. + +LEIPER, R. T. 1907. The etiology and prophylaxis of dracontiasis. + British Med. Journ. 1907, p. 129-132. + +LEISHMAN, W. B. 1910 a. Observations on the mechanism of infection in + tick fever and on the hereditary transmission of _Spirochæta duttoni_ + in the tick. Trans. Soc. Trop. Med. Hyg., iii, p. 77-95. Abstr. in + Bul. Inst. Pasteur, viii, p. 312-313. + +---- 1910 b. On the hereditary transmission and mechanism of infection + in tick fever and on the hereditary transmission of _Spirochæta + duttoni_ in the tick. Lancet., clxxvii, p. 11. + +LINNELL, R. McC. 1914. Notes on a case of death following the sting of a + scorpion. Lancet, 1914, p. 1608-1609. + +LIVINGSTONE, D. 1857. Missionary travels and researches in South Africa. + +LUCAS, H. 1843. (note) _Latrodectus malmignatus_ Bul. Soc. Ent., France, + 1843, p. viii. + +LUDLOW, C. S. 1914. Disease bearing mosquitoes of North and Central + America, the West Indies and the Philippine Islands. War Dept., + Office of Surgeon General. Bul. No. 4, 1-96. + +LUGGER, 1896. Insects injurious in 1896. Agr. Exp. Sta. Bul. 48. p. 33 + to 270. + +MacCALLUM, W. C. 1898. On the hæmatozoan infection of birds. Journ. Exp. + Med. iii, p. 117. + +MacGREGOR, M. E. 1914. The posterior stigmata of dipterous larvæ as a + diagnostic character. Parasitology, vii, p. 176-188. + +MacLOSKIE, G. 1888. The poison apparatus of the mosquito. Amer. + Naturalist, xxii, p. 884-888. + +MALLOCH, J. P. 1913. American black-flies or Buffalo gnats. U. S. Dept. + Agric. Bur. Ent. Tech. Bul. 26, p. 1-72. + +---- 1914. Notes on North American Diptera. Bul. Illinois State Lab. + Nat. Hist., x, p. 213-243. + +MANSON, P. 1911. Tropical diseases: a manual of the diseases of warm + climates. 8^o. London, Cassell & Co. (xx + 876 p.). 4 ed. (1907). + Reprinted. + +MARCHOUX, E. and COUVY, L. 1913. Argas et spirochætes (1 mémoire). Les + granules de Leishman. Ann. Inst. Pasteur, xxvii, p. 450-480. 2 + mémoire. Le virus chez l'acarien. ibid. p. 620-643. + +MARCHOUX, E. and SELIMBENI, A. 1903. La spirillose des poules. Ann. + Inst. Pasteur, xvii, p. 569-580. + +MARCHOUX, E. and SIMOND, P. L. 1905. Études sur la fièvre jaune. Ann. + Inst. Pasteur, xx, pp. 16-40, 104-148, 161-205. + +MARLATT, C. L. 1902. (See Howard, L. O. and Marlatt, C. L.) + +---- 1907. The bed-bug (_Cimex lectularius_ L.) U. S. Dept. Agric., Bur. + Ent., Circ. No. 47, revised ed., 8 pp. + +MARTIN, G. LEBOEUF, and ROUBAUD. 1909. Rapport de la mission d'études de + la maladie du sommeil au Congo français. 4^o. Paris, Masson & Cie. + (vi + 722 p., 8 pls. and map.). + +MAVER, MARIA B. 1911. Transmission of spotted fever by other than + Montana and Idaho ticks. Journ. Infec. Dis., viii, p. 322-326. + +McCLINTIC, T. B. 1912. Investigations of and tick eradications in Rocky + Mountain spotted fever. Publ. Health Repts., Washington, xxvii, p. + 732-760. + +MECKEL, H. 1847. Uber schwarzes Pigment in der Milz und im Blute einer + Geisteskranken. Allgem. Zeitschr. f. Psychiatrie, iv, p. 198-226. + +MEGNI, P. 1906. Les insectes buveurs de sang. 12mo. Paris, Rudeval. (150 + p.). + +MELNIKOFF, N. 1869. Ueber die Jugendzustände der _Tænia cucumerina_. + Arch. f. Naturg., xxxv, p. 62-70. + +MENSE, C. 1913. Handbuch der Tropenkrankheiten. 1 Band. 4^o. Leipzig, + Barth (xv + 295 p.) Entomological parts by A. Eysell, and by Doerr + and Russ. + +MINCHIN, E. A. 1912. An introduction to the study of the Protozoa, with + special reference to the parasitic forms. 8^o. London. Arnold (xi + + 517 p.). + +MITCHELL, EVELYN G. 1907. Mosquito life. 8vo. New York, Putmans. (xxii + + 281 p.). + +MITZMAIN, M. B. 1910. General observations on the bionomics of the + rodent and human flies. U. S. Publ. Health Service. Bul., 38, p. + 1-34. + +---- 1912. The rôle of _Stomoxys calcitrans_ in the transmission of + _Trypanosoma evansi_. Philippine Journ. Sci., vii, p. 475-519, 5 pls. + +---- 1913 a. The biology of _Tabanus striatus_ Fabricius, the horsefly + of the Philippines. ibid., vii, B. p. 197-221. + +---- 1913 b. The mechanical transmission of surra. ibid., viii, sec. B., + p. 223-229. + +---- 1914 a. Experimental insect transmission of anthrax. U. S. Public + Health Repts. xxix, p. 75-77. + +---- 1914 b. I. Collected studies on the insect transmission of + _Trypanosoma evansi_. II. Summary of experiments in the transmission + of anthrax by biting flies. U. S. Pub. Health Service, Hyg. Lab. + Bul., 94, p. 1-48. + +MIYAKE, H. and SCRIBA, J. 1893. Vorläufige Mitteilung über einen neuen + Parasit des Menschen. Berl. klin. Wochenschr., xxx, p. 374. + +MOLLERS, B. 1907. Experimentelle Studien über die Uebertragung des + Rückfallfiebers durch Zecken. Zeitschr. für Hyg. u. + Infektionskrankheiten, lviii, p. 277-286. + +MOTE, D. C. 1914. The cheese-skipper (_Piophila casei_). Ohio Naturalist + xiv, p. 309-310. + +NEIVA, A. 1910. Beiträge zur Biologie der _Conorhinus megistus_ Burm. + Memorias de Institute Oswaldo Cruz., ii, p. 206-212. + +NEVEU-LEMAIRE, M. 1907. Un nouveau cas de parasitisme accidental d'un + myriapode dans le tube digestif de l'homme, C. R. Soc. der Biol., + lxiii p. 305-308. + +---- 1908. Précis de parasitologie humaine. 8vo. Paris, Rudeval. (v + + 712 p.). + +NEWSTEAD, R. 1911. The papataci flies (Phlebotomus) of the Maltese + Islands. Bul. of Ent. Research, ii, p. 47-78, pls. 1-3. + +NICOLL, W. 1911. On the part played by flies in the disposal of the eggs + of parasitic worms. Repts. to the Local Gov't. Board on Publ. Health + and Med. Subjects, n. s. No. 53, p. 13-30. + +NICOLLE, C. 1910, Recherches expérimentales sur la typhus exanthématique + entreprises à l'Institut Pasteur de Tunis pendant l'année 1909. Ann. + Inst. Pasteur, xxiv, p. 243-275. + +---- 1911. Recherches expérimentales sur la typhus exanthématique + entreprises à l'Institut Pasteur de Tunis pendant l'année 1910. + ibid., xxv, p. 1-55, 97-154. + +NICOLLE, C., BLAIZOT, A., and CONSEIL, E. 1912 a. Étiologie de la fièvre + récurrente. Son mode de transmission par le pou. C. R. Acad. Sci., + cliv, p. 1636-1638. + +---- 1912 b. 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Assoc. iii, p. 373-380. + +---- Spotted fever reports 1 and 2. In the 4th Bien. Rept. State Board + of Health, Montana, 1909, p. 87-191. + +RICKETTS, H. T. and WILDER, R. M. 1910. The transmission of the typhus + fever of Mexico (tabardillo) by means of the louse, _Pediculus + vestimenti_. Journ. Am. Med. Assoc. liv. p. 1304. + +RILEY, C. V. and HOWARD, L. O. 1889. A contribution to the literature of + fatal spider-bites. Insect life, Washington, i. p. 204-211. + +RILEY, W. A. 1906. A case of pseudoparasitism by dipterous larvæ. + Canadian Ent. xxxviii, p. 413. + +---- 1910 a. Earlier references to the relation of flies to disease. + Science n. s. xxxi, p. 263-4. + +---- 1910 b. _Dipylidium caninum_ in an American child. Science n. s. + xxxi, p. 349-350. + +---- 1911. The relation of insects to disease. Cornell Countryman ix, P. + 51-55. + +---- 1912 a. Notes on the relation of insects to disease. 8vo. Ithaca, + N. Y. 51 p. + +---- 1912 b. Notes on animal parasites and parasitism. 8vo. Ithaca, N. + Y. 55 p. + +---- 1913. Some sources of laboratory material for work on the relations + of insects to disease. Ent. News. xxiv, p. 172-175. + +---- 1914. Mr. Nott's theory of insect causation of disease. Jour. of + Parasitology. i, p. 37-39. + +ROSENAU, M. J. and BRUES, C. T. 1912. Some experimental observations on + monkeys, concerning the transmission of poliomyelitis through the + agency of _Stomoxys calcitrans_. Monthly Bul. Mass. State Board of + Health. Vol. vii, p. 314-317. + +ROSS, R. 1904. Researches on Malaria. The Nobel Medical Prize Lecture + for 1902, Stockholm, Norstedt & Söner. 89 p. 9 pls. In "Les Prix + Nobel en 1902." + +---- 1910. The prevention of malaria. 4^o. New York. Dutton & Co. (xx + + 669 p.). + +ROTHSCHILD, N. C. 1905 a. North American Ceratophyllus. Novitates + Zoologicæ xii, p. 153-174. + +---- 1905 b. Some further notes on _Pulex canis_ and _P. felis_. ibid. + xii, p. 192-193. + +ROUBAUD, E. 1911. Les Choeromyies. C. R. Acad. Sci. Paris, cliii, p. + 553. + +---- 1913. Recherches sur les Auchméromyies. Bul. Sci. France et Belg. + Paris, p. 105-202. + +SACHS, HANS. 1902. Zur Kentniss der Kreuzspinnengiftes. Beitr. Chem. + hysiol. ii, p. 125-133. Abstr. Centralbl. Bakter. 1. Abth. xxxi., + Referate, p. 788. + +SAMBON, L. W. 1908. Report presented at the International Conference on + Sleeping Sickness. + +---- 1910. Progress report on the investigation of pellagra. Reprinted + from the Journ. Trop. Med. and Hyg.; London; Bale, Sons and + Danielsson, 12^o. 125 p. + +SANDERSON, E. D. 1910. Controlling the black-fly in the White Mountains. + Jour. Econ. Ent. iii, p. 27-29. + +SAUL, E. 1910. Untersuchungen über Beziehungen der Acari zur + Geschwulstätiologie. Centralbl. Bakt. Jena, Abt. 1, lv, Originale, p. + 15-18. + +SAUL, E. 1913. 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Egyptian Gov't + School of Medicine, Cairo, ii, p. 7-44. + + + + +INDEX + + +Abscess, 178 + +Acanthia, 87 + +Acariasis, 58 + +Acarina, 23, 58, 131, 259 + +Acarus dysenteriæ, 132 + +Accidental parasites, 131, 132, 134 + +Aedes, 194, 293 + +Aedes calopus, 182, 201, 205, 206, 208 + +Aedes cantator, 101 + +Aedes sollicitans, 101 + +Aedes tæniorhynchus, 101 + +Aerobic bacteria, 152 + +Æstivo-autumnal, 186 + +African Relapsing Fever, 230 + +Akis spinosa, 177 + +Alternation of Generations, 175 + +Amblyomma, 264 + +Amblyomma americanum, 67 + +Amblyomma cajennense, 67 + +American dog tick, 228 + +AmÅ“boid organism, 189 + +Anisolabis annulipes, 177 + +Anterior poliomyelitis, 241 + +Anopheles, 194, 291 + +Anopheles crucians, 199 + +Anopheles maculipennis, 182 + +Anopheles punctipennis, 198 + +Anopheles quadrimaculatus, 197 + +Anopheline, 192 + +Anthocoris, 279 + +Anthomyiidæ, 300 + +Anthomyia, 138 + +Anthrax, 165 + +Antipruritic treatment, 72 + +Ants, 42 + +Aphiochæta, 295 + +Apis mellifica, 36 + +Arachnida, 258 + +Araneida, 6 + +Argas, 64 + +Argas persicus, 63, 235, 237 + +Argasidæ, 62 + +Argopsylla, 317 + +Argus, 259 + +Arilus, 284 + +Arthropods, poisonous, 6 + +Asopia farinalis, 177 + +Assassin-bugs, 31, 219 + +Auchmeromyia, 117 + +Automeris io, 47 + +Avicularoidea, 12 + + +Babesia, 226 + +Babesia bovis, 223 + +Babesia ovis, 225 + +Babesiosis, 221-222 + +Bacilli, 170 + +Bacillus icteroides, 202, 205 + +Bacillus pestis, 166 + +Bacillus typhosus, 153 + +Back swimmers, 30 + +Bdellolarynx, 304 + +Beauperthuy, Louis Daniel, 2 + +Bed-bug, 86, 88, 90, 173, 219-220 + +Bed-bug, cone-nosed, 92 + +Blister beetles, 54 + +Belostoma, 28, 277 + +Belostoma americana, 31 + +Belostomatidæ, 30 + +Bengalia, 314 + +Bird-spiders, 10 + +Black death, 1, 166 + +Black flies, 33, 104, 247 + +Black heads, 80 + +Blaps mortisaga, 134 + +Blepharoceridæ, 286 + +Boophilus, 264 + +Boophilus annulatus, 67, 223-225 + +Bot-flies, 112 + +Blue bottle flies, 140 + +Brill's disease, 238 + +Brown-tailed moth, 48 + +Bruck, 34 + +Buthus quinquestriatus, 21 + + +Cabbage butterfly, 56 + +Calliphora, 136, 140, 312 + +Calliphora erythrocephala, 141 + +Calobata, 296 + +Camponotinæ, 43 + +Cancer, 254 + +Cantharidin, 54 + +Cantharidin poison, 55 + +Canthariasis, 134 + +Capsidæ, 280 + +Carriers, simple, 4, 144 + +Carriers of disease, 144 + +Carrion's fever, 253 + +Caterpillar rash, 45 + +Cat flea, 172 + +Cattle ticks, 222 + +Causative organism, 170 + +Cellia, 291 + +Centipedes, 25, 257 + +Ceratophyllus, 120, 316 + +Ceratophyllus acutus, 123 + +Ceratophyllus fasciatus, 122, 172, 213 + +Ceratopogon, 108 + +Cheese-fly, 137 + +Cheyletus eruditus, 271 + +Chigger, 60, 70 + +Chigoes, 126 + +Chilopoda, 25, 257 + +Chiracanthium nutrix, 18 + +Chironomidæ, 107 + +Chorioptes, 270 + +Chrysomelid, 55 + +Chrysomyia, 136, 308 + +Chrysomyia macellaria, 117, 140 + +Chrysops, 294 + +Chylous dropsy, 179 + +Chyluria, 178 + +Cicadidæ, 55 + +Cimex L., 278 + +Cimex boueti, 92 + +Cimex columbarius, 92 + +Cimex hemipterus, 91, 220 + +Cimex hirundinis, 92 + +Cimex inodorus, 92 + +Cimex lectularius, 87, 219 + +Citheronia regalis, 44 + +Clinocoris, 87 + +Coleoptera, 134, 274 + +Comedons, 80 + +Complete metamorphosis, 80 + +Compressor muscle, 20 + +Compsomyia, 117 + +Cone-nosed bed-bug, 92 + +Conjunctivitis, nodular, 52 + +Conorhinus, 282 + +Conorhinus megistus, 93, 219-220 + +Conorhinus rubrofasciatus, 220 + +Conorhinus sanguisugus, 32, 92 + +Copra itch, 72 + +Cordylobia, 118 + +Coriscus, 280 + +Coriscus subcoleoptratus, 32 + +Creeping myasis, 112 + +Crustacea, 257 + +Cryptocystis, 176 + +Cryptotoxic, 54-55 + +Cteniza sauvagei, 13 + +Ctenocephalus, 120, 172, 213, 317 + +Culex, 194, 201, 293 + +Culex pipiens, 35, 98 + +Culex quinquefasciatus, 180 + +Culex sollicitans, 200 + +Culex territans, 101 + +Culicidæ, 33, 97 + +Culicin, 34 + +Culicoides, 109, 288 + +Cyclops, 183, 257 + +Cynomyia, 136, 311 + + +Dance, St. Vitus, 8 + +Dancing mania, 8 + +Deer-flies, 110 + +Definitive host, 192 + +Demodecidæ, 78 + +Demodex, 259 + +Demodex folliculorum, 78 + +Dermacentor, 262 + +Dermacentor andersoni, 67, 228 + +Dermacentor occidentalis, 227 + +Dermacentor variabilis, 67 + +Dermacentor venustus, 24, 228 + +Dermanyssidæ, 68 + +Dermanyssus, 266 + +Dermanyssus gallinæ, 68 + +Dermatitis, 72, 77, 85 + +Dermatobia, 115, 298 + +Dermatobia cyaniventris, 163 + +Dermatophilus, 317 + +Dermatophilus penetrans, 60, 126 + +Diamphidia simplex, 55 + +Dimorphism, 65 + +Direct inoculators, 4 + +Diplopoda, 25, 257 + +Diptera, 33, 94, 274 + +Dipterous Larvæ, 135 + +Dipylidium, 175, 221 + +Dipylidium caninum, 4, 175-176 + +Dog flea, 172 + +Dracunculus, 257 + +Dracunculus medinensis, 182 + +Drosophila, 296 + +Dum-dum fever, 220 + +Dysentery, 154 + + +Ear-flies, 110 + +Earwig, 177 + +Echidnophaga, 317 + +Echinorhynchus, 185 + +Elephantiasis, 178-179 + +Empoasca mali, 33 + +Empretia, 46 + +English Plague Commission, 171 + +Epeira diadema, 18 + +Epizootic, 170 + +Eristalis, 137, 295 + +Essential hosts, 4, 165 + +Eumusca, 307 + +European Relapsing Fever, 233 + +Euproctis chrysorrhÅ“a, 48 + +Eusimulium, 286 + + +Facultative parasites, 131 + +Fannia, 136, 138, 145, 300 + +Federal Health Service, 169 + +Fever, lenticular, 237 + African Relapsing, 230, 234 + Carrion's, 253 + dum-dum, 154 + European Relapsing, 233 + pappatici, 96 + red water, 220 + Rocky Mt. Spotted, 226 + three day, 96 + Typhus, 237 + +Filaria, 178, 221 + immitis, 182 + +Filariasis, 178 + +Flannel-moth larvæ, 44 + +Fleas, 119, 166, 213 + cat, 172 + dog, 172 + human, 172, 176 + rodent, 123, 172 + rat, 171 + +Flesope, 125 + +Formaldehyde, 91 + +Fomites, 199, 204 + +Fulgoridæ, 28 + +Fumigation, 320 + + +Gamasid, 68 + +Gangrene, 129 + +Gastrophilus, 113, 297 + +Giant crab spiders, 13 + +Giant water bugs, 30 + +Gigantorhynchus, 185 + +Glossina, 117, 297, 303 + +Glossina morsitans, 214, 217 + palpalis, 215, 217, 218 + +Glyciphagus, 267 + +Grain moth, 69 + +Grocer's itch, 72 + +Guinea-worm, 182 + + +Habronema muscæ, 156, 183 + +Hæmatobia, 166, 304 + irritans, 146 + +Hæmatobosca, 304 + +Hæmatomyidium, 288 + +Hæmatopinus spinulosus, 213 + +Hæmatopota, 294 + +Hæmatosiphon, 279 + +Hæmoglobinuria, 222 + +Hæmozoin, 189 + +Harpactor, 284 + +Harvest mites, 60 + effect of, 59 + +Head-louse, 173 + +Helminthiasis, 138 + +Helophilus, 295 + +Hemiptera, 27, 86, 273-275 + +Heteropodidæ, 13 + +Heuchis sanguinea, 55 + +Hexapod larvæ, 58 + +Hexapoda, 27, 80, 258 + +Hippelates, 297 + +Hippobosca, 285 + +Histiogaster, 269 + spermaticus, 132 + +Homalomyia, 136, 138, 300 + +Honey bee, 36 + poison of, 37 + +Hornets, 43 + +Horn-fly, 137, 304, 308 + +Horse-fly, 110, 165 + +House-fly, 137-139, 144, 183 + control of, 156, 160 + +Human flea, 124 + +Host, definitive, 175 + intermediate, 175 + primary, 175 + +Hyalomma, 264 + ægypticum, 224-225 + +Hydrocyanic Acid Gas, 318 + +Hydrotæa, 300 + +Hymenolepis diminuta, 176 + +Hymenoptera, 36, 275 + +Hypoderma, 113, 298 + diana, 113 + lineata, 113 + +Hypopharynx, 80 + + +Immunity from stings, 39 + +Incomplete metamorphosis, 80 + +Infantile paralysis, 162, 241 + splenic, 220 + Direct inoculation, 164 + +Insects, 258 + blood-sucking, 170 + +Intermediate host, 192, 203 + +Intestinal infestation, 112, 133 + myasis, 137 + +Isosoma, 69 + +Itch, 73-74 + mite, 73 + Norwegian, 77 + +Ixodes, 260 + ricinus, 66, 225 + scapularis, 66 + +Ixodidæ, 64-65 + +Ixodoidea, 62 + + +Janthinosoma lutzi, 116 + +Jigger, 60 + +Johannseniella, 110, 288 + +Journal of Tropical Medicine and Hygiene, 36 + +Julus terrestris, 25 + +June bug, 185 + + +Kala-azar, 220 + +Karakurte, 14 + +Katipo, 14 + +King, A. F. A., 3 + +Kircher, Athanasius, 1, 8 + +Kissing-bug, 31 + + +Labium, 29, 80 + +Labrum, 28, 80 + +Lachnosterna, 185 + +Lælaps, 266 + +Lagoa crispata, 45 + +Lamblia intestinalis, 154 + +Langer, Josef, 37 + +Larder beetles, 135 + +Latrodectus, 12, 14, 17 + mactans, 15 + +Leishmanioses, 220 + +Lenticular fever, 237 + +Lepidoptera, 274 + +Lepidopterous larvæ, 134 + +Leprosy, 252 + +Leptidæ, 112 + +Leptis, 295 + +Leptus, 60, 273 + +Lice, 80 + +Linguatulina, 258 + +Liponyssus, 265 + +LÅ“mopsylla, 172, 317 + +Lone star tick, 228 + +Louse, body, 84 + crab, 85 + dog, 176 + head, 82 + pubic, 85 + +Lucilia, 136, 312 + +Lycosa tarantula, 10 + +Lycosidæ, 10 + +Lyctocoris, 279 + +Lygus pratensis, 33 + +Lymphangitis, 67 + +Lymph scrotum, 178 + +Lyperosia, 304 + +Lyperosiops, 305 + + +Macloskie, 34 + +Maggots, rat-tail, 137 + +Magnes sive de Arte Magnetica, 8 + +Malaria, 186 + +Malmigniatte, 14 + +Mandibles, 28, 80 + +Mange, 73-75 + +Margaropus, 237, 264 + annulatus, 223 + +Masked bed-bug hunter, 32 + +Mastigoproctus giganteus, 19, 80 + +Maxillæ, 28 + +Meal infesting species, 135 + +Melanin granules, 189 + +Melanolestes, 280 + picipes, 32 + +Mena-vodi, 14 + +Mercurialis, 1 + +Merozoites, 190 + +Metamorphosis, 80 + +Miana bug, 63 + +Microgametoblast, 192 + +Midges, 107 + +Migratory ookinete, 192 + +Millipedes, 25, 257 + +Mites, 23, 58 + +Monieziella, 269 + +Mosquitoes, 33, 97, 178, 196, 250 + treatment for bites of, 34, 36, 102 + +Musca, 137, 307 + domestica, 139, 145, 146, 157, 162 + +Muscidæ, 117 + +Muscina, 137, 146, 307 + stabulans, 140 + +Mutualism, 57 + +Myasis, 112, 135 + intestinal, 135-140 + nasal, 141 + +Mycterotypus, 287 + +Myiospila, 146, 307 + +Myriapoda, 225, 132, 257 + + +Nagana, 165, 214 + +Nasal infestation, 114, 133 + +Necrobia, 135 + +Nematode parasite, 182 + +Nepa, 28 + +Nephrophages sanguinarius, 132 + +Nettling insects, 43 + larvæ, poison of, 53 + +Neurasthenia, 89 + +Nits, 86 + +North African Relapsing Fever, 234 + +Norwegian itch, 77 + +No-see-ums, 109 + +NotÅ“dres, 269 + cati, 78 + +Notonecta, 28, 277 + +Notonectidæ, 30 + +Nott, Dr. Josiah, 2 + +Nuttall, 34 + + +Occipital headaches, 138 + +Oecacta, 288 + +Oeciacus, 279 + +Å’sophageal diverticula, 35 + +Oestridæ, 112, 136 + +Oestris ovis, 113 + +Oestrus, 298 + oocyst, 192 + ookinete, 192 + +Opsicoetes personatus, 32 + +Opthalmia, 155 + nodosa, 52 + +Oriental sore, 221 + +Ornithodoros, 65, 260 + moubata, 220, 230 + +Orthotylus flavosparsus, 33 + +Ornithomyia, 286 + +Oroya, 253 + +Oscinus, 297 + +Otiobius, 259 + megnini, 65 + +Otodectes, 271 + + +Pangonia, 294 + +Pappatici fever, 96 + +Parasimulium, 286 + +Parasite, 3, 57, 131, 134, 182 + accidental, 3, 131, 134 + facultative, 3, 57, 131 + nematode, 182 + stationary, 57 + temporary, 57 + true, 3 + +Parasitism, accidental, 134 + +Pathogenic bacteria, 152 + organisms, 144, 164 + +Pawlowsky, 81 + +Pediculoides, 267 + ventricosus, 69, 72 + +Pediculosis, 81 + +Pediculus, 275 + corporis, 84, 233, 238 + humanus, 82, 173 + +Pellagra, 162, 246 + +Pernicious fever, 186 + +Pest, 166 + +Phidippus audax, 19 + +Philæmatomyia, 306 + +Phisalix, 13, 43 + +Phlebotomus, 289 + papatasii, 94 + verrucarum, 254 + vexator, 95 + +Phora, 295 + +Phormia, 136 + +Phormictopus carcerides, 13 + +Phthirus pubis, 85, 275 + +Phortica, 296 + +Pieris brassicæ, 56 + +Piophila, 297 + +Piophila casei, 136, 137 + +Piroplasmosis, 222 + +Plague, 166 + bubonic, 166, 169, 170 + pneumonic, 167 + +Plasmodium, 186 + +Platymetopius acutus, 33 + +Plica palonica, 83 + +Pneumonic, 166 + plague, 167, 173 + +Poisoning by nettling larvæ, 53 + +Poison of spiders, 7 + +Pollenia, 308 + rudis, 146, 147 + +Primary gland, 28 + +Prionurus citrinus, 20 + +Prosimulium, 286 + +Protocalliphora, 136, 312 + +Protozoan blood parasite, 165 + +Pseudo-tubercular, 52 + +Psorophora, 293 + +Psoroptes, 270 + +Psychodidæ, 94 + +Pulex, 120, 124, 126, 172, 317 + cheopis, 172 + irritans, 124 + penetrans, 126 + serraticeps, 120 + +Pulvillus, 150 + +Punkies, 109 + +Pycnosoma, 308 + + +Rasahus, 280 + thoracicus, 32 + +Rat fleas, 120, 124, 171 + +Rat louse, 213 + +Red bugs, 70-72 + +Reduviidæ, 31 + +Reduviolus, 280 + +Reduvius, 282 + personatus, 32 + +Redwater fever, 222 + +Relapsing fever, 230, 233 + +RhinÅ“strus nasalis, 115 + +Rhipicentor, 264 + +Rhipicephalus, 264 + +Rhizoglyphus, 269 + +Rhodnius, 281 + +Rocky Mountain Spotted Fever, 226 + spotted fever tick, 67 + +Russian gad-fly, 115 + + +St. Vitus's or St. John's dance, 8 + +Salivary syringe, 28 + +Sand-flies, 109, 250 + +Sanguinetti, 11 + +Sarcophaga, 136, 142, 143 + +Sarcophila, 302 + +Sarcopsylla, 317 + penetrans, 126 + +Sarcoptes, 270 + minor, 78 + scabiei, 73 + +Sarcoptidæ, 72 + +Scabies, 72, 73, 74, 75 + +Scaurus striatus, 177 + +Schaudinn, 34 + +Schizont, 189, 190 + +Scholeciasis, 134 + +Scolopendra morsitans, 26 + +Scorpions, 20 + poison of, 21 + +Screw worm fly, 140 + +Sepsidæ, 296 + +Sepsis, 136, 297 + +Shipley, 34 + +Sibine, 46 + +Silvius, 294 + +Simple carriers, 4, 144 + +Simuliidæ, 33, 104 + +Simulium, 247, 249, 286, 321 + pictipes, 104 + +Siphonaptera, 119, 274, 316 + +Siphunculata, 80, 275 + +Sitotroga cerealella, 69 + +Skippers, 137 + +Sleeping sickness, 166, 215 + +Snipe-flies, 112 + +Solpugida, 22 + +Spanish fly, 54 + +Spermatozoa, 192 + +Spinose ear-tick, 65 + +Spirochæta, 35 + berberi, 234 + duttoni, 234 + +Spirochætosis, 235 + +Sporozoite, 189 + +Spotted fever, 67, 226 + +Squirrel flea, 123 + +Stable-fly, 137, 160, 163, 165 + +Stegomyia, 182, 293 + calopus, 206 + fasciata, 206 + +Stomoxys, 137, 305 + calcitrans, 117, 146, 160, 161, 165, 242 + +Straw-worm, 69 + +Stygeromyia, 305 + +Sucking stomach, 35 + +Sulphur ointment, 77 + +Surra, 165 + +Symbiosis, 57 + +Symphoromyia, 112, 295 + + +Tabanidæ, 110 + +Tabanus, 110, 166, 294 + striatus, 165 + +Taenia, 175 + +Tapeworm, 4, 176 + +Tarantella, 8 + +Tarantism, 8 + +Tarantula, 10 + +Tarsonemidæ, 69 + +Tarsonemus, 267 + +Tenebrionid beetles, 127 + +Tersesthes, 110, 288 + +Tetanus, 129 + +Tetranychus, 273 + +Texas fever, 220-223 + +Three-day fever, 96 + +Tick, 23, 226 + bites, Treatment of, 68 + fever, 230 + paralysis, 67 + +Treatment, + Bee stings, 36, 41 + Bites of, + Bed-bugs, 90, 93 + Blackflies, 107 + Buffalo flies, 107 + Bugs, 31, 33 + Centipedes, 26, 27 + Chiggers, 127 + Chigoes, 127 + Fleas, 127 + Harvest mites, 61 + Jiggers, 129 + Lice, 83, 85 + Mosquitoes, 34, 36, 102 + Phlebotomus flies, 97 + Sand flies, 96, 107, 109 + Scorpions, 22, 23 + Spiders, 19 + Ticks, 61, 68, 72 + Ticks, ear, 65 + Blister beetle poison, 55 + Brown-tail moth rash, 45 + Cantharidin poison, 55 + Caterpillar rash, 45 + Ear ticks, 65 + House fly control, 156, 160 + Itch, 77 + Itch, grocer's, 72 + Lice, 85 + Nasal myasis, 143 + Rocky Mt. spotted fever, 228, 229 + Rash, caterpillar, 45 + Scabies, 77 + Sleeping sickness control, 218 + Spotted fever, 228, 229 + Stings, bee, 36, 41 + Typhus fever, prophylaxis, 239 + +Trichodectes canis, 176 + +Trichoma, 82 + +Trineura, 295 + +Trochosa singoriensis, 11 + +Trombidium, 60, 273 + +True insects, 80 + +Trypanosoma, 35 + +Trypanosoma, brucei, 165 + +Trypanosoma cruzi, 219 + +Trypanosoma lewisi, 213 + +Trypanosomiases, 212 + +Trypanosomiasis, 165, 219 + +Tsetse flies, 117, 166, 214, 219 + +Tsetse flies disease, 165 + +Tuberculosis, 155 + +Tumbu-fly, 118 + +Tydeus, 271 + +Typhoid, 155 + +Typhoid fever, 154 + +Typhus, 237 + +Typhus fever, 237 + +Tyroglyphus, 72, 131, 268 + +Dr. Tyzzer, 49 + + +Uranotænia, 292 + + +Vancoho, 14 + +Varicose groin glands, 178 + +Verruga peruviana, 253 + +Vescicating insects, 54 + + +Wanzenspritze, 29 + +Warble-flies, 112 + +Wasps, 43 + +Whip-scorpions, 19 + +Wohlfahrtia, 143, 302 + +Wolf-spiders, 10 + +Wyeomyia smithii, 101, 293 + + +Xenopsylla, 172, 317 + +Xenopsylla cheopis, 171, 124 + +Xestopsylla, 317 + + +Yaws, 2 + +Yellow fever, 196, 203, 205 + + + + +TRANSCRIBER'S NOTES + +The following discrepancies in the text are as in the original: + + The inconsistent hyphenation of the following: + assassin-bugs/assassin bugs; + bedbug/bed-bug (and bedbugs); + beekeeper/bee-keeper (and beekeepers); + blackflies/black-flies; + blow-flies/blow flies; + bluebottles/blue-bottles; + bot-flies/bot flies; + bristlelike/bristle-like; + browntail/brown-tail; + coextensive/co-extensive; + deer-flies/deer flies; + dorsocentral/dorso-central; + ectoparasites/ecto-parasites; + endoparasites/endo-parasites; + flesh-fly/flesh fly (and flesh flies); + hotbed/hot-bed; + housefly/house-fly (and houseflies); + horsefly/horse-fly (and horse flies); + horse-manure/horse manure; + midsummer/mid-summer; + preeminently/pre-eminently; + sandfly/sand-fly (and sandflies); + screw-worm fly/screw worm fly; + stable-fly/stable fly; + subequal/sub-equal; + subfamily/sub-family; + subtropical/sub-tropical; + tapeworm/tape-worm (and tapeworms); + today/to-day; + tsetse-flies/tsetse flies; + widespread/wide-spread; + wormlike/worm-like. + + Inconsistent use of diaeresis in Aëdes/Aedes + + Inconsistent spelling of the following: + defence/defense; + disc/disk; + hemolysis/hæmolysis; + hemolytic/hæmolytic; + hexapod/hexopod; + Levaditi/Lavaditi; + metalescent/metallescent; + Mitzmain/Mitzman; + Neveau-Lemaire/Neveau-Lamaire; + offence/offense; + Phthirus/Phthirius + Portschinsky/Portchinsky/Portchinski; + travelled/traveled; + ventra-/ventro-; + Villot/Villet; + Wohlfartia/Wohlfahrtia; + + Inconsistent use of ligatures in Taenia/Tænia + +toxine, insiduous, efficaceous, cyanid are spelt as in the original + +In the first paragraph of chapter XII, "the student might not be lead" +is as in the original. Lead perhaps should be led + +There is no Figure 147 in the original + +In the bibliography, the entry for Vaughan without a text specified is +as in the original + + + + + +End of the Project Gutenberg EBook of Handbook of Medical Entomology, by +William Albert Riley and Oskar Augustus Johanssen + +*** END OF THIS PROJECT GUTENBERG EBOOK HANDBOOK OF MEDICAL ENTOMOLOGY *** + +***** This file should be named 34279-0.txt or 34279-0.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/4/2/7/34279/ + +Produced by Bryan Ness, Iris Schimandle, Brownfox and the +Online Distributed Proofreading Team at http://www.pgdp.net +(This file was produced from images generously made +available by The Internet Archive/American Libraries.) + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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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: Handbook of Medical Entomology + +Author: William Albert Riley + Oskar Augustus Johanssen + +Release Date: November 11, 2010 [EBook #34279] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK HANDBOOK OF MEDICAL ENTOMOLOGY *** + + + + +Produced by Bryan Ness, Iris Schimandle, Brownfox and the +Online Distributed Proofreading Team at http://www.pgdp.net +(This file was produced from images generously made +available by The Internet Archive/American Libraries.) + + + + + +Transcriber's Note: Barring a few obvious typos, the text has been left +as printed. + + + + +[Illustration: Some early medical entomology. Athanasius Kircher's +illustration of the Italian tarantula and the music prescribed as an +antidote for the poison of its bite. (1643).] + + + + +HANDBOOK OF MEDICAL + +ENTOMOLOGY + +WM. A. RILEY, PH.D. + +Professor of Insect Morphology and Parasitology, Cornell University + +and + +O. A. JOHANNSEN, PH.D. + +Professor of Biology, Cornell University + +[Illustration] + +ITHACA, NEW YORK + +THE COMSTOCK PUBLISHING COMPANY + +1915 + + +COPYRIGHT, 1915 + +BY THE COMSTOCK PUBLISHING COMPANY, + +ITHACA, N. Y. + + +Press of W. F. Humphrey +Geneva, N. Y. + + + + +PREFACE + + +The Handbook of Medical Entomology is the outgrowth of a course of +lectures along the lines of insect transmission and dissemination of +diseases of man given by the senior author in the Department of +Entomology of Cornell University during the past six years. More +specifically it is an illustrated revision and elaboration of his "Notes +on the Relation of Insects to Disease" published January, 1912. + +Its object is to afford a general survey of the field, and primarily to +put the student of medicine and entomology in touch with the discoveries +and theories which underlie some of the most important modern work in +preventive medicine. At the same time the older phases of the +subject--the consideration of poisonous and parasitic forms--have not +been ignored. + +Considering the rapid shifts in viewpoint, and the development of the +subject within recent years, the authors do not indulge in any hopes +that the present text will exactly meet the needs of every one +specializing in the field,--still less do they regard it as complete or +final. The fact that the enormous literature of isolated articles is to +be found principally in foreign periodicals and is therefore difficult +of access to many American workers, has led the authors to hope that a +summary of the important advances, in the form of a reference book may +not prove unwelcome to physicians, sanitarians and working +entomologists, and to teachers as a text supplementing lecture work in +the subject. + +Lengthy as is the bibliography, it covers but a very small fraction of +the important contributions to the subject. It will serve only to put +those interested in touch with original sources and to open up the +field. Of the more general works, special acknowledgment should be made +to those of Banks, Brumpt, Castellani and Chalmers, Comstock, Hewitt, +Howard, Manson, Mense, Neveau-Lemaire, Nuttall, and Stiles. + +To the many who have aided the authors in the years past, by suggestions +and by sending specimens and other materials, sincerest thanks is +tendered. This is especially due to their colleagues in the Department +of Entomology of Cornell University, and to Professor Charles W. Howard, +Dr. John Uri Lloyd, Mr. A. H. Ritchie, Dr. I. M. Unger, and Dr. Luzerne +Coville. + +They wish to express indebtedness to the authors and publishers who have +so willingly given permission to use certain illustrations. Especially +is this acknowledgment due to Professor John Henry Comstock, Dr. L. O. +Howard, Dr. Graham-Smith, and Professor G. H. T. Nuttall. Professor +Comstock not only authorized the use of departmental negatives by the +late Professor M. V. Slingerland (credited as M. V. S.), but generously +put at their disposal the illustrations from the MANUAL FOR THE STUDY OF +INSECTS and from the SPIDER BOOK. Figures 5 and 111 are from Peter's +"Der Arzt und die Heilkunst in der deutschen Vergangenheit." It should +be noted that on examining the original, it is found that Gottfried's +figure relates to an event antedating the typical epidemic of dancing +mania. + + WM. A. RILEY. + O. A. JOHANNSEN. + + CORNELL UNIVERSITY, + January, 1915. + + + + +ADDITIONS AND CORRECTIONS + +vi line 11, for Heilkunft read Heilkunst. + +18 line 2, for tarsi read tarsus. + +32 line 21, and legend under fig. 23, for C. (Conorhinus) abdominalis + read Melanolestes abdominalis. + +47 legend under figure for 33c read 34. + +92 line 22 and 25, for sangiusugus read sanguisugus. + +116 legend under fig. 83, for Graham-Smith read Manson. + +136 line 10, from bottom, insert "ring" after "chitin". + +137 line 3, for meditatunda read meditabunda. + +145 line 7, from bottom, for Rs read R_5. + +158 line 20, for have read has. + +212 after the chapter heading insert "continued". + +219 line 10, from bottom, for Cornohinus read Conorhinus. + +266 line 1, fig. 158j refers to the female. + +272 line 5, insert "palpus" before "and leg". + +281 line 6, for discodial read discoidal. + +281 last line, insert "from" before "the". + +284 line 5, for "tubercle of" read "tubercle or". + +305 lines 19, 28, 44, page 306 lines 1, 9, 22, 27, 30, page 307 line 7, + page 309 lines 8, 11, for R_{4+5} read M_{1+2}. + +309 legend under fig. 168 add Bureau of Entomology. + +312 line 36, for "near apex" read "of M_{1+2}". + +313 running head, for Muscidæ read Muscoidea. + +314 line 29, for "distal section" read "distally M_{1+2}". + +315 legend under fig. 172, for Pseudopyrellia read Orthellia, for + Lyperosia read Hæmatobia, for Umbana read urbana. + +323 and 325 legends under the figures, add "After Dr. J. H. Stokes". + +328 line 7 from bottom for Apiochæta read Aphiochæta. + + + + +CONTENTS + + +CHAPTER I + +INTRODUCTION 1-5 + + Early suggestions regarding the transmission of disease by + insects. + The ways in which arthropods may affect the health of man. + + +CHAPTER II + +ARTHROPODS WHICH ARE DIRECTLY POISONOUS 6-56 + + The Araneida, or Spiders. + The tarantulas. Bird spiders. Spiders of the genus + Latrodectus. Other venomous spiders. Summary. + The Pedipalpida, or whip-scorpions. + The Scorpionida, or true scorpions. + The Solpugida, or solpugids. + The Acarina, or mites and ticks. + The Myriapoda, or centipedes and millipedes. + The Hexapoda, or true insects. + Piercing or biting insects poisonous to man. + Hemiptera, or true bugs. + The Notonectidæ or back-swimmers. Belostomidæ or giant + water-bugs. Reduviidæ, or assassin bugs. Other + Hemiptera reported as poisonous to man. + Diptera; the midges, mosquitoes and flies. + Stinging insects. + Apis mellifica, the honey bee. Other stinging forms. + Nettling insects. + Lepidoptera, or butterflies and moths. Relief from + poisoning by nettling larvæ. + Vescicating insects and those possessing other poisons + in their blood plasma. The blister beetles. Other + cryptotoxic insects. + + +CHAPTER III + +PARASITIC ARTHROPODS AFFECTING MAN 57-130 + + Acarina, or mites. + The Trombidiidæ, or harvest mites. + The Ixodoidea, or ticks. + Argasidæ. Ixodidæ. Treatment of tick bites. + The mites. + Dermanyssidæ. Tarsonemidæ. Sarcoptidæ, the itch mites. + Demodecidæ, the follicle mites. + Hexapoda, or true insects. + Siphunculata, or sucking lice. + Hemiptera. + The bed-bug. Other bed-bugs. + Parasitic Diptera, or flies. + Psychodidæ, or moth flies. Phlebotominæ. Culicidæ, or + mosquitoes. Simuliidæ, or black-flies. Chironomidæ, or + midges. Tabanidæ, or horse-flies. Leptidæ or + snipe-flies. Oestridæ, or bot-flies. Muscidæ, the + stable-fly and others. + Siphonaptera, or fleas. + The fleas affecting man, the dog, cat, and rat. + The true chiggers, or chigoes. + +CHAPTER IV + +ACCIDENTAL OR FACULTATIVE PARASITES 131-143 + + Acarina, or mites. + Myriapoda, or centipedes and millipedes. + Lepidopterous larvæ. + Coleoptera, or beetles. + Dipterous larvæ causing myiasis. + Piophila casei, the cheese skipper. Chrysomyia macellaria, + the screw-worm fly. Calliphorinæ, the bluebottles. + Muscinæ, the house or typhoid fly, and others. + Anthomyiidæ, the lesser house-fly and others. + Sarcophagidæ, the flesh-flies. + + +CHAPTER V + +ARTHROPODS AS SIMPLE CARRIERS OF DISEASE 144-163 + + The house or typhoid fly as a carrier of disease. + Stomoxys calcitrans, the stable-fly. + Other arthropods which may serve as simple carriers of + pathogenic organisms. + + +CHAPTER VI + +ARTHROPODS AS DIRECT INOCULATORS OF DISEASE GERMS 164-174 + + Some illustrations of direct inoculations of disease germs + by arthropods. + The rôle of fleas in the transmission of the plague. + + +CHAPTER VII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC ORGANISMS 175-185 + + Insects as intermediate hosts of tape-worms. + Arthropods as intermediate hosts of nematode worms. + Filariasis and mosquitoes. + Other nematode parasites of man and animals. + + +CHAPTER VIII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA 186-211 + + Mosquitoes and malaria. + Mosquitoes and yellow fever. + + +CHAPTER IX + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA 212-229 + + Insects and trypanosomiases. + Fleas and lice as carriers of Trypanosoma lewisi. + Tsetse-flies and nagana. + Tsetse-flies and sleeping sickness in man. + South American trypanosomiasis. + Leishmanioses and insects. + Ticks and diseases of man and animals. + Cattle tick and Texas fever. + Ticks and Rocky Mountain Spotted fever of man. + + +CHAPTER X + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA +(CONTINUED) 230-240 + + Arthropods and Spirochætoses of man and animals. + African relapsing fever of man. + European relapsing fever. + North African relapsing fever of man. + Other types of relapsing fever of man. + Spirochætosis of fowls. + Other spirochæte diseases of animals. + Typhus fever and lice. + + +CHAPTER XI + +SOME POSSIBLE, BUT IMPERFECTLY KNOWN CASES OF +ARTHROPOD TRANSMISSION OF DISEASE 241-256 + + Infantile paralysis, or acute anterior poliomyelitis. + Pellagra. Leprosy. Verruga peruviana. Cancer. + + +CHAPTER XII + +KEYS TO THE ARTHROPODS NOXIOUS TO MAN 257-317 + + Crustacea. + Myriapoda, or centipedes and millipedes. + Arachnida (Orders of). + Acarina or ticks. + Hexapoda (Insecta). + Siphunculata and Hemiptera (lice and true bugs). + Diptera (mosquitoes, midges, and flies). + Siphonaptera (fleas). + + +APPENDIX + +Hydrocyanic acid gas against household insects 318-320 + Proportion of ingredients. A single room as an example. + Fumigating a large house. Precautions. + +Lesions produced by the bite of the black-fly 321-326 + + +BIBLIOGRAPHY 327-340 + + +INDEX 341-348 + + + + +CHAPTER I. + +INTRODUCTION + +EARLY SUGGESTIONS REGARDING THE TRANSMISSION OF DISEASE BY INSECTS + + +Until very recent years insects and their allies have been considered as +of economic importance merely in so far as they are an annoyance or +direct menace to man, or his flocks and herds, or are injurious to his +crops. It is only within the past fifteen years that there has sprung +into prominence the knowledge that in another and much more insiduous +manner, they may be the enemy of mankind, that they may be among the +most important of the disseminators of disease. In this brief period, +such knowledge has completely revolutionized our methods of control of +certain diseases, and has become an important weapon in the fight for +the conservation of health. + +It is nowhere truer than in the case under consideration that however +abrupt may be their coming into prominence, great movements and great +discoveries do not arise suddenly. Centuries ago there was suggested the +possibility that insects were concerned with the spread of disease, and +from time to time there have appeared keen suggestions and logical +hypotheses along this line, that lead us to marvel that the +establishment of the truths should have been so long delayed. + +One of the earliest of these references is by the Italian physician, +Mercurialis, who lived from 1530 to 1607, during a period when Europe +was being ravaged by the dread "black death", or plague. Concerning its +transmission he wrote: "There can be no doubt that flies feed on the +internal secretions of the diseased and dying, then, flying away, they +deposit their excretions on the food in neighboring dwellings, and +persons who eat of it are thus infected." + +It would be difficult to formulate more clearly this aspect of the facts +as we know them to-day, though it must always be borne in mind that we +are prone to interpret such statements in the light of present-day +knowledge. Mercurialis had no conception of the animate nature of +contagion, and his statement was little more than a lucky guess. + +Much more worthy of consideration is the approval which was given to his +view by the German Jesuit, Athanasius Kircher in 1658. One cannot read +carefully his works without believing that long before Leeuwenhook's +discovery, Kircher had seen the larger species of bacteria. Moreover, he +attributed the production of disease to these organisms and formulated, +vaguely, to be sure, a theory of the animate nature of contagion. It has +taken two and a half centuries to accumulate the facts to prove his +hypothesis. + +The theory of Mercurialis was not wholly lost sight of, for in the +medical literature of the eighteenth century there are scattered +references to flies as carriers of disease. Such a view seems even to +have been more or less popularly accepted, in some cases. Gudger (1910), +has pointed out that, as far back as 1769, Edward Bancroft, in "An Essay +on the Natural History of Guiana in South America," wrote concerning the +contagious skin-disease known as "Yaws": "It is usually believed that +this disorder is communicated by the flies who have been feasting on a +diseased object, to those persons who have sores, or scratches, which +are uncovered; and from many observations, I think this is not +improbable, as none ever receive this disorder whose skins are whole." + +Approaching more closely the present epoch, we find that in 1848, Dr. +Josiah Nott, of Mobile, Alabama, published a remarkable article on the +cause of yellow fever, in which he presented "reasons for supposing its +specific cause to exist in some form of insect life." As a matter of +fact, the bearing of Nott's work on present day ideas of the insect +transmission of disease has been very curiously overrated. The common +interpretation of his theory has been deduced from a few isolated +sentences, but his argument appears quite differently when the entire +article is studied. It must be remembered that he wrote at a period +before the epoch-making discoveries of Pasteur and before the +recognition of micro-organisms as factors in the cause of disease. His +article is a masterly refutation of the theory of "malarial" origin of +"all the fevers of hot climates," but he uses the term "insect" as +applicable to the lower forms of life, and specific references to +"mosquitoes," "aphids," "cotton-worms," and others, are merely in the +way of similes. + +But, while Nott's ideas regarding the relation of insects to yellow +fever were vague and indefinite, it was almost contemporaneously that +the French physician, Louis Daniel Beauperthuy argued in the most +explicit possible manner, that yellow fever and various others are +transmitted by mosquitoes. In the light of the data which were available +when he wrote, in 1853, it is not surprising that he erred by thinking +that the source of the virus was decomposing matter which the mosquito +took up and accidentally inoculated into man. Beauperthuy not only +discussed the rôle of mosquitoes in the transmission of disease, but he +taught, less clearly, that house-flies scatter pathogenic organisms. It +seems that Boyce (1909) who quotes extensively from this pioneer work, +does not go too far when he says "It is Dr. Beauperthuy whom we must +regard as the father of the doctrine of insect-borne disease." + +In this connection, mention must be made of the scholarly article by the +American physician, A. F. A. King who, in 1883, brought together an all +but conclusive mass of argument in support of his belief that malaria +was caused by mosquitoes. At about the same time, Finley, of Havana, was +forcefully presenting his view that the mosquito played the chief rôle +in the spread of yellow fever. + +To enter more fully into the general historical discussion is beyond the +scope of this book. We shall have occasion to make more explicit +references in considering various insect-borne diseases. Enough has been +said here to emphasize that the recognition of insects as factors in the +spread of disease was long presaged, and that there were not wanting +keen thinkers who, with a background of present-day conceptions of the +nature of disease, might have been in the front rank of investigators +along these lines. + + +THE WAYS IN WHICH ARTHROPODS MAY AFFECT THE HEALTH OF MAN + +When we consider the ways in which insects and their allies may affect +the health of man, we find that we may treat them under three main +groups: + +A. They may be directly poisonous. Such, for example, are the scorpions, +certain spiders and mites, some of the predaceous bugs, and stinging +insects. Even such forms as the mosquito deserve some consideration from +this viewpoint. + +B. They may be parasitic, living more or less permanently on or in the +body and deriving their sustenance from it. + +Of the parasitic arthropods we may distinguish, first, the _true +parasites_, those which have adopted and become confirmed in the +parasitic habit. Such are the itch mites, the lice, fleas, and the +majority of the forms to be considered as parasitic. + +In addition to these, we may distinguish a group of _accidental_, or +_facultative parasites_, species which are normally free-living, feeding +on decaying substances, but which when accidentally introduced into the +alimentary canal or other cavities of man, may exist there for a greater +or less period. For example, certain fly larvæ, or maggots, normally +feeding in putrifying meat, have been known to occur as accidental or +facultative parasites in the stomach of man. + +C. Finally, and most important, arthropods may be transmitters and +disseminators of disease. In this capacity they may function in one of +three ways; as _simple carriers_, as _direct inoculators_, or as +_essential hosts_ of disease germs. + +As simple carriers, they may, in a wholly incidental manner, transport +from the diseased to the healthy, or from filth to food, pathogenic +germs which cling to their bodies or appendages. Such, for instance, is +the relation of the house-fly to the dissemination of typhoid. + +As direct inoculators, biting or piercing species may take up from a +diseased man or animal, germs which, clinging to the mouth parts, are +inoculated directly into the blood of the insect's next victim. It it +thus that horse-flies may occasionally transmit anthrax. Similarly, +species of spiders and other forms which are ordinarily perfectly +harmless, may accidentally convey and inoculate pyogenic bacteria. + +It is as essential hosts of disease germs that arthropods play their +most important rôle. In such cases an essential part of the life cycle +of the pathogenic organism is undergone in the insect. In other words, +without the arthropod host the disease-producing organism cannot +complete its development. As illustrations may be cited the relation of +the Anopheles mosquito to the malarial parasite, and the relation of the +cattle tick to Texas fever. + +A little consideration will show that this is the most important of the +group. Typhoid fever is carried by water or by contaminated milk, and in +various other ways, as well as by the house-fly. Kill all the +house-flies and typhoid would still exist. On the other hand, malaria is +carried only by the mosquito, because an essential part of the +development of the malarial parasite is undergone in this insect. +Exterminate all of the mosquitoes of certain species and the +dissemination of human malaria is absolutely prevented. + +Once an arthropod becomes an essential host for a given parasite it may +disseminate infection in three different ways: + +1. By infecting man or animals who ingest it. It is thus, for example, +that man, dog, or cat, becomes infected with the double-pored dog +tapeworm, _Dipylidium caninum_. The cysticercoid stage occurs in the +dog louse, or in the dog or cat fleas, and by accidentally ingesting the +infested insect the vertebrate becomes infested. Similarly, _Hymenolepis +diminuta_, a common tapeworm of rats and mice, and occasional in man, +undergoes part of its life cycle in various meal-infesting insects, and +is accidentally taken up by its definitive host. It is very probable +that man becomes infested with _Dracunculus (Filaria) medinensis_ +through swallowing in drinking water, the crustacean, _Cyclops_, +containing the larvæ of this worm. + +2. By infecting man or animals on whose skin or mucous membranes the +insect host may be crushed or may deposit its excrement. The pathogenic +organism may then actively penetrate, or may be inoculated by +scratching. The causative organism of typhus fever is thus transmitted +by the body louse. + +3. By direct inoculation by its bite, the insect host may transfer the +parasite which has undergone development within it. The malarial +parasite is thus transferred by mosquitoes; the Texas fever parasite by +cattle ticks. + + + + +CHAPTER II. + +ARTHROPODS WHICH ARE DIRECTLY POISONOUS + + +Of all the myriads of insects and related forms, a very few are of +direct use to man, some few others have forced his approbation on +account of their wonderful beauty, but the great hordes of them are +loathed or regarded as directly dangerous. As a matter of fact, only a +very small number are in the slightest degree poisonous to man or to the +higher animals. The result is that entomologists and lovers of nature, +intent upon dissipating the foolish dread of insects, are sometimes +inclined to go to the extreme of discrediting all statements of serious +injury from the bites or stings of any species. + +Nevertheless, it must not be overlooked that poisonous forms do exist, +and they must receive attention in a consideration of the ways in which +arthropods may affect the health of man. Moreover, it must be recognized +that "what is one man's meat, is another man's poison," and that in +considering the possibilities of injury we must not ignore individual +idiosyncrasies. Just as certain individuals may be poisoned by what, for +others, are common articles of food, so some persons may be abnormally +susceptible to insect poison. Thus, the poison of a bee sting may be of +varying severity, but there are individuals who are made seriously sick +by a single sting, regardless of the point of entry. Some individuals +scarcely notice a mosquito bite, others find it very painful, and so +illustrations of this difference in individuals might be multiplied. + +In considering the poisonous arthropods, we shall take them up by +groups. The reader who is unacquainted with the systematic relationship +of insects and their allies is referred to Chapter XII. No attempt will +be made to make the lists under the various headings exhaustive, but +typical forms will be discussed. + + +ARANEIDA OR SPIDERS + +Of all the arthropods there are none which are more universally feared +than are the spiders. It is commonly supposed that the majority, if not +all the species are poisonous and that they are aggressive enemies of +man and the higher animals, as well as of lower forms. + +That they really secrete a poison may be readily inferred from the +effect of their bite upon insects and other small forms. Moreover, the +presence of definite and well-developed poison glands can easily be +shown. They occur as a pair of pouches (fig. 1) lying within the +cephalothorax and connected by a delicate duct with a pore on the claw +of the chelicera, or so-called "mandible" on the convex surface of the +claw in such a position that it is not plugged and closed by the flesh +of the victim. + +[Illustration: 1. Head of a spider showing poison gland (_c_) and its +relation to the chelicera (_a_).] + +The glands may be demonstrated by slowly and carefully twisting off a +chelicera and pushing aside the stumps of muscles at its base. By +exercising care, the chitinous wall of the chelicera and its claw may be +broken away and the duct traced from the gland to its outlet. The inner +lining of the sac is constituted by a highly developed glandular +epithelium, supported by a basement membrane of connective tissue and +covered by a muscular layer, (fig. 2). The muscles, which are striated, +are spirally arranged (fig. 1), and are doubtless under control of the +spider, so that the amount of poison to be injected into a wound may be +varied. + +[Illustration: 2. Section through a venom gland of Latrodectus +13-guttatus showing the peritoneal, muscular and epithelial layers. +After Bordas.] + +The poison itself, according to Kobert (1901), is a clear, colorless +fluid, of oily consistency, acid reaction, and very bitter taste. After +the spider has bitten two or three times, its supply is exhausted and +therefore, as in the case of snakes, the poison of the bite decreases +quickly with use, until it is null. To what extent the content of the +poison sacs may contain blood serum or, at least, active principles of +serum, in addition to a specific poison formed by the poison glands +themselves, Kobert regards as an open question. He believes that the +acid part of the poison, if really present, is formed by the glands and +that, in the case of some spiders, the ferment-like, or better, active +toxine, comes from the blood. + +[Illustration: 3. Chelicera of a spider.] + +But there is a wide difference between a poison which may kill an insect +and one which is harmful to men. Certain it is that there is no lack of +popular belief and newspaper records of fatal cases, but the evidence +regarding the possibility of fatal or even very serious results for man +is most contradictory. For some years, we have attempted to trace the +more circumstantial newspaper accounts, which have come to our notice, +of injury by North American species. The results have served, mainly, to +emphasize the straits to which reporters are sometimes driven when there +is a dearth of news. The accounts are usually vague and lacking in any +definite clue for locating the supposed victim. In the comparatively few +cases where the patient, or his physician, could be located, there was +either no claim that the injury was due to spider venom, or there was no +evidence to support the belief. Rarely, there was evidence that a +secondary blood poisoning, such as might be brought about by the prick +of a pin, or by any mechanical injury, had followed the bite of a +spider. Such instances have no bearing on the question of the venomous +nature of these forms. + +[Illustration: 4. The Italian tarantula (Lycosa tarantula). After +Kobert.] + +The extreme to which unreasonable fear of the bites of spiders +influenced the popular mind was evidenced by the accepted explanation of +the remarkable dancing mania, or tarantism, of Italy during the Middle +Ages. This was a nervous disorder, supposed to be due to the bite of a +spider, the European tarantula (fig. 4), though it was also, at times, +attributed to the bite of the scorpion. In its typical form, it was +characterized by so great a sensibility to music that under its +influence the victims indulged in the wildest and most frenzied dancing, +until they sank to the ground utterly exhausted and almost lifeless. The +profuse perspiring resulting from these exertions was supposed to be the +only efficacious remedy for the disease. Certain forms of music were +regarded as of especial value in treating this tarantism, and hence the +name of "tarantella" was applied to them. Our frontispiece, taken from +Athanasius Kircher's _Magnes sive de Arte Magnetica_, 1643 ed., +represents the most commonly implicated spider and illustrates some of +what Fabre has aptly designated as "medical choreography." + +The disease was, in reality, a form of hysteria, spreading by sympathy +until whole communities were involved, and was paralleled by the +outbreaks of the so-called St. Vitus's or St. John's dance, which swept +Germany at about the same time (fig. 5). The evidence that the spider +was the cause of the first is about as conclusive as is that of the +demoniacal origin of the latter. The true explanation of the outbreaks +is doubtless to be found in the depleted physical and mental condition +of the people, resulting from the wars and the frightful plagues which +devastated all Europe previous to, and during these times. An +interesting discussion of these aspects of the question is to be found +in Hecker. + +[Illustration: 5. Dancing mania. Illustration from Johann Ludwig +Gottfried's Chronik. 1632.] + +So gross has been the exaggeration and so baseless the popular fear +regarding spiders that entomologists have been inclined to discredit all +accounts of serious injury from their bites. Not only have the most +circumstantial of newspaper accounts proved to be without foundation but +there are on record a number of cases where the bite of many of the +commoner species have been intentionally provoked and where the effect +has been insignificant. Some years ago the senior author personally +experimented with a number of the largest of our northern species, and +with unexpected results. The first surprise was that the spiders were +very unwilling to bite and that it required a considerable effort to get +them to attempt to do so. In the second place, most of those +experimented with were unable to pierce the skin of the palm or the back +of the hand, but had to be applied to the thin skin between the fingers +before they were able to draw blood. Unfortunately, no special attempt +was made to determine, at the time, the species experimented with, but +among them were _Theridion tepidariorum_, _Miranda aurantia_ +(_Argiopa_), _Metargiope trifasciata_, _Marxia stellata_, _Aranea +trifolium_, _Misumena vatia_, and _Agelena nævia_. In no case was the +bite more severe than a pin prick and though in some cases the sensation +seemed to last longer, it was probably due to the fact that the mind was +intent upon the experiment. + +[Illustration: 6. An American tarantula (Eurypelma hentzii). Natural +size. After Comstock.] + +Similar experiments were carried out by Blackwell (1855), who believed +that in the case of insects bitten, death did not result any more +promptly than it would have from a purely mechanical injury of equal +extent. He was inclined to regard all accounts of serious injury to man +as baseless. The question cannot be so summarily dismissed, and we shall +now consider some of the groups which have been more explicitly +implicated. + + +THE TARANTULAS.--In popular usage, the term "tarantula" is loosely +applied to any one of a number of large spiders. The famous tarantulas +of southern Europe, whose bites were supposed to cause the dancing +mania, were Lycosidæ, or wolf-spiders. Though various species of this +group were doubtless so designated, the one which seems to have been +most implicated was _Lycosa tarantula_ (L.), (fig. 4). On the other +hand, in this country, though there are many Lycosidæ, the term +"tarantula" has been applied to members of the superfamily Avicularoidea +(fig. 6), including the bird-spiders. + +Of the Old World Lycosidæ there is no doubt that several species were +implicated as the supposed cause of the tarantism. In fact, as we have +already noted, the blame was sometimes attached to a scorpion. However, +there seems to be no doubt that most of the accounts refer to the spider +known as _Lycosa tarantula_. + +There is no need to enter into further details here regarding the +supposed virulence of these forms, popular and the older medical +literature abound in circumstantial accounts of the terrible effects of +the bite. Fortunately, there is direct experimental evidence which bears +on the question. + +Fabre induced a common south European wolf-spider, _Lycosa narbonensis_, +to bite the leg of a young sparrow, ready to leave the nest. The leg +seemed paralyzed as a result of the bite, and though the bird seemed +lively and clamored for food the next day, on the third day it died. A +mole, bitten on the nose, succumbed after thirty-six hours. From these +experiments Fabre seemed justified in his conclusion that the bite of +this spider is not an accident which man can afford to treat lightly. +Unfortunately, there is nothing in the experiments, or in the symptoms +detailed, to exclude the probability that the death of the animals was +the result of secondary infection. + +As far back as 1693, as we learn from the valuable account of Kobert, +(1901), the Italian physician, Sanguinetti allowed himself to be bitten +on the arm by two tarantulas, in the presence of witnesses. The +sensation was equivalent to that from an ant or a mosquito bite and +there were no other phenomena the first day. On the second day the wound +was inflamed and there was slight ulceration. It is clear that these +later symptoms were due to a secondary infection. These experiments have +been repeated by various observers, among whom may be mentioned Leon +Dufour, Josef Erker and Heinzel, and with the similar conclusion that +the bite of the Italian tarantula ordinarily causes no severe symptoms. +In this conclusion, Kobert, though firmly convinced of the poisonous +nature of some spiders, coincides. He also believes that striking +symptoms may be simulated or artificially induced by patients in order +to attract interest, or because they have been assured that the bite, +under all circumstances, caused tarantism. + +The so-called Russian tarantula, _Trochosa singoriensis_ (fig. 7), is +much larger than the Italian species, and is much feared. Kobert carried +out a series of careful experiments with this species and his results +have such an important bearing on the question of the venomous nature of +the tarantula that we quote his summary. Experimenting first on nearly a +hundred living specimens of _Trochosa singoriensis_ from Crimea he says +that: + +"The tarantulas, no matter how often they were placed on the skin, +handled, and irritated, could not be induced to bite either myself, the +janitor, or the ordinary experimental animals. The objection that the +tarantulas were weak and indifferent cannot stand, for as soon as I +placed two of them on the shaved skin of a rabbit, instead of an attack +on the animal, there began a furious battle between the two spiders, +which did not cease until one of the two was killed." + +[Illustration: 7. Trochosa singoriensis. After Kobert.] + +"Since the spiders would not bite, I carefully ground up the fresh +animals in physiological salt solution, preparing an extract which must +have contained, in solution, all of the poisonous substance of their +bodies. While in the case of _Latrodectus_, as we shall see, less than +one specimen sufficed to yield an active extract, I have injected the +filtered extract of six fresh Russian tarantulas, of which each one was +much heavier than an average _Latrodectus_, subcutaneously and into the +jugular vein of various cats without the animals dying or showing any +special symptoms. On the basis of my experiments I can therefore only +say that the quantity of the poison soluble in physiological salt +solution, even when the spiders are perfectly fresh and well nourished, +is very insignificant. That the poison of the Russian tarantula is not +soluble in physiological salt solution, is exceedingly improbable. +Moreover, I have prepared alcoholic extracts and was unable to find them +active. Since the Russian spider exceeds the Italian in size and in +intensity of the bite, it seems very improbable to me that the +pharmacological test of the Italian tarantula would yield essentially +other results than those from the Russian species." + +To the AVICULAROIDEA belong the largest and most formidable appearing of +the spiders and it is not strange that in the New World they have fallen +heir to the bad reputation, as well as to the name of the tarantula of +Europe. In this country they occur only in the South or in the far West, +but occasionally living specimens are brought to our northern ports in +shipments of bananas and other tropical produce, and are the source of +much alarm. It should be mentioned, however, that the large spider most +frequently found under such circumstances is not a tarantula at all, but +one of the Heteropodidæ, or giant crab-spiders, (fig. 8). + +[Illustration: 8. The giant crab-spider or banana spider (Heteropoda +venatoria). Natural size. After Comstock.] + +In spite of their prominence and the fear which they arouse there are +few accurate data regarding these American tarantulas. It has often been +shown experimentally that they can kill small birds and mammals, though +it is doubtful if these form the normal prey of any of the species, as +has been claimed. There is no question but that the mere mechanical +injury which they may inflict, and the consequent chances of secondary +infection, justify, in part, their bad reputation. In addition to the +injury from their bite, it is claimed that the body hairs of several of +the South American species are readily detached and are urticating. + +Recently, Phisalix (1912) has made a study of the physiological effects +of the venom of two Avicularoidea, _Phormictopus carcerides_ Pocock, +from Haiti and _Cteniza sauvagei_ Rossi, from Corsica. The glands were +removed aseptically and ground up with fine, sterilized sand in +distilled water. The resultant liquid was somewhat viscid, colorless, +and feebly alkaline. Injected into sparrows and mice the extract of +_Phormictopus_ proved very actively poisonous, that from a single spider +being sufficient to kill ten sparrows or twenty mice. It manifested +itself first and, above all, as a narcotic, slightly lowering the +temperature and paralyzing the respiration. Muscular and cardiac +weakening, loss of general sensibility, and the disappearance of +reflexes did not occur until near the end. The extract from _Cteniza_ +was less active and, curiously enough, the comparative effect on +sparrows and on mice was just reversed. + + +SPIDERS OF THE GENUS LATRODECTUS.--While most of the popular accounts of +evil effects from the bites of spiders will not stand investigation, it +is a significant fact that, the world over, the best authenticated +records refer to a group of small and comparatively insignificant +spiders belonging to the genus _Latrodectus_, of the family Theridiidæ. +The dread "Malmigniatte" of Corsica and South Europe, the "Karakurte" of +southeastern Russia, the "Katipo" of New Zealand, the "Mena-vodi" and +"Vancoho" of Madagascar, and our own _Latrodectus mactans_, all belong +to this genus, and concerning all of these the most circumstantial +accounts of their venomous nature are given. These accounts are not mere +fantastic stories by uneducated natives but in many cases are reports +from thoroughly trained medical men. + +The symptoms produced are general, rather than local. As summarized by +Kobert (1901) from a study of twenty-two cases treated in 1888, in the +Kherson (Russia) Government Hospital and Berislaw (Kherson) District +Hospital the typical case, aside from complications, exhibits the +following symptoms. The victim suddenly feels the bite, like the sting +of a bee. Swelling of the barely reddened spot seldom follows. The +shooting pains, which quickly set in, are not manifested at the point of +injury but localized at the joints of the lower limb and in the region +of the hip. The severity of the pain forces the victim to the hospital, +in spite of the fact that they otherwise have a great abhorrence of it. +The patient is unable to reach the hospital afoot, or, at least, not +without help, for there is usually inability to walk. The patient, even +if he has ridden, reaches the hospital covered with cold sweat and +continues to perspire for a considerable period. His expression +indicates great suffering. The respiration may be somewhat dyspnoeic, +and a feeling of oppression in the region of the heart is common. There +is great aversion to solid food, but increasing thirst for milk and tea. +Retention of urine, and constipation occur. Cathartics and, at night, +strong narcotics are desired. Warm baths give great relief. After three +days, there is marked improvement and usually the patient is dismissed +after the fifth. This summary of symptoms agrees well with other +trustworthy records. + +It would seem, then, that Riley and Howard (1889), who discussed a +number of accounts in the entomological literature, were fully justified +in their statement that "It must be admitted that certain spiders of the +genus _Latrodectus_ have the power to inflict poisonous bites, which may +(probably exceptionally and depending upon exceptional conditions) bring +about the death of a human being." + +And yet, until recently the evidence bearing on the question has been +most conflicting. The eminent arachnologist, Lucas, (1843) states that +he himself, had been repeatedly bitten by the Malmigniatte without any +bad effects. Dr. Marx, in 1890, gave before the Entomological Society of +Washington, an account of a series of experiments to determine whether +the bite of _Latrodectus mactans_ is poisonous or not. He described the +poison glands as remarkably small[A] and stated that he had introduced +the poison in various ways into guinea-pigs and rabbits without +obtaining any satisfactory results. Obviously, carefully conducted +experiments with the supposed venom were needed and fortunately they +have been carried out in the greatest detail by Kobert (1901). + +This investigator pointed out that there were two factors which might +account for the discrepancies in the earlier experiments. In the first +place, the poison of spiders, as of snakes, might be so exhausted after +two or three bites that further bites, following directly, might be +without visible effect. Secondly, the application of the poison by means +of the bite, is exceedingly inexact, since even after the most careful +selection of the point of application, the poison might in one instance +enter a little vein or lymph vessel, and in another case fail to do so. +Besides, there would always remain an incalculable and very large amount +externally, in the nonabsorptive epithelium. While all of these factors +enter into the question of the effect of the bite in specific instances, +they must be as nearly as possible obviated in considering the question +of whether the spiders really secrete a venom harmful to man. + +Kobert therefore sought to prepare extracts which would contain the +active principles of the poison and which could be injected in definite +quantities directly into the blood of the experimental animal. For this +purpose various parts of the spiders were rubbed up in a mortar with +distilled water, or physiological salt solution, allowed to stand for an +hour, filtered, and then carefully washed, by adding water drop by drop +for twenty-four hours. The filtrate and the wash-water were then united, +well mixed and, if necessary, cleared by centrifuging or by exposure to +cold. The mixture was again filtered, measured, and used, in part, for +injection and, in part, for the determination of the organic materials. + +Such an extract was prepared from the cephalothoraces of eight dried +specimens of the Russian _Latrodectus_ and three cubic centimeters of +this, containing 4.29 mg. of organic material, were injected into the +jugular vein of a cat weighing 2450 grams. The previously very active +animal was paralyzed and lay in whatever position it was placed. The +sensibility of the skin of the extremities and the rump was so reduced +that there was no reaction from cutting or sticking. There quickly +followed dyspnoea, convulsions, paralysis of the respiratory muscles +and of the heart. In twenty-eight minutes the cat was dead, after having +exhibited exactly the symptoms observed in severe cases of poisoning of +man from the bite of this spider. + +These experiments were continued on cats, dogs, guinea pigs and various +other animals. Not only extracts from the cephalothorax, but from other +parts of the body, from newly hatched spiders, and from the eggs were +used and all showed a similar virulence. Every effort was made to avoid +sources of error and the experiments, conducted by such a recognized +authority in the field of toxicology, must be accepted as conclusively +showing that this spider and, presumably, other species of the genus +_Latrodectus_ against which the clinical evidence is quite parallel, +possess a poison which paralyzes the heart and central nervous system, +with or without preliminary stimulus of the motor center. If the +quantity of the poison which comes into direct contact with the blood is +large, there may occur hæmolysis and thrombosis of the vessels. + +On the other hand, check experiments were carried out, using similar +extracts of many common European spiders of the genera _Tegenaria_, +_Drassus_, _Agelena_, _Eucharia_ and _Argyroneta_, as well as the +Russian tarantula, _Lycosa singoriensis_. In no other case was the +effect on experimental animals comparable to the _Latrodectus_ extract. + +Kobert concludes that in its chemical nature the poison is neither an +alkaloid, nor a glycoside, nor an acid, but a toxalbumen, or poisonous +enzyme which is very similar to certain other animal poisons, notably +that of the scorpion. + +[Illustration: 9. Latrodectus mactans; (_a_) female, x 3; (_b_) venter +of female; (_c_) dorsum of male. After Comstock.] + +The genus _Latrodectus_ is represented in the United States by at least +two species, _L. mactans_ and _L. geometricus_. Concerning _L. mactans_ +there are very circumstantial accounts of serious injury and even death +in man[B]. _Latrodectus mactans_ is coal black, marked with red or +yellow or both. It has eight eyes, which are dissimilar in color and +are distinctly in front of the middle of the thorax, the lateral eyes of +each side widely separate. The tarsus of the fourth pair of legs has a +number of curved setæ in a single series. It has on the ventral side of +its abdomen an hour-glass shaped spot. The full-grown female is about +half an inch in length. Its globose abdomen is usually marked with one +or more red spots dorsally along the middle line. The male is about half +as long but has in addition to the dorsal spots, four pairs of stripes +along the sides. Immature females resemble the male in coloring (fig. +9). + +Regarding the distribution of _Latrodectus mactans_, Comstock states +that: "Although it is essentially a Southern species, it occurs in +Indiana, Ohio, Pennsylvania, New Hampshire, and doubtless other of the +Northern States." _L. geometricus_ has been reported from California. + + +OTHER VENOMOUS SPIDERS--While conclusive evidence regarding the venomous +nature of spiders is meager and relates almost wholly to that of the +genus _Latrodectus_, the group is a large one and we are not justified +in dismissing arbitrarily, all accounts of injury from their bites. +Several species stand out as especially needing more detailed +investigation. + +_Chiracanthium nutrix_ is a common European species of the family +Clubionidæ, concerning which there is much conflicting testimony. Among +the reports are two by distinguished scientists whose accounts of +personal experiences cannot be ignored. A. Forel allowed a spider of +this species to bite him and not only was the pain extreme, but the +general symptoms were so severe that he had to be helped to his house. +The distinguished arachnologist, Bertkau reports that he, himself, was +bitten and that an extreme, burning pain spread almost instantaneously +over the arm and into the breast. There were slight chills the same day +and throbbing pain at the wound lasted for days. While this particular +species is not found in the United States, there are two other +representatives of the genus and it is possible that they possess the +same properties. We are unaware of any direct experimental work on the +poison. + +_Epeira diadema_, of Europe, belongs to a wholly different group, that +of the orb-weavers, but has long been reputed venomous. Kobert was able +to prepare from it an extract whose effects were very similar to that +prepared from _Latrodectus_, though feebler in its action. Under +ordinary circumstances this spider is unable to pierce the skin of man +and though Kobert's results seem conclusive, the spider is little to be +feared. + +_Phidippus audax_ (_P. tripunctatus_) is one of our largest Attids, or +jumping spiders. The late Dr. O. Lugger describes a case of severe +poisoning from the bite of this spider and though details are lacking, +it is quite possible that this and other large species of the same +group, which stalk their prey, may possess a more active poison than +that of web-building species. + + +SUMMARY--It is clearly established that our common spiders are not to be +feared and that the stories regarding their virulence are almost wholly +without foundation. On the other hand, the chances of secondary +infection from the bites of some of the more powerful species are not to +be ignored. + +Probably all species possess a toxin secreted by the poison gland, +virulent for insects and other normal prey of the spiders, but with +little or no effect on man. + +There are a very few species, notably of the genus _Latrodectus_, and +possibly including the European _Chiracanthium nutrix_ and _Epeira +diadema_, which possess, in addition, a toxalbumen derived from the +general body tissue, which is of great virulence and may even cause +death in man and the higher animals. + +[Illustration: 10. A whip-scorpion (Mastigoproctus giganteus). Half +natural size. After Comstock.] + + +THE PEDIPALPIDA OR WHIP-SCORPIONS + +The tailed whip-scorpions, belonging to the family Thelyphonidæ, are +represented in the United States by the giant whip-scorpion +_Mastigoproctus giganteus_ (fig. 10), which is common in Florida, Texas +and some other parts of the South. In Florida, it is locally known as +the "grampus" or "mule-killer" and is very greatly feared. There is no +evidence that these fears have any foundation, and Dr. Marx states that +there is neither a poison gland nor a pore in the claw of the +chelicera. + + +THE SCORPIONIDA, OR TRUE SCORPIONS + +The true scorpions are widely distributed throughout warm countries and +everywhere bear an evil reputation. According to Comstock (1912), about +a score of species occur in the Southern United States. These are +comparatively small forms but in the tropics members of this group may +reach a length of seven or eight inches. They are pre-eminently +predaceous forms, which lie hidden during the day and seek their prey by +night. + +[Illustration: 11. A true scorpion. After Comstock.] + +The scorpions (fig. 11) possess large pedipalpi, terminated by strongly +developed claws, or chelæ. They may be distinguished from all other +Arachnids by the fact that the distinctly segmented abdomen is divided +into a broad basal region of seven segments and a terminal, slender, +tail-like division of five distinct segments. + +The last segment of the abdomen, or telson, terminates in a +ventrally-directed, sharp spine, and contains a pair of highly developed +poison glands. These glands open by two small pores near the tip of the +spine. Most of the species when running carry the tip of the abdomen +bent upward over the back, and the prey, caught and held by the +pedipalpi, is stung by inserting the spine of the telson and allowing it +to remain for a time in the wound. + +The glands themselves have been studied in _Prionurus citrinus_ by +Wilson (1904). He found that each gland is covered by a sheet of muscle +on its mesal and dorsal aspects, which may be described as the +_compressor muscle_. The muscle of each side is inserted by its edge +along the ventral inner surface of the chitinous wall of the telson, +close to the middle line, and by a broader insertion laterally. A layer +of fine connective tissue completely envelops each gland and forms the +basis upon which the secreting cells rest. The secreting epithelium is +columnar; and apparently of three different types of cells. + +1. The most numerous have the appearance of mucous cells, resembling the +goblet cells of columnar mucous membranes. The nucleus, surrounded by a +small quantity of protoplasm staining with hæmatoxylin, lies close to +the base of the cell. + +2. Cells present in considerable numbers, the peripheral portions of +which are filled with very numerous fine granules, staining with acid +dyes such as methyl orange. + +3. Cells few in number, filled with very large granules, or irregular +masses of a substance staining with hæmatoxylin. + +The poison, according to Kobert (1893), is a limpid, acid-reacting +fluid, soluble in water but insoluble in absolute alcohol and ether. +There are few data relative to its chemical nature. Wilson (1901) states +that a common Egyptian species, _Buthus quinquestriatus_, has a specific +gravity of 1.092, and contains 20.3% of solids and 8.4% ash. + +The venom of different species appears to differ not only quantitatively +but qualitatively. The effects of the bite of the smaller species of the +Southern United States may be painful but there is no satisfactory +evidence that it is ever fatal. On the other hand, certain tropical +species are exceedingly virulent and cases of death of man from the bite +are common. + +In the case of _Buthus quinquestriatus_, Wilson (1904) found the +symptoms in animals to be hypersecretion, salivation and lachrymation, +especially marked, convulsions followed by prolonged muscular spasm; +death from asphyxia. The temperature shows a slight, rarely +considerable, rise. Rapid and considerable increase of blood-pressure +(observed in dogs) is followed by a gradual fall with slowing of the +heart-beat. The coagulability of the blood is not affected. + +An interesting phase of Wilson's work was the experiments on desert +mammals. The condition under which these animals exist must frequently +bring them in contact with scorpions, and he found that they possess a +degree of immunity to the venom sufficient at least to protect them from +the fatal effects of the sting. + +As far as concerns its effect on man, Wilson found that much depended +upon the age. As high as 60 per cent of the cases of children under +five, resulted fatally. Caroroz (1865), states that in a Mexican state +of 15,000 inhabitants, the scorpions were so abundant and so much feared +that the authorities offered a bounty for their destruction. A result +was a large number of fatalities, over two hundred per year. Most of the +victims were children who had attempted to collect the scorpions. + +The treatment usually employed in the case of bites by the more +poisonous forms is similar to that for the bite of venomous snakes. +First, a tight ligature is applied above the wound so as to stop the +flow of blood and lymph from that region. The wound is then freely +excised and treated with a strong solution of permanganate of potash, or +with lead and opium lotion. + +In recent years there have been many attempts to prepare an antivenom, +or antiserum comparable to what has been used so effectively in the case +of snake bites. The most promising of these is that of Todd (1909), +produced by the immunization of suitable animals. This antivenom proved +capable of neutralizing the venom when mixed _in vitro_ and also acts +both prophylactically and curatively in animals. Employed curatively in +man, it appears to have a very marked effect on the intense pain +following the sting, and the evidence so far indicates that its prompt +use greatly reduces the chance of fatal results. + + +THE SOLPUGIDA, OR SOLPUGIDS + +The SOLPUGIDA are peculiar spider-like forms which are distinguished +from nearly all other arachnids by the fact that they possess no true +cephalothorax, the last two leg-bearing segments being distinct, +resembling those of the abdomen in this respect. The first pair of legs +is not used in locomotion but seemingly functions as a second pair of +pedipalpi. Figure 12 illustrates the striking peculiarities of the +group. They are primarily desert forms and occur in the warm zones of +all countries. Of the two hundred or more species, Comstock lists twelve +as occurring in our fauna. These occur primarily in the southwest. + +[Illustration: 12. A solpugid (Eremobates cinerea). After Comstock.] + +The Solpugida have long borne a bad reputation and, regarding virulence, +have been classed with the scorpions. Among the effects of their bites +have been described painful swelling, gangrene, loss of speech, cramps, +delirium, unconsciousness and even death. Opposed to the numerous loose +accounts of poisoning, there are a number of careful records by +physicians and zoölogists which indicate clearly that the effects are +local and though they may be severe, they show not the slightest symptom +of direct poisoning. + +More important in the consideration of the question is the fact that +there are neither poison glands nor pores in the fangs for the exit of +any poisonous secretion. This is the testimony of a number of prominent +zoölogists, among whom is Dr. A. Walter, who wrote to Kobert at length +on the subject and whose conclusions are presented by him. + +However, it should be noted that the fangs are very powerful and are +used in such a manner that they may inflict especially severe wounds. +Thus, there may be more opportunity for secondary infection than is +usual in the case of insect wounds. + +The treatment of the bite of the Solpugida is, therefore, a matter of +preventing infection. The wound should be allowed to bleed freely and +then washed out with a 1:3000 solution of corrosive sublimate, and, if +severe, a wet dressing of this should be applied. If infection takes +place, it should be treated in the usual manner, regardless of its +origin. + + +THE ACARINA, OR MITES AND TICKS + +A number of the parasitic Acarina evidently secrete a specific poison, +presumably carried by the saliva, but in most cases its effect on man is +insignificant. There is an abundant literature dealing with the +poisonous effect of the bite of these forms, especially the ticks, but +until recently it has been confused by failure to recognize that various +species may transmit diseases of man, rather than produce injury through +direct poisoning. We shall therefore discuss the Acarina more especially +in subsequent chapters, dealing with parasitism and with disease +transmission. + +Nevertheless, after the evidence is sifted, there can be no doubt that +the bites of certain ticks may occasionally be followed by a direct +poisoning, which may be either local or general in its effects. Nuttall +(1908) was unable to determine the cause of the toxic effect, for, in +_Argas persicus_, the species most often implicated, he failed to get +the slightest local or general effect on experimental animals, from the +injection of an emulsion prepared by crushing three of the ticks. + +It seems clearly established that the bite of certain ticks may cause a +temporary paralysis, or even complete paralysis, involving the organs of +respiration or the heart, and causing death. In 1912, Dr. I. U. Temple, +of Pendleton, Oregon, reported several cases of what he called "acute +ascending paralysis" associated with the occurrence of ticks on the head +or the back of the neck. A typical severe case was that of a six year +old child, who had retired in her usual normal health. The following +morning upon arising she was unable to stand on her feet. She exhibited +paralysis extending to the knees, slight temperature, no pain, sensory +nerves normal, motor nerves completely paralyzed, reflexes absent. The +following day the paralysis had extended to the upper limbs, and before +night of the third day the nerves of the throat (hypoglossal) were +affected. The thorax and larynx were involved, breathing was labored, +she was unable to swallow liquids, phonation was impossible and she +could only make low, guttural sounds. At this stage, two ticks, fully +distended with blood, were found over the junction of the spinal column +with the occipital bones in the hollow depression. They were removed by +the application of undiluted creoline. Though the child's life was +despaired of, by the following morning she was very much improved. By +evening she was able to speak. The paralysis gradually receded, +remaining longest in the feet, and at the end of one week the patient +was able to go home. + +There was some doubt as to the exact species of tick implicated in the +cases which Dr. Temple reported, although the evidence pointed strongly +to _Dermacentor venustus_.[C] Somewhat later, Hadwen (1913) reported +that "tick paralysis" occurs in British Columbia, where it affects not +only man, but sheep and probably other animals. It is caused by the +bites of _Dermacentor venustus_ and was experimentally produced in lambs +and a dog (Hadwen and Nuttall, 1913). It is only when the tick begins to +engorge or feed rapidly, some days after it has become attached, that +its saliva produces pathogenic effects. + +Ulceration following tick bite is not uncommon. In many of the instances +it is due to the file-like hypostome, with its recurved teeth, being +left in the wound when the tick is forcibly pulled off. + + +THE MYRIAPODA, OR CENTIPEDES AND MILLIPEDES + +The old class, Myriapoda includes the DIPLOPODA, or millipedes, and the +CHILOPODA, or centipedes. The present tendency is to raise these groups +to the rank of classes. + + +The Diplopoda + +The DIPLOPODA, or millipedes (fig. 13), are characterized by the +presence of two pairs of legs to a segment. The largest of our local +myriapods belong to this group. They live in moist places, feeding +primarily on decaying vegetable matter, though a few species +occasionally attack growing plants. + +[Illustration: 13. A millipede. After Comstock.] + +The millipedes are inoffensive and harmless. _Julus terrestris_, and +related species, when irritated pour out over the entire body a +yellowish secretion which escapes from cutaneous glands. It is volatile, +with a pungent odor, and Phisalix (1900) has shown that it is an active +poison when injected into the blood of experimental animals. This, +however, does not entitle them to be considered as poisonous arthropods, +in the sense of this chapter, any more than the toad can be considered +poisonous to man because it secretes a venom from its cutaneous glands. + + +The Chilopoda + +[Illustration: 14. Two common centipedes. + +(_a_) Lithobius forficatus. After Comstock. + +(_b_) Scutigera forceps. Natural size; after Howard.] + +The CHILOPODA, or centipedes (fig. 14), unlike the millipedes, are +predaceous forms, and possess well developed poison glands for killing +their prey. These glands are at the base of the first pair of legs +(fig. 15), which are bent forward so as to be used in holding their +prey. The legs terminate in a powerful claw, at the tip of which is the +outlet of the poison glands. + +The poison is a limpid, homogeneous, slightly acid fluid, which +precipitates in distilled water. Briot (1904) extracted it from the +glands of _Scolopendra morsitans_, a species common in central France, +and found that it was actively venomous for the ordinary experimental +animals. A rabbit of two kilograms weight received an injection of three +cubic centimeters in the vein of the ear and died in a minute. A white +rat, weighing forty-eight grams, received one and a half cubic +centimeters in the hind leg. There was an almost immediate paralysis of +the leg and marked necrosis of the tissues. + +[Illustration: 15. Mandible of Scolopendra cingulata showing venom +gland. After Dubosq.] + +As for the effect on man, there is little foundation for the fear with +which centipedes are regarded. Our native species produce, at most, +local symptoms,--sometimes severe local pain and swelling,--but there is +no authentic record of fatal results. In the tropics, some of the +species attain a large size, _Scolopendra gigantea_ reaching a length of +nearly a foot. These forms are justly feared, and there is good evidence +that death sometimes, though rarely, results from their bite. + +One of the most careful accounts of death from the sting of the scorpion +is that of Linnell, (1914), which relates to a comparatively small +Malayan species, unfortunately undetermined. The patient, a coolie, aged +twenty, was admitted to a hospital after having been stung two days +previously on the left heel. For cure, the other coolies had made him +eat the head of the scorpion. On admission, the patient complained of +"things creeping all over the body". Temp. 102.8°. On the fourth day he +had paralysis of the legs, and on the fifth day motor paralysis to the +umbilicus, sensation being unaltered. On the sixth day there was +retention of the urine and on the ninth day (first test after third day) +sugar was present. On the thirteenth day the patient became comatose, +but could be roused to eat and drink. The temperature on the following +day fell below 95° and the patient was still comatose. Death fifteenth +day. + +Examination of the spinal (lumbar) cord showed acute disseminated +myelitis. In one part there was an acute destruction of the anterior +horn and an infiltration of round cells. In another portion Clarke's +column had been destroyed. The perivascular sheaths were crowded with +small round cells and the meninges were congested. Some of the cells of +the anterior horn were swollen and the nuclei eccentric; chromatolysis +had occurred in many of them. + +As for treatment, Castellani and Chalmers (1910), recommend bathing the +part well with a solution of ammonia (one in five, or one in ten). After +bathing, apply a dressing of the same alkali or, if there is much +swelling and redness, an ice-bag. If necessary, hypodermic injections of +morphine may be given to relieve the pain. At a later period +fomentations may be required to reduce the local inflammation. + + +THE HEXAPODA OR TRUE INSECTS + +There are a number of HEXAPODA, or true insects, which are, in one way +or another, poisonous to man. These belong primarily to the orders +Hemiptera, or true bugs; Lepidoptera, or butterflies and moths (larval +forms); Diptera, or flies; Coleoptera, or beetles; and Hymenoptera, or +ants, bees, and wasps. There are various ways in which they may be +poisonous. + +1. _Piercing_ or _biting_ forms may inject an irritating or poisonous +saliva into the wound caused by their mouth-parts. + +2. _Stinging forms_ may inject a poison, from glands at the caudal end +of the abdomen, into wounds produced by a specially modified ovipositer, +the _sting_. + +3. _Nettling_ properties may be possessed by the hairs of the insect. + +4. _Vescicating_, or _poisonous blood plasma_, or _body fluids_ are +known to exist in a large number of species and may, under exceptional +circumstances, affect man. + +For convenience of discussion, we shall consider poisonous insects under +these various headings. In this, as in the preceding discussion, no +attempt will be made to give an exhaustive list of the poisonous forms. +Typical instances will be selected and these will be chosen largely from +North American species. + + +PIERCING OR BITING INSECTS POISONOUS TO MAN + + +HEMIPTERA + +Several families of the true bugs include forms which, while normally +inoffensive, are capable of inflicting painful wounds on man. In these, +as in all of the Hemiptera, the mouth-parts are modified to form an +organ for piercing and sucking. This is well shown by the accompanying +illustration (fig. 16). + +The upper lip, or _labrum_, is much reduced and immovable, the lower +lip, or _labium_, is elongated to form a jointed sheath, within which +the lance-like mandibles and maxillæ are enclosed. The mandibles are +more or less deeply serrate, depending on the species concerned. + +[Illustration: 16. Beak of hemipteron.] + +The poison is elaborated by the salivary glands, excepting, possibly, in +_Belostoma_ where Locy is inclined to believe that it is secreted by the +maxillary glands. The salivary glands of the Hemiptera have been the +subject of much study but the most recent, comprehensive work has been +done by Bugnion and Popoff, (1908 and 1910) to whose text the reader is +referred for details. + +The Hemiptera have two pairs of salivary glands: the _primary gland_, of +which the efferent duct leads to the salivary syringe, and the +_accessory gland_, of which the very long and flexuous duct empties into +the primary duct at its point of insertion. Thus, when one observes the +isolated primary gland it appears as though it had efferent ducts +inserted at the same point. In _Nepa_ and the _Fulgoridæ_ there are two +accessory glands and therefore apparently three ducts at the same point +on the primary gland. The _ensemble_ differs greatly in appearance in +different species but we shall show here Bugnion and Popoff's figure of +the apparatus of _Notonecta maculata_, a species capable of inflicting a +painful bite on man (fig. 17). + +[Illustration: 17. Salivary glands of Notonecta maculata. After Bugnion +and Popoff.] + +[Illustration: 18. Pharyngeal syringe or salivary pump of Fulgora +maculata. After Bugnion and Popoff.] + +[Illustration: 19. Heteroptera, (_a_) Melanolestes picipes; (_b_) +Notonecta undulata; (_c_, _d_) Aradus robustus (_c_) adult, (_d_) nymph, +much enlarged; (_e_) Arilus cristatus; (_f_) Belostoma americana; (_g_) +Nabis (Coriscus) subcoleoptratus, enlarged; (_h_) Cimex lectularius, +(_i_) Oeciacus vicarius, much enlarged; (_j_) Lyctocoris fitchii, much +enlarged. After Lugger.] + +Accessory to the salivary apparatus there is on the ventral side of the +head, underneath the pharynx, a peculiar organ which the Germans have +called the "Wanzenspritze," or syringe. The accompanying figure of the +structure in _Fulgora maculata_ (fig. 18) shows its relation to the +ducts of the salivary glands and to the beak. It is made up of a +dilatation forming the body of the pump, in which there is a chitinous +piston. Attached to the piston is a strong retractor muscle. The +function of the salivary pump is to suck up the saliva from the salivary +ducts and to force it out through the beak. + +Of the Hemiptera reported as attacking man, we shall consider briefly +the forms most frequently noted. + + +The NOTONECTIDÆ, or _back swimmers_, (fig. 19_b_) are small, aquatic +bugs that differ from all others in that they always swim on their +backs. They are predaceous; feeding on insects and other small forms. +When handled carelessly they are able to inflict a painful bite, which +is sometimes as severe as the sting of a bee. In fact, they are known in +Germany as "Wasserbienen." + + +The BELOSTOMATIDÆ, or _giant water bugs_, (fig. 19_f_) include the +largest living Hemiptera. They are attracted to lights and on account of +the large numbers which swarm about the electric street lamps in some +localities they have received the popular name "electric light bugs." +Our largest representatives in the northern United States belong to the +two genera _Belostoma_ and _Banacus_, distinguished from each other by +the fact that _Belostoma_ has a groove on the under side of the femur of +the front leg, for the reception of the tibia. + +The salivary glands of Belostoma were figured by Leidy (1847) and later +were studied in more detail by Locy (1884). There are two pairs of the +glands, those of one pair being long and extending back as far as the +beginning of the abdomen, while the others are about one-fourth as long. +They lie on either side of the oesophagus. On each side of the +oesophagus there is a slender tube with a sigmoid swelling which may +serve as a poison reservoir. In addition to this salivary system, there +is a pair of very prominent glands on the ventral side of the head, +opening just above the base of the beak. These Locy has called the +"cephalic glands" and he suggests that they are the source of the +poison. They are the homologues of the maxillary glands described for +other Hemiptera, and it is by no means clear that they are concerned +with the production of venom. It seems more probable that in +_Belostoma_, as in other Hemiptera, it is produced by the salivary +glands, though the question is an open one. + +The Belostomatidæ feed not only on insects, but on small frogs, fish, +salamanders and the like. Matheson (1907) has recorded the killing of a +good-sized bird by _Belostoma americana_. A woodpecker, or flicker, was +heard to utter cries of distress, and fluttered and fell from a tree. On +examination it was found that a bug of this species had inserted its +beak into the back part of the skull and was apparently busily engaged +in sucking the blood or brains of the bird. Various species of +_Belostoma_ have been cited as causing painful bites in man. We can +testify from personal experience that the bite of _Belostoma americana_ +may almost immediately cause severe, shooting pains that may extend +throughout the arm and that they may be felt for several days. + +[Illustration: 20. Reduvius (Opsicoetus) personatus. (×2).] + +Relief from the pain may be obtained by the use of dilute ammonia, or a +menthol ointment. In the not uncommon case of secondary infection the +usual treatment for that should be adopted. + +[Illustration: 21. (_a_) Reduvius personatus, nymph. Photograph by M. V. +S.] + + +The REDUVIIDÆ, or _assassin-bugs_ are capable of inflicting very painful +wounds, as most collectors of Hemiptera know to their sorrow. Some +species are frequently to be found in houses and outhouses and Dr. +Howard suggests that many of the stories of painful spider bites relate +to the attack of these forms. + +[Illustration: 21. (_b_) Reduvius personatus, adult (×2) Photograph by +M. V. S.] + +An interesting psychological study was afforded in the summer of 1899, +by the "kissing-bug" scare which swept over the country. It was reported +in the daily papers that a new and deadly bug had made its appearance, +which had the unpleasant habit of choosing the lips or cheeks for its +point of attack on man. So widespread were the stories regarding this +supposedly new insect that station entomologists all over the country +began to receive suspected specimens for identification. At Cornell +there were received, among others, specimens of stone-flies, may-flies +and even small moths, with inquiries as to whether they were +"kissing-bugs." + +[Illustration: 22. Rasahus biguttatus. (×2). After Howard.] + +Dr. L. O. Howard has shown that the scare had its origin in newspaper +reports of some instances of bites by either _Melanolestes picipes_ +(fig. 19a) or _Opsicoetes personatus_ (fig. 20), in the vicinity of +Washington, D. C. He then discusses in considerable detail the more +prominent of the Reduviidæ which, with greater or less frequency pierce +the skin of human beings. These are _Opsicoetes personatus_, +_Melanolestes picipes_, _Coriscus subcoleoptratus_ (fig. 19_g_), +_Rasahus thoracicus_, _Rasahus biguttatus_ (fig. 22), _Conorhinus +sanguisugus_ (fig. 71), and _Melanolestes abdominalis_ (fig. 23). + +[Illustration: 23. Melanolestes abdominalis (×2). After Marlatt.] + +One of the most interesting of these species is _Reduvius personatus_, +(= _Opsicoetus personatus_), which is popularly known as the "masked +bed-bug hunter." It owes this name to the fact that the immature nymphs +(fig. 21) have their bodies and legs completely covered by dust and +lint, and that they are supposed to prey upon bed-bugs. LeConte is +quoted by Howard as stating that "This species is remarkable for the +intense pain caused by its bite. I do not know whether it ever willingly +plunges its rostrum into any person, but when caught, or unskilfully +handled it always stings. In this case the pain is almost equal to the +bite of a snake, and the swelling and irritation which result from it +will sometimes last for a week." + +A species which very commonly attacks man is _Conorhinus sanguisugus_, +the so-called "big bed-bug" of the south and southern United States. It +is frequently found in houses and is known to inflict an exceedingly +painful bite. As in the case of a number of other predaceous Hemiptera, +the salivary glands of these forms are highly developed. The effect of +the bite on their prey and, as Marlatt has pointed out, the constant and +uniform character of the symptoms in nearly all cases of bites in man, +clearly indicate that their saliva contains a specific substance. No +satisfactory studies of the secretions have been made. On the other +hand, Dr. Howard is doubtless right in maintaining that the very serious +results which sometimes follow the bite are due to the introduction of +extraneous poison germs. This is borne out by the symptoms of most of +the cases cited in literature and also by the fact that treatment with +corrosive sublimate, locally applied to the wound, has yielded favorable +results. + + +OTHER HEMIPTERA REPORTED AS POISONOUS TO MAN--A large number of other +Hemiptera have been reported as attacking man. Of these, there are +several species of Lygæidæ, Coreidæ, and Capsidæ. Of the latter, _Lygus +pratensis_, the tarnished plant-bug, is reported by Professor Crosby as +sucking blood. _Orthotylus flavosparsus_ is another Capsid which has +been implicated. _Empoasca mali_ and _Platymetopius acutus_ of the +Jassidæ have also been reported as having similar habits. + +Whenever the periodical cicada or "seventeen-year locust" becomes +abundant, the newspapers contain accounts of serious results from its +bites. The senior author has made scores of attempts to induce this +species to bite and only once successfully. At that time the bite was in +no wise more severe than a pin-prick. A student in our department +reports a similar experience. There is no case on record which bears +evidence of being worthy of any credence, whatsoever. + +Under the heading of poisonous Hemiptera we might consider the bed-bugs +and the lice. These will be discussed later, as parasites and as +carriers of disease, and therefore need only be mentioned here. + + +DIPTERA + +Several species of blood-sucking Diptera undoubtedly secrete a saliva +possessing poisonous properties. Chief among these are the Culicidæ, or +mosquitoes, and the Simuliidæ, or black-flies. As we shall consider +these forms in detail under the heading of parasitic species and +insects transmitting disease, we shall discuss here only the poison of +the mosquitoes. + +It is well known that mosquitoes, when they bite, inject into the wound +a minute quantity of poison. The effect of this varies according to the +species of mosquito and also depends very much on the susceptibility of +the individual. Soon after the bite a sensation of itching is noticed +and often a wheal, or eminence, is produced on the skin, which may +increase to a considerable swelling. The scratching which is induced may +cause a secondary infection and thus lead to serious results. Some +people seem to acquire an immunity against the poison. + +The purpose of this irritating fluid may be, as Reaumur suggested, to +prevent the coagulation of the blood and thus not only to cause it to +flow freely when the insect bites but to prevent its rapid coagulation +in the stomach. Obviously, it is not developed as a protective fluid, +and its presence subjects the group to the additional handicap of the +vengeance of man. + +[Illustration: 24. Diagram of a longitudinal section of a mosquito.] + +As to the origin of the poison, there has been little question, until +recent years, that it was a secretion from the salivary glands. +Macloskie (1888) showed that each gland is subdivided into three lobes, +the middle of which differs from the others in having evenly granulated +contents and staining more deeply than the others (fig. 24). This middle +lobe he regarded as the source of the poison. Bruck, (1911), by the use +of water, glycerine, chloroform, and other fluids, extracted from the +bodies of a large number of mosquitoes a toxine which he calls +_culicin_. This he assumes comes from the salivary glands. Animal +experimentation showed that this extract possessed hemolytic powers. +Inoculated into the experimenter's own skin it produced lesions which +behaved exactly as do those of mosquito bites. + +Similarly, most writers on the subject have concurred with the view that +the salivary glands are the source of the poison. However, recent work, +especially that of Nuttall and Shipley (1903), and Schaudinn (1904), has +shown that the evidence is by no means conclusive. Nuttall dissected out +six sets (thirty-six acini) of glands from freshly killed _Culex +pipiens_ and placed them in a drop of salt solution. The drop was +allowed to dry, it being thought that the salt crystals would facilitate +the grinding up of the glands with the end of a small glass rod, this +being done under microscopic control. After grinding up, a small drop of +water was added of the size of the original drop of saline, and an equal +volume of human blood taken from the clean finger-tip was quickly mixed +therewith, and the whole drawn up into a capillary tube. Clotting was +not prevented and no hemolysis occurred. Salivary gland emulsion added +to a dilute suspension of corpuscles did not lead to hemolysis. This +experiment was repeated a number of times, with slight modification, but +with similar results. The data obtained from the series "do not support +the hypothesis that the salivary glands, at any rate in _Culex pipiens_, +contain a substance which prevents coagulation." + +Much more detailed, and the more important experiments made along this +line, are those of Schaudinn (1904). The results of these experiments +were published in connection with a technical paper on the alternation +of generations and of hosts in _Trypanosoma_ and _Spirochæta_, and for +this reason seem to have largely escaped the notice of entomologists. +They are so suggestive that we shall refer to them in some detail. + +Schaudinn observed that the three oesophageal diverticula (commonly, +but incorrectly, known as the "sucking stomach") (fig. 24) usually +contain large bubbles of gas and in addition, he always found yeast +cells. On the ground of numerous observations, Schaudinn was convinced +that these yeast plants are normal and constant commensals of the +insect. He regarded them as the cause of the gas bubbles to be found in +diverticula. It was found that as the insect fed, from time to time the +abdomen underwent convulsive contractions which resulted in the emptying +of the oesophageal diverticula and the salivary glands through blood +pressure. + +In order to test the supposed toxic action of the salivary glands, +Schaudinn repeatedly introduced them under his skin and that of his +assistant, in a drop of salt solution, and never obtained a suggestion +of the irritation following a bite of the insect, even though the glands +were carefully rubbed to fragments after their implantation. Like +Nuttall, he failed to get satisfactory evidence that the secretion of +the salivary glands retarded coagulation of the blood. + +He then carefully removed the oesophageal diverticula with their +content of yeast and introduced them into an opening in the skin of the +hand. Within a few seconds there was noticeable the characteristic +itching irritation of the mosquito bite; and in a short time there +appeared reddening and typical swelling. This was usually much more +severe than after the usual mosquito bite, and the swelling persisted +and itched longer. This was because by the ordinary bite of the mosquito +most of the yeast cells are again sucked up, while in these experiments +they remained in the wound. These experiments were repeated a number of +times on himself, his assistant and others, and always with the same +result. From them Schaudinn decided that the poisonous action of the +mosquito bite is caused by an enzyme from a commensal fungus. These +conclusions have not, as yet, been satisfactorily tested. + +Relief from the effect of the mosquito bite may be obtained by bathing +the swellings with weak ammonia or, according to Howard, by using moist +soap. The latter is to be rubbed gently on the puncture and is said to +speedily allay the irritation. Howard also quotes from the _Journal of +Tropical Medicine and Hygiene_ to the effect that a few drops of a +solution of thirty to forty grains of iodine to an ounce of saponated +petroleum rubbed into the mosquito bite, or wasp sting, allay the pain +instantaneously. + +Methods of mosquito control will be discussed later, in considering +these insects as parasites and as carriers of disease. + + +STINGING INSECTS + +The stinging insects all belong to the order HYMENOPTERA. In a number of +families of this group the ovipositor is modified to form a sting and is +connected with poison-secreting glands. We shall consider the apparatus +of the honey-bee and then make briefer reference to that of other forms. + + +APIS MELLIFICA, THE HONEY BEE--The sting of the worker honey-bee is +situated within a so-called sting chamber at the end of the abdomen. +This chamber is produced by the infolding of the greatly reduced and +modified eighth, ninth and tenth abdominal segments into the seventh.[D] +From it the dart-like sting can be quickly exserted. + +The sting (fig. 25) is made up of a central shaft, ventro-laterad of +which are the paired _lancets_, or darts, which are provided with sharp, +recurved teeth. Still further laterad lie the paired whitish, +finger-like _sting palpi_. Comparative morphological as well as +embryological studies have clearly established that these three parts +correspond to the three pairs of gonopophyses of the ovipositor of more +generalized insects. + +[Illustration: 25. Sting of a honey bee. _Psn Sc_, base of acid poison +gland; _B Gl_, alkaline poison gland; _Stn Plp_, sting palpi; _Sh B_, +bulb of sting; _Sh A_, basal arm; _Lct_, lancets or darts; _Sh s_, shaft +of sting. Modified from Snodgrass.] + +[Illustration: 26. Poison apparatus of a honey bee. Modified from +Snodgrass.] + +An examination of the internal structures (fig. 26) reveals two distinct +types of poison glands, the acid-secreting and the alkaline-secreting +glands, and a prominent poison reservoir. In addition, there is a small +pair of accessory structures which have been called lubricating glands, +on account of the supposed function of their product. The acid-secreting +gland empties into the distal end of the poison reservoir which in turn +pours the secretion into the muscular bulb-like enlargement at the base +of the shaft. The alkaline secreting gland empties into the bulb ventrad +of the narrow neck of the reservoir. + +The poison is usually referred to as formic acid. That it is not so +easily explained has been repeatedly shown and is evidenced by the +presence of the two types of glands. Carlet maintains that the product +of either gland is in itself innocent,--it is only when they are +combined that the toxic properties appear. + +The most detailed study of the poison of the honey-bee is that of Josef +Langer (1897), who in the course of his work used some 25,000 bees. +Various methods of obtaining the active poison for experimental purposes +were used. For obtaining the pure secretion, bees were held in the +fingers and compressed until the sting was exserted, when a clear drop +of the poison was visible at its tip. This was then taken up in a +capillary tube or dilute solutions obtained by dipping the tip of the +sting into a definite amount of distilled water. + +An aqueous solution of the poison was more readily obtained by pulling +out the sting and poison sacs by means of forceps, and grinding them up +in water. The somewhat clouded fluid was then filtered one or more +times. For obtaining still greater quantities, advantage was taken of +the fact that while alcohol coagulates the poison, the active principle +remains soluble in water. Hence the stings with the annexed glands where +collected in 96 per cent alcohol, after filtering off of the alcohol +were dried at 40° C., then rubbed to a fine powder and this was +repeatedly extracted with water. Through filtering of this aqueous +extract there was obtained a yellowish-brown fluid which produced the +typical reactions, according to concentration of the poison. + +The freshly expelled drop of poison is limpid, of distinct acid +reaction, tastes bitter and has a delicate aromatic odor. On +evaporation, it leaves a sticky residue, which at 100 degrees becomes +fissured, and suggests dried gum arabic. The poison is readily soluble +in water and possesses a specific gravity of 1.1313. On drying at room +temperature, it leaves a residue of 30 per cent, which has not lost in +poisonous action or in solubility. In spite of extended experiments, +Langer was unable to determine the nature of the active principle. He +showed that it was not, as had been supposed, an albuminous body, but +rather an organic base. + +The pure poison, or the two per cent aqueous solution, placed on the +uninjured skin showed absolutely no irritating effect, though it +produced a marked reaction on the mucus membrane of the nose or eye. A +single drop of one-tenth per cent aqueous solution of the poison brought +about a typical irritation in the conjunctiva of the rabbit's eye. On +the other hand, the application of a drop of the poison, or its +solution, to the slightest break in the skin, or by means of a needle +piercing the skin, produced typical effects. There is produced a local +necrosis, in the neighborhood of which there is infiltration of +lymphocytes, oedema, and hyperæmia. + +The effect of the sting on man (fig. 27) is usually transitory but there +are some individuals who are made sick for hours, by a single sting. +Much depends, too, on the place struck. It is a common experience that +an angry bee will attempt to reach the eye of its victim and a sting on +the lid may result in severe and prolonged swelling. In the case of a +man stung on the cheek, Legiehn observed complete aphonia and a breaking +out of red blotches all over the body. A sting on the tongue has been +known to cause such collateral oedema as to endanger life through +suffocation. Cases of death of man from the attacks of bees are rare but +are not unknown. Such results are usually from a number of stings but, +rarely, death has been known to follow a single sting, entering a blood +vessel of a particularly susceptible individual. + +[Illustration: Effect of bee stings. After Root.] + +It is clearly established that partial immunity from the effects of the +poison may be acquired. By repeated injections of the venom, mice have +been rendered capable of bearing doses that certainly would have killed +them at first. It is a well-known fact that most bee-keepers become +gradually hardened to the stings, so that the irritation and the +swelling become less and less. Some individuals have found this immunity +a temporary one, to be reacquired each season. A striking case of +acquired immunity is related by the Roots in their "A B C and X Y Z of +Bee Culture." The evidence in the case is so clear that it should be +made more widely available and hence we quote it here. + +A young man who was determined to become a bee-keeper, was so +susceptible to the poison that he was most seriously affected by a +single sting, his body breaking out with red blotches, breathing growing +difficult, and his heart action being painfully accelerated. "We finally +suggested taking a live bee and pressing it on the back of his hand +until it merely pierced his skin with the sting, then immediately +brushing off both bee and sting. This was done and since no serious +effect followed, it was repeated inside of four or five days. This was +continued for some three or four weeks, when the patient began to have a +sort of itching sensation all over his body. The hypodermic injections +of bee-sting poison were then discontinued. At the end of a month they +were repeated at intervals of four or five days. Again, after two or +three weeks the itching sensation came on, but it was less pronounced. +The patient was given a rest of about a month, when the doses were +repeated as before." By this course of treatment the young man became so +thoroughly immunized that neither unpleasant results nor swelling +followed the attacks of the insects and he is able to handle bees with +the same freedom that any experienced bee-keeper does. + +In an interesting article in the _Entomological News_ for November, +1914, J. H. Lovell calls attention to the fact that "There has been a +widespread belief among apiarists that a beekeeper will receive more +stings when dressed in black than when wearing white clothing. A large +amount of evidence has been published in the various bee journals +showing beyond question that honey-bees under certain conditions +discriminate against black. A few instances may be cited in +illustration. Of a flock of twelve chickens running in a bee-yard seven +black ones were stung to death, while five light colored ones escaped +uninjured. A white dog ran among the bee-hives without attracting much +attention, while at the same time a black dog was furiously assailed by +the bees. Mr. J. D. Byer, a prominent Canadian beekeeper, relates that a +black and white cow, tethered about forty feet from an apiary, was one +afternoon attacked and badly stung by bees. On examination it was found +that the black spots had five or six stings to one on the white. All +noticed this fact, although no one was able to offer any explanation. A +white horse is in much less danger of being stung, when driven near an +apiary, than a black one. It has, indeed, been observed repeatedly that +domestic animals of all kinds, if wholly or partially black, are much +more liable to be attacked by bees, if they wander among the hives, than +those which are entirely white." + +In order to test the matter experimentally, the following series of +experiments was performed. In the language of the investigator: + +"On a clear, warm day in August I dressed wholly in white with the +exception of a black veil. Midway on the sleeve of my right arm there +was sewed a band of black cloth ten inches wide. I then entered the +bee-yard and, removing the cover from one of the hives, lifted a piece +of comb with both hands and gently shook it. Instantly many of the bees +flew to the black band, which they continued to attack as long as they +were disturbed. Not a single bee attempted to sting the left sleeve, +which was of course entirely white, and very few even alighted upon it." + +"This experiment was repeated a second, third and fourth time; in each +instance with similar results. I estimated the number of bees on the +band of black cloth at various moments was from thirty to forty; it was +evident from their behavior that they were extremely irritable. To the +left white sleeve and other portions of my clothing they paid very +little attention; but the black veil was very frequently attacked." + +"A few days later the experiments were repeated, but the band of black +cloth, ten inches wide, was sewed around my left arm instead of around +the right arm as before. When the bees were disturbed, after the hive +cover had been removed, they fiercely attacked the band of black cloth +as in the previous experiences; but the right white sleeve and the white +suit were scarcely noticed. At one time a part of the black cloth was +almost literally covered with furiously stinging bees, and the black +veil was assailed by hundreds. The bees behaved in a similar manner when +a second hive on the opposite side of the apiary was opened." + +"A white veil which had been procured for this purpose, was next +substituted for the black veil. The result was most surprising, for, +whereas in the previous experiments hundreds of bees had attacked the +black veil, so few flew against the white veil as to cause me no +inconvenience. Undoubtedly beekeepers will find it greatly to their +advantage to wear white clothing when working among their colonies of +bees and manipulating the frames of the hives." + +When a honey-bee stings, the tip of the abdomen, with the entire sting +apparatus, is torn off and remains in the wound. Here the muscles +continue to contract, for some minutes, forcing the barbs deeper and +deeper into the skin, and forcing out additional poison from the +reservoir. + +Treatment, therefore, first consists in removing the sting without +squeezing out additional poison. This is accomplished by lifting and +scraping it out with a knife-blade or the fingernail instead of grasping +and pulling it out. Local application of alkalines, such as weak +ammonia, are often recommended on the assumption that the poison is an +acid to be neutralized on this manner, but these are of little or no +avail. They should certainly not be rubbed in, as that would only +accelerate the absorption of the poison. The use of cloths wrung out in +hot water and applied as hot as can be borne, affords much relief in the +case of severe stings. The application of wet clay, or of the end of a +freshly cut potato is sometimes helpful. + +In extreme cases, where there is great susceptibility, or where there +may have been many stings, a physician should be called. He may find +strychnine injections or other treatment necessary, if general symptoms +develop. + +[Illustration: 28. The poison apparatus of Formica. Wheeler, after +Forel.] + + +OTHER STINGING FORMS--Of the five thousand, or more, species of bees, +most possess a sting and poison apparatus and some of the larger forms +are capable of inflicting a much more painful sting than that of the +common honey-bee. In fact, some, like the bumble bees, possess the +advantage that they do not lose the sting from once using it, but are +capable of driving it in repeatedly. In the tropics there are found many +species of stingless bees but these are noted for their united efforts +to drive away intruders by biting. Certain species possess a very +irritating saliva which they inject into the wounds. + +The ants are not ordinarily regarded as worthy of consideration under +the heading of "stinging insects" but as a matter of fact, most of them +possess well developed stings and some of them, especially in the +tropics, are very justly feared. Even those which lack the sting possess +well-developed poison glands and the parts of the entire stinging +apparatus, in so far as it is developed in the various species, may +readily be homologized with those of the honey-bee. + +The ants lacking a sting are those of the subfamily CAMPONOTINÆ, which +includes the largest of our local species. It is an interesting fact +that some of these species possess the largest poison glands and +reservoir (fig. 28) and it is found that when they attack an enemy they +bring the tip of the abdomen forward and spray the poison in such a way +that it is introduced into the wound made by the powerful mandibles. + +[Illustration: 29. A harmless, but much feared larva, the "tomato worm." +Natural size. Photograph by M. V. S.] + +More feared than any of the other Hymenoptera are the hornets and wasps. +Of these there are many species, some of which attain a large size and +are truly formidable. Phisalix (1897), has made a study of the venom of +the common hornet and finds that, like the poison of the honey-bee, it +is neither an albuminoid nor an alkaloid. Its toxic properties are +destroyed at 120° C. Phisalix also says that the venom is soluble in +alcohol. If this be true, it differs in this respect from that of the +bee. An interesting phase of the work of Phisalix is that several of her +experiments go to show that the venom of hornets acts as a vaccine +against that of vipers. + + +NETTLING INSECTS + +So far, we have considered insects which possess poison glands connected +with the mouth-parts or a special sting and which actively inject their +poison into man. There remain to be considered those insects which +possess poisonous hairs or body fluids which, under favorable +circumstances, may act as poisons. To the first of these belong +primarily the larvæ of certain Lepidoptera. + + +LEPIDOPTERA + +[Illustration: 30. Another innocent but much maligned caterpillar, the +larva of the Regal moth. Photograph by M. V. S.] + +When we consider the reputedly poisonous larvæ of moths and butterflies, +one of the first things to impress us is that we cannot judge by mere +appearance. Various species of Sphingid, or hawk-moth larvæ, bear at the +end of the body a chitinous horn, which is often referred to as a +"sting" and regarded as capable of inflicting dangerous wounds. It would +seem unnecessary to refer to this absurd belief if it were not that each +summer the newspapers contain supposed accounts of injury from the +"tomato worm" (fig. 29) and others of this group. The grotesque, spiny +larva (fig. 30) of one of our largest moths, _Citheronia regalis_ is +much feared though perfectly harmless, and similar instances could be +multiplied. + +[Illustration: 31. The flannel moth (Lagoa crispata). (_a_) Poisonous +larva.] + +[Illustration: 31. (_b_) Adult. Enlarged. Photographs by M. V. S.] + +But if the larvæ are often misjudged on account of their ferocious +appearance, the reverse may be true. A group of most innocent looking +and attractive caterpillars is that of the flannel-moth larvæ, of which +_Lagoa crispata_ may be taken as an example. Its larva (fig. 31) has a +very short and thick body, which is fleshy and completely covered and +hidden by long silken hairs of a tawny or brown color, giving a convex +form to the upper side. Interspersed among these long hairs are +numerous short spines connected with underlying hypodermal poison +glands. These hairs are capable of producing a marked nettling effect +when they come in contact with the skin. This species is found in our +Atlantic and Southern States. Satisfactory studies of its poisonous +hairs and their glands have not yet been made. + +[Illustration: 32 The poisonous saddle back caterpillar. Empretia +(Sibine) stimulea. Photograph by M. V. S.] + +[Illustration: 33_a_. Io moth larvæ on willow. Photograph by M. V. S.] + +_Sibine stimulea (Empretia stimulea)_, or the saddle-back caterpillar +(fig. 32), is another which possesses nettling hairs. This species +belongs to the group of Eucleidæ, or slug caterpillars. It can be +readily recognized by its flattened form, lateral, bristling spines and +by the large green patch on the back resembling a saddle-cloth, while +the saddle is represented by an oval, purplish-brown spot. The small +spines are venomous and affect some persons very painfully. The larva +feeds on the leaves of a large variety of forest trees and also on +cherry, plum, and even corn leaves. It is to be found throughout the +Eastern and Southern United States. + +[Illustration: 33_b_. Io moth. Full grown larva. Photograph by M. V. S.] + +[Illustration: 34. Io moth. Adult. Photograph by M. V. S.] + +_Automeris io_ is the best known of the nettling caterpillars. It is the +larva of the Io moth, one of the Saturniidæ. The mature caterpillar, +(fig. 33), which reaches a length of two and one-half inches, is of a +beautiful pale green with sublateral stripes of cream and red color and +a few black spines among the green ones. The green radiating spines give +the body a mossy appearance. They are tipped with a slender chitinous +hair whose tip is readily broken off in the skin and whose poisonous +content causes great irritation. Some individuals are very susceptible +to the poison, while others are able to handle the larvæ freely without +any discomfort. The larvæ feed on a wide range of food plants. They are +most commonly encountered on corn and on willow, because of the +opportunities for coming in contact with them. + +[Illustration: 35. Larva of brown-tail moth. (Natural size). Photograph +by M. V. S.] + +The larvæ of the brown-tail moth (_Euproctis chrysorrhoea_) (fig. 35 +and 36), where they occur in this country, are, on account of their +great numbers, the most serious of all poisonous caterpillars. It is not +necessary here, to go into details regarding the introduction of this +species from Europe into the New England States. This is all available +in the literature from the United States Bureau of Entomology and from +that of the various states which are fighting the species. Suffice to +say, there is every prospect that the pest will continue to spread +throughout the Eastern United States and Canada and that wherever it +goes it will prove a direct pest to man as well as to his plants. + +Very soon after the introduction of the species there occurred in the +region where it had gained a foothold, a mysterious dermatitis of man. +The breaking out which usually occurred on the neck or other exposed +part of the body was always accompanied by an intense itching. It was +soon found that this dermatitis was caused by certain short, barbed +hairs of the brown-tail caterpillars and that not only the caterpillars +but their cocoons and even the adult female moths might harbor these +nettling hairs and thus give rise to the irritation. In many cases the +hairs were wafted to clothing on the line and when this was worn it +might cause the same trouble. Still worse, it was found that very +serious internal injury was often caused by breathing or swallowing the +poisonous hairs. + +[Illustration: 36. Browntail moths. One male and two females. Photograph +by M. V. S.] + +The earlier studies seemed to indicate that the irritation was purely +mechanical in origin, the result of the minute barbed hairs working into +the skin in large numbers. Subsequently, however, Dr. Tyzzer (1907) +demonstrated beyond question that the trouble was due to a poison +contained in the hairs. In the first place, it is only the peculiar +short barbed hairs which will produce the dermatitis when rubbed on the +skin, although most of the other hairs are sharply barbed. Moreover, it +was found that in various ways the nettling properties could be +destroyed without modifying the structure of the hairs. This was +accomplished by baking for one hour at 110° C, by warming to 60° C in +distilled water, or by soaking in one per cent. or in one-tenth per +cent. of potassium hydrate or sodium hydrate. The most significant part +of his work was the demonstration of the fact that if the nettling +hairs are mingled with blood, they immediately produce a change in the +red corpuscles. These at once become coarsely crenated, and the roleaux +are broken up in the vicinity of the hair (fig. 37_b_). The corpuscles +decrease in size, the coarse crenations are transformed into slender +spines which rapidly disappear, leaving the corpuscles in the form of +spheres, the light refraction of which contrasts them sharply with the +normal corpuscles. The reaction always begins at the basal sharp point +of the hair. It could not be produced by purely mechanical means, such +as the mingling of minute particles of glass wool, the barbed hairs of a +tussock moth, or the other coarser hairs of the brown-tail, with the +blood. + +[Illustration: 37. (_a_) Ordinary hairs and three poison hairs of +subdorsal and lateral tubercles of the larva of the browntail moth. +Drawing by Miss Kephart.] + +The question of the source of the poison has been studied in our +laboratory by Miss Cornelia Kephart. She first confirmed Dr. Tyzzer's +general results and then studied carefully fixed specimens of the larvæ +to determine the distribution of the hairs and their relation to the +underlying tissues. + +[Illustration: 37. (_b_) Effect of the poison on the blood corpuscles of +man. After Tyzzer.] + +The poison hairs (fig. 37), are found on the subdorsal and lateral +tubercles (fig. 38), in bunches of from three to twelve on the minute +papillæ with which the tubercles are thickly covered. The underlying +hypodermis is very greatly thickened, the cells being three or four +times the length of the ordinary hypodermal cells and being closely +crowded together. Instead of a pore canal through the cuticula for each +individual hair, there is a single pore for each papillæ on a tubercle, +all the hairs of the papilla being connected with the underlying cells +through the same pore canal, (figs. 39 and 40). + +[Illustration: 38. Cross section of the larva of the browntail moth +showing the tubercles bearing the poison hairs. Drawing by Miss +Kephart.] + +The hypodermis of this region is of two distinct types of cells. First, +there is a group of slender fusiform cells, one for each poison hair on +the papilla, which are the trichogen, or hair-formative cells. They are +crowded to one side and towards the basement membrane by a series of +much larger, and more prominent cells (fig. 40), of which there is a +single one for each papilla. These larger cells have a granular +protoplasm with large nuclei and are obviously actively secreting. They +are so characteristic in appearance as to leave no question but that +they are the true poison glands. + +[Illustration: 39. Epithelium underlying poison hairs of the larva of +the browntail moth. Drawing by Miss Kephart.] + +Poisonous larvæ of many other species have been reported from Europe and +especially from the tropics but the above-mentioned species are the more +important of those occurring in the United States and will serve as +types. It should be noted in this connection that through some curious +misunderstanding Goeldi (1913) has featured the larva of _Orgyia +leucostigma_, the white-marked tussock moth, as the most important of +the poisonous caterpillars of this country. Though there are occasional +reports of irritation from its hairs such cases are rare and there is no +evidence that there is any poison present. Indeed, subcutaneous +implantation of the hairs leads to no poisoning, but merely to temporary +irritation. + +[Illustration: 40. Same as figure 39, on larger scale.] + +Occasionally, the hairs of certain species of caterpillars find +lodgement in the conjunctiva, cornea, or iris of the eye of man and give +rise to the condition known as _opthalmia nodosa_. The essential feature +of this trouble is a nodular conjunctivitis which simulates tuberculosis +of the conjunctiva and hence has been called _pseudo-tubercular_. It may +be distinguished microscopically by the presence of the hairs. + +[Illustration: 41. (_a_) Nodular conjunctivitis in the eye of a child. +De Schweinitz and Shumway.] + +Numerous cases of opthalmia nodosa are on record. Of those from this +country, one of the most interesting is reported by de Schweinitz and +Shumway (1904). It is that of a child of fifteen years whose eye had +become inflamed owing to the presence of some foreign body. Downward and +inward on the bulbar conjunctiva were a number of flattened, +grayish-yellow nodules, between which was a marked congestion of the +conjunctival and episcleral vessels (fig. 41_a_). Twenty-seven nodules +could be differentiated, those directly in the center of the collection +being somewhat confluent and assuming a crescentic and circular +appearance. The nodules were excised and, on sectioning, were found to +be composed of a layer of spindle cells and round cells, outside of +which the tissue was condensed into a capsule. The interior consisted of +epithelioid cells, between which was a considerable intercellular +substance. Directly in the center of a certain number of nodules was +found the section of a hair (fig. 41_b_). The evidence indicated that +the injury had resulted from playing with caterpillars of one of the +Arctiid moths, _Spilosoma virginica_. Other reported cases have been +caused by the hairs of larvæ of _Lasiocampa rubi_, _L. pini_, +_Porthetria dispar_, _Psilura monacha_ and _Cnethocampa processionea_. + +[Illustration: 41_b_. Section through one of the nodules showing the +caterpillar hair. De Schweinitz and Shumway.] + + +RELIEF FROM POISONING BY NETTLING LARVÆ--The irritation from nettling +larvæ is often severe and, especially in regions where the brown-tail +abounds, inquiries as to treatment arise. In general, it may be said +that cooling lotions afford relief, and that scratching, with the +possibilities of secondary infection, should be avoided, in so far as +possible. + +Among the remedies usually at hand, weak solutions of ammonia, or a +paste of ordinary baking soda are helpful. Castellani and Chalmers +recommend cleaning away the hairs by bathing the region with an alkaline +lotion, such as two per cent solution of bicarbonate of soda, and then +applying an ointment of ichthyol (10%). + +In the brown-tail district, there are many proprietary remedies of which +the best ones are essentially the following, as recommended by Kirkland +(1907): + + Carbolic acid ½ drachm. + Zinc oxide ½ oz. + Lime water 8 oz. + +Shake thoroughly and rub well into the affected parts. + +In some cases, and especially where there is danger of secondary +infection, the use of a weak solution of creoline (one teaspoonful to a +quart of water), is to be advised. + + +VESCICATING INSECTS AND THOSE POSSESSING OTHER POISONS IN THEIR BLOOD +PLASMA + +We have seen that certain forms, for example, the poisonous spiders, not +only secrete a toxine in their poison glands, but that such a substance +may be extracted from other parts of their body, or even their eggs. +There are many insects which likewise possess a poisonous blood plasma. +Such forms have been well designated by Taschenberg as _cryptotoxic_ +([Greek: kryptos] = hidden). We shall consider a few representative +forms. + +[Illustration: 42_a_. Blister beetle.] + +[Illustration: 42_b_. An American blister beetle. Meloe angusticollis. +Photograph by M. V. S.] + + +THE BLISTER BEETLES--Foremost among the cryptotoxic insects are the +_Meloidæ_ or "blister beetles," to which the well-known "Spanish fly" +(fig. 42_a_), formerly very generally used in medical practice, belongs. +The vescicating property is due to the presence in the blood plasma of a +peculiar, volatile, crystalline substance known as _cantharidin_, which +is especially abundant in the reproductive organs of the beetle. +According to Kobert, the amount of this varies in different species from +.4 or .5% to 2.57% of the dry weight of the beetle. + +While blister beetles have been especially used for external +application, they are also at times used internally as a stimulant and a +diuretic. The powder or extract was formerly much in vogue as an +aphrodisiac, and formed the essential constituent of various philters, +or "love powders". It is now known that its effects on the reproductive +organs appear primarily after the kidneys have been affected to such an +extent as to endanger life, and that many cases of fatal poison have +been due to its ignorant use. + +There are many cases on record of poisoning and death due to internal +use, and in some instances from merely external application. There are +not rarely cases of poisoning of cattle from feeding on herbage bearing +a large number of the beetles and authentic cases are known of human +beings who have been poisoned by eating the flesh of such cattle. Kobert +states that the beetles are not poisonous to birds but that the flesh of +birds which have fed on them is poisonous to man, and that if the flesh +of chickens or frogs which have fed on the cantharidin be fed to cats it +causes in them the same symptoms as does the cantharidin. + +Treatment of cases of cantharidin poison is a matter for a skilled +physician. Until he can be obtained, emetics should be administered and +these should be followed by white of egg in water. Oils should be +avoided, as they hasten the absorption of the poison. + + +OTHER CRYPTOTOXIC INSECTS--Though the blister beetles are the best known +of the insects with poisonous blood plasma, various others have been +reported and we shall refer to a few of the best authenticated. + +One of the most famous is the Chrysomelid beetle, _Diamphidia simplex_, +the body fluids of whose larvæ are used by certain South African bushmen +as an arrow poison. Its action is due to the presence of a toxalbumin +which exerts a hæmolytic action on the blood, and produces inflammation +of the subcutaneous connective tissue and mucous membranes. Death +results from general paralysis. Krause (1907) has surmised that the +active principle may be a bacterial toxin arising from decomposition of +the tissues of the larva, but he presents no support of this view and it +is opposed by all the available evidence. + +In China, a bug, _Heuchis sanguinea_, belonging to the family Cicadidæ, +is used like the Meloidæ, to produce blistering, and often causes +poisoning. It has been assumed that its vescicating properties are due +to cantharidin, but the presence of this substance has not been +demonstrated. + +Certain Aphididæ contain a strongly irritating substance which produces, +not merely on mucous membranes but on outer skin, a characteristic +inflammation. + +It has been frequently reported that the larvæ of the European cabbage +butterfly, _Pieris brassicæ_, accidentally eaten by cows, horses, ducks, +and other domestic animals, cause severe colic, attempts to vomit, +paralysis of the hind legs, salivation, and stomatitis. On _postmortem_ +there are to be found hæmorrhagic gastro-enteritis, splenitis, and +nephritis. Kobert has recently investigated the subject and has found a +poisonous substance in the blood of not only the larvæ but also the +pupæ. + + +FOOTNOTES: + +[A] This is diametrically opposed to the findings of Bordas (1905) in +the case of the European _Latrodectus 13-guttatus_, whose glands are +"much larger than those of other spiders." From a considerable +comparative study, we should also unhesitatingly make this statement +regarding the glands of our American species, _L. mactans_. + +[B] Dr. E. H. Coleman (Kellogg, 1915) has demonstrated its virulence by +a series of experiments comparable with those of Kobert. + +[C] According to Stiles, the species occurring in the Northwest which is +commonly identified as _D. venustus_ should be called _D. andersoni_ +(see footnote, chapter 12). + +[D] It should be remembered that in all the higher Hymenoptera the first +abdominal segment is fused with the thorax and that what is apparently +the sixth segment is, in reality, the seventh. + + + + +CHAPTER III + +PARASITIC ARTHROPODA AFFECTING MAN + + +The relation of insects to man as simple parasites has long been +studied, and until very recent years the bulk of the literature of +medical entomology referred to this phase of the subject. This is now +completely overshadowed by the fact that so many of these parasitic +forms are more than simple parasites, they are transmitters of other +microscopic parasites which are pathogenic to man. Yet the importance of +insects as parasites still remains and must be considered in a +discussion of the relation of insects to the health of man. In taking up +the subject we shall first consider some general features of the +phenomenon of animal parasitism. + +Parasitism is an adaptation which has originated very often among living +organisms and in widely separated groups. It would seem simple to define +what is meant by a "parasite" but, in reality, the term is not easily +limited. It is often stated that a parasite is "An organism which lives +at the expense of another," but this definition is applicable to a +predatory species or, in its broadest sense, to all organisms. For our +purpose we may say with Braun: "A parasite is an organism which, for the +purpose of obtaining food, takes up its abode, temporarily or +permanently, on or within another living organism". + +Thus, parasitism is a phase of the broad biological phenomenon of +_symbiosis_, or living together of organisms. It is distinguished from +_mutualism_, or symbiosis in the narrow sense, by the fact that only one +party to the arrangement obtains any advantage, while the other is to a +greater or less extent injured. + +Of parasites we may distinguish on the basis of their location on or in +the host, _ecto-parasites_, which live outside of the body; and +_endoparasites_, which live within the body. On account of their method +of breathing the parasitic arthropods belong almost exclusively to the +first of these groups. + +On the basis of relation to their host, we find _temporary parasites_, +those which seek the host only occasionally, to obtain food; and the +_stationary_ or _permanent_ parasites which, at least during certain +stages, do not leave their host. + +_Facultative parasites_ are forms which are not normally parasitic, but +which, when accidentally ingested, or otherwise brought into the body, +are able to exist for a greater or less period of time in their unusual +environment. These are generally called in the medical literature +"pseudoparasites" but the term is an unfortunate one. + +We shall now take up the different groups of arthropods, discussing the +more important of the parasitic forms attacking man. The systematic +relationship of these forms, and key for determining important species +will be found in Chapter XII. + + +ACARINA OR MITES + +The ACARINA, or _mites_, form a fairly natural group of arachnids, +characterized, in general, by a sac-like, unsegmented body which is +generally fused with the cephalothorax. The mouth-parts have been united +to form a beak or rostrum. + +The representatives of this group undergo a marked metamorphosis. +Commonly, the larvæ on hatching from the egg, possess but three pairs of +legs, and hence are called _hexapod larvæ_. After a molt, they transform +into nymphs which, like the adult, have four pairs of legs and are +called _octopod nymphs_. These after a period of growth, molt one or +more times and, acquiring external sexual organs, become adult. + +Most of the mites are free-living, but there are many parasitic species +and as these have originated in widely separated families, the Acarina +form an especially favorable group for study of the origin of +parasitism. Such a study has been made by Ewing (1911), who has reached +the following conclusions: + +"We have strong evidence indicating that the parasitic habit has +originated independently at least eleven times in the phylogeny of the +Ararina. Among the zoophagous parasites, the parasitic habit has been +developed from three different types of free-living Acarina: (a) +predaceous forms, (b) scavengers, (c) forms living upon the juices of +plants." + +Ewing also showed that among the living forms of Acarina we can trace +out all the stages of advancing parasitism, semiparasitism, facultative +parasitism, even to the fixed and permanent type, and finally to +endoparasitism. + +Of the many parasitic forms, there are several species which are serious +parasites of man and we shall consider the more important of these. +Infestation by mites is technically known as _acariasis_. + +[Illustration: 43. Effect of the harvest mites on the skin of man. +Photograph by J. C. Bradley.] + + +The Trombidiidæ, or Harvest Mites + +In many parts of this country it is impossible for a visitor to go into +the fields and, particularly, into berry patches and among tall weeds +and grass in the summer or early fall without being affected by an +intolerable itching, which is followed, later, by a breaking out of +wheals, or papules, surrounded by a bright red or violaceous aureola, +(fig. 43). It is often regarded as a urticaria or eczema, produced by +change of climate, an error in diet, or some condition of general +health. + +Sooner or later, the victim finds that it is due to none of these, but +to the attacks of an almost microscopic red mite, usually called +"jigger" or "chigger" in this country. As the term "chigger" is applied +to one of the true fleas, _Dermatophilus penetrans_, of the tropics, +these forms are more correctly known as "harvest mites." Natives of an +infested region may be so immune or accustomed to its attacks as to be +unaware of its presence, though such immunity is by no means possessed +by all who have been long exposed to the annoyance. + +[Illustration: 44. Harvest mites. (Larvæ of Trombidium). After C. V. +Riley.] + +The harvest mites, or chiggers, attacking man are larval forms, +possessing three pairs of legs (fig. 44). Their systematic position was +at first unknown and they were classed under a special genus _Leptus_, a +name which is very commonly still retained in the medical literature. It +is now known that they are the larval forms of various species of the +genus _Trombidium_, a group of predaceous forms, the adults of which +feed primarily on insects and their eggs. In this country the species +best known are those to be found late in summer, as larvæ at the base of +the wings of houseflies or grasshoppers. + +There is much uncertainty as to the species of the larvæ attacking man +but it is clear that several are implicated. Bruyant has shown that in +France the larvæ _Trombidium inapinatum_ and _Trombidium holosericeum_ +are those most frequently found. The habit of attacking man is abnormal +and the larvæ die after entering the skin. Normally they are parasitic +on various insects. + +Most recent writers agree that, on man, they do not bore into the skin, +as is generally supposed, but enter a hair follicle or sebaceous gland +and from the bottom of this, pierce the cutis with their elongate +hypopharynx. According to Braun, there arises about the inserted +hypopharynx a fibrous secretion--the so-called "beak" which is, in +reality, a product of the host. Dr. J. C. Bradley, however, has made +careful observations on their method of attack, and he assures us that +the mite ordinarily remains for a long time feeding on the surface of +the skin, where it produces the erythema above described. During this +time it is not buried in the skin but is able to retreat rapidly into it +through a hair follicle or sweat gland. The irritation from the mites +ceases after a few days, but not infrequently the intolerable itching +leads to so much scratching that secondary infection follows. + +Relief from the irritation may be afforded by taking a warm salt bath as +soon as possible after exposure or by killing the mites by application +of benzine, sulphur ointment or carbolized vaseline. When they are few +in number, they can be picked out with a sterile needle. + +Much may be done in the way of warding off their attacks by wearing +gaiters or close-woven stockings extending from ankle to the knee. Still +more efficacious is the sprinkling of flowers of sulphur in the +stockings and the underclothes from a little above the knee, down. The +writers have known this to make it possible for persons who were +especially susceptible to work with perfect comfort in badly infested +regions. Powdered naphthalene is successfully used in the same way and +as Chittenden (1906) points out, is a safeguard against various forms of +man-infesting tropical insect pests. + +The question of the destruction of the mites in the field is sometimes +an important one, and under some conditions, is feasible. Chittenden +states that much can be accomplished by keeping the grass, weeds, and +useless herbage mowed closely, so as to expose the mites to the sun. He +believes that in some cases good may be done by dusting the grass and +other plants, after cutting, with flowers of sulphur or by spraying with +dilute kerosene emulsion in which sulphur has been mixed. More recently +(1914) he calls attention to the value of cattle, and more especially +sheep, in destroying the pests by tramping on them and by keeping the +grass and herbage closely cropped. + + +IXODOIDEA OR TICKS + +Until recently, the ticks attracted comparatively little attention from +entomologists. Since their importance as carriers of disease has been +established, interest in the group has been enormously stimulated and +now they rank second only to the mosquitoes in the amount of detailed +study that has been devoted to them. + +[Illustration: 45_a_. Argus persicus. Capitulum of male. After Nuttall +and Warburton.] + +The ticks are the largest of the Acarina. They are characterized by the +fact that the hypostome, or "tongue" (fig. 45) is large and file-like, +roughened by sharp teeth. They possess a breathing pore on each side of +the body, above the third or fourth coxæ (fig. 45_b_). + +[Illustration: 45_b_. Left spiracle of nymph of _Argas persicus_. After +Nuttall and Warburton.] + +There are two distinct families of the IXODOIDEA, differing greatly in +structure, life-history and habits. These are the ARGASIDÆ and the +IXODIDÆ. We shall follow Nuttall (1908) in characterizing these two +families and in pointing out their biological differences, and shall +discuss briefly the more important species which attack man. The +consideration of the ticks as carriers of disease will be reserved for a +later chapter. + + +Argasidæ + +In the ticks belonging to the family ARGASIDÆ, there is comparatively +little sexual dimorphism, while this is very marked in the Ixodidæ. The +capitulum, or so-called "head" is ventral, instead of terminal; the +palpi are leg-like, with the segments subequal; the scutum, or dorsal +shield, is absent; eyes, when present, are lateral, on supracoxal folds. +The spiracles are very small; coxæ unarmed; tarsi without ventral +spurs, and the pulvilli are absent or rudimentary. + +In habits and life history the Argasidæ present striking +characteristics. In the first place, they are long-lived, a factor which +counts for much in the maintenance of the species. They are intermittent +feeders, being comparable with the bed-bug in this respect. There are +two or more nymphal stages, and they may molt after attaining maturity. +The female lays comparatively few eggs in several small batches. + +Nuttall (1911) concludes that "The Argasidæ represent the relatively +primitive type of ticks because they are less constantly parasitic than +are the Ixodidæ. Their nymphs and adults are rapid feeders and chiefly +infest the habitat of their hosts. * * * Owing to the Argasidæ infesting +the habitats of their hosts, their resistance to prolonged starvation +and their rapid feeding habits, they do not need to bring forth a large +progeny, because there is less loss of life in the various stages, as +compared with the Ixodidæ, prior to their attaining maturity." + +[Illustration: 46. Argus persicus. Dorsal and ventral aspects. (×4). +After Hassell.] + +Of the Argasidæ, we have in the United States, several species which +have been reported as attacking man. + +_Argas persicus_, the famous "Miana bug" (fig. 46), is a very widely +distributed species, being reported from Europe, Asia, Africa, and +Australia. It is everywhere preeminently a parasite of fowls. According +to Nuttall it is specifically identical with _Argas americanus_ Packard +or _Argas miniatus_ Koch, which is commonly found on fowls in the United +States, in the South and Southwest. Its habits are comparable to those +of the bed-bug. It feeds intermittently, primarily at night, and instead +of remaining on its host, it then retreats to cracks and crevices. +Hunter and Hooker (1908) record that they have found the larva to remain +attached for five or eight days before dropping. Unlike the Ixodidæ, the +adults oviposit frequently. + +[Illustration: 47. Otiobius (Ornithodoros) megnini, head of nymph. After +Stiles.] + +[Illustration: 48. Otiobius (Ornithodoros) megnini, male. (_a_) dorsal, +(_b_) ventral aspect. After Nuttall and Warburton.] + +The most remarkable feature of the biology of this species is the great +longevity, especially of the adult. Hunter and Hooker report keeping +larvæ confined in summer in pill boxes immediately after hatching for +about two months while under similar conditions those of the Ixodid, +_Boophilus annulatus_ lived for but two or three days. Many writers have +recorded keeping adults for long periods without food. We have kept +specimens in a tin box for over a year and a half and at the end of that +time a number were still alive. Laboulliene kept unfed adults for over +three years. In view of the effectiveness of sulphur in warding off the +attacks of Trombidiidæ, it is astonishing to find that Lounsbury has +kept adults of _Argas persicus_ for three months in a box nearly filled +with flowers of sulphur, with no apparent effect on them. + +We have already called attention to the occasional serious effects of +the bites of this species. While such reports have been frequently +discredited there can be no doubt that they have foundation in fact. The +readiness with which this tick attacks man, and the extent to which old +huts may be infested makes it especially troublesome. + +_Otiobius (Ornithodoros) megnini_, the "spinose ear-tick" (figs. 47, +48), first described from Mexico, as occurring in the ears of horses, is +a common species in our Southwestern States and is recorded by Banks as +occurring as far north as Iowa. + +The species is remarkable for the great difference between the spiny +nymph stage and the adult. The life history has been worked out by +Hooker (1908). Seed ticks, having gained entrance to the ear, attach +deeply down in the folds, engorge, and in about five days, molt; as +nymphs with their spinose body they appear entirely unlike the larvæ. As +nymphs they continue feeding sometimes for months. Finally the nymph +leaves the host, molts to form the unspined adult, and without further +feeding is fertilized and commences oviposition. + +The common name is due to the fact that in the young stage the ticks +occur in the ear of their hosts, usually horses or cattle. Not +uncommonly it has been reported as occurring in the ear of man and +causing very severe pain. Stiles recommends that it be removed by +pouring some bland oil into the ear. + +Banks (1908) reports three species of _Ornithodoros_--_O. turicata_, +_coriaceus_ and _talaje_--as occurring in the United States. All of these +attack man and are capable of inflicting very painful bites. + + +Ixodidæ + +The ticks belonging to the family IXODIDÆ (figs. 49 and 50) exhibit a +marked sexual dimorphism. The capitulum is anterior, terminal, instead +of ventral as in the Argasidæ; the palpi are relatively rigid (except in +the subfamily Ixodinæ), with rudimentary fourth segment; scutum present; +eyes, when present, dorsal, on side of scutum. The spiracles are +generally large, situated well behind the fourth coxæ; coxæ generally +with spurs; pulvilli always present. + +In habits and life history the typical Ixodidæ differ greatly from the +Argasidæ. They are relatively short-lived, though some recent work +indicates that their longevity has been considerably under-estimated. +Typically, they are permanent feeders, remaining on the host, or hosts, +during the greater part of their life. They molt twice only, on leaving +the larval and the nymphal stages. The adult female deposits a single, +large batch of eggs. Contrasting the habits of the Ixodidæ to those of +the Argasidæ, Nuttall (1911) emphasizes that the Ixodidæ are more highly +specialized parasites. "The majority are parasitic on hosts having no +fixed habitat and consequently all stages, as a rule, occur upon the +host." + +[Illustration: 49. Ixodes ricinus; male, ventral aspect. After Braun and +Luehe.] + +As mere parasites of man, apart from their power to transmit disease, +the Ixodidæ are much less important than the Argasidæ. Many are reported +as occasionally attacking man and of these the following native species +may be mentioned. + +[Illustration: 50. Ixodes ricinus, var. scapularis, female. Capitulum +and scutum; ventral aspect of capitulum; coxæ; tarsus 4; spiracle; +genital and anal grooves. After Nuttall and Warburton.] + +_Ixodes ricinus_, the European castor bean tick (figs. 49, 50), is a +species which has been often reported from this country but Banks (1908) +has shown that, though it does occur, practically all of the records +apply to _Ixodes scapularis_ or _Ixodes cookei_. In Europe, _Ixodes +ricinus_ is very abundant and very commonly attacks man. At the point +of penetration of the hypostome there is more or less inflammation but +serious injury does not occur unless there have been introduced +pathogenic bacteria or, unless the tick has been abruptly removed, +leaving the capitulum in the wound. Under the latter circumstances, +there may be an abscess formed about the foreign body and occasionally, +serious results have followed. Under certain conditions the tick, in +various stages, may penetrate under the skin and produce a tumor, within +which it may survive for a considerable period of time. + +_Ixodes cookei_ is given by Banks as "common on mammals in the Eastern +States as far west as the Rockies." It is said to affect man severely. + +_Amblyomma americanum_, (fig. 158c), the "lone star tick," is widely +distributed in the United States. Its common name is derived from the +single silvery spot on the scutum of the female. Hunter and Hooker +regard this species as, next to _Boophilus annulatus_, the most +important tick in the United States. Though more common on cattle, it +appears to attack mammals generally, and "in portions of Louisiana and +Texas it becomes a pest of considerable importance to moss gatherers and +other persons who spend much time in the forests." + +_Amblyomma cajennense_, noted as a pest of man in central and tropical +America, is reported from various places in the south and southwestern +United States. + +_Dermacentor variabilis_ is a common dog tick of the eastern United +States. It frequently attacks man, but the direct effects of its bite +are negligible. + +The "Rocky Mountain spotted fever tick" (_Dermacentor andersoni_ +according to Stiles, _D. venustus_ according to Banks) is, from the +viewpoint of its effects on man, the most important of the ticks of the +United States. This is because, as has been clearly established, it +transmits the so-called "spotted fever" of man in our northwestern +states. This phase of the subject will be discussed later and it need +merely be mentioned here, that this species has been reported as causing +painful injuries by its bites. Dr. Stiles states that he has seen cases +of rather severe lymphangitis and various sores and swellings developing +from this cause. In one case, of an individual bitten near the elbow, +the arm became very much swollen and the patient was confined in bed for +several days. The so-called tick paralysis produced by this species is +discussed in a preceding chapter. + +There are many other records of various species of ticks attacking man, +but the above-mentioned will serve as typical and it is not necessary to +enter into greater detail. + + +TREATMENT OF TICK BITES--When a tick attaches to man the first thing to +be done is to remove it without leaving the hypostome in the wound to +fester and bring about secondary effects. This is best accomplished by +applying to the tick's body some substance which will cause it to more +readily loosen its hold. Gasoline or petroleum, oil or vaseline will +serve. For removing the spinose ear-tick, Stiles recommends pouring some +bland oil into the ear. Others have used effectively a pledget of cotton +soaked in chloroform. + +In general, the treatment recommended by Wellman for the bites of +_Ornithodoros moubata_ will prove helpful. It consists of prolonged +bathing in very hot water, followed by the application of a strong +solution of bicarbonate of soda, which is allowed to dry upon the skin. +He states that this treatment is comforting. For severe itching he +advises smearing the bites with vaseline, which is slightly impregnated +with camphor or menthol. Medical aid should be sought when complications +arise. + +The DERMANYSSIDÆ are Gamasid mites which differ from others of the group +in that they are parasitic on vertebrates. None of the species normally +attack man, but certain of them, especially the poultry mite, may be +accidental annoyances. + +[Illustration: 51. Dermanyssus gallinæ, female. After Delafond.] + +_Dermanyssus gallinæ_ (fig. 51), the red mite of poultry, is an +exceedingly common and widespread parasite of fowls. During the day it +lives in cracks and crevices of poultry houses, under supports of +roosts, and in litter of the food and nests, coming out at night to +feed. They often attack people working in poultry houses or handling +and plucking infested fowls. They may cause an intense pruritis, but +they do not produce a true dermatosis, for they do not find conditions +favorable for multiplication on the skin of man. + + +Tarsonemidæ + +The representatives of the family TARSONEMIDÆ are minute mites, with the +body divided into cephalothorax and abdomen. There is marked sexual +dimorphism. The females possess stigmata at the anterior part of the +body, at the base of the rostrum, and differ from all other mites in +having on each side, a prominent clavate organ between the first and +second legs. The larva, when it exists, is hexapodous and resembles the +adult. A number of the species are true parasites on insects, while +others attack plants. Several of them may be accidental parasites of +man. + +[Illustration: 52. Pediculoides ventricosus, female. After Webster.] + +[Illustration: 53. Pediculoides ventricosus, gravid female. (×80). +After Webster.] + +_Pediculoides ventricosus_ (fig. 52 and 53) is, of all the Tarsonemidæ +reported, the one which has proved most troublesome to man. It is a +predaceous species which attacks a large number of insects but which has +most commonly been met with by man through its fondness for certain +grain-infesting insects, notably the Angoumois grain moth, _Sitotroga +cerealella_, and the wheat straw-worm, _Isosoma grande_. In recent years +it has attracted much attention in the United States and its +distribution and habits have been the object of detailed study by +Webster (1901). + +[Illustration: 54. Pediculoides ventricosus, male. After Braun.] + +There is a very striking sexual dimorphism in this species. The +non-gravid female is elongate, about 200µ by 70µ (fig. 52), with the +abdomen slightly striated longitudinally. The gravid female (fig. 53) +has the abdomen enormously swollen, so that it is from twenty to a +hundred times greater than the rest of the body. The species is +viviparous and the larvæ undergo their entire growth in the body of the +mother. They emerge as sexually mature males and females which soon +pair. The male (fig. 54) is much smaller, reaching a length of only 320µ +but is relatively broad, 80µ, and angular. Its abdomen is very greatly +reduced. + +As far back as 1850 it was noted as causing serious outbreaks of +peculiar dermatitis among men handling infested grain. For some time the +true source of the difficulty was unknown and it was even believed that +the grain had been poisoned. Webster has shown that in this country (and +probably in Europe as well) its attacks have been mistaken for those of +the red bugs or "chiggers" (larval Trombiidæ). More recently a number of +outbreaks of a mysterious "skin disease" were traced to the use of straw +mattresses, which were found to be swarming with these almost +microscopic forms which had turned their attentions to the occupants of +the beds. Other cases cited were those of farmers running wheat through +a fanning mill, and of thrashers engaged in feeding unthrashed grain +into the cylinder of the machine. + +[Illustration: 55. Lesions produced by the attacks of Pediculoides +ventricosus. After Webster.] + +The medical aspects of the question have been studied especially by +Schamberg and Goldberger and from the latter's summary (1910) we derive +the following data. Within twelve to sixteen hours after exposure, +itching appears and in severe cases, especially where exposure is +continued night after night by sleeping on an infested bed, the itching +may become almost intolerable. Simultaneously, there appears an eruption +which characteristically consists of wheals surrounded by a vesicle +(fig. 55). The vesicle as a rule does not exceed a pin head in size but +may become as large as a pea. Its contents rapidly become turbid and in +a few hours it is converted into a pustule. The eruption is most +abundant on the trunk, slight on the face and extremities and almost +absent on the feet and hands. In severe cases there may be +constitutional disturbances marked, at the outset, by chilliness, +nausea, and vomiting, followed for a few days by a slight elevation of +temperature, with the appearance of albumin in the urine. In some cases +the eruption may simulate that of chicken-pox or small-pox. + +Treatment for the purpose of killing the mites is hardly necessary as +they attach feebly to the surface and are readily brushed off by +friction of the clothes. "Antipruritic treatment is always called for; +warm, mildly alkaline baths or some soothing ointment, such as zinc +oxide will be found to fulfil this indication." Of course, reinfestation +must be guarded against, by discarding, or thoroughly fumigating +infested mattresses, or by avoiding other sources. Goldberger suggests +that farm laborers who must work with infested wheat or straw might +protect themselves by anointing the body freely with some bland oil or +grease, followed by a change of clothes and bath as soon as their work +is done. We are not aware of any experiments to determine the effect of +flowers of sulphur, but their efficiency in the case of "red bugs" +suggests that they are worth a trial against _Pediculoides_. + +Various species of TYROGLYPHIDÆ (fig. 150_f_) may abound on dried fruits +and other products and attacking persons handling them, may cause a +severe dermatitis, comparable to that described above for _Pediculoides +ventricosus_. Many instances of their occurrence as such temporary +ectoparasites are on record. Thus, workers who handle vanilla pods are +subject to a severe dermatitis, known as vanillism, which is due to the +attacks of _Tyroglyphus siro_, or a closely related species. The +so-called "grocer's itch" is similarly caused by mites infesting various +products. Castellani has shown that in Ceylon, workers employed in the +copra mills, where dried cocoanut is ground up for export, are much +annoyed by mites, which produce the so-called "copra itch." The skin of +the hands, arms and legs, and sometimes of the whole body, except the +face, is covered by fairly numerous, very pruriginous papules, often +covered by small, bloody crusts due to scratching. The condition is +readily mistaken for scabies. It is due to the attacks of _Tyroglyphus +longior castellanii_ which occur in enormous numbers in some samples of +the copra. + + +Sarcoptidæ + +The SARCOPTIDÆ are minute whitish mites, semi-globular in shape, with a +delicate transversely striated cuticula. They lack eyes and tracheæ. The +mouth-parts are fused at the base to form a cone which is usually +designated as the head. The legs are short and stout, and composed of +five segments. The tarsi may or may not possess a claw and may terminate +in a pedunculated sucker, or simple long bristle, or both. The presence +or absence of these structures and their distribution are much used in +classification. The mites live on or under the skin of mammals and +birds, where they produce the disease known as scabies, mange, or itch. +Several species of the Sarcoptidæ attack man but the most important of +these, and the one pre-eminent as the "itch mite" is _Sarcoptes +scabiei_. + +The female of _Sarcoptes scabiei_, of man, is oval and yellowish white; +the male more rounded and of a somewhat reddish tinge, and much smaller. +The body is marked by transverse striæ which are partly interrupted on +the back. There are transverse rows of scales, or pointed spines, and +scattered bristles on the dorsum. + +[Illustration: 56_a_. Sarcoptes scabiei, male. (×100). After +Fürstenberg.] + +The male (fig. 56) which is from 200-240µ in length, and 150-200µ in +breadth, possesses pedunculated suckers on each pair of legs except the +third, which bears, instead, a long bristle. The female (fig. 56) +300-450µ in length and 250-350µ in breadth, has the pedunculated suckers +on the first and second pairs of legs, only, the third and fourth +terminating in bristles. + +[Illustration: 56_b_. Sarcoptes scabiei, female. (×100.) After +Fürstenberg.] + +The mite lives in irregular galleries from a few millimeters to several +centimeters in length, which it excavates in the epidermis (fig. 57). It +works especially where the skin is thin, such as between the fingers, in +the bend of the elbows and knees, and in the groin, but it is by no +means restricted to these localities. The female, alone, tunnels into +the skin; the males remain under the superficial epidermal scales, and +seldom are found, as they die soon after mating. + +As she burrows into the skin the female deposits her eggs, which measure +about 150 × 100µ. Fürstenberg says that each deposits an average of +twenty-two to twenty-four eggs, though Gudden reports a single burrow as +containing fifty-one. From these there develop after about seven days, +the hexapod larvæ. These molt on the sixteenth day to form an octopod +nymph, which molts again the twenty-first day. At the end of the fourth +week the nymphs molt to form the sexually mature males and the so-called +pubescent females. These pair, the males die, and the females again cast +their skin, and become the oviparous females. Thus the life cycle is +completed in about twenty-eight days. + +The external temperature exercises a great influence on the development +of the mites and thus, during the winter, the areas of infestation not +only do not spread, but they become restricted. As soon as the +temperature rises, the mites increase and the infestation becomes much +more extensive. + +[Illustration: 57. Sarcoptes scabiei. Diagrammatic representation of the +course in the skin of man.] + +In considering the possible sources of infestation, and the chances of +reinfestation after treatment, the question of the ability of the mite +to live apart from its host is a very important one. Unfortunately there +are few reliable data on this subject. Gerlach found that, exposed in +the dry, warm air of a room they became very inactive within twenty-four +hours, that after two days they showed only slight movement, and that +after three or four days they could not be revived by moisture and +warming. The important fact was brought out that in moist air, in folded +soiled underwear, they survived as long as ten days. Bourguignon found +that under the most favorable conditions the mites of _Sarcoptes scabiei +equi_ would live for sixteen days. + +The disease designated the "itch" or "scabies," in man has been known +from time immemorial, but until within less than a hundred years it was +almost universally attributed to malnutrition, errors of diet, or "bad +blood." This was in spite of the fact that the mite was known to Mouffet +and that Bonomo had figured both the adult and the egg and had declared +the mite the sole cause of the disease. In 1834 the Corsican medical +student, Francis Renucci, demonstrated the mite before a clinic in Saint +Louis Hospital in Paris and soon thereafter there followed detailed +studies of the life history of the various itch mites of man and +animals. + +[Illustration: 58. Scabies on the hand. From portfolio of Dermochromes +by permission of Rebman & Co., of New York. Publishers.] + +The disease is a cosmopolitan one, being exceedingly abundant in some +localities. Its spread is much favored where large numbers of people are +crowded together under insanitary conditions and hence it increases +greatly during wars and is widely disseminated and abundant immediately +afterwards. Though more commonly to be met with among the lower classes, +it not infrequently appears among those of the most cleanly, careful +habits, and it is such cases that are most liable to wrong diagnosis by +the physician. + +Infection occurs solely through the passage, direct or indirect, of the +young fertilized females to the skin of a healthy individual. The adult, +oviparous females do not quit their galleries and hence do not serve to +spread the disease. The young females move about more or less at night +and thus the principal source of infestation is through sleeping in the +same bed with an infested person, or indirectly through bedclothes, or +even towels or clothing. Diurnal infestation through contact or clothing +is exceptional. Many cases are known of the disease being contracted +from animals suffering from scabies, or mange. + +When a person is exposed to infestation, the trouble manifests itself +after eight or ten days, though there usually elapses a period of twenty +to thirty days before there is a suspicion of anything serious. The +first symptom is an intense itching which increases when the patient is +in bed. When the point of irritation is examined the galleries may +usually be seen as characteristic sinuous lines, at first whitish in +color but soon becoming blackish because of the contained eggs and +excrement. The galleries, which may not be very distinct in some cases, +may measure as much as four centimeters in length. Little vesicles, of +the size of a pin head are produced by the secretions of the feeding +mite; they are firm, and projecting, and contain a limpid fluid. Figures +58 and 59 show the typical appearance of scabies on the hands, while +figure 60 shows a severe general infestation. The intolerable itching +induces scratching and through this various complications may arise. The +lesions are not normally found on the face and scalp, and are rare on +the back. + +[Illustration: 59. Scabies on the hand. After Duhring.] + +[Illustration: 60. Generalized infection of Scabies. After Morrow.] + +Formerly, scabies was considered a very serious disease, for its cause +and method of treatment were unknown, and potentially it may continue +indefinitely. Generation after generation of the mites may develop and +finally their number become so great that the general health of the +individual is seriously affected. Now that the true cause of the disease +is known, it is easily controlled. + +Treatment usually consists in softening the skin by friction with soap +and warm water, followed by a warm bath, and then applying some +substance to kill the mites. Stiles gives the following directions, +modified from Bourguignon's, as "a rather radical guide, to be modified +according to facilities and according to the delicacy of the skin or +condition of the patient": + +1. The patient, stripped naked, is energetically rubbed all over (except +the head) for twenty minutes, with green soap and warm water. 2. He is +then placed in a warm bath for thirty minutes, during which time the +rubbing is continued. 3. The parasiticide is next rubbed in for twenty +minutes and is allowed to remain on the body for four or five hours; in +the meantime the patient's clothes are sterilized, to kill the eggs or +mites attached to them. 4. A final bath is taken to remove the +parasiticide. + +The parasiticide usually relied on is the officinal sulphur ointment of +the United States pharmacopoeia. When infestation is severe it is +necessary to repeat treatment after three or four days in order to kill +mites which have hatched from the eggs. + +The above treatment is too severe for some individuals and may, of +itself, produce a troublesome dermatitis. We have seen cases where the +treatment was persisted in and aggravated the condition because it was +supposed to be due to the parasite. For delicate-skinned patients the +use of balsam of Peru is very satisfactory, and usually causes no +irritation whatever. Of course, sources of reinfection should be +carefully guarded against. + +_Sarcoptes scabiei crustosæ_, which is a distinct variety, if not +species, of the human itch mite, is the cause of so-called Norwegian +itch. This disease is very contagious, and is much more resistant than +the ordinary scabies. Unlike the latter, it may occur on the face and +scalp. + +_Sarcoptes scabiei_ not only attacks man but also occurs on a large +number of mammals. Many species, based on choice of host, and minute +differences in size and secondary characters, have been established, but +most students of the subject relegate these to varietal rank. Many of +them readily attack man, but they have become sufficiently adapted to +their normal host so that they are usually less persistent on man. + +_Notoedres cati_ (usually known as _Sarcoptes minor_) is a species of +itch mites which produce an often fatal disease of cats. The body is +rounded and it is considerably smaller than _Sarcoptes scabiei_, the +female (fig. 61) measuring 215-230µ long and 165-175µ wide; the males +145-150µ by 120-125µ. The most important character separating +_Notoedres_ from _Sarcoptes_ is the position of the anus, which is +dorsal instead of terminal. The mite readily transfers to man but does +not persist, the infestation usually disappearing spontaneously in about +two weeks. Infested cats are very difficult to cure, unless treatment is +begun at the very inception of the outbreak, and under ordinary +circumstances it is better to kill them promptly, to avoid spread of the +disease to children and others who may be exposed. + +[Illustration: 61. Notoedres cati, male and female. After Railliet.] + + +Demodecidæ + +The DEMODECIDÆ are small, elongate, vermiform mites which live in the +hair follicles of mammals. The family characteristics will be brought +out in the discussion of the species infesting man, _Demodex +folliculorum_. + +[Illustration: 62. Demodex folliculorum. (×200) After Blanchard.] + +_Demodex folliculorum_ (fig. 62) is to be found very commonly in the +hair follicles and sebaceous glands of man. It is vermiform in +appearance, and with the elongate abdomen transversely striated so as to +give it the appearance of segmentation. The female is 380-400µ long by +45µ; the male 300µ by 40µ. The three-jointed legs, eight in number, are +reduced to mere stubs in the adult. The larval form is hexopod. These +mites thus show in their form a striking adaptation to their +environment. In the sebaceous glands and hair follicles they lie with +their heads down (fig. 63). Usually there are only a few in a gland, but +Gruby has counted as many as two hundred. + +[Illustration: 63. Demodex folliculorum. Section through skin showing +the mites in situ. Magnification of Nos. 1, 2, 6 and 7, ×150; Nos, 3, +4, 5, ×450. After Megnin.] + +The frequency with which they occur in man is surprising. According to +European statistics they are found in 50 per cent to 60 per cent or even +more. Gruby found them in forty out of sixty persons examined. These +figures are very commonly quoted, but reliable data for the United +States seem to be lacking. Our studies indicate that it is very much +less common in this country than is generally assumed. + +The Demodex in man does not, as a rule, cause the slightest +inconvenience to its host. It is often stated that they give rise to +comedons or "black-heads" but there is no clear evidence that they are +ever implicated. Certain it is that they are not the usual cause. A +variety of the same, or a very closely related species of _Demodex_, on +the dog gives rise to the very resistant and often fatal follicular +mange. + + +HEXAPODA OR TRUE INSECTS + +The HEXAPODA, or true insects, are characterized by the fact that the +adult possesses three pairs of legs. The body is distinctly segmented +and is divided into head, thorax, and abdomen. + +The mouth-parts in a generalized form, consist of an upper lip, or +_labrum_, which is a part of the head capsule, and a central unpaired +_hypopharynx_, two _mandibles_, two _maxillæ_ and a lower lip, or +_labium_, made up of the fused pair of second maxillæ. These parts may +be greatly modified, dependent upon whether they are used for biting, +sucking, piercing and sucking, or a combination of biting and sucking. + +Roughly speaking, insects may be grouped into those which undergo +_complete metamorphosis_ and those which have _incomplete +metamorphosis_. They are said to undergo complete metamorphosis when the +young form, as it leaves the egg, bears no resemblance to the adult. For +example, the maggot changes to a quiescent pupa and from this emerges +the winged active fly. They undergo incomplete metamorphosis, when the +young insect, as it leaves the egg, resembles the adult to a greater or +less extent, and after undergoing a certain number of molts becomes +sexually mature. + +Representatives of several orders have been reported as accidental or +faculative parasites of man, but the true parasites are restricted to +four orders. These are the Siphunculata; the Hemiptera, the Diptera and +the Siphonaptera. + + +SIPHUNCULATA + +The order SIPHUNCULATA was established by Meinert to include the true +sucking lice. These are small wingless insects, with reduced +mouth-parts, adapted for sucking; thorax apparently a single piece due +to indistinct separation of its three segments: the compound eyes +reduced to a single ommatidium on each side. The short, powerful legs +are terminated by a single long claw. Metamorphosis incomplete. + +There has been a great deal of discussion regarding the structure of the +mouth-parts, and the relationships of the sucking lice, and the +questions cannot yet be regarded as settled. The conflicting views are +well represented by Cholodkovsky (1904 and 1905) and by Enderlein +(1904). + +[Illustration: 64. Pediculus showing the blind sac (_b_) containing the +mouth parts (_a_) beneath the alimentary canal (_p_). After Pawlowsky.] + +Following Graber, it is generally stated that the mouth-parts consist of +a short tube furnished with hooks in front, which constitutes the lower +lip, and that within this is a delicate sucking tube derived from the +fusion of the labrum and the mandibles. Opposed to this, Cholodkovsky +and, more recently, Pawlowsky, (1906), have shown that the piercing +apparatus lies in a blind sac under the pharynx and opening into the +mouth cavity (fig. 64). It does not form a true tube but a furrow with +its open surface uppermost. Eysell has shown that, in addition, there is +a pair of chitinous rods which he regards as the homologues of the +maxillæ. + +When the louse feeds, it everts the anterior part of the mouth cavity, +with its circle of hooks. The latter serve for anchoring the bug, and +the piercing apparatus is then pushed out. + +[Illustration: 65. Pediculus humanus, ventral aspect of male. (×10)] + +Most writers have classed the sucking lice as a sub-order of the +Hemiptera, but the more recent anatomical and developmental studies +render this grouping untenable. An important fact, bearing on the +question, is that, as shown by Gross, (1905), the structure of the +ovaries is radically different from that of the Hemiptera. + +Lice infestation and its effects are known medically as _pediculosis_. +Though their continued presence is the result of the grossest neglect +and filthiness, the original infestation may be innocently obtained and +by people of the most careful habits. + +Three species commonly attack man. Strangely enough, there are very few +accurate data regarding their life history. + +_Pediculus humanus_ (fig. 65), the head louse, is the most widely +distributed. It is usually referred to in medical literature as +_Pediculus capitis_, but the Linnean specific name has priority. In +color it is of a pale gray, blackish on the margins. It is claimed by +some authors that the color varies according to the color of the skin of +the host. The abdomen is composed of seven distinct segments, bearing +spiracles laterally. There is considerable variation in size. The males +average 1.8 mm. and the females 2.7 mm. in length. + +The eggs, fifty to sixty in number, stick firmly to the hairs of the +host and are known as nits. They are large and conspicuous, especially +on dark hair and are provided with an operculum, or cap, at the free +end, where the nymphs emerge. They hatch in about six days and about the +eighteenth day the young lice are sexually mature. + +[Illustration: 66. Pediculosis of the head. The illustration shows the +characteristic indications of the presence of lice, viz: the occipital +eczema gluing the hairs together, the swollen cervical glands, and the +porrigo, or eruption of contagious pustules upon the neck. After Fox.] + +The head lice live by preference on the scalp of their host but +occasionally they are found on the eyelashes and beard, or in the pubic +region. They may also occur elsewhere on the body. The penetration of +the rostrum into the skin and the discharge of an irritating saliva +produce a severe itching, accompanied by the formation of an eczema-like +eruption (fig. 66). When the infestation is severe, the discharge from +the pustules mats down the hair, and scabs are formed, under which the +insects swarm. "If allowed to run, a regular carapace may form, called +_trichoma_, and the head exudes a foetid odor. Various low plants may +grow in the trichoma, the whole being known as _plica +palonica_."--Stiles. + +Sources of infestation are various. School children may obtain the lice +from seatmates, by wearing the hats or caps of infested mates, or by the +use, in common, of brushes and combs. They may be obtained from infested +beds or sleeper berths. Stiles reports an instance in which a large +number of girls in a fashionable boarding school developed lousiness a +short time after traveling in a sleeping car. + +Treatment is simple, for the parasites may readily be controlled by +cleanliness and washing the head with a two per cent solution of +carbolic acid or even kerosene. The latter is better used mixed with +equal parts of olive oil, to avoid irritation. The treatment should be +applied at night and followed the next morning by a shampoo with soap +and warm water. It is necessary to repeat the operation in a few days. +Xylol, used pure, or with the addition of five per cent of vaseline, is +also very efficacious. Of course, the patient must be cautioned to stay +away from a lighted lamp or fire while using either the kerosene or +xylol. While these treatments will kill the eggs or nits, they will not +remove them from the hairs. Pusey recommends repeated washings with +vinegar or 25 per cent of acetic acid in water, for the purpose of +loosening and removing the nits. + +Treatment of severe infestations in females is often troublesome on +account of long hair. For such cases the following method recommended by +Whitfield (1912) is especially applicable: + +The patient is laid on her back on the bed with her head over the edge, +and beneath the head is placed a basin on a chair so that the hair lies +in the basin. A solution of 1 in 40 carbolic acid is then poured over +the hair into the basin and sluiced backwards and forwards until the +whole of the hair is thoroughly soaked with it. It is especially +necessary that care should be taken to secure thorough saturation of the +hair over the ears and at the nape of the neck, since these parts are +not only the sites of predilection of the parasites but they are apt to +escape the solution. This sluicing is carried out for ten minutes by the +clock. At the end of ten minutes the hair is lifted from the basin and +allowed to drain, but is not dried or even thoroughly wrung out. The +whole head is then swathed with a thick towel or better, a large piece +of common house flannel, which is fastened up to form a sort of turban, +and is allowed to remain thus for an hour. It can then be washed or +simply allowed to dry, as the carbolic quickly disperses. At the end of +this period every pediculus and what is better, every ovum is dead and +no relapse will occur unless there is exposure to fresh contagion. +Whitfield states that there seem to be no disadvantages in this method, +which he has used for years. He has never seen carboluria result from +it, but would advise first cutting the hair of children under five years +of age. + +_Pediculus corporis_ (= _P. vestimenti_) the body louse, is larger than +the preceding species, the female measuring 3.3 mm., and the male 3 mm. +in length. The color is a dirty white, or grayish. _P. corporis_ has +been regarded by some authorities as merely a variety of _P. humanus_ +but Piaget maintains there are good characters separating the two +species. + +The body louse lives in the folds and seams of the clothing of its host, +passing to the skin only when it wishes to feed. Brumpt states that he +has found enormous numbers of them in the collars of glass-ware or +grains worn by certain naked tribes in Africa. + +Exact data regarding the life-history of this species have been +supplied, in part, by the work of Warburton (1910), cited by Nuttall. He +found that _Pediculus corporis_ lives longer than _P. humanus_ under +adverse conditions. This is doubtless due to its living habitually on +the clothing, whereas _humanus_ lives upon the head, where it has more +frequent opportunities of feeding. He reared a single female upon his +own person, keeping the louse enclosed in a cotton-plugged tube with a +particle of cloth to which it could cling. The tube was kept next to his +body, thus simulating the natural conditions of warmth and moisture +under which the lice thrive. The specimen was fed twice daily, while it +clung to the cloth upon which it rested. Under these conditions she +lived for one month. Copulation commenced five days after the female had +hatched and was repeated a number of times, sexual union lasting for +hours. The female laid one hundred and twenty-four eggs within +twenty-five days. + +The eggs hatched after eight days, under favorable conditions, such as +those under which the female was kept. They did not hatch in the cold. +Eggs kept near the person during the day and hung in clothing by the +bedside at night, during the winter, in a cold room, did not hatch until +the thirty-fifth day. When the nymphs emerge from the eggs, they feed at +once, if given a chance to do so. They are prone to scatter about the +person and abandon the fragment of cloth to which the adult clings. + +The adult stage is reached on the eleventh day, after three molts, about +four days apart. Adults enter into copulation about the fifth day and as +the eggs require eight days for development, the total cycle, under +favorable conditions, is about twenty-four days. Warburton's data differ +considerably from those commonly quoted and serve to emphasize the +necessity for detailed studies of some of the commonest of parasitic +insects. + +Body lice are voracious feeders, producing by their bites and the +irritating saliva which they inject, rosy elevations and papules which +become covered with a brownish crust. The intense itching provokes +scratching, and characteristic white scars (fig. 67) surrounded by +brownish pigment (fig. 68) are formed. The skin may become thickened and +take on a bronze tinge. This melanoderma is especially marked in the +region between the shoulders but it may become generalized, a prominent +characteristic of "vagabond's disease." According to Dubre and Beille, +this melanoderma is due to a toxic substance secreted by the lice, which +indirectly provokes the formation of pigment. + +[Illustration: 67. Pediculosis in man caused by the body louse. After +Morrow.] + +Control measures, in the case of the body louse, consist in boiling or +steaming the clothes or in some cases, sterilizing by dry heat. The +dermatitis may be relieved by the use of zinc-oxide ointment, to which +Pusey recommends that there be added, on account of their parasiticidal +properties, sulphur and balsam of Peru, equal parts, 15 to 30 grains to +the ounce. + +_Phthirius pubis_ (= _P. inguinalis_), the pubic louse, or so-called +"crab louse," differs greatly from the preceding in appearance. It is +characterized by its relatively short head which fits into a broad +depression in the thorax. The latter is broad and flat and merges into +the abdomen. The first pair of legs is slender and terminated by a +straight claw. The second and third pairs of legs are thicker and are +provided with powerful claws fitted for clinging to hairs. The females +(fig. 69) measure 1.5 to 2 mm. in length by 1.5 mm. in breadth. The male +averages a little over half as large. The eggs, or nits, are fixed at +the base of the hairs. Only a few, ten to fifteen are deposited by a +single female, and they hatch in about a week's time. The young lice +mature in two weeks. + +The pubic louse usually infests the hairs of the pubis and the perineal +region. It may pass to the arm pits or even to the beard or moustache. +Rarely, it occurs on the eyelids, and it has even been found, in a very +few instances, occurring in all stages, on the scalp. Infestation may be +contracted from beds or even from badly infested persons in a crowd. We +have seen several cases which undoubtedly were due to the use of public +water closets. It produces papular eruption and an intense pruritis. +When abundant, there occurs a grayish discoloration of the skin which +Duguet has shown is due to a poisonous saliva injected by the louse, as +is the melanoderma caused by the body louse. + +[Illustration: 68. Melanoderma caused by the body louse. From Portfolio +of Dermochromes, by permission of Rebman & Co., New York, Publishers.] + +The pubic louse may be exterminated by the measures recommended for the +head louse, or by the use of officinal mercurial ointment. + +[Illustration: 69. Phthirius pubis. Ventral aspect of female. (×12).] + + +HEMIPTERA + +Several species of HEMIPTERA-HETEROPTERA are habitual parasites of man, +and others occur as occasional or accidental parasites. Of all these, +the most important and widespread are the bed-bugs, belonging to the +genus _Cimex_ (= _Acanthia_). + + +THE BED-BUGS--The bed-bugs are characterized by a much flattened oval +body, with the short, broad head unconstricted behind, and fitting into +the strongly excavated anterior margin of the thorax. The compound eyes +are prominent, simple eyes lacking. Antennæ four-jointed, the first +segment short, the second long and thick, and the third and fourth +slender. The tarsi are short and three segmented. + +It is often assumed in the literature of the subject that there is but a +single species of _Cimex_ attacking man, but several such species are to +be recognized. These are distinguishable by the characters given in +Chapter XII. We shall consider especially _Cimex lectularius_, the most +common and widespread species. + +_Cimex lectularius_ (= _Acanthia lectularia_, _Clinocoris +lectularius_), is one of the most cosmopolitan of human parasites but, +like the lice, it has been comparatively little studied until recent +years, when the possibility that it may be concerned with the +transmission of various diseases has awakened interest in the details of +its life-history and habits. + +[Illustration: 70. Cimex lectularius adult and eggs. Photograph by M. V. +S.] + +The adult insect (fig. 70) is 4-5 mm. long by 3 mm. broad, reddish brown +in color, with the beak and body appendages lighter in color. The short, +broad and somewhat rectangular head has no neck-like constriction but +fits into the broadly semilunar prothorax. The four segmented labium or +proboscis encloses the lancet-like maxillæ and mandibles. The distal of +the four antennal segments is slightly club-shaped. The prothorax is +characteristic of the species, being deeply incised anteriorly and with +its thin lateral margins somewhat turned up. The mesothorax is +triangular, with the apex posteriorly, and bears the greatly atrophied +first pair of wings. There is no trace of the metathoracic pair. The +greatly flattened abdomen has eight visible segments, though in reality +the first is greatly reduced and has been disregarded by most writers. +The body is densely covered with short bristles and hairs, the former +being peculiarly saber-shaped structures sharply toothed at the apex and +along the convex side (fig. 159_b_). + +The peculiar disagreeable odor of the adult bed-bug is due to the +secretion of the stink glands which lie on the inner surface of the +mesosternum and open by a pair of orifices in front of the metacoxæ, +near the middle line. In the nymphs, the thoracic glands are not +developed but in the abdomen there are to be found three unpaired dorsal +stink glands, which persist until the fifth molt, when they become +atrophied and replaced by the thoracic glands. The nymphal glands occupy +the median dorsal portion of the abdomen, opening by paired pores at the +anterior margin of the fourth, fifth and sixth segments. The secretion +is a clear, oily, volatile fluid, strongly acid in reaction. Similar +glands are to be found in most of the Hemiptera-Heteroptera and their +secretion is doubtless protective, through being disagreeable to the +birds. In the bed-bug, as Marlatt points out, "it is probably an +illustration of a very common phenomenon among animals, i.e., the +persistence of a characteristic which is no longer of any special value +to the possessor." In fact, its possession is a distinct disadvantage to +the bed-bug, as the odor frequently reveals the presence of the bugs, +before they are seen. + +The eggs of the bed-bug (fig. 70) are pearly white, oval in outline, +about a millimeter long, and possess a small operculum or cap at one +end, which is pushed off when the young hatches. They are laid +intermittently, for a long period, in cracks and crevices of beds and +furniture, under seams of mattresses, under loose wall paper, and +similar places of concealment of the adult bugs. Girault (1905) observed +a well-fed female deposit one hundred and eleven eggs during the +sixty-one days that she was kept in captivity. She had apparently +deposited some of her eggs before being captured. + +The eggs hatch in six to ten days, the newly emerged nymphs being about +1.5 mm. in length and of a pale yellowish white color. They grow slowly, +molting five times. At the last molt the mesathoracic wing pads appear, +characteristic of the adult. The total length of the nymphal stage +varies greatly, depending upon conditions of food supply, temperature +and possibly other factors. Marlatt (1907) found under most favorable +conditions a period averaging eight days between molting which, added to +an equal egg period, gave a total of about seven weeks from egg to adult +insect. Girault (1912) found the postembryonic period as low as +twenty-nine days and as high as seventy days under apparently similar +and normal conditions of food supply. Under optimum and normal +conditions of food supply, beginning August 27, the average nymphal life +was 69.9 days; average number of meals 8.75 and the molts 5. Under +conditions allowing about half the normal food supply the average +nymphal life was from 116.9 to 139 days. Nymphs starved from birth lived +up to 42 days. We have kept unfed nymphs, of the first stage, alive in a +bottle for 75 days. The interesting fact was brought out that under +these conditions of minimum food supply there were sometimes six molts +instead of the normal number. + +The adults are remarkable for their longevity, a factor which is of +importance in considering the spread of the insect and methods of +control. Dufour (1833) (not De Geer, as often stated) kept specimens for +a year, in a closed vial, without food. This ability, coupled with their +willingness to feed upon mice, bats, and other small mammals, and even +upon birds, accounts for the long periods that deserted houses and camps +may remain infested. There is no evidence that under such conditions +they are able to subsist on the starch of the wall paper, juices of +moistened wood, or the moisture in the accumulations of dust, as is +often stated. + +There are three or four generations a year, as Girault's breeding +experiments have conclusively shown. He found that the bed-bug does not +hibernate where the conditions are such as to allow it to breed and that +breeding is continuous unless interrupted by the lack of food or, during +the winter, by low temperature. + +Bed-bugs ordinarily crawl from their hiding places and attack the face +and neck or uncovered parts of the legs and arms of their victims. If +undisturbed, they will feed to repletion. We have found that the young +nymph would glut itself in about six minutes, though some individuals +fed continuously for nine minutes, while the adult required ten to +fifteen minutes for a full meal. When gorged, it quickly retreats to a +crack or crevice to digest its meal, a process which requires two or +three days. The effect of the bite depends very greatly on the +susceptibility of the individual attacked. Some persons are so little +affected that they may be wholly ignorant of the presence of a large +number of bugs. Usually the bite produces a small hard swelling, or +wheal, whitish in color. It may even be accompanied by an edema and a +disagreeable inflammation, and in such susceptible individuals the +restlessness and loss of sleep due to the presence of the insects may be +a matter of considerable importance. Stiles (1907) records the case of a +young man who underwent treatment for neurasthenia, the diagnosis being +agreed upon by several prominent physicians; all symptoms promptly +disappeared, however, immediately following a thorough fumigation of +his rooms, where nearly a pint of bed-bugs were collected. + +It is natural to suppose that an insect which throughout its whole life +is in such intimate relationship with man should play an important rôle +in the transmission of disease. Yet comparatively little is definitely +known regarding the importance of the bed-bug in this respect. It has +been shown that it is capable of transmitting the bubonic plague, and +South American trypanosomiasis. Nuttall succeeded in transmitting +European relapsing fever from mouse to mouse by its bite. It has been +claimed that Oriental sore, tuberculosis, and even syphilis may be so +carried. These phases of the subject will be considered later. + +The sources of infestation are many, and the invasion of a house is not +necessarily due to neglect, though the continued presence of the pests +is quite another matter. In apartments and closely placed houses they +are known to invade new quarters by migration. They are frequently to be +met with in boat and sleeper berths, and even the plush seats of day +coaches, whence a nucleus may be carried in baggage to residences. They +may be brought in the laundry or in clothes of servants. + +Usually they are a great scourge in frontier settlements and it is +generally believed that they live in nature under the bark of trees, in +lumber, and under similar conditions. This belief is founded upon the +common occurrence of bugs resembling the bed-bug, in such places. As a +matter of fact, they are no relation to bed-bugs but belong to +plant-feeding forms alone (fig. 19 _c_, _d_). + +It is also often stated that bed-bugs live in poultry houses, in +swallows nests, and on bats, and that it is from these sources that they +gain access to dwellings. These bugs are specifically distinct from the +true bed-bug, but any of them may, rarely, invade houses. Moreover, +chicken houses are sometimes thoroughly infested with the true _Cimex +lectularius_. + +Control measures consist in the use of iron bedsteads and the reduction +of hiding places for the bugs. If the infestation is slight they may be +exterminated by a vigilant and systematic hunt, and by squirting +gasoline or alcohol into cracks and crevices of the beds, and furniture. +Fumigation must be resorted to in more general infestations. + +The simplest and safest method of fumigation is by the use of flowers of +sulphur at the rate of two pounds to each one thousand cubic feet of +room space. The sulphur should be placed in a pan, a well made in the +top of the pile and a little alcohol poured in, to facilitate burning. +The whole should be placed in a larger pan and surrounded by water so as +to avoid all danger of fire. Windows should be tightly closed, beds, +closets and drawers opened, and bedding spread out over chairs in order +to expose them fully to the fumes. As metal is tarnished by the sulphur +fumes, ornaments, clocks, instruments, and the like should be removed. +When all is ready the sulphur should be fired, the room tightly closed +and left for twelve to twenty-four hours. Still more efficient in large +houses, or where many hiding places favor the bugs, is fumigation with +hydrocyanic acid gas. This is a deadly poison and must be used under +rigid precautions. Through the courtesy of Professor Herrick, who has +had much experience with this method, we give in the Appendix, the clear +and detailed directions taken from his bulletin on "Household Insects." + +Fumigation with formaldehyde gas, either from the liquid or "solid" +formalin, so efficient in the case of contagious diseases, is useless +against bed-bugs and most other insects. + + +OTHER BED-BUGS--_Cimex hemipterus_ (= _C. rotundatus_) is a tropical and +subtropical species, occurring in both the old and new world. Patton and +Cragg state that it is distributed throughout India, Burma, Assam, the +Malay Peninsula, Aden, the Island of Mauritius, Reunion, St. Vincent and +Porto Rico. "It is widely distributed in Africa, and is probably the +common species associated there with man." Brumpt also records it for +Cuba, the Antilles, Brazil, and Venezuela. + +This species, which is sometimes called the Indian bed-bug, differs from +_C. lectularius_ in being darker and in having a more elongate abdomen. +The head also is shorter and narrower, and the prothorax has rounded +borders. + +It has the same habits and practically the same life cycle as _Cimex +lectularius_. Mackie, in India, has found that it is capable of +transmitting the Asiatic type of recurrent fever. Roger suggested that +it was also capable of transmitting Kala-azar and Patton has described +in detail the developmental stages of _Leishmania_, the causative +organism of Kala-azar, in the stomach of this bug, but Brumpt declares +that the forms described are those of a common, non-pathogenic +flagellate to be found in the bug, and have nothing to do with the +human disease. Brumpt has shown experimentally that _Cimex hemipterus_ +may transmit _Trypanosoma cruzi_ in its excrement. + +_Cimex boueti_, occurring in French Guinea, is another species attacking +man. Its habits and general life history are the same as for the above +species. It is 3 to 4.5 mm. in length, has vestigial elytra, and much +elongated antennæ and legs. The extended hind legs are about as long as +the body. + +_Cimex columbarius_, a widely distributed species normally living in +poultry houses and dove cotes, _C. inodorus_, infesting poultry in +Mexico, _C. hirundinis_, occurring in the nests of swallows in Europe +and _Oeciacus vicarius_ (fig. 19i) occurring in swallows' nests in this +country, are species which occasionally infest houses and attack man. + +[Illustration: 71. Conorhinus sanguisugus.] + +_Conorhinus sanguisugus_, the cone-nosed bed-bug. We have seen in our +consideration of poisonous insects, that various species of Reduviid +bugs readily attack man. Certain of these are nocturnal and are so +commonly found in houses that they have gained the name, of "big +bed-bugs." The most noted of these, in the United States, is _Conorhinus +sanguisugus_ (fig. 71), which is widely distributed in our Southern +States. + +Like its near relatives, _Conorhinus sanguisugus_ is carnivorous in +habit and feeds upon insects as well as upon mammalian and human blood. +It is reported as often occurring in poultry houses and as attacking +horses in barns. The life history has been worked out in considerable +detail by Marlatt, (1902), from whose account we extract the following. + +[Illustration: 72. Beak of Conorhinus sanguisugus. After Marlatt.] + +The eggs are white, changing to yellow and pink before hatching. The +young hatch within twenty days and there are four nymphal stages. In all +these stages the insect is active and predaceous, the mouth-parts (fig. +72) being powerfully developed. The eggs are normally deposited, and the +early stages are undoubtedly passed, out of doors, the food of the +immature forms being other insects. Immature specimens are rarely found +indoors. It winters both in the partly grown and adult stage, often +under the bark of trees or in any similar protection, and only in its +nocturnal spring and early summer flights does it attack men. Marlatt +states that this insect seems to be decidedly on the increase in the +region which it particularly infests,--the plains region from Texas +northward and westward. In California a closely related species of +similar habits is known locally as the "monitor bug." + +The effect of the bite of the giant bed-bug on man is often very severe, +a poisonous saliva apparently being injected into the wound. We have +discussed this phase of the subject more fully under the head of +poisonous insects. + +_Conorhinus megistus_ is a Brazilian species very commonly attacking +man, and of special interest since Chagas has shown that it is the +carrier of a trypanosomiasis of man. Its habits and life history have +been studied in detail by Neiva, (1910). + +This species is now pre-eminently a household insect, depositing its +eggs in cracks and crevices in houses, though this is a relatively +recent adaptation. The nymphs emerge in from twenty to forty days, +depending upon the temperature. There are five nymphal stages, and as in +the case of true bed-bugs, the duration of these is very greatly +influenced by the availability of food and by temperature. Neiva reckons +the entire life cycle, from egg to egg, as requiring a minimum of three +hundred and twenty-four days. + +The nymphs begin to suck blood in three to five days after hatching. +They usually feed at night and in the dark, attacking especially the +face of sleeping individuals. The bite occasions but little pain. The +immature insects live in cracks and crevices in houses and invade the +beds which are in contact with walls, but the adults are active flyers +and attack people sleeping in hammocks. The males as well as the females +are blood suckers. + +Like many blood-sucking forms, _Conorhinus megistus_ can endure for long +periods without food. Neiva received a female specimen which had been +for fifty-seven days alive in a tightly closed box. They rarely feed on +two consecutive days, even on small quantities of blood, and were never +seen to feed on three consecutive days. + +Methods of control consist in screening against the adult bugs, and the +elimination of crevices and such hiding places of the nymphs. Where the +infestation is considerable, fumigation with sulphur is advisable. + + +PARASITIC DIPTERA OR FLIES + +Of the DIPTERA or two-winged flies, many species occasionally attack +man. Of these, a few are outstanding pests, many of them may also serve +to disseminate disease, a phase of our subject which will be considered +later. We shall now consider the most important of the group from the +viewpoint of their direct attacks on man. + + +Psychodidæ or Moth-Flies + +The PSYCHODIDÆ or Moth-flies, include a few species which attack man, +and at least one species, _Phlebotomus papatasii_, is known to transmit +the so-called "three-day fever" of man. Another species is supposed to +be the vector of Peruvian verruga. + +The family is made up of small, sometimes very small, nematocerous +Diptera, which are densely covered with hairs, giving them a moth-like +appearance. The wings are relatively large, oval or lanceolate in shape, +and when at rest are held in a sloping manner over the abdomen, or are +held horizontally in such a way as to give the insect a triangular +outline. Not only is the moth-like appearance characteristic, but the +venation of the wings (fig. 163, d) is very peculiar and, according to +Comstock, presents an extremely generalized form. All of the +longitudinal veins separate near the base of the wing except veins R_2 +and R_3 and veins M_1 and M_2. Cross veins are wanting in most cases. + +Comparatively little is known regarding the life-history and habits of +the Psychodidæ, but one genus, _Phlebotomus_, contains minute, +blood-sucking species, commonly known as sand-flies. The family is +divided into two subfamilies, the PSYCHODINÆ and the PHLEBOTOMINÆ. The +second of these, the PHLEBOTOMINÆ, is of interest to us. + + +THE PHLEBOTOMINÆ--The Phlebotominæ differ from the Psychodinæ in that +the radical sector branches well out into the wing rather than at the +base of the wing. They are usually less hairy than the Psychodinæ. The +ovipositor is hidden and less strongly chitinized. The species attacking +man belong to the genus _Phlebotomus_, small forms with relatively +large, hairy wings which are held upright, and with elongate proboscis. +The mandibles and maxillæ are serrated and fitted for biting. + +According to Miss Summers (1913) there are twenty-nine known species of +the genus _Phlebotomus_, five European, eleven Asiatic, seven African +and six American. One species only, _Phlebotomus vexator_, has been +reported for the United States. This was described by Coquillett, +(1907), from species taken on Plummer's Island, Maryland. It measures +only 1.5 mm. in length. As it is very probable that this species is much +more widely distributed, and that other species of these minute flies +will be found to occur in our fauna, we quote Coquillett's description. + +_Phlebotomus vexator_, Coq.: Yellow, the mesonotum brown, hairs chiefly +brown; legs in certain lights appear brown, but are covered with a white +tomentum; wings hyaline, unmarked; the first vein (R_1) terminates +opposite one-fifth of the length of the first submarginal cell (cell +R_2); this cell is slightly over twice as long as its petiole; terminal, +horny portion of male claspers slender, bearing many long hairs; the +apex terminated by two curved spines which are more than one-half as +long as the preceding part, and just in front of these are two similar +spines, while near the middle of the length of this portion is a fifth +spine similar to the others. Length 1.5 mm. + +The life-history of the Phlebotomus flies has been best worked out for +the European _Phlebotomus papatasii_ and we shall briefly summarize the +account of Doerr and Russ (1913) based primarily on work on this +species. The European Phlebotomus flies appear at the beginning of the +warm season, a few weeks after the cessation of the heavy rains and +storms of springtime. They gradually become more abundant until they +reach their first maximum, which in Italy is near the end of July +(Grassi). They then become scarcer but reach a second maximum in +September. At the beginning of winter they vanish completely, +hibernating individuals not being found. + +After fertilization there is a period of eight to ten days before +oviposition. The eggs are then deposited, the majority in a single mass +covered by a slimy secretion from the sebaceous glands. The larvæ emerge +in fourteen to twenty days. There is uncertainty as to the length of +larval life, specimens kept in captivity remaining fifty or more days +without transforming. Growth may be much more rapid in nature. The larvæ +do not live in fluid media but in moist detritus in dark places. Marett +believes that they live chiefly on the excrement of pill-bugs (Oniscidæ) +and lizards. Pupation always occurs during the night. The remnants of +the larval skin remain attached to the last two segments of the +quiescent pupa and serve to attach it to the stone on which it lives. +The pupal stage lasts eleven to sixteen days, the adult escaping at +night. + +Only the females suck blood. They attack not only man but all +warm-blooded animals and, according to recent workers, also cold-blooded +forms, such as frogs, lizards, and larvæ. Indeed, Townsend (1914) +believes that there is an intimate relation between _Phlebotomus_ and +lizards, or other reptiles the world over. The Phlebotomus passes the +daylight hours within the darkened recesses of the loose stone walls and +piles of rock in order to escape wind and strong light. Lizards inhabit +the same places, and the flies, always ready to suck blood in the +absence of light and wind, have been found more prone to suck reptilian +than mammalian blood. + +On hot summer nights, when the wind is not stirring, the Phlebotomus +flies, or sand-flies, as they are popularly called, invade houses and +sleeping rooms in swarms and attack the inmates. As soon as light begins +to break the flies either escape to the breeding places, or cool, dark +places protected from the wind, or a part of them remain in the rooms, +hiding behind pictures, under garments, and in similar places. Wherever +the Phlebotomus flies occur they are an intolerable nuisance. On account +of their small size they can easily pass through the meshes of ordinary +screens and mosquito curtains. They attack silently and inflict a very +painful, stinging bite, followed by itching. The ankles, dorsum of the +feet, wrists, inner elbow, knee joint and similar places are favorite +places of attack, possibly on account of their more delicate skin. + +Special interest has been attracted to these little pests in recent +years, since it has been shown that they transmit the European +"pappatici fever" or "three day fever." More recently yet, it appears +that they are the carriers of the virus of the Peruvian "verruga." This +phase of the subject will be discussed later. + +Control measures have not been worked out. As Newstead says, "In +consideration of the facts which have so far been brought to light +regarding the economy of Phlebotomus, it is clearly evident that the +task of suppressing these insects is an almost insurmountable one. Had +we to deal with insects as large and as accessible as mosquitoes, the +adoption of prophylactic measures would be comparatively easy, but owing +to the extremely minute size and almost flea-like habits of the adult +insects, and the enormous area over which the breeding-places may occur, +we are faced with a problem which is most difficult of solution." For +these reasons, Newstead considers that the only really prophylactic +measures which can at present be taken, are those which are considered +as precautionary against the bites of the insects. + +Of repellents, he cites as one of the best a salve composed of the +following: + + Ol. Anisi 3 grs. + Ol. Eucalypti 3 grs. + Ol. Terebenth 3 grs. + Unq. Acid Borac. + +Of sprays he recommends as the least objectionable and at the same time +one of the most effective, formalin. "The dark portions and angles of +sleeping apartments should be sprayed with a one per cent. solution of +this substance every day during the season in which the flies are +prevalent. A fine spraying apparatus is necessary for its application +and an excessive amount must not be applied. It is considered an +excellent plan also to spray the mosquito curtains regularly every day +towards sunset; nets thus treated are claimed to repel the attacks of +these insects." This effectiveness of formalin is very surprising for, +as we have seen, it is almost wholly ineffective against bed-bugs, +mosquitoes, house flies and other insects, where it has been tried. + +A measure which promises to be very effective, where it can be adopted, +is the use of electric fans so placed as to produce a current of air in +the direction of the windows of sleeping apartments. On account of the +inability of the Phlebotomus flies to withstand even slight breezes, it +seems very probable that they would be unable to enter a room so +protected. + + +Culicidæ or Mosquitoes + +From the medical viewpoint, probably the most interesting and important +of the blood-sucking insects are the mosquitoes. Certainly this is true +of temperate zones, such as those of the United States. The result is +that no other group of insects has aroused such widespread interest, or +has been subjected to more detailed study than have the mosquitoes, +since their rôle as carriers of disease was made known. There is an +enormous literature dealing with the group, but fortunately for the +general student, this has been well summarized by a number of workers. +The most important and helpful of the general works are those of Howard +(1901), Smith (1904), Blanchard (1905), Mitchell (1907), and especially +of Howard, Dyar, and Knab, whose magnificent monograph is still in +course of publication. + +Aside from their importance as carriers of disease, mosquitoes are +notorious as pests of man, and the earlier literature on the group is +largely devoted to references to their enormous numbers and their +blood-thirstiness in certain regions. They are to be found in all parts +of the world, from the equator to the Arctic and Antarctic regions. +Linnæus, in the "Flora Lapponica," according to Howard, Dyar and Knab, +"dwells at some length upon the great abundance of mosquitoes in Lapland +and the torments they inflicted upon man and beast. He states that he +believes that nowhere else on earth are they found in such abundance and +he compares their numbers to the dust of the earth. Even in the open, +you cannot draw your breath without having your mouth and nostrils +filled with them; and ointments of tar and cream or of fish grease are +scarcely sufficient to protect even the case-hardened cuticle of the +Laplander from their bite. Even in their cabins, the natives cannot take +a mouthful of food or lie down to sleep unless they are fumigated almost +to suffocation." In some parts of the Northwestern and Southwestern +United States it is necessary to protect horses working in the fields by +the use of sheets or burlaps, against the ferocious attacks of these +insects. It is a surprising fact that even in the dry deserts of the +western United States they sometimes occur in enormous numbers. + +Until comparatively recent years, but few species of mosquitoes were +known and most of the statements regarding their life-history were based +upon the classic work of Reaumur (1738) on the biology of the rain +barrel mosquito, _Culex pipiens_. In 1896, Dr. Howard refers to +twenty-one species in the United States, now over fifty are known; +Giles, in 1900, gives a total of two hundred and forty-two for the world +fauna, now over seven hundred species are known. We have found eighteen +species at Ithaca, N. Y. + +All of the known species of mosquitoes are aquatic in the larval stage, +but in their life-histories and habits such great differences occur that +we now know that it is not possible to select any one species as typical +of the group. For our present purpose we shall first discuss the general +characteristics and structure of mosquitoes, and shall then give the +life-history of a common species, following this by a brief +consideration of some of the more striking departures from what have +been supposed to be the typical condition. + +The CULICIDÆ are slender, nematocerous Diptera with narrow wings, +antennæ plumose in the males, and usually with the proboscis much longer +than the head, slender, firm and adapted for piercing in the female. +The most characteristic feature is that the margins of the wings and, in +most cases, the wing veins possess a fringe of scale-like hairs. These +may also cover in part, or entirely, the head, thorax, abdomen and legs. +The females, only, suck blood. + +On account of the importance of the group in this country and the +desirability of the student being able to determine material in various +stages, we show in the accompanying figures the characters most used in +classification. + +The larvæ (fig. 73) are elongate, with the head and thorax sharply +distinct. The larval antennæ are prominent, consisting of a single +cylindrical and sometimes curved segment. The outer third is often +narrower and bears at its base a fan-shaped tuft of hairs, the +arrangement and abundance of which is of systematic importance. About +the mouth are the so-called rotary mouth brushes, dense masses of long +hairs borne by the labrum and having the function of sweeping food into +the mouth. The form and arrangement of thoracic, abdominal, and anal +tufts of hair vary in different species and present characteristics of +value. On either side of the eighth abdominal segment is a patch of +scales varying greatly in arrangement and number and of much value in +separating species. Respiration is by means of tracheæ which open at the +apex of the so-called anal siphon, when it is present. In addition, +there are also one or two pairs of tracheal gills which vary much in +appearance in different species. On the ventral side of the anal siphon +is a double row of flattened, toothed spines whose number and shape are +likewise of some value in separating species. They constitute the comb +or pecten. + +[Illustration: 73. Culex larva showing details of external structure.] + +The pupa (fig. 139, b) unlike that of most insects, is active, though it +takes no food. The head and thorax are not distinctly separated, but the +slender flexible abdomen in sharply marked off. The antennæ, +mouth-parts, legs, and wings of the future adult are now external, but +enclosed in chitinous cases. On the upper surface, near the base of the +wings are two trumpets, or breathing tubes, for the pupal spiracles are +towards the anterior end instead of at the caudal end, as in the larva. +At the tip of the abdomen is a pair of large chitinous swimming paddles. + +As illustrative of the life cycle of a mosquito we shall discuss the +development of a common house mosquito, _Culex pipiens_, often referred +to in the Northern United States as the rain barrel mosquito. Its life +cycle is often given as typical for the entire group, but, as we have +already emphasized, no one species can serve this purpose. + +The adults of _Culex pipiens_ hibernate throughout the winter in +cellars, buildings, hollow trees, or similar dark shelters. Early in the +spring they emerge and deposit their eggs in a raft-like mass. The +number of eggs in a single mass is in the neighborhood of two hundred, +recorded counts varying considerably. A single female may deposit +several masses during her life time. The duration of the egg stage is +dependent upon temperature. In the warm summer time the larvæ may emerge +within a day. The larvæ undergo four molts and under optimum conditions +may transform into pupæ in about a week's time. Under the same favorable +conditions, the pupal stage may be completed in a day's time. The total +life cycle of _Culex pipiens_, under optimum conditions, may thus be +completed in a week to ten days. This period may be considerably +extended under less favorable conditions of temperature and food supply. + +_Culex pipiens_ breeds continuously throughout the summer, developing in +rain barrels, horse troughs, tin cans, or indeed in any standing water +about houses, which lasts for a week or more. The catch basins of sewers +furnish an abundant supply of the pests under some conditions. Such +places, the tin gutters on residences, and all possible breeding places +must be considered in attempts to exterminate this species. + +Other species of mosquitoes may exhibit radical departures from _Culex +pipiens_ in life-history and habits. To control them it is essential +that the biological details be thoroughly worked out for, as Howard, +Dyar, and Knab have emphasized, "much useless labor and expense can be +avoided by an accurate knowledge of the habits of the species." For a +critical discussion of the known facts the reader is referred to their +monograph. We shall confine ourselves to a few illustrations. + +The majority of mosquitoes in temperate climates hibernate in the egg +stage, hatching in the spring or even mild winter days in water from +melting snow. It is such single-brooded species which appear in +astounding numbers in the far North. Similarly, in dry regions the eggs +may stand thorough dessication, and yet hatch out with great promptness +when submerged by the rains. "Another provision to insure the species +against destruction in such a case, exists in the fact * * * that not +all the eggs hatch, a part of them lying over until again submerged by +subsequent rains." In temperate North America, a few species pass the +winter in the larval state. An interesting illustration of this is +afforded by _Wyeomia smithii_, whose larvæ live in pitcher plants and +are to be found on the coldest winter days imbedded in the solid ice. +Late in the spring, the adults emerge and produce several broods during +the summer. + +In the United States, one of the most important facts which has been +brought out by the intensive studies of recent years is that certain +species are migratory and that they can travel long distances and become +an intolerable pest many miles from their breeding places. This was +forcibly emphasized in Dr. Smith's work in New Jersey, when he found +that migratory mosquitoes, developing in the salt marshes along the +coast, are the dominant species largely responsible for the fame of the +New Jersey mosquito. The species concerned are _Aedes sollicitans_, _A. +cantator_ and _A. tæniorhynchus_. Dr. Smith decided that the first of +these might migrate at least forty miles inland. It is obvious that +where such species are the dominant pest, local control measures are a +useless waste of time and money. Such migratory habits are rare, +however, and it is probable that the majority of mosquitoes do not fly +any great distance from their breeding places. + +While mosquitoes are thought of primarily as a pest of man, there are +many species which have never been known to feed upon human or mammalian +blood, no matter how favorable the opportunity. According to Howard, +Dyar, and Knab, this is true of _Culex territans_, one of the common +mosquitoes in the summer months in the Northern United States. There are +some species, probably many, in which the females, like the males, are +plant feeders. In experimental work, both sexes are often kept alive for +long periods by feeding them upon ripe banana, dried fig, raisins, and +the like, and in spite of sweeping assertions that mosquitoes must have +a meal of blood in order to stimulate the ovaries to development, some +of the common blood-sucking species, notably _Culex pipiens_, have been +bred repeatedly without opportunity to feed upon blood. + +The effect of the bite varies greatly with different species and depends +upon the susceptibility of the individual bitten. Some persons are +driven almost frantic by the attacks of the pests when their companions +seem almost unconscious of any inconvenience. Usually, irritation and +some degree of inflammation appear shortly following the bite. Not +infrequently a hardened wheal or even a nodule forms, and sometimes +scratching leads to secondary infection and serious results. + +The source of the poison is usually supposed to be the salivary glands +of the insect. As we have already pointed out, (p. 34), Macloskie +believed that one lobe of the gland, on each side, was specialized for +forming the poison, while a radically different view is that of +Schaudinn, who believed that the irritation is due to the expelled +contents of the oesophageal diverticula, which contain a gas and a +peculiar type of fungi or bacteria. In numerous attempts, Schaudinn was +unable to produce any irritation by applying the triturated salivary +glands to a wound, but obtained the typical result when he used the +isolated diverticula. + +The irritation of the bite may be relieved to some extent by using +ammonia water, a one per cent. alcoholic solution of menthol, or +preparations of cresol, or carbolic acid. Dr. Howard recommends rubbing +the bite gently with a piece of moist toilet soap. Castellani and +Chalmers recommend cleansing inflamed bites with one in forty carbolic +lotion, followed by dressing with boracic ointment. Of course, +scratching should be avoided as much as possible. + +Repellents of various kinds are used, for warding off the attacks of the +insects. We have often used a mixture of equal parts of oil of +pennyroyal and kerosene, applied to the hands and face. Oil of +citronella is much used and is less objectionable to some persons. A +recommended formula is, oil of citronella one ounce, spirits of camphor +one ounce, oil of cedar one-half ounce. A last resort would seem to be +the following mixture recommended by Howard, Dyar, and Knab for use by +hunters and fishermen in badly infested regions, against mosquitoes and +blackflies. + +Take 2¼ lbs. of mutton tallow and strain it. While still hot add ½ +lb. black tar (Canadian tar). Stir thoroughly and pour into the +receptacle in which it is to be contained. When nearly cool stir in +three ounces of oil of citronella and 1¼ oz. of pennyroyal. + +At night the surest protection is a good bed net. There are many types +of these in use, but in order to be serviceable and at the same time +comfortable it should be roomy and hung in such a way as to be stretched +tightly in every direction. We prefer one suspended from a broad, square +frame, supported by a right-angled standard which is fastened to the +head of the bed. It must be absolutely free from rents or holes and +tucked in securely under the mattress or it will serve merely as a +convenient cage to retain mosquitoes which gain an entrance. While such +nets are a convenience in any mosquito ridden community, they are +essential in regions where disease-carrying species abound. Screening of +doors, windows and porches, against the pests is so commonly practiced +in this country that its importance and convenience need hardly be +urged. + +Destruction of mosquitoes and prevention of breeding are of fundamental +importance. Such measures demand first, as we have seen, the correct +determination of the species which is to be dealt with, and a knowledge +of its life-history and habits. If it prove to be one of the migratory +forms, it is beyond mere local effort and becomes a problem demanding +careful organization and state control. An excellent illustration of the +importance and effectiveness of work along these lines is afforded by +that in New Jersey, begun by the late Dr. John B. Smith and being pushed +with vigor by his successor, Dr. Headlee. + +In any case, there is necessity for community action. Even near the +coast, where the migratory species are dominant, there are the local +species which demand attention and which cannot be reached by any +measures directed against the species of the salt marshes. The most +important of local measures consist in the destruction of breeding +places by filling or draining ponds and pools, clearing up of more +temporary breeding places, such as cans, pails, water barrels and the +like. Under conditions where complete drainage of swamps is +impracticable or undesirable, judicious dredging may result in a pool or +series of steep-sided pools deep enough to maintain a supply of fish, +which will keep down the mosquito larvæ. Where water receptacles are +needed for storage of rain water, they should be protected by careful +screening or a film of kerosene over the top of the water, renewed every +two weeks or so, so as to prevent mosquitoes from depositing their eggs. +When kerosene is used, Water drawn from the bottom of the receptacle +will not be contaminated by it to any injurious extent. Where ponds +cannot be drained much good will be accomplished by spraying kerosene +oil on the surface of the water, or by the introduction of fish which +will feed on the larvæ. + +Detailed consideration of the most efficient measures for controlling +mosquitoes is to be found in Dr. Howard's Bulletin No. 88 of the Bureau +of Entomology, "Preventive and remedial work against mosquitoes" or, in +more summarized form, in Farmers' Bulletin No. 444. One of these should +be obtained by any person interested in the problems of mosquito control +and public health. + +[Illustration: 74. Mouth parts of Simulium. After Grünberg.] + + +The Simuliidæ, or Black Flies + +The SIMULIIDÆ, or black flies, are small, dark, or black flies, with a +stout body and a hump-back appearance. The antennæ are short but +eleven-segmented, the wings broad, without scales or hairs, and with the +anterior veins stout but the others very weak. The mouth-parts (fig. 74) +are fitted for biting. + +The larvæ of the Simuliidæ (fig. 75) are aquatic and, unlike those of +mosquitoes, require a well ærated, or swiftly running water. Here they +attach to stones, logs, or vegetation and feed upon various +micro-organisms. They pupate in silken cocoons open at the top. Detailed +life-histories have not been worked out for most of the species. We +shall consider as typical that of _Simulium pictipes_, an inoffensive +species widely distributed in the Eastern United States, which has been +studied especially at Ithaca, N.Y. (Johannsen, 1903). + +[Illustration: 75. Larva of Simulium, (×8). After Garman.] + +The eggs are deposited in a compact yellowish layer on the surface of +rock, on the brinks of falls and rapids where the water is flowing +swiftly. They are elongate ellipsoidal in shape, about .4 by .18 mm. As +myriads of females deposit in the same place the egg patches may be +conspicuous coatings of a foot or much more in diameter. When first laid +they are enveloped in a yellowish white slime, which becomes darker, +until finally it becomes black just before the emerging of the larvæ. +The egg stage lasts a week. + +The larvæ (fig. 75) are black, soft skinned, somewhat cylindrical in +shape, enlarged at both ends and attenuated in the middle. The posterior +half is much stouter than the anterior part and almost club-shaped. The +head bears two large fan-shaped organs which aid in procuring food. +Respiration is accomplished by means of three so-called blood gills +which are pushed out from the dorsal part of the rectum. The larvæ occur +in enormous numbers, in moss-like patches. If removed from their natural +habitat and placed in quiet water they die within three or four hours. +Fastened to the rock by means of a disk-like sucker at the caudal end of +the body, they ordinarily assume an erect position. They move about on +the surface of the rocks, to a limited extent, with a looping gait +similar to that of a measuring worm, and a web is secreted which +prevents their being washed away by the swiftly flowing water. They feed +chiefly upon algæ and diatoms. + +The complete larval stage during the summer months occupies about four +weeks, varying somewhat with the temperature and velocity of the water. +At the end of this period they spin from cephalic glands, boot-shaped +silken cocoons within which they pupate. The cocoon when spun is firmly +attached to the rock and also to adjacent cocoons. Clustered +continuously over a large area and sometimes one above another, they +form a compact, carpet-like covering on the rocks, the reddish-brown +color of which is easily distinguishable from the jet-black appearance +of the larvæ. The pupal stage lasts about three weeks. The adult fly, +surrounded by a bubble of air, quickly rises to the surface of the water +and escapes. The adults (fig. 76) are apparently short lived and thus +the entire life cycle, from egg to egg is completed in approximately +eight weeks. + +[Illustration: 76. Simulium venustum, (×8). After Garman.] + +In the case of _Simulium pictipes_ at Ithaca, N. Y., the first brood of +adults emerges early in May and successive generations are produced +throughout the summer and early autumn. This species winters in the +larval condition. Most of the other species of _Simulium_ which have +been studied seem to be single brooded. + +While _Simulium pictipes_ does not attack man, there are a number of the +species which are blood-sucking and in some regions they are a veritable +scourge. In recent years the greatest interest in the group has been +aroused by Sambon's hypothesis that they transmit pellagra from man to +man. This has not been established, and, indeed, seems very doubtful, +but the importance of these insects as pests and the possibility that +they may carry disease make it urgent that detailed life-histories of +the hominoxious species be worked out. + +As pests a vivid account of their attacks is in Agassiz's "Lake +Superior" (p. 61), quoted by Forbes (1912). + +"Neither the love of the picturesque, however, nor the interests of +science, could tempt us into the woods, so terrible were the black +flies. This pest of flies which all the way hither had confined our +ramblings on shore pretty closely to the rocks and the beach, and had +been growing constantly worse, here reached its climax. Although +detained nearly two days, * * * we could only sit with folded hands, or +employ ourselves in arranging specimens, and such other operations as +could be pursued in camp, and under the protection of a 'smudge.' One, +whom scientific ardor tempted a little way up the river in a canoe, +after water plants, came back a frightful spectacle, with blood-red +rings round his eyes, his face bloody, and covered with punctures. The +next morning his head and neck were swollen as if from an attack of +erysipelas." + +There are even well authenticated accounts on record of death of humans +from the attacks of large swarms of these gnats. In some regions, and +especially in the Mississippi Valley in this country, certain species of +black flies have been the cause of enormous losses to farmers and +stockmen, through their attacks on poultry and domestic animals. C. V. +Riley states that in 1874 the loss occasioned in one county in Tennessee +was estimated at $500,000. + +The measures of prevention and protection against these insects have +been well summarized by Forbes (1912). They are of two kinds: "the use +of repellents intended to drive away the winged flies, and measures for +the local destruction of the aquatic larvæ. The repellents used are +either smudges, or surface applications made to keep the flies from +biting. The black-fly will not endure a dense smoke, and the well-known +mosquito smudge seems to be ordinarily sufficient for the protection of +man. In the South, leather, cloth, and other materials which will make +the densest and most stifling smoke, are often preserved for this use in +the spring. Smudges are built in pastures for the protection of stock, +and are kept burning before the doors of barns and stables. As the +black-flies do not readily enter a dark room, light is excluded from +stables as much as possible during the gnat season. If teams must be +used in the open field while gnats are abroad, they may be protected +against the attacks of the gnats by applying cotton-seed oil or axle +grease to the surface, especially to the less hairy parts of the +animals, at least twice a day. A mixture of oil and tar and, indeed, +several other preventives, are of practical use in badly infested +regions; but no definite test or exact comparison has been made with any +them in a way to give a record of the precise results." + +"It is easy to drive the flies from houses or tents by burning pyrethrum +powder inside; this either kills the flies or stupifies them so that +they do not bite for some time thereafter." * * * "Oil of tar is +commonly applied to the exposed parts of the body for the purpose of +repelling the insects, and this preparation is supplied by the Hudson +Bay Company to its employees. Minnesota fishermen frequently grease +their faces and hands with a mixture of kerosene and mutton tallow for +the same purpose." We have found a mixture of equal parts of kerosene +and oil of pennyroyal efficient. + +Under most circumstances very little can be done to destroy this insect +in its early stage, but occasionally conditions are such that a +larvicide can be used effectively. Weed (1904), and Sanderson (1910) +both report excellent results from the use of phinotas oil, a +proprietary compound. The first-mentioned also found that in some places +the larvæ could be removed by sweeping them loose in masses with stiff +stable brooms and then catching them downstream on wire netting +stretched in the water. + + +Chironomidæ or Midges + +The flies of this family, commonly known as midges, resemble mosquitoes +in form and size but are usually more delicate, and the wing-veins, +though sometimes hairy, are not fringed with scales. The venation is +simpler than in the mosquitoes and the veins are usually less distinct. + +These midges, especially in spring or autumn, are often seen in immense +swarms arising like smoke over swamps and producing a humming noise +which can be heard for a considerable distance. At these seasons they +are frequently to be found upon the windows of dwellings, where they are +often mistaken for mosquitoes. + +The larvæ are worm-like, but vary somewhat in form in the different +genera. Most of them are aquatic, but a few live in the earth, in +manure, decaying wood, under bark, or in the sap of trees, especially in +the sap which collects in wounds. + +[Illustration: 77. Culicoides guttipennis; (_a_) adult, (×15); (_b_) +head of same; (_c_) larva; (_d_) head; (_e_) pupa. After Pratt.] + +Of the many species of CHIRONOMIDÆ, (over eight hundred known), the vast +majority are inoffensive. The sub-family Ceratopogoninæ, however, forms +an exception, for some of the members of this group, known as sandflies, +or punkies, suck blood and are particularly troublesome in the +mountains, along streams, and at the seashore. Most of these have been +classed under the genus _Ceratopogon_, but the group has been broken up +into a number of genera and _Ceratopogon_, in the strict sense, is not +known to contain any species which sucks the blood of vertebrates. + +THE CERATOPOGONINÆ--The Ceratopogoninæ are among the smallest of the +Diptera, many of them being hardly a millimeter long and some not even +so large. They are Chironomidæ in which the thorax is not prolonged over +the head. The antennæ are filiform with fourteen (rarely thirteen) +segments in both sexes, those of the male being brush-like. The basal +segment is enlarged, the last segment never longer than the two +preceding combined, while the last five are sub-equal to, or longer than +the preceding segment. The legs are relatively stouter than in the other +Chironomidæ. The following three genera of this subfamily are best known +as blood suckers in this country. + +Of the genus _Culicoides_ there are many species occurring in various +parts of the world. A number are known to bite man and animals and it is +probable that all are capable of inflicting injury. In some localities +they are called punkies, in others, sand-flies, a name sometimes also +applied to the species of _Simulium_ and _Phlebotomus_. Owing to their +very small size they are known by some tribes of Indians as No-see-ums. +The larvæ are found in ponds, pools, water standing in hollow tree +stumps, and the like. Though probably living chiefly in fresh water, we +have found a species occurring in salt water. The larvæ are small, +slender, legless, worm-like creatures (fig. 77_c_) with small brown head +and twelve body segments. The pupæ (fig. 77_e_) are slender, more +swollen at the anterior end and terminating in a forked process. They +float nearly motionless in a vertical position, the respiratory tubes in +contact with the surface film. The adults are all small, rarely +exceeding 2¼ mm. in length. The wings are more or less covered with +erect setulæ or hairs and in many species variously spotted and marked +with iridescent blotches. The antennæ have fourteen segments, the palpi +usually five. The wing venation and mouth-parts are shown in figures 77 +and 78. Of the twenty or more species of this genus occurring in the +United States the following are known to bite: _C. cinctus_, _C. +guttipennis_, _C. sanguisuga_, _C. stellifer_, _C. variipennis_, _C. +unicolor_. + +[Illustration: 78. Culicoides guttipennis; mouth parts of adult. After +Pratt.] + +One of the most widely distributed and commonest species in the Eastern +States is _C. guttipennis_ (fig. 77a). It is black with brown legs, a +whitish ring before the apex of each femur and both ends of each tibia; +tarsi yellow, knobs of halteres yellow. Mesonotum opaque, brown, two +vittæ in the middle, enlarging into a large spot on the posterior half, +also a curved row of three spots in front of each wing, and the narrow +lateral margins, light gray pruinose. Wings nearly wholly covered with +brown hairs, gray, with markings as shown in the figure. Length one mm. + +_Johannseniella_ Will. is a wide-spread genus related to the foregoing. +Its mouth-parts are well adapted for piercing and it is said to be a +persistent blood sucker, particularly in Greenland. This genus is +distinguished from _Culicoides_ by its bare wings, the venation (fig. +163, c), and the longer tarsal claws. There are over twenty North +American species. + +[Illustration: 79. Chrysops univittatus, (×4). After Osborn.] + +In the Southwestern United States, _Tersesthes torrens_ Towns. occurs, a +little gnat which annoys horses, and perhaps man also, by its bite. It +is related to _Culicoides_ but differs in the number of antennal +segments and in its wing venation (fig. 163, e). The fly measures but +two mm. in length and is blackish in color. The antennæ of the female +have thirteen segments, the palpi but three, of which the second is +enlarged and swollen. + + +Tabanidæ or Horse-Flies + +The TABANIDÆ,--horse-flies, ear-flies, and deer-flies,--are well-known +pests of cattle and horses and are often extremely annoying to man. The +characteristics of the family and of the principal North American genera +are given in the keys of Chapter XII. There are over 2500 recorded +species. As in the mosquitoes, the females alone are blood suckers. The +males are flower feeders or live on plant juices. This is apparently +true also of the females of some of the genera. + +The eggs are deposited in masses on water plants or grasses and sedges +growing in marshy or wet ground. Those of a common species of _Tabanus_ +are illustrated in figure 80, _a_. They are placed in masses of several +hundred, light colored when first deposited but turning black. In a week +or so the cylindrical larvæ, tapering at both ends (fig. 80, _b_), +escape to the water, or damp earth, and lead an active, carnivorous +life, feeding mainly on insect larvæ, and worms. In the forms which have +been best studied the larval life is a long one, lasting for months or +even for more than a year. Until recently, little was known concerning +the life-histories of this group, but the studies of Hart (1895), and +Hine (1903 +) have added greatly to the knowledge concerning North +American forms. + +Many of the species attack man with avidity and are able to inflict +painful bites, which may smart for hours. In some instances the wound is +so considerable that blood will continue to flow after the fly has left. +We have seen several cases of secondary infection following such bites. + +[Illustration: 80. (_a_) Eggs of Tabanus. Photograph by J. T. Lloyd.] + +[Illustration: 80. (_b_) Larva of Tabanus. Photograph by M. V. S.] + +The horse-flies have been definitely convicted of transferring the +trypanosome of surra from diseased to healthy animals and there is good +evidence that they transfer anthrax. The possibility of their being +important agents in the conveyal of human diseases should not be +overlooked. Indeed, Leiper has recently determined that a species of +_Chrysops_ transfers the blood parasite _Filaria diurna_. + + +Leptidæ or Snipe-Flies + +The family LEPTIDÆ is made up of moderate or large sized flies, +predaceous in habit. They are sufficiently characterized in the keys of +Chapter XII. Four blood-sucking species belonging to three genera have +been reported. Of these _Symphoromyia pachyceras_ is a western species. +Dr. J. C. Bradley, from personal experience, reports it as a vicious +biter. + +[Illustration: 80. (_c_) Mouth parts of Tabanus. After Grünberg.] + + +Oestridæ or Bot-flies + +To the family OESTRIDÆ belong the bot and warble-flies so frequently +injurious to animals. The adults are large, or of medium size, heavy +bodied, rather hairy, and usually resemble bees in appearance. + +The larvæ live parasitically in various parts of the body of mammals, +such as the stomach (horse bot-fly), the subcutaneous connective tissue +(warble-fly of cattle), or the nasal passage (sheep bot-fly or head +maggot). + +There are on record many cases of the occurrence of the larvæ of +Oestridæ as occasional parasites of man. A number of these have been +collected and reviewed in a thesis by Mme. Pètrovskaia (1910). The +majority of them relate to the following species. + +_Gastrophilus hæmorrhoidalis_, the red tailed bot-fly, is one of the +species whose larvæ are most commonly found in the stomach of the horse. +Schoch (1877) cites the case of a woman who suffered from a severe case +of chronic catarrh of the stomach, and who vomited, and also passed from +the anus, larvæ which apparently belonged to this species. Such cases +are exceedingly rare but instances of subcutaneous infestation are +fairly numerous. In the latter type these larvæ are sometimes the cause +of the peculiar "creeping myasis." This is characterized at its +beginning by a very painful swelling which gradually migrates, producing +a narrow raised line four to twenty-five millimeters broad. When the +larva is mature, sometimes after several months, it becomes stationary +and a tumor is formed which opens and discharges the larva along with +pus and serum. + +_Gastrophilus equi_ is the most widespread and common of the horse +bot-flies. Portschinsky reports it as commonly causing subcutaneous +myasis of man in Russia. + +_Hypoderma bovis_ (= _Oestrus bovis_), and _Hypoderma lineata_ are the +so-called warble-flies of cattle. The latter species is the more common +in North America but Dr. C. G. Hewitt has recently shown that _H. bovis_ +also occurs. Though warbles are very common in cattle in this country, +the adult flies are very rarely seen. They are about half an inch in +length, very hairy, dark, and closely resemble common honey-bees. + +They deposit their eggs on the hairs of cattle and the animals in +licking themselves take in the young larvæ. These pass out through the +walls of the oesophagus and migrate through the tissues of the animal, +to finally settle down in the subcutaneous tissue of the back. The +possibility of their entering directly through the skin, especially in +case of infestation of man, is not absolutely precluded, although it is +doubtful. + +For both species of _Hypoderma_ there are numerous cases on record of +their occurrence in man. Hamilton (1893) saw a boy, six years of age, +who had been suffering for some months from the glands on one side of +his neck being swollen and from a fetid ulceration around the back teeth +of the lower jaw of the same side. Three months' treatment was of no +avail and the end seemed near; one day a white object, which was seen to +move, was observed in the ulcer at the root of the tongue, and on being +extracted was recognized as a full grown larva of _Hypoderma_. It was of +usual tawny color, about half an inch long when contracted, about one +third that thickness, and quite lively. The case resulted fatally. The +boy had been on a dairy farm the previous fall, where probably the egg +(or larva) was in some way taken into his mouth, and the larva found +between the base of the tongue and the jaw suitable tissue in which to +develop. + +Topsent (1901) reports a case of "creeping myasis" caused by _H. +lineata_ in the skin of the neck and shoulders of a girl eight years of +age. The larva travelled a distance of nearly six and a half inches. The +little patient suffered excruciating pain in the place occupied by the +larva. + +_Hypoderma diana_ infests deer, and has been known to occur in man. + +_Oestris ovis_, the sheep bot-fly, or head maggot, is widely distributed +in all parts of the world. In mid-summer the flies deposit living +maggots in the nostrils of sheep. These larvæ promptly pass up the nasal +passages into the frontal and maxillary sinuses, where they feed on the +mucous to be found there. In their migrations they cause great +irritation to their host, and when present in numbers may cause vertigo, +paroxysms, and even death. Portschinsky in an important monograph on +this species, has discussed in detail its relation to man. He shows that +it is not uncommon for the fly to attack man and that the minute living +larvæ are deposited in the eyes, nostrils, lips, or mouth. A typical +case in which the larvæ were deposited in the eye was described by a +German oculist Kayser, in 1905. A woman brought her six year old +daughter to him and said that the day before, about noontime, a flying +insect struck the eye of the child and that since then she had felt a +pain which increased towards evening. In the morning the pain ceased but +the eye was very red. She was examined at about noon, at which time she +was quiet and felt no pain. She was not sensitive to light, and the only +thing noticed was a slight congestion and accumulation of secretion in +the corner of the right eye. A careful examination of the eye disclosed +small, active, white larvæ that crawled out from the folds of the +conjunctiva and then back and disappeared. Five of these larvæ were +removed and although an uncomfortable feeling persisted for a while, the +eye became normal in about three weeks. + +Some of the other recorded cases have not resulted so favorably, for the +eyesight has been seriously affected or even lost. + +According to Edmund and Etienne Sergent (1907), myasis caused by the +larvæ of _Oestris ovis_ is very common among the shepherds in Algeria. +The natives say that the fly deposits its larvæ quickly, while on the +wing, without pause. The greatest pain is caused when these larvæ +establish themselves in the nasal cavities. They then produce severe +frontal headaches, making sleep impossible. This is accompanied by +continuous secretion from the nasal cavities and itching pains in the +sinuses. If the larvæ happen to get into the mouth, the throat becomes +inflamed, swallowing is painful, and sometimes vomiting results. The +diseased condition may last for from three to ten days or in the case of +nasal infection, longer, but recovery always follows. The natives remove +the larvæ from the eye mechanically by means of a small rag. When the +nose is infested, tobacco fumigations are applied, and in case of throat +infestation gargles of pepper, onion, or garlic extracts are used. + +_Rhinoestrus nasalis_, the Russian gad-fly, parasitizes the +nasopharyngeal region of the horse. According to Portschinsky, it not +infrequently attacks man and then, in all the known cases deposits its +larvæ in the eye, only. This is generally done while the person is +quiet, but not during sleep. The fly strikes without stopping and +deposits its larva instantaneously. Immediately after, the victim +experiences lancinating pains which without intermission increase in +violence. There is an intense conjunctivitis and if the larvæ are not +removed promptly the envelopes of the eye are gradually destroyed and +the organ lost. + +[Illustration: 81. Larvæ of Dermatobia cyaniventris. After Blanchard.] + +[Illustration: 82. Young larva of Dermatobia cyaniventris. After +Surcouf.] + +_Dermatobia cyaniventris_--This fly (fig. 83) is widely distributed +throughout tropical America, and in its larval stage is well known as a +parasite of man. The larvæ (figs. 81 and 82) which are known as the "ver +macaque," "torcel," "ver moyocuil" or by several other local names, +enter the skin and give rise to a boil-like swelling, open at the top, +and comparable with the swelling produced by the warble fly larvæ, in +cattle. They cause itching and occasional excruciating pain. When +mature, nearly an inch in length, they voluntarily leave their host, +drop to the ground and complete their development. The adult female is +about 12 mm. in length. The face is yellow, the frons black with a +grayish bloom; antennæ yellow, the third segment four times as long as +the second, the arista pectinate. The thorax is bluish black with +grayish bloom; the abdomen depressed, brilliant metallescent blue with +violet tinge. The legs are yellowish, the squamæ and wings brownish. + +The different types of larvæ represented in figure 81 were formerly +supposed to belong to different species but Blanchard regards them as +merely various stages of the same species. It is only very recently that +the early stage and the method by which man becomes infested were made +known. + +[Illustration: 83. Dermatobia cyaniventris (×1¾). After Manson.] + +[Illustration: 84. Mosquito carrying eggs of Dermatobia cyaniventris. +After Surcouf.] + +About 1900, Blanchard observed the presence of packets of large-sized +eggs under the abdomen of certain mosquitoes from Central America; and +in 1910, Dr. Moralès, of Costa Rica, declared that the Dermatobia +deposited its eggs directly under the abdomen of the mosquito and that +they were thus carried to vertebrates. Dr. Nunez Tovar observed the +mosquito carriers of the eggs and placing larvæ from this source on +animals, produced typical tumors and reared the adult flies. It remained +for Surcouf (1913) to work out the full details. He found that the +Dermatobia deposits its eggs in packets covered by a very viscid +substance, on leaves. These become attached to mosquitoes of the species +_Janthinosoma lutzi_ (fig. 84) which walk over the leaves. The eggs +which adhere to the abdomen, remain attached and are thus transported. +The embryo develops, but the young larva (fig. 82) remains in the egg +until it has opportunity to drop upon a vertebrate fed upon by the +mosquito. + + +Muscidæ + +The following MUSCIDÆ, characterized elsewhere, deserve special mention +under our present grouping of parasitic species. Other important species +will be considered as facultative parasites. + +_Stomoxys calcitrans_, the stable-fly, or the biting house-fly, is often +confused with _Musca domestica_ and therefore is discussed especially in +our consideration of the latter species as an accidental carrier of +disease. Its possible relation to the spread of infantile paralysis is +also considered later. + +The _tsetse flies_, belonging to the genus _Glossina_, are African +species of blood-sucking Muscidæ which have attracted much attention +because of their rôle in transmitting various trypanosome diseases of +man and animals. They are characterized in Chapter XII and are also +discussed in connection with the diseases which they convey. + +_Chrysomyia macellaria_, (= _Compsomyia_), the "screw worm"-fly is one +of the most important species of flies directly affecting man, in North +America. It is not normally parasitic, however, and hence will be +considered with other facultative parasites in Chapter IV. + +[Illustration: 85. Larva of Auchmeromyia luteola. After Graham-Smith.] + +[Illustration: 86. Auchmeromyia luteola (×4). After Graham-Smith.] + +_Auchmeromyia luteola_, the Congo floor maggot. This is a muscid of +grewsome habits, which has a wide distribution throughout Africa. The +fly (fig. 86) deposits its eggs on the ground of the huts of the +natives. The whitish larvæ (fig. 85) on hatching are slightly flattened +ventrally, and each segment bears posteriorly three foot-pads +transversely arranged. At night the larvæ find their way into the low +beds or couches of the natives and suck their blood. The adult flies do +not bite man and, as far as known, the larvæ do not play any rôle in the +transmission of sleeping sickness or other diseases. + +This habit of blood-sucking by muscid larvæ is usually referred to as +peculiar to _Auchmeromyia luteola_ but it should be noted that the larvæ +of _Protocalliphora_ frequent the nests of birds and feed upon the +young. Mr. A. F. Coutant has studied especially the life-history and +habits of _P. azurea_, whose larvæ he found attacking young crows at +Ithaca, N.Y. He was unable to induce the larvæ to feed on man. + +[Illustration: 87. Cordylobia anthropophaga (×3). After Fülleborn.] + +[Illustration: 88. Larva of Cordylobia anthropophaga. After Blanchard.] + +_Cordylobia anthropophaga_, (_Ochromyia anthropophaga_), or Tumbu-fly +(fig. 87) is an African species whose larvæ affect man much as do those +of _Dermatobia cyniventris_, of Central and South America. The larva +(fig. 88), which is known as "ver du Cayor" because it was first +observed in Cayor, in Senegambia, develops in the skin of man and of +various animals, such as dogs, cats, and monkeys. It is about 12 mm. in +length, and of the form of the larvæ of other muscids. Upon the +intermediate segments are minute, brownish recurved spines which give to +the larva its characteristic appearance. The life-history is not +satisfactorily worked out, but Fuller (1914), after reviewing the +evidence believes that, as a rule, it deposits its young in the sleeping +places of man and animals, whether such be a bed, a board, the floor, or +the bare ground. In the case of babies, the maggots may be deposited on +the scalp. The minute maggots bore their way painlessly into the skin. +As many as forty parasites have been found in one individual and one +author has reported finding more than three hundred in a spaniel puppy. +Though their attacks are at times extremely painful, it is seldom that +any serious results follow. + + +THE SIPHONAPTERA OR FLEAS + +The SIPHONAPTERA, or fleas (fig. 89) are wingless insects, with highly +chitinized and laterally compressed bodies. The mouth-parts are formed +for piercing and sucking. Compound eyes are lacking but some species +possess ocelli. The metamorphosis is complete. + +This group of parasites, concerning which little was known until +recently, has assumed a very great importance since it was learned that +fleas are the carriers of bubonic plague. Now over four hundred species +are known. Of these, several species commonly attack man. The most +common hominoxious species are _Pulex irritans_, _Xenopsylla cheopis_, +_Ctenocephalus canis_, _Ctenocephalus felis_, _Ceratophyllus fasciatus_ +and _Dermatophilus penetrans_, but many others will feed readily on +human blood if occasion arises. + +[Illustration: 89. Xenopsylla cheopis, male (×25). After Jordan and +Rothschild.] + +We shall treat in this place of the general biology and habits of the +hominoxious forms and reserve for the systematic section the discussion +of the characteristics of the different genera. + +The most common fleas infesting houses in the Eastern United States are +the cosmopolitan dog and cat fleas, _Ctenocephalus canis_ (fig. 90) and +_C. felis_. Their life cycles will serve as typical. These two species +have until recently been considered as one, under the name _Pulex +serraticeps_. See figure 92. + +[Illustration: 90. Dog flea (×15). After Howard.] + +The eggs are oval, slightly translucent or pearly white, and measure +about .5 mm. in their long diameter. They are deposited loosely in the +hairs of the host and readily drop off as the animal moves around. +Howard found that these eggs hatch in one to two days. The larvæ are +elongate, legless, white, worm-like creatures. They are exceedingly +active, and avoid the light in every way possible. They cast their first +skin in from three to seven days and their second in from three to four +days. They commenced spinning in from seven to fourteen days after +hatching and the imago appeared five days later. Thus in summer, at +Washington, the entire life cycle may be completed in about two weeks. +(cf. fig. 91, 92). + +Strickland's (1914) studies on the biology of the rat flea, +_Ceratophyllus fasciatus_, have so important a general bearing that we +shall cite them in considerable detail. + +[Illustration: 91. Larva of Xenopsylla cheopis. After Bacot and +Ridewood.] + +He found, to begin with, that there is a marked inherent range in the +rate of development. Thus, of a batch of seventy-three eggs, all laid in +the same day and kept together under the same conditions, one hatched in +ten days; four in eleven days; twenty-five in twelve days; thirty-one in +thirteen days; ten in fourteen days; one in fifteen days; and one in +sixteen days. Within these limits the duration of the egg period seems +to depend mainly on the degree of humidity. The incubation period is +never abnormally prolonged as in the case of lice, (Warburton) and +varying conditions of temperature and humidity have practically no +effect on the percentage of eggs which ultimately hatch. + +The same investigator found that the most favorable condition for the +larva is a low temperature, combined with a high degree of humidity; and +that the presence of rubbish in which the larva may bury itself is +essential to its successful development. When larvæ are placed in a +bottle containing either wood-wool soiled by excrement, or with feathers +or filter paper covered with dried blood they will thrive readily and +pupate. They seem to have no choice between dried blood and powdered rat +feces for food, and also feed readily on flea excrement. They possess +the curious habit of always devouring their molted skins. + +[Illustration: 92. Head and pronotum of (_a_) dog flea; (_b_) of cat +flea; (_c_) of hen flea. After Rothschild. (_d_) Nycteridiphilus +(Ischnopsyllus) hexactenus. After Oudemans.] + +An important part of Strickland's experiments dealt with the question of +duration of the pupal stage under the influence of temperature and with +the longevity and habits of the adult. In October, he placed a batch of +freshly formed cocoons in a small dish that was kept near a white rat in +a deep glass jar in the laboratory. Two months later one small and +feeble flea had emerged, but no more until February, four months after +the beginning of the experiment. Eight cocoons were then dissected and +seven more found to contain the imago fully formed but in a resting +state. The remainder of the batch was then placed at 70° F. for one +night, near a white rat. The next day all the cocoons were empty and the +fleas were found on the white rat. + +Thus, temperature greatly influences the duration of the pupal period, +which in _Ceratophyllus fasciatus_ averages seventeen days. Moreover, +when metamorphosis is complete a low temperature will cause the imago to +remain within the cocoon. + +Sexually mature and ovipositing fleas, he fed at intervals and kept +alive for two months, when the experiment was discontinued. In the +presence of rubbish in which they could bury themselves, unfed rat fleas +were kept alive for many months, whereas in the absence of any such +substratum they rarely lived a month. In the former case, it was found +that the length of life is influenced to some degree by the temperature +and humidity. In an experiment carried out at 70° F. and 45 per cent +humidity, the fleas did not live for more than four months, while in an +experiment at 60° F. and 70 per cent humidity they lived for at least +seventeen months. There was no indication that fleas kept under these +conditions sucked moisture from surrounding objects, and those kept in +bell jars, with an extract of flea-rubbish on filter paper, did not live +any longer than those which were not so supplied. + +Curiously enough, although the rat is the normal host of _Ceratophyllus +fasciatus_, it was found that when given the choice these fleas would +feed upon man in preference to rats. However, none of the fleas laid +eggs unless they fed on rat blood. + +The experiments of Strickland on copulation and oviposition in the rat +flea showed that fleas do not copulate until they are sexually mature +and that, at least in the case of _Ceratophyllus fasciatus_, the +reproductive organs are imperfectly developed for some time (more than a +week) after emerging from the pupa. When mature, copulation takes place +soon after the fleas have fed on their true host--the rat--but not if +they have fed on a facultative host only, such as man. Copulation is +always followed by oviposition within a very short time. + +The effect of the rat's blood on the female with regard to egg-laying, +Strickland concludes, is stimulating rather than nutritive, as fleas +that were without food for many months were observed to lay eggs +immediately after one feed. Similarly, the male requires the stimulus of +a meal of rat's blood before it displays any copulatory activity. + +Mitzmain (1910) has described in detail the act of biting on man, as +observed in the squirrel flea, _Ceratophyllus acutus_. "The flea when +permitted to walk freely on the arm selects a suitable hairy space where +it ceases abruptly in its locomotion, takes a firm hold with the tarsi, +projects its proboscis, and prepares to puncture the skin. A puncture is +drilled by the pricking epipharynx, the saw-tooth mandibles +supplementing the movement by lacerating the cavity formed. The two +organs of the rostrum work alternately, the middle piece boring, while +the two lateral elements execute a sawing movement. The mandibles, owing +to their basal attachments, are, as is expressed by the advisory +committee on plague investigations in India (_Journal of Hygiene_, vol. +6, No. 4, p. 499), 'capable of independent action, sliding up and down +but maintaining their relative positions and preserving the lumen of the +aspiratory channel.' The labium doubles back, the V-shaped groove of +this organ guiding the mandibles on either side." + +"The action of the proboscis is executed with a forward movement of the +head and a lateral and downward thrust of the entire body. As the +mouth-parts are sharply inserted, the abdomen rises simultaneously. The +hind and middle legs are elevated, resembling oars. The forelegs are +doubled under the thorax, the tibia and tarsi resting firmly on the +epidermis serve as a support for the body during the feeding. The +maxillary palpi are retracted beneath the head and thorax. The labium +continues to bend, at first acting as a sheath for the sawing mandibles, +and as these are more deeply inserted, it bends beneath the head with +the elasticity of a bow, forcing the mandibles into the wound until the +maxillæ are embedded in the skin of the victim. When the proboscis is +fully inserted, the abdomen ceases for a time its lateral swinging." + +"The acute pain of biting is first felt when the mandibles have not +quite penetrated and subsequently during each distinct movement of the +abdomen. The swinging of the abdomen gradually ceases as it becomes +filled with blood. The sting of the biting gradually becomes duller and +less sensitive as feeding progresses. The movements of the elevated +abdomen grow noticeably feebler as the downward thrusts of the springy +bow-like labium becomes less frequent." + +"As the feeding process advances one can discern through the translucent +walls of the abdomen a constant flow of blood, caudally from the +pharynx, accompanied by a peristaltic movement. The end of the meal is +signified in an abrupt manner. The flea shakes its entire body, and +gradually withdraws its proboscis by lowering the abdomen and legs and +violently twisting the head." + +"When starved for several days the feeding of the rat fleas is conducted +in a rather vigorous manner. As soon as the proboscis is buried to the +full length the abdomen is raised and there ensues a gradual lateral +swaying motion, increasing the altitude of the raised end of the abdomen +until it assumes the perpendicular. The flea is observed at this point +to gain a better foothold by advancing the fore tarsi, and then, +gradually doubling back the abdomen, it turns with extreme agility, +nearly touching with its dorsal side the skin of the hand upon which it +is feeding. Meanwhile, the hungry parasite feeds ravenously." + +"It is interesting to note the peculiar nervous action which the rodent +fleas exhibit immediately when the feeding process is completed or when +disturbed during the biting. Even while the rostrum is inserted to the +fullest the parasite shakes its head spasmodically; in a twinkling the +mouth is withdrawn and then the flea hops away." + +A habit of fleas which we shall see is of significance in considering +their agency in the spread of bubonic plague, is that of ejecting blood +from the anus as they feed. + +Fleas are famous for their jumping powers, and in control measures it is +of importance to determine their ability along this line. It is often +stated that they can jump about four inches, or, according to the Indian +Plague Commission _Xenopsylla cheopis_ cannot hop farther than five +inches. Mitzmain (1910) conducted some careful experiments in which he +found that the human flea, _Pulex irritans_, was able to jump as far as +thirteen inches on a horizontal plane. The mean average of five +specimens permitted to jump at will was seven and three-tenths inches. +The same species was observed to jump perpendicularly to a height of at +least seven and three-fourths inches. Other species were not able to +equal this record. + +The effect of the bite of fleas on man varies considerably according to +the individual susceptibility. According to Patton and Cragg, this was +borne out in a curious manner by the experiments of Chick and Martin. +"In these, eight human hosts were tried; in seven, little or no +irritation was produced, while in one quite severe inflammation was set +up around each bite." Of two individuals, equally accustomed to the +insects, going into an infested room, one may be literally tormented by +them while the other will not notice them. Indeed it is not altogether +a question of susceptibility, for fleas seem to have a special +predilection for certain individuals. The typical itching wheals +produced by the bites are sometimes followed, especially after +scratching, by inflammatory papules. + +The itching can be relieved by the use of lotions of carbolic acid (2-3 +per cent), camphor, menthol lotion, or carbolated vaseline. If forced to +sleep in an infested room, protection from attacks can be in a large +measure gained by sprinkling pyrethrum, bubach, or California insect +powder between the sheets. The use of camphor, menthol, or oil of +eucalyptus, or oil of pennyroyal is also said to afford protection to a +certain extent. + +In the Eastern United States the occurrence of fleas as household pests +is usually due to infested cats and dogs which have the run of the +house. We have seen that the eggs are not attached to the host but drop +to the floor when they are laid. Verrill, cited by Osborn, states that +on one occasion he was able to collect fully a teaspoonful of eggs from +the dress of a lady in whose lap a half-grown kitten had been held for a +short time. Patton and Cragg record seeing the inside of a hat in which +a kitten had spent the night, so covered with flea eggs that it looked +"as if it had been sprinkled with sugar from a sifter." It is no wonder +that houses in which pets live become overrun with the fleas. + +One of the first control measures, then, consists in keeping such +animals out of the house or in rigorously keeping them free from fleas. +The latter can best be accomplished by the use of strong tar soap or +Armour's "Flesope," which may be obtained from most druggists. The use +of a three per cent solution of creolin, approximately four teaspoonfuls +to a quart of warm water, has also been recommended. While this is +satisfactory in the case of dogs, it is liable to sicken cats, who will +lick their fur in an effort to dry themselves. Howard recommends +thoroughly rubbing into the fur a quantity of pyrethrum powder. This +partially stupifies the fleas which should be promptly swept up and +burned. + +He also recommends providing a rug for the dog or cat to sleep on and +giving this rug a frequent shaking and brushing, afterwards sweeping up +and burning the dust thus removed. + +Since the larvæ of fleas are very susceptible to exposure, the use of +bare floors, with few rugs, instead of carpets or matting, is to be +recommended. Thorough sweeping, so as to allow no accumulation of dust +in cracks and crevices will prove efficient. If a house is once +infested it may be necessary to thoroughly scrub the floors with hot +soapsuds, or to spray them with gasoline. If the latter method is +adopted, care must be taken to avoid the possibility of fire. + +To clear a house of fleas Skinner recommends the use of flake +naphthalene. In a badly infested house he took one room at a time, +scattering on the floor five pounds of flake naphthalene, and closed it +for twenty-four hours. It proved to be a perfect and effectual remedy +and very inexpensive, as the naphthalene could be swept up and +transferred to other rooms. Dr. Skinner adds, "so far as I am concerned, +the flea question is solved and if I have further trouble I know the +remedy. I intend to keep the dog and cat." + +The late Professor Slingerland very effectively used hydrocyanic acid +gas fumigation in exterminating fleas in houses. In one case, where +failure was reported, he found on investigation that the house had +become thoroughly reinfested from pet cats, which had been left +untreated. Fumigation with sulphur is likewise efficient. + +The fact that adult fleas are usually to be found on the floor, when not +on their hosts, was ingeniously taken advantage of by Professor S. H. +Gage in ridding an animal room at Cornell University of the pests. He +swathed the legs of a janitor with sticky fly-paper and had him walk +back and forth in the room. Large numbers of the fleas were collected in +this manner. + +In some parts of the southern United States hogs are commonly infested +and in turn infest sheds, barns and even houses. Mr. H. E. Vick informs +us that it is a common practice to turn sheep into barn-lots and sheds +in the spring of the year to collect in their wool, the fleas which +abound in these places after the hogs have been turned out. + +It is a common belief that adult fleas are attracted to fresh meat and +that advantage of this can be taken in trapping them. Various workers, +notably Mitzman (1910), have shown that there is no basis for such a +belief. + + +THE TRUE CHIGGERS--The chigoes, or true chiggers, are the most +completely parasitic of any of the fleas. Of the dozen or more known +species, one commonly attacks man. This is _Dermatophilus penetrans_, +more commonly known as _Sarcopsylla penetrans_ or _Pulex penetrans_. + +This species occurs in Mexico, the West Indies, Central and South +America. There are no authentic records of its occurrence in the United +States although, as Baker has pointed out, there is no reason why it +should not become established in Florida and Texas. It is usually +believed that Brazil was its original home. Sometime about the middle of +the nineteenth century it was introduced into West Africa and has spread +across that continent. + +The males and the immature females of _Dermatophilus penetrans_ (fig. +93) closely resemble those of other fleas. They are very active little +brown insects about 1-1.2 mm. in size, which live in the dust of native +huts and stables, and in dry, sandy soil. In such places they often +occur in enormous numbers and become a veritable plague. + +[Illustration: 93. Dermatophilus penetrans. Much enlarged. After +Karsten.] + +They attack not only man but various animals. According to Castellani +and Chalmers, "Perhaps the most noted feature is the way in which it +attacks pigs. On the Gold Coast it appeared to be largely kept in +existence by these animals. It is very easily captured in the free state +by taking a little pig with a pale abdomen, and placing it on its back +on the ground on which infected pigs are living. After watching a few +moments, a black speck will appear on the pig's abdomen, and quickly +another and another. These black specks are jiggers which can easily be +transferred to a test tube. On examination they will be found to be +males and females in about equal numbers." + +Both the males and females suck blood. That which characterizes this +species as distinguished from other fleas attacking man is that when the +impregnated female attacks she burrows into the skin and there swells +until in a few days she has the size and appearance of a small pea (fig. +94). Where they are abundant, hundreds of the pests may attack a single +individual (fig. 95). Here they lie with the apex of the abdomen +blocking the opening. According to Fülleborn (1908) they do not +penetrate beneath the epidermis. The eggs are not laid in the flesh of +the victim, as is sometimes stated, but are expelled through this +opening. The female then dies, withers and falls away or is expelled by +ulceration. According to Brumpt, she first quits the skin and then, +falling to the ground, deposits her eggs. The subsequent development in +so far as known, is like that of other fleas. + +[Illustration: 94. Dermatophilus penetrans, gravid female. After +Moniez.] + +[Illustration: 95. Chiggers in the sole of foot of man. Manson's +Tropical Diseases. Permission of Cassell and Co.] + +The chigoe usually enters between the toes, the skin about the roots of +the nails, or the soles of the feet, although it may attack other parts +of the body. Mense records the occurrence in folds of the epidermis, as +in the neighborhood of the anus. They give rise to irritation and unless +promptly and aseptically removed there often occurs pus formation and +the development of a more or less serious abscess. Gangrene and even +tetanus may ensue. + +Treatment consists in the careful removal of the insect, an operation +more easily accomplished a day or two after its entrance, than at first, +when it is unswollen. The ulcerated point should then be treated with +weak carbolic acid, or tincture of iodine, or dusted thoroughly with an +antiseptic powder. + +[Illustration: 96. Echidnophaga gallinacea.] + +[Illustration: 97. Echidnophaga gallinacea infesting head of chicken. +After Enderlein.] + +Castellani and Chalmers recommend as prophylactic measures, keeping the +house clean and keeping pigs, poultry, and cattle away therefrom. "High +boots should be used, and especial care should be taken not to go to a +ground floor bathroom with bare feet. The feet, especially the toes, and +under the nails, should be carefully examined every morning to see if +any black dots can be discovered, when the jigger should be at once +removed, and in this way suppuration will be prevented. It is +advisable, also, to sprinkle the floors with carbolic lotion, Jeyes' +fluid, or with pyrethrum powder, or with a strong infusion of native +tobacco, as recommended by Law and Castellani." + +_Echidnophaga gallinacea_ (fig. 96) is a widely distributed Hectopsyllid +attacking poultry (fig. 97). It occurs in the Southern and Southwestern +United States and has been occasionally reported as attacking man, +especially children. It is less highly specialized than _Dermatophilus +penetrans_, and does not ordinarily cause serious trouble in man. + + + + +CHAPTER IV + +ACCIDENTAL OR FACULTATIVE PARASITES + + +In addition to the many species of Arthropods which are normally +parasitic on man and animals, there is a considerable number of those +which may be classed as _accidental_ or _facultative_ parasites. + +Accidental or facultative parasites are species which are normally +free-living, but which are able to exist as parasites when accidentally +introduced into the body of man or other animal. A wide range of forms +is included under this grouping. + + +ACARINA + +A considerable number of mites have been reported as accidental or even +normal, endoparasites of man, but the authentic cases are comparatively +few. + +In considering such reports it is well to keep in mind von Siebold's +warning that in view of the universal distribution of mites one should +be on his guard. In vessels in which animal and other organic fluids and +moist substances gradually dry out, mites are very abundantly found. If +such vessels are used without very careful preliminary cleaning, for the +reception of evacuations of the sick, or for the reception of parts +removed from the body, such things may be readily contaminated by mites, +which have no other relation whatever to them. + +Nevertheless, there is no doubt but that certain mites, normally +free-living, have occurred as accidental parasites of man. Of these the +most commonly met with is _Tyroglyphus siro_, the cheese-mite. + +_Tyroglyphus siro_ is a small mite of a whitish color. The male measures +about 500µ long by 250µ wide, the female slightly larger. They live in +cheese of almost any kind, especially such as is a little decayed. "The +individuals gather together in winter in groups or heaps in the hollows +and chinks of the cheese and there remain motionless. As soon as the +temperature rises a little, they gnaw away at the cheese and reduce it +to a powder. The powder is composed of excrement having the appearance +of little grayish microscopic balls; eggs, old and new, cracked and +empty; larvæ, nymphs, and perfect mites, cast skins and fragments of +cheese, to which must be added numerous spores of microscopic +fungi."--Murray. + +_Tyroglyphus siro_, and related species, have been found many times in +human feces, under conditions which preclude the explanation that the +contamination occurred outside of the body. They have been supposed to +be the cause of dysentery, or diarrhoea, and it is probable that the +_Acarus dysenteriæ_ of Linnæus, and Latreille, was this species. +However, there is little evidence that the mites cause any noteworthy +symptoms, even when taken into the body in large numbers. + +_Histiogaster spermaticus_ (fig. 152) is a Tyroglyphid mite which was +reported by Trouessart (1902) as having been found in a cyst in the +groin, adherent to the testis. When the cyst was punctured, it yielded +about two ounces of opalescent fluid containing spermatozoa and numerous +mites in all stages of development. The evidence indicated that a +fecundated female mite had been introduced into the urethra by means of +an unclean catheter. Though Trouessart reported the case as that of a +Sarcoptid, Banks places the genus _Histiogaster_ with the Tyroglyphidæ. +He states that our species feeds on the oyster-shell bark louse, +possibly only after the latter is dead, and that in England a species +feeds within decaying reeds. + +_Nephrophages sanguinarius_ is a peculiarly-shaped, angular mite which +was found by Miyake and Scriba (1893) for eight successive days in the +urine of a Japanese suffering from fibrinuria. Males, .117 mm. long by +.079 mm. wide, females .36 mm. by. 12 mm., and eggs were found both in +the spontaneously emitted urine and in that drawn by means of a +catheter. All the mites found were dead. The describers regarded this +mite as a true endoparasite, but it is more probable that it should be +classed as an accidental parasite. + + +MYRIAPODA + +There are on record a number of cases of myriapods occurring as +accidental parasites of man. The subject has been treated in detail by +Blanchard (1898 and 1902), who discussed forty cases. Since then at +least eight additions have been made to the list. + +Neveau-Lamaire (1908) lists thirteen species implicated, representing +eight different genera. Of the _Chilognatha_ there are three, _Julus +terrestris_, _J. londinensis_ and _Polydesmus complanatus_. The +remainder are _Chilopoda_, namely, _Lithobius forficatus_, _L. +malenops_, _Geophilus carpophagus_, _G. electricus_, _G. similis_, _G. +cephalicus_, _Scutigera coleoptrata_, _Himantarium gervaisi_, +_Chætechelyne vesuviana_ and _Stigmatogaster subterraneus_. + +The majority of the cases relate to infestation of the nasal fossæ, or +the frontal sinus, but intestinal infestation also occurs and there is +one recorded case of the presence of a species in _Julus_ (fig. 13) in +the auditory canal of a child. + +In the nose, the myriapods have been known to live for months and +according to some records, even for years. The symptoms caused by their +presence are inflammation, with or without increased flow of mucus, +itching, more or less intense headache, and at times general symptoms +such as vertigo, delirium, convulsions, and the like. These symptoms +disappear suddenly when the parasites are expelled. + +In the intestine of man, myriapods give rise to obscure symptoms +suggestive of infestation by parasitic worms. In a case reported by +Verdun and Bruyant (1912), a child twenty months of age had been +affected for fifteen days by digestive disturbances characterized by +loss of appetite, nausea and vomiting. The latter had been particularly +pronounced for three days, when there was discovered in the midst of the +material expelled a living myriapod of the species _Chætechelyne +vesuviana_. Anthelminthics had been administered without result. In some +of the other cases, the administration of such drugs had resulted in the +expulsion of the parasite through the anus. + +One of the extreme cases on record is that reported by Shipley (1914). +Specimens of _Geophilus gorizensis_ (= _G. subterraneus_) "were vomited +and passed by a woman of 68 years of age. Some of the centipedes emerged +through the patient's nose, and it must be mentioned that she was also +suffering from a round worm. One of her doctors was of the opinion that +the centipedes were certainly breeding inside the lady's intestines, and +as many as seven or eight, sometimes more, were daily leaving the +alimentary canal." + +"According to her attendant's statements those centipedes had left the +body in some hundreds during a period of twelve or eighteen months. +Their presence produced vomiting and some hæmatemesis, and treatment +with thymol, male-fern and turpentine had no effect in removing the +creatures." + +The clinical details, as supplied by Dr. Theodore Thompson were as +follows: + +"Examined by me July, 1912, her tongue was dry and glazed. There was +bleeding taking place from the nose and I saw a living centipede she had +just extracted from her nostril. Her heart, lungs and abdomen appeared +normal. She was not very wasted, and did not think she had lost much +flesh, nor was there any marked degree of anemia." + +Shipley gives the following reasons for believing it impossible that +these centipedes could have multiplied in the patient's intestine. "The +breeding habits of the genus _Geophilus_ are peculiar, and ill adapted +for reproducing in such a habitat. The male builds a small web or nest, +in which he places his sperm, and the female fertilizes herself from +this nest or web, and when the eggs are fertilized they are again laid +in a nest or web in which they incubate and in two or three weeks hatch +out. The young _Geophilus_ differ but very little from the adult, except +in size. It is just possible, but improbable, that a clutch of eggs had +been swallowed by the host when eating some vegetables or fruit, but +against this is the fact that the _Geophilus_ does not lay its eggs upon +vegetables or fruit, but upon dry wood or earth. The egg-shell is very +tough and if the eggs had been swallowed the egg-shells could certainly +have been detected if the dejecta were examined. The specimens of the +centipede showed very little signs of being digested, and it is almost +impossible to reconcile the story of the patient with what one knows of +the habits of the centipedes." + +In none of the observed cases have there been any clear indications as +to the manner of infestation. It is possible that the myriapods have +been taken up in uncooked fruit or vegetables. + + +LEPIDOPTEROUS LARVÆ + +SCHOLECIASIS--Hope (1837) brought together six records of infestation of +man by lepidopterous larvæ and proposed to apply the name scholeciasis +to this type of parasitism. The clearest case was that of a young boy +who had repeatedly eaten raw cabbage and who vomited larvæ of the +cabbage butterfly, _Pieris brassicæ_. Such cases are extremely rare, and +there are few reliable data relative to the subject. In this connection +it may be noted that Spuler (1906) has described a moth whose larvæ live +as ectoparasites of the sloth. + + +COLEOPTERA + +CANTHARIASIS--By this term Hope designated instances of accidental +parasitism by the larvæ or adults of beetles. Reports of such cases are +usually scouted by parasitologists but there seems no good basis for +wholly rejecting them. Cobbold refers to a half dozen cases of +accidental parasitism by the larvæ of _Blaps mortisaga_. In one of +these cases upwards of 1200 larvæ and several perfect insects were said +to have been passed _per annum_. French (1905) reports the case of a man +who for a considerable period voided adult living beetles of the species +_Nitidula bipustulata_. Most of the other cases on record relate to the +larvæ of _Dermestidæ_ (larder beetles _et al._) or _Tenebrionidæ_ (meal +infesting species). Infestation probably occurs through eating raw or +imperfectly cooked foods containing eggs or minute larvæ of these +insects. + +[Illustration: 98. Larva of Piophila casei. Caudal aspect of larva. +Posterior stigmata.] + +Brumpt cites a curious case of accidental parasitism by a coleopterous +larva belonging to the genus _Necrobia_. This larva was extracted from a +small tumor, several millimeters long, on the surface of the conjunctiva +of the eye. The larvæ of this genus ordinarily live in decomposing flesh +and cadavers. + + +DIPTEROUS LARVÆ + +[Illustration: 99. Piophila casei. After Graham-Smith.] + +MYASIS--By this term (spelled also myiasis, and myiosis), is meant +parasitism by dipterous larvæ. Such parasitism may be normal, as in the +cases already described under the heading parasitic Diptera, or it may +be facultative, due to free-living larvæ being accidentally introduced +into wounds or the body-cavities of man. Of this latter type, there is a +multitude of cases on record, relating to comparatively few species. The +literature of the subject, like that relating to facultative parasitism +in general, is unsatisfactory, for most of the determinations of species +have been very loose. Indeed, so little has been known regarding the +characteristics of the larvæ concerned that in many instances they could +not be exactly determined. Fortunately, several workers have undertaken +comparative studies along this line. The most comprehensive publication +is that of Banks (1912), entitled "The structure of certain dipterous +larvæ, with particular reference to those in human food." + +Without attempting an exhaustive list, we shall discuss here the more +important species of Diptera whose larvæ are known to cause myasis, +either external or internal. The following key will serve to determine +those most likely to be encountered. The writers would be glad to +examine specimens not readily identifiable, if accompanied by exact data +relative to occurrence. + +_a._ Body more or less flattened, depressed; broadest in the middle, + each segment with dorsal, lateral, and ventral fleshy processes, of + which the laterals, at least, are more or less spiniferous (fig. + 101). _Fannia_ (= _Homalomyia_). + + In _F. canicularis_ the dorsal processes are nearly as long as the + laterals; in _F. scalaris_ the dorsal processes are short spinose + tubercles. + +_aa._ Body cylindrical, or slender conical tapering toward the head; + without fleshy lateral processes (fig. 105). + +_b._ With the posterior stigmata at the end of shorter or longer + tubercles, or if not placed upon tubercles, then not in pit; usually + without a "marginal button" and without a chitinous ring surrounding + the three slits; the slits narrowly or broadly oval, not bent (fig. + 171 i). _Acalyptrate muscidæ_ and some species of _Anthomyiidæ_. To + this group belong the cheese skipper (_Piophila casei_, figs. 98, + 99), the pomace-fly (_Drosophila ampelophila_), the apple maggot + (_Rhagoletis pomonella_), the cherry fruit fly (_Rhagoletis + cingulata_), the small dung fly (_Sepsis violacea_, fig. 170), the + beet leaf-miner (_Pegomyia vicina_, fig. 171 i), the cabbage, bean + and onion maggots (_Phorbia_ spp.) et. al. + +_bb._ Posterior stigmata of various forms, if the slits are narrowly + oval (fig. 171) then they are surrounded by a chitin ring which may + be open ventro-mesally. + +_c._ Integument leathery and usually strongly spinulose; larvæ + hypodermatic or endoparasitic. Bot flies (fig. 171, f, g, + k).--_Oestridæ_ + +_cc._ Integument not leathery and (except in _Protocalliphora_) spinulæ + restricted to transverse patches near the incisures of the segments. + +_d._ The stigmal plates in a pit; the lip-like margin of the pit with a + number of fleshy tubercles; chitin ring of the stigma not complete; + open ventro-mesally, button absent (fig. 171 e). Flesh + flies.--_Sarcophaga_ + +_dd._ Stigmata not in a pit. + +_e._ The chitin ring open ventra-mesally; button absent (fig. 171 c). + Screw-worm fly. _Chrysomyia_ + +_ee._ The chitin ring closed. + +_f._ Slits of the posterior stigmata straight; marginal "button" present + (fig. 171 b); two distinct mouth hooks, fleshy tubercles around the + anal area. _Phormia_ (fig. 171 f), _Lucilia_ and _Calliphora_ (fig. + 172, a, b), _Protocalliphora_ (fig. 171, j), _Cynomyia_ (fig. 171, + a). Blow flies, bluebottle flies. _Calliphorinæ_ + +_ff._ Slits of the posterior stigmata sinuous or bent. Subfamily + Muscinæ. + +_g._ Slits of the posterior stigmata bent; usually two mouth hooks. + _Muscina stabulans_ (fig. 171, l), _Muscina similis_, _Myiospila + meditabunda_ (fig. 172, i), and some of the higher _Anthomyiidæ_. + +_gg._ Slits of the posterior stigmata sinuous; mouth hooks usually + consolidated into one. The house-fly (_Musca domestica_ fig. 171, + d), the stable fly (_Stomoxys calcitrans_), the horn fly (_Lyperosia + irritans_), _Pyrellia_, _Pseudopyrellia_, _Morellia_, _Mesembrina_. + _Polietes_, et. al. (fig. 172 in part). + +_Eristalis_--The larvæ of _Eristalis_ are the so-called rat-tailed +maggots, which develop in foul water. In a few instances these larvæ +have been known to pass through the human alimentary canal uninjured. +Hall and Muir (1913) report the case of a boy five years of age, who had +been ailing for ten weeks and who was under treatment for indigestion +and chronic constipation. For some time he had vomited everything he +ate. On administration of a vermifuge he voided one of the rat-tailed +maggots of _Eristalis_. He admitted having drunk water from a ditch full +of all manner of rotting matter. It was doubtless through this that he +became infested. It is worth noting that the above described symptoms +may have been due to other organisms or substances in the filthy water. + +_Piophila casei_, the cheese-fly (fig. 99), deposits its eggs not only +in old cheeses, but on ham, bacon, and other fats. The larvæ (fig. 98) +are the well-known cheese skippers, which sometimes occur in great +abundance on certain kinds of cheese. Indeed, some people have a +comfortable theory that such infested cheese is especially good. Such +being the case, it is small wonder that this species has been repeatedly +reported as causing intestinal myasis. Thebault (1901) describes the +case of a girl who, shortly after consuming a large piece of badly +infested cheese, became ill and experienced severe pains in the region +of the navel. Later these extended through the entire alimentary canal, +the excrement was mixed with blood and she suffered from vertigo and +severe headaches. During the four following days the girl felt no +change, although the excretion of the blood gradually diminished and +stopped. On the fourth day she voided two half-digested larvæ and, +later, seven or eight, of which two were alive and moving. + +That these symptoms may be directly attributed to the larvæ, or +"skippers," has been abundantly shown by experimental evidence. +Portschinsky cites the case of a dog fed on cheese containing the larvæ. +The animal suffered much pain and its excrement contained blood. On +_post mortem_ it was found that the small intestine throughout almost +its entire length was marked by bloody bruises. The papillæ on these +places were destroyed, although the walls were not entirely perforated. +In the appendix were found two or three dead larvæ. Alessandri (1910) +has likewise shown that the larvæ cause intestinal lesions. + +According to Graham-Smith, Austen (1912) has recorded a case of myasis +of the nose, attended with a profuse watery discharge of several weeks +duration and pain, due to the larvæ of _Piophila casei_. + + +ANTHYOMYIIDÆ--The characteristic larvæ of two species of _Fannia_ (= +_Homalomyia_ or _Anthomyia_, in part) (fig. 101) are the most commonly +reported of dipterous larvæ causing intestinal myasis. Hewitt (1912) has +presented a valuable study of the bionomics and of the larvæ of these +flies, a type of what is needed for all the species concerned in myasis. +We have seen two cases of their having been passed in stools, without +having caused any special symptoms. In other instances their presence in +the alimentary canal has given rise to symptoms vaguely described as +those of tapeworm infestation, or helminthiasis. More specifically, they +have been described as causing vertigo, severe headache, nausea and +vomiting, severe abdominal pains, and in some instances, bloody +diarrhoea. + +[Illustration: 100. Fannia canicularis (×4). After Graham-Smith.] + +One of the most striking cases is that reported by Blankmeyer (1914), of +a woman whose illness began fourteen years previously with nausea and +vomiting. After several months of illness she began passing larvæ and +was compelled to resort to enemas. Three years previous to the report, +she noticed frequent shooting pains in the rectal region and at times +abdominal tenderness was marked. There was much mucus in the stools and +she "experienced the sensation of larvæ crawling in the intestine." +Occipital headaches were marked, with remissions, and constipation +became chronic. The appetite was variable, there was a bad taste in the +mouth, tongue furred and ridged, and red at the edges. Her complexion +was sallow, and general nervousness was marked. As treatment, there were +given doses of magnesium sulphate before breakfast and at 4 P. M., with +five grain doses of salol four times a day. The customary parasiticides +yielded no marked benefit. At the time of the report the patient passed +from four to fifty larvæ per day, and was showing some signs of +improvement. The nausea had disappeared, her nervousness was less +evident, and there was a slight gain in weight. + +The case was complicated by various other disorders, but the symptoms +given above seem to be in large part attributable to the myasis. There +is nothing in the case to justify the assumption that larvæ were +continuously present, for years. It seems more reasonable to suppose +that something in the habits of the patient favored repeated +infestation. Nevertheless, a study of the various cases of intestinal +myasis caused by these and other species of dipterous larvæ seems to +indicate that the normal life cycle may be considerably prolonged under +the unusual conditions. + +The best authenticated cases of myasis of the urinary passage have been +due to larvæ of _Fannia_. Chevril (1909) collected and described twenty +cases, of which seven seemed beyond doubt. One of these was that of a +woman of fifty-five who suffered from albuminuria, and urinated with +much difficulty, and finally passed thirty to forty larvæ of _Fannia +canicularis_. + +It is probable that infestation usually occurs through eating partially +decayed fruit or vegetables on which the flies have deposited their +eggs. Wellman points out that the flies may deposit their eggs in or +about the anus of persons using outside privies and Hewitt believes that +this latter method of infection is probably the common one in the case +of infants belonging to careless mothers. "Such infants are sometimes +left about in an exposed and not very clean condition, in consequence of +which flies are readily attracted to them and deposit their eggs." + +[Illustration: 101. Larva of Fannia scalaris.] + + +MUSCINÆ--The larvæ of the common house-fly, _Musca domestica_, are +occasionally recorded as having been passed with the feces or vomit of +man. While such cases may occur, it is probable that in most instances +similar appearing larvæ of other insects have been mistakenly +identified. + +_Muscina stabulans_ is regarded by Portschinsky (1913) as responsible +for many of the cases of intestinal myasis attributed to other species. +He records the case of a peasant who suffered from pains in the lower +part of the breast and intestines, and whose stools were mixed with +blood. From November until March he had felt particularly ill, being +troubled with nausea and vomiting in addition to the pain in his +intestines. In March, his physician prescribed injections of a +concentrated solution of tannin, which resulted in the expulsion of +fifty living larvæ of _Muscina stabulans_. Thereafter the patient felt +much better, although he suffered from intestinal catarrh in a less +severe form. + +[Illustration: 102 Muscina stabulans (×4). After Graham-Smith.] + + +CALLIPHORINÆ--Closely related to the Sarcophagidæ are the +_Calliphorinæ_, to which group belong many of the so-called "blue +bottle" flies. Their larvæ feed upon dead animals, and upon fresh and +cooked meat. Those of _Protocalliphora_, already mentioned, are +ectoparasitic on living nestling birds. Larva of _Lucilia_, we have +taken from tumors on living turtles. To this sub-family belongs also +_Aucheromyia luteola_, the Congo floor maggot. Some of these, and at +least the last mentioned, are confirmed, rather than faculative +parasites. Various species of Calliphorinæ are occasionally met with as +facultative parasites of man. + +[Illustration: 103. Lucilia cæsar, (×3). After Howard.] + +_Chrysomyia macellaria_, the screw worm fly (fig. 107), is the fly which +is responsible for the most serious cases of human myasis in the United +States. It is widely distributed in the United States but is especially +abundant in the south. While the larvæ breed in decaying matter in +general, they so commonly breed in the living flesh of animals that they +merit rank as true parasites. The females are attracted to open wounds +of all kinds on cattle and other animals and quickly deposit large +numbers of eggs. Animals which have been recently castrated, dehorned, +or branded, injured by barbed wire, or even by the attacks of ticks are +promptly attacked in the regions where the fly abounds. Even the navel +of young calves or discharges from the vulva of cows may attract the +insect. + +[Illustration: 104. Calliphora erythrocephala, (×6). After +Graham-Smith.] + +Not infrequently the fly attacks man, being attracted by an offensive +breath, a chronic catarrh, or a purulent discharge from the ears. Most +common are the cases where the eggs are deposited in the nostrils. The +larvæ, which are hatched in a day or two, are provided with strong +spines and proceed to bore into the tissues of the nose, even down into +or through the bone, into the frontal sinus, the pharynx, larynx, and +neighboring parts. + +Osborn (1896) quotes a number of detailed accounts of the attacks of the +_Chrysomyia_ on man. A vivid picture of the symptomology of rhinal +myasis caused by the larvæ of this fly is given by Castellani and +Chalmers: "Some couple of days after a person suffering from a chronic +catarrh, foul breath, or ozæna, has slept in the open or has been +attacked by a fly when riding or driving,--_i.e._, when the hands are +engaged--signs of severe catarrh appear, accompanied with inordinate +sneezing and severe pain over the root of the nose or the frontal bone. +Quickly the nose becomes swollen, and later the face also may swell, +while examination of the nose may show the presence of the larvæ. Left +untreated, the patient rapidly becomes worse, and pus and blood are +discharged from the nose, from which an offensive odor issues. Cough +appears as well as fever, and often some delirium. If the patient lives +long enough, the septum of the nose may fall in, the soft and hard +palates may be pierced, the wall of the pharynx may be destroyed. By +this time, however, the course of the disease will have become quite +evident by the larvæ dropping out of the nose, and if the patient +continues to live all the larvæ may come away naturally." + +For treatment of rhinal myasis these writers recommend douching the nose +with chloroform water or a solution of chloroform in sweet milk (10-20 +per cent), followed by douches of mild antiseptics. Surgical treatment +may be necessary. + +[Illustration: 105. Larva of a flesh fly (Sarcophaga). Caudal aspect. +Anterior stigmata. Pharyngeal skeleton.] + + +SARCOPHAGIDÆ--The larvæ (fig. 105) of flies of this family usually feed +upon meats, but have been found in cheese, oleomargerine, pickled +herring, dead and living insects, cow dung and human feces. Certain +species are parasitic in insects. Higgins (1890) reported an instance of +"hundreds" of larvæ of _Sarcophaga_ being vomited by a child eighteen +months of age. There was no doubt as to their origin for they were +voided while the physician was in the room. There are many other reports +of their occurrence in the alimentary canal. We have recorded elsewhere +(Riley, 1906) a case in which some ten or twelve larvæ of _Sarcophaga_ +were found feeding on the diseased tissues of a malignant tumor. The +tumor, a melanotic sarcoma, was about the size of a small walnut, and +located in the small of the back of an elderly lady. Although they had +irritated and caused a slight hæmorrhage, neither the patient nor others +of the family knew of their presence. Any discomfort which they had +caused had been attributed to the sarcomatous growth. The infestation +occurred in mid-summer. It is probable that the adult was attracted by +the odor of the discharges and deposited the living maggots upon the +diseased tissues. + +[Illustration: 106. A flesh fly (Sarcophaga), (×4). After Graham-Smith.] + +According to Küchenmeister, _Sarcophaga carnaria_ (fig. 106), attracted +by the odor, deposits its eggs and larvæ in the vagina of girls and +women when they lie naked in hot summer days upon dirty clothes, or when +they have a discharge from the vagina. In malignant inflammations of the +eyes the larvæ even nestle under the eyelids and in Egypt, for example, +produce a very serious addition to the effects of small-pox upon the +cornea, as according to Pruner, in such cases perforation of the cornea +usually takes place. + +[Illustration: 107. Chrysomyia macellaria, (×3).] + +_Wohlfartia magnifica_ is another Sarcophagid which commonly infests man +in the regions where it is abundant. It is found in all Europe but is +especially common in Russia, where Portschinsky has devoted much +attention to its ravages. It deposits living larvæ in wounds, the nasal +fossæ, the ears and the eyes, causing injuries even more revolting than +those described for _Chrysomyia_. + + + + +CHAPTER V + +ARTHROPODS AS SIMPLE CARRIERS OF DISEASE + + +The fact that certain arthropods are poisonous, or may affect the health +of man as direct parasites has always received attention in the medical +literature. We come now to the more modern aspect of our subject,--the +consideration of insects and other arthropods as transmitters and +disseminators of disease. + +The simplest way in which arthropods may function in this capacity is as +_simple carriers_ of pathogenic organisms. It is conceivable that any +insect which has access to, and comes in contact with such organisms and +then passes to the food, or drink, or to the body of man, may in a +wholly accidental and incidental manner convey infection. That this +occurs is abundantly proved by the work of recent years. We shall +consider as typical the case against the house-fly, which has attracted +so much attention, both popular and scientific. The excellent general +treatises of Hewitt (1910), Howard (1911), and Graham-Smith (1913), and +the flood of bulletins and popular literature render it unnecessary to +consider the topic in any great detail. + + +THE HOUSE-FLY AS A CARRIER OF DISEASE + +Up to the past decade the house-fly has usually been regarded as a mere +pest. Repeatedly, however, it had been suggested that it might +disseminate disease. We have seen that as far back as the sixteenth +century, Mercurialis suggested that it was the agent in the spread of +bubonic plague, and in 1658, Kircher reiterated this view. In 1871, +Leidy expressed the opinion that flies were probably a means of +communicating contagious diseases to a greater degree than was generally +suspected. From what he had observed regarding gangrene in hospitals, he +thought flies should be carefully excluded from wounds. In the same +year, the editor of the _London Lancet_, referring to the belief that +they play a useful rôle in purifying the air said, "Far from looking +upon them as dipterous angels dancing attendance on Hygeia, regard them +rather in the light of winged sponges spreading hither and thither to +carry out the foul behests of Contagion." + +These suggestions attracted little attention from medical men, for it is +only within very recent years that the charges have been supported by +direct evidence. Before considering this evidence, it is necessary that +we define what is meant by "house-fly" and that we then consider the +life-history of the insect. + +There are many flies which are occasionally to be found in houses, but +according to various counts, from 95 per cent to 99 per cent of these in +warm weather in the Eastern United States belong to the one species +_Musca domestica_ (fig. 108). This is the dominant house-fly the world +over and is the one which merits the name. It has been well +characterized by Schiner (1864), whose description has been freely +translated by Hewitt, as follows: + +"Frons of male occupying a fourth part of the breadth of the head. +Frontal stripe of female narrow in front, so broad behind that it +entirely fills up the width of the frons. The dorsal region of the +thorax dusty grey in color with four equally broad longitudinal stripes. +Scutellum gray with black sides. The light regions of the abdomen +yellowish, transparent, the darkest parts at least at the base of the +ventral side yellow. The last segment and a dorsal line blackish brown. +Seen from behind and against the light, the whole abdomen shimmering +yellow, and only on each side of the dorsal line on each segment a dull +transverse band. The lower part of the face silky yellow, shot with +blackish brown. Median stripe velvety black. Antennæ brown. Palpi black. +Legs blackish brown. Wings tinged with pale gray with yellowish base. +The female has a broad velvety back, often reddishly shimmering frontal +stripe, which is not broader at the anterior end than at the bases of +the antennæ, but become so very much broader above that the light +dustiness of the sides is entirely obliterated. The abdomen gradually +becoming darker. The shimmering areas on the separate segments generally +brownish. All the other parts are the same as in the male." + +The other species of flies found in houses in the Eastern United States +which are frequently mistaken for the house or typhoid fly may readily +be distinguished by the characters of the following key: + +_a._ Apical cell (R_5) of the wide wing open, i.e., the bounding veins + parallel or divergent (fig. 100). Their larvæ are flattened, the + intermediate body segments each fringed with fleshy, more or less + spinose, processes. _Fannia_ + + b. Male with the sides of the second and third abdominal segments + translucent yellowish. The larva with three pairs of nearly equal + spiniferous appendages on each segment, arranged in a + longitudinal series and in addition two pairs of series of smaller + processes (fig. 100) _F. canicularis_ + + bb. Male with blackish abdomen, middle tibia with a tubercle beyond + the middle. The larva with spiniferous appendages of which the + dorsal and ventral series are short, the lateral series long and + feathered (fig. 101) _F. scalaris_ + +aa. Apical cell (R) of the wing more or less narrowed in the margin; i. + e., the bounding veins more or less converging (fig. 108). + + b. The mouth-parts produced and pointed, fitted for piercing. + + c. Palpi much shorter than the proboscis; a brownish gray fly, its + thorax with three rather broad whitish stripes; on each border + of the middle stripe and on the mesal borders of the lateral + stripes is a blackish brown line. Abdomen yellowish brown; on + the second, third and fourth segments are three brown spots + which may be faint or even absent. The larvæ live in dung. The + stable-fly (fig. 110) _Stomoxys calcitrans_ + + cc. Palpi nearly as long as the proboscis. Smaller species than the + house-fly. The horn-fly (fig. 167) _Hæmatobia irritans_ + + bb. Mouth-parts blunt, fitted for lapping. + + c. Thorax, particularly on the sides and near the base of the wings + with soft golden yellow hairs among the bristles. This fly is + often found in the house in very early spring or even in the + winter. The cluster-fly, _Pollenia rudis_ + + cc. Thorax without golden yellow hairs among the bristles. + + d. The last segment of the vein M with an abrupt angle. (fig. + 108). The larvæ live in manure, etc. House-fly, _Musca + domestica_ + + dd. The last segment of vein M with a broad, gentle curve (fig. + 102). + + e. Eyes microscopically hairy; each abdominal segment with two + spots. Larvæ in dung. _Myiospila meditabunda_ + + ee. Eyes bare; abdomen gray and brown marbled. _Muscina_ + + f. With black legs and palpi. _M. assimilis_ + + ff. With legs more or less yellowish; palpi yellow. Larvæ in + decaying vegetable substances, dung, etc. _M. stabulans_ + +It is almost universally believed that the adults of _Musca domestica_ +hibernate, remaining dormant throughout the winter in attics, around +chimneys, and in sheltered but cold situations. This belief has been +challenged by Skinner (1913), who maintains that all the adult flies die +off during the fall and early winter and that the species is carried +over in the pupal stage, and in no other way. The cluster-fly, _Pollenia +rudis_, undoubtedly does hibernate in attics and similar situations and +is often mistaken for the house-fly. In so far as concerns _Musca +domestica_, the important question as to hibernation in the adult stage +is an open one. Many observations by one of the writers (Johannsen) tend +to confirm Dr. Skinner's conclusion, in so far as it applies to +conditions in the latitude of New York State. Opposed, is the fact that +various experimenters, notably Hewitt (1910) and Jepson (1909) wholly +failed to carry pupæ through the winter. + +[Illustration: 108. The house or typhoid fly (Musca domestica (×4)). +After Howard.] + +The house-fly breeds by preference in horse manure. Indeed, Dr. Howard, +whose extensive studies of the species especially qualify him for +expressing an opinion on the subject, has estimated that under ordinary +city and town conditions, more than ninety per cent of the flies present +in houses have come from horse stables or their vicinity. They are not +limited to such localities, by any means, for it has been found that +they would develop in almost any fermenting organic substance. Thus, +they have been bred from pig, chicken, and cow manure, dirty waste +paper, decaying vegetation, decaying meat, slaughter-house refuse, +sawdust-sweepings, and many other sources. A fact which makes them +especially dangerous as disease-carriers is that they breed readily in +human excrement. + +The eggs are pure white, elongate ovoid, somewhat broader at the +anterior end. They measure about one millimeter (1-25 inch) in length. +They are deposited in small, irregular clusters, one hundred and twenty +to one hundred and fifty from a single fly. A female may deposit as many +as four batches in her life time. The eggs hatch in from eight to +twenty-four hours. + +The newly hatched larva, or maggot (fig. 108), measures about two +millimeters (1-12 inch) in length. It is pointed at the head end and +blunt at the opposite end, where the spiracular openings are borne. It +grows rapidly, molts three times and reaches maturity in from six to +seven days, under favorable conditions. + +The pupal stage, like that of related flies, is passed in the old larval +skin which, instead of being molted, becomes contracted and heavily +chitinized, forming the so-called _puparium_ (fig. 108). The pupal stage +may be completed in from three to six days. + +Thus during the warm summer months a generation of flies may be produced +in ten to twelve days. Hewitt at Manchester, England, found the minimum +to be eight days but states that larvæ bred in the open air in horse +manure which had an average daily temperature of 22.5° C., occupied +fourteen to twenty days in their development, according to the air +temperature. + +After emergence, a period of time must elapse before the fly is capable +of depositing eggs. This period has been tuned the _preoviposition_ +period. Unfortunately we have few exact data regarding this period. +Hewitt found that the flies became sexually mature in ten to fourteen +days after their emergence from the pupal state and four days after +copulation they began to deposit their eggs; in other words the +preoviposition stage was fourteen days or longer. Griffith (1908) found +this period to be ten days. Dr. Howard believes that the time "must +surely be shorter, and perhaps much shorter, under midsummer conditions, +and in the freedom of the open air." He emphasizes that the point is of +great practical importance, since it is during this period that the +trapping and other methods of destroying the adult flies, will prove +most useful. + +Howard estimates that there may be nine generations of flies a year +under outdoor conditions in places comparable in climate to Washington. +The number may be considerably increased in warmer climates. + +The rate at which flies may increase under favorable conditions is +astounding. Various writers have given estimates of the numbers of flies +which may develop as the progeny of a single individual, providing all +the eggs and all the individual flies survived. Thus, Howard estimates +that from a single female, depositing one hundred and twenty eggs on +April 15th, there may be by September 10th, 5,598,720,000,000 adults. +Fortunately, living forms do not produce in any such mathematical manner +and the chief value of the figures is to illustrate the enormous +struggle for existence which is constantly taking place in nature. + +Flies may travel for a considerable distance to reach food and shelter, +though normally they pass to dwellings and other sources of food supply +in the immediate neighborhood of their breeding places. Copeman, Howlett +and Merriman (1911) marked flies by shaking them in a bag containing +colored chalk. Such flies were repeatedly recovered at distances of +eight to one thousand yards and even at a distance of seventeen hundred +yards, nearly a mile. + +Hindle and Merriman (1914) continued these experiments on a large scale +at Cambridge, England. They "do not think it likely that, as a rule, +flies travel more than a quarter of a mile in thickly-housed areas." In +one case a single fly was recovered at a distance of 770 yards but a +part of this distance was across open fen-land. The surprising fact was +brought out that flies tend to travel either _against_ or across the +wind. The actual direction followed may be determined either directly by +the action of the wind (positive anemotropism), or indirectly owing to +the flies being attracted by any odor that it may convey from a source +of food. They conclude that it is likely that the chief conditions +favoring the disposal of flies are fine weather and a warm temperature. +The nature of the locality is another considerable factor. Hodge (1913) +has shown that when aided by the wind they may fly to much greater +distances over the water. He reports that at Cleveland, Ohio, the cribs +of the water works, situated a mile and a quarter, five miles, and six +miles out in Lake Erie are invaded by a regular plague of flies when the +wind blows from the city. Investigation showed that there was absolutely +nothing of any kind in which flies could breed on the crib. + +The omnivorous habits of the house-fly are matters of everyday +observation. From our view point, it is sufficient to emphasize that +from feeding on excrement, on sputum, on open sores, or on putrifying +matter, the flies may pass to the food or milk upon the table or to +healthy mucous membranes, or uncontaminated wounds. There is nothing in +its appearance to tell whether the fly that comes blithely to sup with +you is merely unclean, or whether it has just finished feeding upon +dejecta teeming with typhoid bacilli. + +[Illustration: 100. Pulvillus of foot of house-fly, showing glandular +hairs.] + +The method of feeding of the house-fly has an important bearing on the +question of its ability to transmit pathogenic organisms. Graham-Smith +(1910) has shown that when feeding, flies frequently moisten soluble +substances with "vomit" which is regurgitated from the crop. This is, of +course, loaded with bacteria from previous food. When not sucked up +again these drops of liquid dry, and produce round marks with an opaque +center and rim and an intervening less opaque area. Fly-specks, then, +consist of both vomit spots and feces. Graham-Smith shows a photograph +of a cupboard window where, on an area six inches square, there were +counted eleven hundred and two vomit marks and nine fecal deposits. + +From a bacteriologist's viewpoint a discussion of the possibility of a +fly's carrying bacteria would seem superfluous. Any exposed object, +animate or inanimate, is contaminated by bacteria and will transfer them +if brought into contact with suitable culture media, whether such +substance be food, or drink, open wounds, or the sterile culture media +of the laboratory. A needle point may convey enough germs to produce +disease. Much more readily may the house-fly with its covering of hairs +and its sponge-like pulvilli (fig. 109) pick up and transfer bits of +filth and other contaminated material. + +For popular instruction this inevitable transfer of germs by the +house-fly is strikingly demonstrated by the oft copied illustration of +the tracks of a fly on a sterile culture plate. Two plates of gelatine +or, better, agar medium are prepared. Over one of these a fly (with +wings clipped) is allowed to walk, the other is kept as a check. Both +are put aside at room temperature, to be examined after twenty-four to +forty-eight hours. At the end of that time, the check plate is as clear +as ever, the one which the fly has walked is dotted with colonies of +bacteria and fungi. The value in the experiment consists in emphasizing +that by this method we merely render visible what is constantly +occurring in nature. + +A comparable experiment which we use in our elementary laboratory work +is to take three samples of _clean_ (preferably, sterile) fresh milk in +sterile bottles. One of them is plugged with a pledget of cotton, into +the second is dropped a fly from the laboratory and into the third is +dropped a fly which has been caught feeding upon garbage or other filth. +After a minute or two the flies are removed and the vials plugged as was +number one. The three are then set aside at room temperature. When +examined after twenty-four hours the milk in the first vial is either +still sweet or has a "clean" sour odor; that of the remaining two is +very different, for it has a putrid odor, which is usually more +pronounced in the case of sample number three. + +Several workers have carried out experiments to determine the number of +bacteria carried by flies under natural conditions. One of the most +extended and best known of these is the series by Esten and Mason +(1908). These workers caught flies from various sources in a sterilized +net, placed them in a sterile bottle and poured over them a known +quantity of sterilized water, in which they were shaken so as to wash +the bacteria from their bodies. They found the number of bacteria on a +single fly to range from 550 to 6,600,000. Early in the fly season the +numbers of bacteria on flies are comparatively small, while later the +numbers are comparatively very large. The place where flies live also +determines largely the numbers that they carry. The lowest number, 550, +was from a fly caught in the bacteriological laboratory, the highest +number, 6,600,000 was the average from eighteen swill-barrel flies. +Torrey (1912) made examination of "wild" flies from a tenement house +district of New York City. He found "that the surface contamination of +these 'wild' flies may vary from 570 to 4,400,000 bacteria per insect, +and the intestinal bacterial content from 16,000 to 28,000,000." + +Less well known in this country is the work of Cox, Lewis, and Glynn +(1912). They examined over four hundred and fifty naturally infected +house-flies in Liverpool during September and early October. Instead of +washing the flies they were allowed to swim on the surface of sterile +water for five, fifteen, or thirty minutes, thus giving natural +conditions, where infection occurs from vomit and dejecta of the flies, +as well as from their bodies. They found, as might be expected, that +flies from either insanitary or congested areas of the city contain far +more bacteria than those from the more sanitary, less congested, or +suburban areas. The number of aerobic bacteria from the former varied +from 800,000 to 500,000,000 per fly and from the latter from 21,000 to +100,000. The number of intestinal forms conveyed by flies from +insanitary or congested areas was from 10,000 to 333,000,000 as compared +with from 100 to 10,000 carried by flies from the more sanitary areas. + +Pathogenic bacteria and those allied to the food poisoning group were +only obtained from the congested or moderately congested areas and not +from the suburban areas, where the chances of infestation were less. + +The interesting fact was brought out that flies caught in milk shops +apparently carry and obtain more bacteria than those from other shops +with exposed food in a similar neighborhood. The writers explained this +as probably due to the fact that milk when accessible, especially during +the summer months, is suitable culture medium for bacteria, and the +flies first inoculate the milk and later reinoculate themselves, and +then more of the milk, so establishing a vicious circle. + +They conclude that in cities where food is plentiful flies rarely +migrate from the locality in which they are bred, and consequently the +number of bacteria which they carry depends upon the general standard +of cleanliness in that locality. Flies caught in a street of modern, +fairly high class, workmen's dwellings forming a sanitary oasis in the +midst of a slum area, carried far less bacteria than those caught in the +adjacent neighborhood. + +Thus, as the amount of dirt carried by flies in any particular locality, +measured in the terms of bacteria, bears a definite relation to the +habits of the people and to the state of the streets, it demonstrates +the necessity of efficient municipal and domestic cleanliness, if the +food of the inhabitants is to escape pollution, not only with harmless +but also with occasional pathogenic bacteria. + +The above cited work is of a general nature, but, especially in recent +years, many attempts have been made to determine more specifically the +ability of flies to transmit pathogenic organisms. The critical reviews +of Nuttall and Jepson (1909), Howard (1911), and Graham-Smith (1913) +should be consulted by the student of the subject. We can only cite here +a few of the more striking experiments. + +Celli (1888) fed flies on pure cultures of _Bacillus typhosus_ and +declared that he was able to recover these organisms from the intestinal +contents and excrement. + +Firth and Horrocks (1902), cited by Nuttall and Jepson, "kept _Musca +domestica_ (also bluebottles) in a large box measuring 4 × 3 × 3 feet, +with one side made of glass. They were fed on material contaminated with +cultures of _B. typhosus_. Agar plates, litmus, glucose broth and a +sheet of clean paper were at the same time exposed in the box. After a +few days the plates and broth were removed and incubated with a positive +result." Graham-Smith (1910) "carried out experiments with large numbers +of flies kept in gauze cages and fed for eight hours on emulsions of _B. +typhosus_ in syrup. After that time the infested syrup was removed and +the flies were fed on plain syrup. _B. typhosus_ was isolated up to 48 +hours (but not later) from emulsions of their feces and from plates over +which they walked." + +Several other workers, notably Hamilton (1903), Ficker (1903), +Bertarelli (1910) Faichnie (1909), and Cochrane (1912), have isolated +_B. typhosus_ from "wild" flies, naturally infected. The papers of +Faichnie and of Cochrane we have not seen, but they are quoted in +_extenso_ by Graham-Smith (1913). + +On the whole, the evidence is conclusive that typhoid germs not only may +be accidentally carried on the bodies of house-flies but may pass +through their bodies and be scattered in a viable condition in the feces +of the fly for at least two days after feeding. Similar, results have +been reached in experiments with cholera, tuberculosis and yaws, the +last-mentioned being a spirochæte disease. Darling (1913) has shown that +murrina, a trypanosome disease of horses and mules in the Canal zone is +transmitted by house-flies which feed upon excoriated patches of +diseased animals and then pass to cuts and galls of healthy animals. + +Since it is clear that flies are abundantly able to disseminate viable +pathogenic bacteria, it is important to consider whether they have +access to such organisms in nature. A consideration of the method of +spread of typhoid will serve to illustrate the way in which flies may +play an important rôle. + +Typhoid fever is a specific disease caused by _Bacillus typhosus_, and +by it alone. The causative organism is to be found in the excrement and +urine of patients suffering from the disease. More than that, it is +often present in the dejecta for days, weeks, or even months and years, +after the individual has recovered from the disease. Individuals so +infested are known as "typhoid carriers" and they, together with those +suffering from mild cases, or "walking typhoid," are a constant menace +to the health of the community in which they are found. + +Human excrement is greedily visited by flies, both for feeding and for +ovipositing. The discharges of typhoid patients, or of chronic +"carriers," when passed in the open, in box privies, or camp latrines, +or the like, serve to contaminate myriads of the insects which may then +spread the germ to human food and drink. Other intestinal diseases may +be similarly spread. There is abundant epidæmiological evidence that +infantile diarrhoea, dysentery, and cholera may be so spread. + +Stiles and Keister (1913) have shown that spores of _Lamblia +intestinalis_, a flagellate protozoan living in the human intestine, may +be carried by house-flies. Though this species is not normally +pathogenic, one or more species of _Entamoeba_ are the cause of a type +of a highly fatal tropical dysentery. Concerning it, and another +protozoan parasite of man, they say, "If flies can carry _Lamblia_ +spores measuring 10 to 7µ, and bacteria that are much smaller, and +particles of lime that are much larger, there is no ground to assume +that flies may not carry _Entamoeba_ and _Trichomonas_ spores." + +Tuberculosis is one of the diseases which it is quite conceivable may be +carried occasionally. The sputum of tubercular patients is very +attractive to flies, and various workers, notably Graham-Smith, have +found that _Musca domestica_ may distribute the bacillus for several +days after feeding on infected material. + +A type of purulent opthalmia which is very prevalent in Egypt is often +said to be carried by flies. Nuttall and Jepson (1909) consider that the +evidence regarding the spread of this disease by flies is conclusive and +that the possibility of gonorrhoeal secretions being likewise conveyed +cannot be denied. + +Many studies have been published, showing a marked agreement between the +occurrence of typhoid and other intestinal diseases and the prevalence +of house-flies. The most clear-cut of these are the studies of the Army +Commission appointed to investigate the cause of epidemics of enteric +fever in the volunteer camps in the Southern United States during the +Spanish-American War. Though their findings as presented by Vaughan +(1909), have been quoted very many times, they are so germane to our +discussion that they will bear repetition: + +"Flies swarmed over infected fecal matter in the pits and fed upon the +food prepared for the soldiers in the mess tents. In some instances +where lime had recently been sprinkled over the contents of the pits, +flies with their feet whitened with lime were seen walking over the +food." Under such conditions it is no wonder that "These pests had +inflicted greater loss upon American soldiers than the arms of Spain." + +Similar conditions prevailed in South Africa during the Boer War. Seamon +believes that very much of the success of the Japanese in their fight +against Russia was due to the rigid precautions taken to prevent the +spread of disease by these insects and other means. + +Veeder has pointed out that the characteristics of a typical fly-borne +epidemic of typhoid are that it occurs in little neighborhood epidemics, +extending by short leaps from house to house, without regard to water +supply or anything else in common. It tends to follow the direction of +prevailing winds (cf. the conclusions of Hindle and Merriman). It occurs +during warm weather. Of course, when the epidemic is once well under +way, other factors enter into its spread. + +In general, flies may be said to be the chief agency in the spread of +typhoid in villages and camps. In cities with modern sewer systems they +are less important, though even under the best of such conditions, they +are important factors. Howard has emphasized that in such cities there +are still many uncared-for box privies and that, in addition, the +deposition of feces overnight in uncared-for waste lots and alleys is +common. + +Not only unicellular organisms, such as bacteria and protozoa, but also +the eggs, embryos and larvæ of parasitic worms have been found to be +transported by house-flies. Ransom (1911) has found that _Habronema +muscæ_, a nematode worm often found in adult flies, is the immature +stage of a parasite occurring in the stomach of the horse. The eggs or +embryos passing out with the feces of the horse, are taken up by fly +larvæ and carried over to the imago stage. + +Grassi (1883), Stiles (1889), Calandruccio (1906), and especially Nicoll +(1911), have been the chief investigators of the ability of house-flies +to carry the ova and embryos of human intestinal parasites. Graham-Smith +(1913) summarizes the work along this line as follows: + +"It is evident from the investigations that have been quoted that +house-flies and other species are greatly attracted to the ova of +parasitic worms contained in feces and other materials, and make great +efforts to ingest them. Unless the ova are too large they often succeed, +and the eggs are deposited uninjured in their feces, in some cases up to +the third day at least. The eggs may also be carried on their legs or +bodies. Under suitable conditions, food and fluids may be contaminated +with the eggs of various parasitic worms by flies, and in one case +infection of the human subject has been observed. Feces containing +tape-worm segments may continue to be a source of infection for as long +as a fortnight. Up to the present, however, there is no evidence to show +what part flies play in the dissemination of parasitic worms under +natural conditions." + +Enough has been said to show that the house-fly must be dealt with as a +direct menace to public health. Control measures are not merely matters +of convenience but are of vital importance. + +Under present conditions the speedy elimination of the house-fly is +impossible and the first thing to be considered is methods of protecting +food and drink from contamination. The first of these methods is the +thorough screening of doors and windows to prevent the entrance of +flies. In the case of kitchen doors, the flies, attracted by odors, are +likely to swarm onto the screen and improve the first opportunity for +gaining an entrance. This difficulty can be largely avoided by +screening-in the back porch and placing the screen door at one end +rather than directly before the door. + +The use of sticky fly paper to catch the pests that gain entrance to the +house is preferable to the various poisons often used. Of the latter, +formalin (40 per cent formaldehyde) in the proportion of two +tablespoonfuls to a pint of water is very efficient, if all other +liquids are removed or covered, so that the flies must depend on the +formalin for drink. The mixture is said to be made more attractive by +the addition of sugar or milk, though we have found the plain solution +wholly satisfactory, under proper conditions. It should be emphasized +that this formalin mixture is not perfectly harmless, as so often +stated. There are on record cases of severe and even fatal poisoning +from the accidental drinking of solutions. + +When flies are very abundant in a room they can be most readily gotten +rid of by fumigation with sulphur, or by the use of pure pyrethrum +powder either burned or puffed into the air. Herrick (1913) recommends +the following method: "At night all the doors and windows of the kitchen +should be closed; fresh powder should be sprinkled over the stove, on +the window ledges, tables, and in the air. In the morning flies will be +found lying around dead or stupified. They may then be swept up and +burned." This method has proved very efficaceous in some of the large +dining halls in Ithaca. + +The writers have had little success in fumigating with the vapors of +carbolic acid, or carbolic acid and gum camphor, although these methods +will aid in driving flies from a darkened room. + +All of these methods are but makeshifts. As Howard has so well put it, +"the truest and simplest way of attacking the fly problem is to prevent +them from breeding, by the treatment or abolition of all places in which +they can breed. To permit them to breed undisturbed and in countless +numbers, and to devote all our energy to the problem of keeping them out +of our dwellings, or to destroy them after they have once entered in +spite of all obstacles, seems the wrong way to go about it." + +We have already seen that _Musca domestica_ breeds in almost any +fermenting organic material. While it prefers horse manure, it breeds +also in human feces, cow dung and that of other animals, and in refuse +of many kinds. To efficiently combat the insect, these breeding places +must be removed or must be treated in some such way as to render them +unsuitable for the development of the larvæ. Under some conditions +individual work may prove effective, but to be truly efficient there +must be extensive and thorough coöperative efforts. + +Manure, garbage, and the like should be stored in tight receptacles and +carted away at least once a week. The manure may be carted to the fields +and spread. Even in spread manure the larvæ may continue their +development. Howard points out that "it often happens that after a lawn +has been heavily manured in early summer the occupants of the house will +be pestered with flies for a time, but finding no available breeding +place these disappear sooner or later. Another generation will not breed +in the spread manure." + +Hutchinson (1914) has emphasized that the larvæ of houseflies have +deeply engrained the habit of migrating in the prepupal stage and has +shown that this offers an important point of attack in attempts to +control the pest. He has suggested that maggot traps might be developed +into an efficient weapon in the warfare against the house-fly. Certain +it is that the habit greatly simplifies the problem of treating the +manure for the purpose of killing the larvæ. + +There have been many attempts to find some cheap chemical which would +destroy fly larvæ in horse manure without injuring the bacteria or +reducing the fertilizing values of the manure. The literature abounds in +recommendations of kerosene, lime, chloride of lime, iron sulphate, and +other substances, but none of them has met the situation. The whole +question has been gone into thoroughly by Cook, Hutchinson and Scales +(1914), who tested practically all of the substances which have been +recommended. They find that by far the most effective, economical, and +practical of the substances is borax in the commercial form in which it +is available throughout the country. + +"Borax increases the water-soluble nitrogen, ammonia and alkalinity of +manure and apparently does not permanently injure the bacterial flora. +The application of manure treated with borax at the rate of 0.62 pound +per eight bushels (10 cubic feet) to soil does not injure the plants +thus far tested, although its cumulative effect, if any, has not been +determined." + +As their results clearly show that the substances so often recommended +are inferior to borax, we shall quote in detail their directions for +treating manure so as to kill fly eggs and maggots. + +"Apply 0.62 pound borax or 0.75 pound calcined colemanite to every 10 +cubic feet (8 bushels) of manure immediately on its removal from the +barn. Apply the borax particularly around the outer edges of the pile +with a flour sifter or any fine sieve, and sprinkle two or three gallons +of water over the borax-treated manure. + +"The reason for applying the borax to the fresh manure immediately after +its removal from the stable is that the flies lay their eggs on the +fresh manure, and borax, when it comes in contact with the eggs, +prevents their hatching. As the maggots congregate at the outer edge of +the pile, most of the borax should be applied there. The treatment +should be repeated with each addition of fresh manure, but when the +manure is kept in closed boxes, less frequent applications will be +sufficient. When the calcined colemanite is available, it may be used at +the rate of 0.75 pound per 10 cubic feet of manure, and is a cheaper +means of killing the maggots. In addition to the application of borax to +horse manure to kill fly larvæ, it may be applied in the same proportion +to other manures, as well as to refuse and garbage. Borax may also be +applied to the floors and crevices in barns, stables, markets, etc., as +well as to street sweepings, and water should be added as in the +treatment of horse manure. After estimating the amount of material to be +treated and weighing the necessary amount of borax, a measure may be +used which will hold the proper amount, thus avoiding the subsequent +weighings. + +"While it can be safely stated that no injurious action will follow the +application of manure treated with borax at the rate of 0.62 pound for +eight bushels, or even larger amounts in the case of some plants, +nevertheless the borax-treated manure has not been studied in connection +with the growth of all crops, nor has its cumulative effect been +determined. It is therefore recommended that not more than 15 tons per +acre of the borax-treated manure should be applied to the field. As +truckmen use considerably more than this amount, it is suggested that +all cars containing borax-treated manure be so marked, and that +public-health officials stipulate in their directions for this treatment +that not over 0.62 pound for eight bushels of manure be used, as it has +been shown that larger amounts of borax will injure most plants. It is +also recommended that all public-health officials and others, in +recommending the borax treatment for killing fly eggs and maggots in +manure, warn the public against the injurious effects of large amounts +of borax on the growth of plants." + +"The amount of manure from a horse varies with the straw or other +bedding used, but 12 or 15 bushels per week represent the approximate +amount obtained. As borax costs from five to six cents per pound in +100-pound lots in Washington, it will make the cost of the borax +practically one cent per horse, per day. And if calcined colemanite is +purchased in large shipments the cost should be considerably less." + +Hodge (1910) has approached the problem of fly extermination from +another viewpoint. He believes that it is practical to trap flies out of +doors during the preoviposition period, when they are sexually immature, +and to destroy such numbers of them that the comparatively few which +survive will not be able to lay eggs in sufficient numbers to make the +next generation a nuisance. To the end of capturing them in enormous +numbers he has devised traps to be fitted over garbage cans, into stable +windows, and connected with the kitchen window screens. Under some +conditions this method of attack has proved very satisfactory. + +One of the most important measures for preventing the spread of disease +by flies is the abolition of the common box privy. In villages and rural +districts this is today almost the only type to be found. It is the +chief factor in the spread of typhoid and other intestinal diseases, as +well as intestinal parasites. Open and exposed to myriads of flies which +not only breed there but which feed upon the excrement, they furnish +ideal conditions for spreading contamination. Even where efforts are +made to cover the contents with dust, or ashes, or lime, flies may +continue to breed unchecked. Stiles and Gardner have shown that +house-flies buried in a screened stand-pipe forty-eight inches under +sterile sand came to the surface. Other flies of undetermined species +struggled up through seventy-two inches of sand. + +So great is the menace of the ordinary box privy that a number of +inexpensive and simple sanitary privies have been designed for use where +there are not modern sewer systems. Stiles and Lumsden (1911) have given +minute directions for the construction of one of the best types, and +their bulletin should be obtained by those interested. + +Another precaution which is of fundamental importance in preventing the +spread of typhoid, is that of disinfecting all discharges from patients +suffering with the disease. For this purpose, quick-lime is the cheapest +and is wholly satisfactory. In chamber vessels it should be used in a +quantity equal to that of the discharge to be treated. It should be +allowed to act for two hours. Air-slaked lime is of no value whatever. +Chloride of lime, carbolic acid, or formalin may be used, but are more +expensive. Other intestinal diseases demand similar precautions. + + +STOMOXYS CALCITRANS, THE STABLE-FLY--It is a popular belief that +house-flies bite more viciously just before a rain. As a matter of +fact, the true house-flies never bite, for their mouth-parts are not +fitted for piercing. The basis of the misconception is the fact that a +true biting fly, _Stomoxys calcitrans_ (fig. 110), closely resembling +the house-fly, is frequently found in houses and may be driven in in +greater numbers by muggy weather. From its usual habitat this fly is +known as the "stable-fly" or, sometimes as the "biting house-fly." + +_Stomoxys calcitrans_ may be separated from the house-fly by the use of +the key on p. 145. It may be more fully characterized as follows: + +The eyes of the male are separated by a distance equal to one-fourth of +the diameter of the head, in the female by one-third. The frontal stripe +is black, the cheeks and margins of the orbits silvery-white. The +antennæ are black, the arista feathered on the upper side only. The +proboscis is black, slender, fitted for piercing and projects forward in +front of the head. The thorax is grayish, marked by four conspicuous, +more or less complete black longitudinal stripes; the scutellum is +paler; the macrochætæ are black. The abdomen is gray, dorsally with +three brown spots on the second and third segments and a median spot on +the fourth. These spots are more pronounced in the female. The legs are +black, the pulvilli distinct. The wings are hyaline, the vein M_{1+2} +less sharply curved than in the house-fly, the apical cell being thus +more widely open (cf. fig. 110). Length 7 mm. + +[Illustration: 110. Stomoxys calcitrans; adult, larva, puparium and +details, (×5). After Howard.] + +This fly is widely distributed, being found the world over. It was +probably introduced into the United States, but has spread to all parts +of the country. Bishopp (1913) regards it as of much more importance as +a pest of domestic animals in the grain belt than elsewhere in the +United States. The life-history and habits of this species have assumed +a new significance since it has been suggested that it may transmit the +human diseases, infantile paralysis and pellagra. In this country, the +most detailed study of the fly is that of Bishopp (1913) whose data +regarding the life cycle are as follows: + +The eggs like those of the house-fly, are about one mm. in length. Under +a magnifying glass they show a distinct furrow along one side. When +placed on any moist substance they hatch in from one to three days after +being deposited. + +The larva or maggots (fig. 110) have the typical shape and actions of +most maggots of the Muscid group. They can be distinguished from those +of the house-fly as the stigma-plates are smaller, much further apart, +with the slits less sinuous. Development takes place fairly rapidly when +the proper food conditions are available and the growth is completed +within eleven to thirty or more days. + +The pupa (fig. 110), like that of related flies, undergoes its +development within the contracted and hardened last larval skin, or +puparium. This is elongate oval, slightly thicker towards the head end, +and one-sixth to one-fourth of an inch in length. The pupal stage +requires six to twenty days, or in cool weather considerably longer. + +The life-cycle of the stable-fly is therefore considerably longer than +that of _Musca domestica_. Bishopp found that complete development might +be undergone in nineteen days, but that the average period was somewhat +longer, ranging from twenty-one to twenty-five days, where conditions +are very favorable. The longest period which he observed was forty-three +days, though his finding of full grown larvæ and pupæ in straw during +the latter part of March, in Northern Texas, showed that development may +require about three months, as he considered that these stages almost +certainly developed from eggs deposited the previous December. + +The favorite breeding place, where available, seems to be straw or +manure mixed with straw. It also breeds in great numbers in +horse-manure, in company with _Musca domestica_. + +Newstead considers that in England the stable-fly hibernates in the +pupal stage. Bishopp finds that in the southern part of the United +States there is no true hibernation, as the adults have been found to +emerge at various times during the winter. He believes that in the +northern United States the winter is normally passed in the larval and +pupal stages, and that the adults which have been observed in heated +stables in the dead of winter were bred out in refuse within the warm +barns and were not hibernating adults. + +Graham-Smith (1913) states that although the stable-fly frequents stable +manure, it is probably not an important agent in distributing the +organisms of intestinal diseases. Bishopp makes the important +observation that "it has never been found breeding in human excrement +and does not frequent malodorous places, which are so attractive to the +house-fly. Hence it is much less likely to carry typhoid and other germs +which may be found in such places." + +Questions of the possible agency of _Stomoxys calcitrans_ in the +transmission of infantile paralysis and of pellagra, we shall consider +later. + + +OTHER ARTHROPODS WHICH MAY SERVE AS SIMPLE CARRIERS OF PATHOGENIC +ORGANISMS--It should be again emphasized that any insect which has +access to, and comes in contact with, pathogenic organisms and then +passes to the food, or drink, or the body of man, may serve as a simple +carrier of disease. In addition to the more obvious illustrations, an +interesting one is the previously cited case of the transfer of +_Dermatobia cyaniventris_ by a mosquito (fig. 81-84). Darling (1913) has +shown that in the tropics, the omnipresent ants may be important factors +in the spread of disease. + + + + +CHAPTER VI + +ARTHROPODS AS DIRECT INOCULATORS OF DISEASE GERMS + + +We have seen that any insect which, like the house-fly, has access to +disease germs and then comes into contact with the food or drink of man, +may serve to disseminate disease. Moreover, it has been clearly +established that a contaminated insect, alighting upon wounded or +abraded surfaces, may infect them. These are instances of mere +accidental, mechanical transfer of pathogenic organisms. + +Closely related are the instances of direct inoculation of disease germs +by insects and other arthropods. In this type, a blood-sucking species +not only takes up the germs but, passing to a healthy individual, it +inserts its contaminated mouth-parts and thus directly inoculates its +victim. In other words, the disease is transferred just as blood +poisoning may be induced by the prick of a contaminated needle, or as +the laboratory worker may inoculate an experimental animal. + +Formerly, it was supposed that this method of the transfer of disease by +arthropods was a very common one and many instances are cited in the +earlier literature of the subject. It is, however, difficult to draw a +sharp line between such cases and those in which, on the one hand, the +arthropod serves as a mere passive carrier or, on the other hand, serves +as an essential host of the pathogenic organism. More critical study of +the subject has led to the belief that the importance of the rôle of +arthropods as direct inoculators has been much overestimated. + +The principal reason for regarding this phase of the subject as +relatively unimportant, is derived from a study of the habits of the +blood-sucking species. It is found that, in general, they are +intermittent feeders, visiting their hosts at intervals and then +abstaining from feeding for a more or less extended period, while +digesting their meal. In the meantime, most species of bacteria or of +protozoan parasites with which they might have contaminated their +mouth-parts, would have perished, through inability to withstand drying. + +In spite of this, it must be recognized that this method of transfer +does occur and must be reckoned with in any consideration of the +relations of insects to disease. We shall first cite some general +illustrations and shall then discuss the rôle of fleas in the spreading +of bubonic plague, an illustration which cannot be regarded as typical, +since it involves more than mere passive carriage. + + +SOME ILLUSTRATIONS OF DIRECT INOCULATION OF DISEASE GERMS BY ARTHROPODS + +In discussing poisonous arthropods, we have already emphasized that +species which are of themselves innocuous to man, may occasionally +introduce bacteria by their bite or sting and thus cause more or less +severe secondary symptoms. That such cases should occur, is no more than +is to be expected. The mouth-parts or the sting of the insect are not +sterile and the chances of their carrying pyogenic organisms are always +present. + +More strictly falling in the category of transmission of disease germs +by direct inoculation are the instances where the insect, or related +form, feeds upon a diseased animal and passes promptly to a healthy +individual which it infects. Of such a nature are the following: + +Various species of biting flies are factors in the dissemination of +anthrax, an infectious and usually fatal disease of animals and, +occasionally, of man. That the bacteria with which the blood of diseased +animals teem shortly before death might be transmitted by such insects +has long been contended, but the evidence in support of the view has +been unsatisfactory. Recently, Mitzmain (1914) has reported a series of +experiments which show conclusively that the disease may be so conveyed +by a horse-fly, _Tabanus striatus_, and by the stable-fly, _Stomoxys +calcitrans_. + +Mitzmain's experiments were tried with an artificially infected guinea +pig, which died of the disease upon the third day. The flies were +applied two and one-half hours, to a few minutes, before the death of +the animal. With both species the infection was successfully transferred +to healthy guinea pigs by the direct method, in which the flies were +interrupted while feeding on the sick animal. The evidence at hand does +not warrant the conclusion that insect transmission is the rule in the +case of this disease. + +The nagana, or tsetse-fly disease of cattle is the most virulent disease +of domestic animals in certain parts of Africa. It is caused by a +protozoan blood parasite, _Trypanosoma brucei_, which is conveyed to +healthy animals by the bite of _Glossina morsitans_ and possibly other +species of tsetse-flies. The flies remain infective for forty-eight +hours after feeding on a diseased animal. The insect also serves as an +essential host of the parasite. + +Surra, a similar trypanosomiasis affecting especially horses and mules, +occurs in southern Asia, Malaysia, and the Philippines where the +tsetse-flies are not to be found. It is thought to be spread by various +species of blood-sucking flies belonging to the genera _Stomoxys_, +_Hæmatobia_, and _Tabanus_. Mitzmain (1913) demonstrated that in the +Philippines it is conveyed mechanically by _Tabanus striatus_. + +The sleeping sickness of man, in Africa, has also been supposed to be +directly inoculated by one, or several, species of tsetse-flies. It is +now known that the fly may convey the disease for a short time after +feeding, but that there is then a latent period of from fourteen to +twenty-one days, after which it again becomes infectious. This indicates +that in the meantime the parasite has been undergoing some phase of its +life-cycle and that the fly serves as an intermediate host. We shall +therefore consider it more fully under that grouping. + +These are a few of the cases of direct inoculation which may be cited as +of the simpler type. We shall next consider the rôle of the flea in the +dissemination of the bubonic plague, an illustration complicated by the +fact that the bacillus multiples within the insect and may be indirectly +inoculated. + + +THE RÔLE OF FLEAS IN THE TRANSMISSION OF THE PLAGUE + +The plague is a specific infectious disease caused by _Bacillus pestis_. +It occurs in several forms, of which the bubonic and the pneumonic are +the most common. According to Wyman, 80 per cent of the human cases are +of the bubonic type. It is a disease which, under the name of oriental +plague, the pest, or the black death, has ravaged almost from time +immemorial the countries of Africa, Asia, and Europe. The record of its +ravages are almost beyond belief. In 542 A. D. it caused in one day ten +thousand deaths in Constantinople. In the 14th century it was introduced +from the East and prevailed throughout Armenia, Asia Minor, Egypt and +Northern Africa and Europe. Hecker estimates that one-fourth of the +population of Europe, or twenty-five million persons, died in the +epidemic of that century. From then until the 17th century it was almost +constantly present in Europe, the great plague of London, in 1665 +killing 68,596 out of a population of 460,000. Such an epidemic would +mean for New York City a proportionate loss of over 600,000 in a single +year. It is little wonder that in the face of such an appalling disaster +suspicion and credulity were rife and the wildest demoralization ensued. + +During the 14th century the Jews were regarded as responsible for the +disease, through poisoning wells, and were subjected to the most +incredible persecution and torture. In Milan the visitation of 1630 was +credited to the so-called anointers,--men who were supposed to spread +the plague by anointing the walls with magic ointment--and the most +horrible tortures that human ingenuity could devise were imposed on +scores of victims, regardless of rank or of public service (fig. 112, +a). Manzoni's great historical novel, "The Betrothed" has well pictured +conditions in Italy during this period. + +[Illustration: 111. A contemporaneous engraving of the pest hospital in +Vienna in 1679. After Peters.] + +In modern times the plague is confined primarily to warm climates, a +condition which has been brought about largely through general +improvement in sanitary conditions. + +At present, the hotbed of the disease is India, where there were +1,040,429 deaths in 1904 and where in a period of fifteen years, ending +with January 1912, there were over 15,000,000 deaths. The reported +deaths in that country for 1913 totaled 198,875. + +During the winter of 1910-11 there occurred in Manchuria and North China +a virulent epidemic of the pneumonic plague which caused the death of +nearly 50,000 people. The question as to its origin and means of spread +will be especially referred to later. + +[Illustration: 112 a. A medieval method of combating the plague. The +persecution of the anointers in Milan in 1630. From a copy of "Il +processi originale degli untori" in the library of Cornell University.] + +Until recent years, the plague had not been known to occur in the New +World but there were outbreaks in Brazil and Hawaii in 1899, and in 1900 +there occurred the first cases in San Francisco. In California there +were 125 cases in the period 1900-04; three cases in the next three +years and then from May 1907 to March 1908, during the height of the +outbreak, 170 cases. Since that time there have been only sporadic +cases, the last case reported being in May 1914. Still more recent were +the outbreaks in the Philippine Islands, Porto Rico, and Cuba. + +On June 24, 1914, there was recognized a case of human plague in New +Orleans. The Federal Health Service immediately took charge, and +measures for the eradication of the disease were vigorously enforced. Up +to October 10, 1914 there had been reported 30 cases of the disease in +man, and 181 cases of plague in rats. + +[Illustration: 112 b. The modern method of combating the plague. A day's +catch of rats in the fight against plague in San Francisco. Courtesy of +Review of Reviews.] + +The present-day methods of combating bubonic plague are well illustrated +by the fight in San Francisco. Had it not been for the strenuous and +radical anti-plague campaign directed by the United States Marine +Hospital Service we might have had in our own country an illustration of +what the disease can accomplish. On what newly acquired knowledge was +this fight based? + +The basis was laid in 1894, when the plague bacillus was first +discovered. All through the centuries, before and during the Christian +era, down to 1894, the subject was enveloped in darkness and there had +been a helpless, almost hopeless struggle in ignorance on the part of +physicians, sanitarians, and public health officials against the ravages +of this dread disease. Now its cause, method of propagation and means to +prevent its spread are matters of scientific certainty. + +After the discovery of the causative organism, one of the first advances +was the establishment of the identity of human plague and that of +rodents. It had often been noted that epidemics of the human disease +were preceded by great epizootics among rats and mice. So well +established was this fact that with the Chinese, unusual mortality among +these rodents was regarded as foretelling a visitation of the human +disease. That there was more than an accidental connection between the +two was obvious when Yersin, the discoverer of _Bacillus pestis_, +announced that during an epidemic the rats found dead in the houses and +in the streets almost always contain the bacillus in great abundance in +their organs, and that many of them exhibit veritable buboes. + +Once it was established that the diseases were identical, the attention +of the investigators was directed to a study of the relations between +that of rats and of humans, and evidence accumulated to show that the +bubonic plague was primarily a disease of rodents and that in some +manner it was conveyed from them to man. + +There yet remained unexplained the method of transfer from rat to man. +As long ago as the 16th century, Mercuralis suggested that house-flies +were guilty of disseminating the plague but modern investigation, while +blaming the fly for much in the way of spreading disease, show that it +is an insignificant factor in this case. + +Search for blood-sucking insects which would feed on both rodents and +man, and which might therefore be implicated, indicated that the fleas +most nearly met the conditions. At first it was urged that rat fleas +would not feed upon man and that the fleas ordinarily attacking man +would not feed upon rats. More critical study of the habits of fleas +soon showed that these objections were not well-founded. Especially +important was the evidence that soon after the death of their host, rat +fleas deserted its body and might then become a pest in houses where +they had not been noticed before. + +Attention was directed to the fact that while feeding, fleas are in the +habit of squirting blood from the anus and that in the case of those +which had fed upon rats and mice dying of the plague, virulent plague +bacilli were to be found in such blood. Liston (1905) even found, and +subsequent investigations confirmed, that the plague bacilli multiply in +the stomach of the insect and that thus the blood ejected was richer in +the organisms than was that of the diseased animal. It was found that a +film of this infected blood spread out under the body of the flea and +that thus the bacilli might be inoculated by the bite of the insect and +by scratching. + +Very recently, Bacot and Martin (1914) have paid especial attention to +the question of the mechanism of the transmission of the plague bacilli +by fleas. They believe that plague infested fleas regurgitate blood +through the mouth, and that under conditions precluding the possibility +of infection by dejecta, the disease may be thus transmitted. The +evidence does not seem sufficient to establish that this is the chief +method of transmission. + +Conclusive experimental proof that fleas transmit the disease is further +available from a number of sources. The most extensive series of +experiments is that of the English Plague Commission in India, which +reported in 1906 that: + +On thirty occasions a healthy rat contracted plague in sequence of +living in the neighborhood of a plague infected rat under circumstances +which prevented the healthy rat coming in contact with either the body +or excreta of the diseased animal. + +In twenty-one experiments out of thirty-eight, healthy rats living in +flea-proof cages contracted plague when exposed to rat fleas +(_Xenopsylla cheopis_), collected from rats dead or dying of septicæmic +plague. + +Close contact of plague-infected with healthy animals, if fleas are +excluded, does not give rise to an epizootic among the latter. As the +huts were never cleaned out, close contact included contact with feces +and urine of infected animals, and contact with, and eating of food +contaminated with feces and urine of infected animals, as well as pus +from open plague ulcers. Close contact of young, even when suckled by +plague-infected mothers, did not give the disease to the former. + +If fleas are present, then the epizootic, once started, spreads from +animal to animal, the rate of progress being in direct proportion to the +number of fleas. + +Aerial infection was excluded. Thus guinea-pigs suspended in a cage two +feet above the ground did not contract the disease, while in the same +hut those animals allowed to run about and those placed two inches above +the floor became infected. It had previously been found that a rat flea +could not hop farther than about five inches. + +Guinea pigs and monkeys were placed in plague houses in pairs, both +protected from soil contact infection and both equally exposed to aerial +infection, but one surrounded with a layer of tangle-foot paper and the +other surrounded with a layer of sand. The following observations were +made: + +(_a_) Many fleas were caught in the tangle-foot, a certain proportion of +which were found on dissection to contain in their stomachs abundant +bacilli microscopically identical with plague bacilli. Out of +eighty-five human fleas dissected only one contained these bacilli, +while out of seventy-seven rat fleas twenty-three were found thus +infected. + +(_b_) The animals surrounded with tangle-foot in no instance developed +plague, while several (24 per cent) of the non-protected animals died of +the disease. + +Thus, the experimental evidence that fleas transmit the plague from rat +to rat, from rats to guinea pigs, and from rats to monkeys is +indisputable. There is lacking direct experimental proof of its transfer +from rodents to man but the whole chain of indirect evidence is so +complete that there can be no doubt that such a transfer does occur so +commonly that in the case of bubonic plague it must be regarded as the +normal method. + +Rats are not the only animals naturally attacked by the plague but as +already suggested, it occurs in various other rodents. In California the +disease has spread from rats to ground squirrels (_Otospermophilus +beecheyi_), a condition readily arising from the frequency of +association of rats with the squirrels in the neighborhood of towns, and +from the fact that the two species of fleas found on them are also found +on rats. While the danger of the disease being conveyed from squirrels +to man is comparatively slight, the menace in the situation is that the +squirrels may become a more or less permanent reservoir of the disease +and infect rats, which may come into more frequent contact with man. + +The tarbagan (_Arctomys bobac_), is a rodent found in North Manchuria, +which is much prized for its fur. It is claimed that this animal is +extremely susceptible to the plague and there is evidence to indicate +that it was the primary source of the great outbreak of pneumonic plague +which occurred in Manchuria and North China during the winter of +1910-11. + +Of fleas, any species which attacks both rodents and man may be an agent +in the transmission of the plague. We have seen that in India the +species most commonly implicated is the rat flea, _Xenopsylla cheopis_, +(= _Loemopsylla_ or _Pulex cheopis_) (fig. 89). This species has also +been found commonly on rats in San Francisco. The cat flea, +_Ctenocephalus felis_, the dog flea, _Ctenocephalus canis_, the human +flea, _Pulex irritans_, the rat fleas, _Ceratophyllus fasciatus_ and +_Ctenopsyllus musculi_ have all been shown to meet the conditions. + +But, however clear the evidence that fleas are the most important agent +in the transfer of plague, it is a mistake fraught with danger to assume +that they are the only factor in the spread of the disease. The +causative organism is a bacillus and is not dependent upon any insect +for the completion of its development. + +Therefore, any blood-sucking insect which feeds upon a plague infected +man or animal and then passes to a healthy individual, conceivably might +transfer the bacilli. Verjbitski (1908) has shown experimentally that +bed-bugs may thus convey the disease. Hertzog found the bacilli in a +head-louse, _Pediculus humanus_, taken from a child which had died from +the plague, and McCoy found them in a louse taken from a plague-infected +squirrel. On account of their stationary habits, the latter insects +could be of little significance in spreading the disease. + +Contaminated food may also be a source of danger. While this source, +formerly supposed to be the principal one, is now regarded as +unimportant, there is abundant experimental evidence to show that it +cannot be disregarded. It is believed that infection in this way can +occur only when there is some lesion in the alimentary canal. + +Still more important is the proof that in pneumonic plague the patient +is directly infective and that the disease is spread from man to man +without any intermediary. Especially conclusive is the evidence obtained +by Drs. Strong and Teague during the Manchurian epidemic of 1910-11. +They found that during coughing, in pneumonic plague cases, even when +sputum visible to the naked eye is not expelled, plague bacilli in large +numbers may become widely disseminated into the surrounding air. By +exposing sterile plates before patients who coughed a single time, very +numerous colonies of the bacillus were obtained. + +But the great advance which has been made rests on the discovery that +bubonic plague is in the vast majority of cases transmitted by the flea. +The pneumonic type forms a very small percentage of the human cases and +even with it, the evidence indicates that the original infection is +derived from a rodent through the intermediary of the insect. + +So modern prophylactic measures are directed primarily against the rat +and fleas. Ships coming from infected ports are no longer disinfected +for the purpose of killing the plague germs, but are fumigated to +destroy the rats and the fleas which they might harbor. When anchored at +infected ports, ships must observe strenuous precautions to prevent the +ingress of rats. Cargo must be inspected just before being brought on +board, in order to insure its freedom from rats. Even lines and hawsers +must be protected by large metal discs or funnels, for rats readily run +along a rope to reach the ship. Once infested, the ship must be +thoroughly fumigated, not only to avoid carrying the disease to other +ports but to obviate an outbreak on board. + +When an epidemic begins, rats must be destroyed by trapping and +poisoning. Various so-called biological poisons have not proved +practicable. Sources of food supply should be cut off by thorough +cleaning up, by use of rat-proof garbage cans and similar measures. Hand +in hand with these, must go the destruction of breeding places, and the +rat-proofing of dwellings, stables, markets, warehouses, docks and +sewers. All these measures are expensive, and a few years ago would have +been thought wholly impossible to put into practice but now they are +being enforced on a large scale in every fight against the disease. + +Rats and other rodents are regularly caught in the danger zone and +examined for evidence of infection, for the sequence of the epizootic +and of the human disease is now understood. In London, rats are +regularly trapped and poisoned in the vicinity of the principal docks, +to guard against the introduction of infected animals in shipping. +During the past six years infected rats have been found yearly, thirteen +having been found in 1912. In Seattle, Washington, seven infected rats +were found along the water front in October, 1913, and infected ground +squirrels are still being found in connection with the anti-plague +measures in California. + +The procedure during an outbreak of the human plague was well +illustrated by the fight in San Francisco. The city was districted, and +captured rats, after being dipped in some fluid to destroy the fleas, +were carefully tagged to indicate their source, and were sent to the +laboratory for examination. If an infected rat was found, the officers +in charge of the work in the district involved were immediately notified +by telephone, and the infected building was subjected to a thorough +fumigation. In addition, special attention was given to all the +territory in the four contiguous blocks. + +By measures such as these, this dread scourge of the human race is being +brought under control. Incidentally, the enormous losses due to the +direct ravages of rats are being obviated and this alone would justify +the expenditure many times over of the money and labor involved in the +anti-rat measures. + + + + +CHAPTER VII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC ORGANISMS + + +We now have to consider the cases in which the arthropod acts as the +essential host of a pathogenic organism. In other words, cases in which +the organism, instead of being passively carried or merely accidentally +inoculated by the bite of its carrier, or vector, is taken up and +undergoes an essential part of its development within the arthropod. + +[Illustration: 113. Dipylidium caninum. The double pored tapeworm of the +dog.] + +In some cases, the sexual cycle of the parasite is undergone in the +arthropod, which then serves as the _definitive_ or _primary host_. In +other cases, it is the asexual stage of the parasite which is undergone, +and the arthropod then acts as the _intermediate host_. This distinction +is often overlooked and all the cases incorrectly referred to as those +in which the insect or other arthropod acts as intermediate host. + +We have already emphasized that this is the most important way in which +insects may transmit disease, for without them the particular organisms +concerned could never complete their development. Exterminate the +arthropod host and the life cycle of the parasite is broken, the disease +is exterminated. + +As the phenomenon of alternation of generations, as exhibited by many of +the parasitic protozoa, is a complicated one and usually new to the +student, we shall first take up some of the grosser cases illustrated by +certain parasitic worms. There is the additional reason that these were +the first cases known of arthropod transmission of pathogenic organisms. + + +INSECTS AS INTERMEDIATE HOSTS OF TAPEWORMS + +A number of tapeworms are known to undergo their sexual stage in an +insect or other arthropod. Of these at least two are occasional +parasites of man. + +_Dipylidium caninum_ (figs. 113 and 114), more generally known as +_Taenia cucumerina_ or _T. elliptica_, is the commonest intestinal +parasite of pet dogs and cats. It is occasionally found as a human +parasite, 70 per cent of the cases reported being in young children. + +In 1869, Melnikoff found in a dog louse, _Trichodectes canis_, some +peculiar bodies which Leuckart identified as the larval form of this +tapeworm. The worm is, however, much more common in dogs and cats than +is the skin parasite, and hence it appears that the _Trichodectes_ could +not be the only intermediate host. In 1888, Grassi found that it could +also develop in the cat and dog fleas, _Ctenocephalus felis_ and _C. +canis_, and in the human flea, _Pulex irritans_. + +[Illustration: 114. Dipylidium caninum. Rostrum evaginated and +invaginated. After Blanchard.] + +[Illustration: 115. Dipylidium caninum. Immature cysticercoid. After +Grassi and Rovelli.] + +The eggs, scattered among the hairs of the dog or cat, are ingested by +the insect host and in its body cavity they develop into pyriform +bodies, about 300µ in length, almost entirely destitute of a bladder, +but in the immature stage provided with a caudal appendage (fig. 115). +Within the pear-shaped body (fig. 116) are the invaginated head and +suckers of the future tapeworm. This larval form is known as a +cysticercoid, in contradistinction to the bladder-like cysticercus of +many other cestodes. It is often referred to in literature as +_Cryptocystis trichodectis_ Villot. + +As many as fifty of the cysticercoids have been found in the body cavity +of a single flea. When the dog takes up an infested flea or louse, by +biting itself, or when the cat licks them up, the larvæ quickly develop +into tapeworms, reaching sexual maturity in about twenty days in the +intestine of their host. Puppies and kittens are quickly infested when +suckling a flea-infested mother, the developing worms having been found +in the intestines of puppies not more than five or six days old. + +[Illustration: 116. Dipylidium caninum. Cysticercoid. After Villet.] + +Infestation of human beings occurs only through accidental ingestion of +an infested flea. It is natural that such cases should occur largely in +children, where they may come about in some such way as illustrated in +the accompanying figures 117 and 118. + +_Hymenolepis diminuta_, very commonly living in the intestine of mice +and rats, is also known to occur in man. Its cysticercoid develops in +the body cavity of a surprising range of meal-infesting insects. Grassi +and Rovelli (abstract in Ransom, 1904) found it in the larvæ and adult +of a moth, _Asopia farinalis_, in the earwig, _Anisolabis annulipes_, +the Tenebrionid beetles _Akis spinosa_ and _Scaurus striatus_. Grassi +considers that the lepidopter is the normal intermediate host. The +insect takes up the eggs scattered by rats and mice. It has been +experimentally demonstrated that man may develop the tapeworm by +swallowing infested insects. Natural infection probably occurs by +ingesting such insects with cereals, or imperfectly cooked foods. + +[Illustration: 117. One way in which Dipylidium infection in children +may occur. After Blanchard.] + +_Hymenolepis lanceolata_, a parasite of geese and ducks, has been +reported once for man. The supposed cysticercoid occurs in various small +crustaceans of the family Cyclopidæ. + +[Illustration: 118. The probable method by which Dipylidium infection +usually occurs.] + +Several other cestode parasites of domestic animals are believed to +develop their intermediate stage in certain arthropods. Among these may +be mentioned: + +_Choanotænia infundibulformis_, of chickens, developing in the house-fly +(Grassi and Rovelli); + +_Davainea cesticillus_, of chickens, in some lepidopter or coleopter +(Grassi and Rovelli); + +_Hymenolepis anatina_, _H. gracilis_, _H. sinuosa_, _H. coronula_ and +_Fimbriaria fasciolaris_, all occurring in ducks, have been reported as +developing in small aquatic crustaceans. In these cases, cysticercoids +have been found which, on account of superficial characters, have been +regarded as belonging to the several species, but direct experimental +evidence is scant. + + +ARTHROPODS AS INTERMEDIATE HOSTS OF NEMATODE WORMS + + +FILARIASIS AND MOSQUITOES--A number of species of Nematode worms +belonging to the genus _Filaria_, infest man and other vertebrates and +in the larval condition are to be found in the blood. Such infestation +is known as _filariasis_. The sexually mature worms are to be found in +the blood, the lymphatics, the mesentery and subcutaneous connective +tissue. In the cases best studied it has been found that the larval +forms are taken up by mosquitoes and undergo a transformation before +they can attain maturity in man. + +The larvæ circulating in the blood are conveniently designated as +microfilariæ. In this stage they are harmless and only one species, +_Filaria bancrofti_, appears to be of any great pathological +significance at any stage. + +_Filaria bancrofti_ in its adult state, lives in the lymphatics of man. +Though often causing no injury it has been clearly established that they +and their eggs may cause various disorders due to stoppage of the +lymphatic trunks (fig. 119). Manson lists among other effects, abscess, +varicose groin glands, lymph scrotum, chyluria, and elephantiasis. + +The geographical distribution of this parasite is usually given as +coextensive with that of elephantiasis, but it is by no means certain +that it is the only cause of this disease and so actual findings of the +parasites are necessary. Manson reports that it is "an indigenous +parasite in almost every country throughout the tropical and subtropical +world, as far north as Spain in Europe and Charlestown in the United +States, and as far south as Brisbane in Australia." In some sections, +fully 50 per cent of the natives are infested. Labredo (1910) found +17.82 per cent infestation in Havana. + +[Illustration: 119. Elephantiasis in Man. From "New Sydenham Society's +Atlas."] + +The larval forms of _Filaria bancrofti_ were first discovered in 1863, +by Demarquay, in a case of chylous dropsy. They were subsequently noted +under similar conditions, by several workers, and by Wücherer in the +urine of twenty-eight cases of tropical chyluria, but in 1872 Lewis +found that the blood of man was the normal habitat, and gave them the +name _Filaria sanguinis hominis_. The adult worm was found in 1876 by +Bancroft, and in 1877, Cobbold gave it the name _Filaria bancrofti_. It +has since been found repeatedly in various parts of the lymphatic +system, and its life-history has been the subject of detailed studies by +Manson (1884), Bancroft (1899), Low (1900), Grassi and Noé (1900), Noé +(1901) and Fülleborn (1910). + +The larvæ as they exist in the circulating blood, exhibit a very active +wriggling movement, without material progression. They may exist in +enormous numbers, as many as five or six hundred swarming in a single +drop of blood. This is the more surprising when we consider that they +measure about 300µ × 8µ, that is, their width is equal to the diameter +of the red blood corpuscle of their host and their length over +thirty-seven times as great. + +Their organs are very immature and the structure obscure. When they have +quieted down somewhat in a preparation it may be seen that at the head +end there is a six-lipped and very delicate prepuce, enclosing a short +"fang" which may be suddenly exserted and retracted. Completely +enclosing the larva is a delicate sheath, which is considerably longer +than the worm itself. To enter into further details of anatomy is beyond +the scope of this discussion and readers interested are referred to the +work of Manson and of Fülleborn. + +One of the most surprising features of the habits of these larvæ is the +periodicity which they exhibit in their occurrence in the peripheral +blood. If a preparation be made during the day time there may be no +evidence whatever of filarial infestation, whereas a preparation from +the same patient taken late in the evening or during the night may be +literally swarming with the parasites. Manson quotes Mackenzie as having +brought out the further interesting fact that should a "filarial subject +be made to sleep during the day and remain awake at night, the +periodicity is reversed; that is to say, the parasites come into the +blood during the day and disappear from it during the night." There have +been numerous attempts to explain this peculiar phenomenon of +periodicity but in spite of objections which have been raised, the most +plausible remains that of Manson, who believes that it is an adaptation +correlated with the life-habits of the liberating agent of the parasite, +the mosquito. + +The next stages in the development of _Filaria nocturna_ occur in +mosquitoes, a fact suggested almost simultaneously by Bancroft and +Manson in 1877, and first demonstrated by the latter very soon +thereafter. The experiments were first carried out with _Culex +quinquefasciatus_ (= _fatigans_) as a host, but it is now known that a +number of species of mosquitoes, both anopheline and culicine, may serve +equally well. + +When the blood of an infested individual is sucked up and reaches the +stomach of such a mosquito, the larvæ, by very active movements, escape +from their sheaths and within a very few hours actively migrate to the +body cavity of their new host and settle down primarily in the thoracic +muscles. There in the course of sixteen to twenty days they undergo a +metamorphosis of which the more conspicuous features are the formation +of a mouth, an alimentary canal and a trilobed tail. At the same time +there is an enormous increase in size, the larvæ which measured .3 mm. +in the blood becoming 1.5 mm. in length. This developmental period may +be somewhat shortened in some cases and on the other hand may be +considerably extended. The controlling factor seems to be the one of +temperature. + +The transformed larvæ then reenter the body cavity and finally the +majority of them reach the interior of the labium (fig. 120). A few +enter the legs and antennæ, and the abdomen, but these are wanderers +which, it is possible, may likewise ultimately reach the labium, where +they await the opportunity to enter their human host. + +It was formerly supposed that when the infested mosquito punctured the +skin of man, the mature larvæ were injected into the circulation. The +manner in which this occurred was not obvious, for when the insect feeds +it inserts only the stylets, the labium itself remaining on the surface +of the skin. Fülleborn has cleared up the question by showing that at +this time the filariæ escape and, like the hookworm, actively bore into +the skin of their new host. + +[Illustration: 120. Filaria in the muscles and labium of Culex. After +Blanchard.] + +Once entered, they migrate to the lymphatics and there quickly become +sexually mature. The full grown females measure 85-90 mm. in length by +.24-.28 mm. in diameter, while the males are less than half this size, +being about 40 mm. by .1 mm. Fecundation occurs and the females will be +found filled with eggs in various stages of development, for they are +normally viviparous. + +_Filaria philippinensis_ is reported by Ashburn and Craig (1907) as a +common blood filaria in the Philippine Islands. As they describe it, it +differs from _Filaria bancrofti_ primarily in that it does not exhibit +periodicity. Its development has been found to occur in _Culex +quinquefasciatus_, where it undergoes metamorphosis in about fourteen or +fifteen days. There is doubt as to the species being distinct from +_bancrofti_. + +Several other species occur in man and are thought to be transferred by +various insects, among which have been mentioned Tabanidæ and +tsetse-flies, but there is no experimental proof in support of such +conjectures. + +_Filaria immitis_ is a dangerous parasite of the dog, the adult worm +living in the heart and veins of this animal. It is one of the species +which has been clearly shown to undergo its development in the mosquito, +particularly in _Anopheles maculipennis_ and _Aedes calopus_ (= +Stegomyia). The larval form occurs in the peripheral blood, especially +at night. When taken up by mosquitoes they differ from _Filaria +bancrofti_ in that they undergo their development in the Malpighian +tubules rather than in the thoracic muscles. In about twelve days they +have completed their growth in the tubules, pierce the distal end, and +pass to the labium. This species occurs primarily in China and Japan, +but is also found in Europe and in the United States. It is an +especially favorable species for studying the transformations in the +mosquito. + +[Illustration: 121. Dracunculus medinensis; female; mouth; embryo. After +Bastian and Leuckart.] + +_Filariæ_ are also commonly found in birds, and in this country this is +the most available source of laboratory material. We have found them +locally (Ithaca, N. Y.) in the blood of over sixty per cent of all the +crows examined, at any season of the year, and have also found them in +English sparrows. + +In the crows, they often occur in enormous numbers, as many as two +thousand having been found in a single drop of the blood of the most +heavily infested specimen examined. For study, a small drop of blood +should be mounted on a clean slide and the coverglass rung with vaseline +or oil to prevent evaporation. In this way they can be kept for hours. + +Permanent preparations may be made by spreading out the blood in a film +on a perfectly clean slide and staining. This is easiest done by +touching the fresh drop of blood with the end of a second slide which is +then held at an angle of about 45° to the first slide and drawn over it +without pressure. Allow the smear to dry in the air and stain in the +usual way with hæmatoxylin. + + +OTHER NEMATODE PARASITES OF MAN AND ANIMALS DEVELOPING IN ARTHROPODS + +_Dracunculus medinensis_ (fig. 121), the so-called guinea-worm, is a +nematode parasite of man which is widely distributed in tropical Africa, +Asia, certain parts of Brazil and is occasionally imported into North +America. + +The female worm is excessively long and slender, measuring nearly three +feet in length and not more than one-fifteenth of an inch in diameter. +It is found in the subcutaneous connective tissue and when mature +usually migrates to some part of the leg. Here it pierces the skin and +there is formed a small superficial ulcer through which the larvæ reach +the exterior after bursting the body of the mother. + +[Illustration: 122. Cyclops, the intermediate host of Dracunculus.] + +Fedtschenko (1879) found that when these larvæ reach the water they +penetrate the carapace of the little crustacean, _Cyclops_ (fig. 122). +Here they molt several times and undergo a metamorphosis. Fedtschenko, +in Turkestan, found that these stages required about five weeks, while +Manson who confirmed these general results, found that eight or nine +weeks were required in the cooler climate of England. + +Infection of the vertebrate host probably occurs through swallowing +infested cyclops in drinking water. Fedtschenko was unable to +demonstrate this experimentally and objection has been raised against +the theory, but Leiper (1907), and Strassen (1907) succeeded in +infesting monkeys by feeding them on cyclops containing the larvæ. + +_Habronema muscæ_ is a worm which has long been known in its larval +stage, as a parasite of the house-fly. Carter found them in 33 per cent +of the house-flies examined in Bombay during July, 1860, and since that +time they have been shown to be very widely distributed. Italian workers +reported them in 12 per cent to 30 per cent of the flies examined. +Hewitt reported finding it rarely in England. In this country it was +first reported by Leidy who found it in about 20 per cent of the flies +examined at Philadelphia, Pa. Since then it has been reported by several +American workers. We have found it at Ithaca, N. Y., but have not made +sufficient examinations to justify stating percentage. Ransom (1913) +reports it in thirty-nine out of one hundred and thirty-seven flies, or +28 per cent. + +[Illustration: 123. An Echinorhynchid, showing the spinose retractile +proboscis.] + +[Illustration: 124. June beetle (Lachnosterna).] + +[Illustration: Larva] + +Until very recently the life-history of this parasite was unknown but +the thorough work of Ransom (1911, 1913) has shown clearly that the +adult stage occurs in the stomach of horses. The embryos, produced by +the parent worms in the stomach of the horse, pass out with the feces +and enter the bodies of fly larvæ which are developing in the manure. In +these they reach their final stage of larval development at about the +time the adult flies emerge from the pupal stage. In the adult fly they +are commonly found in the head, frequently in the proboscis, but they +occur also in the thorax and abdomen. Infested flies are accidentally +swallowed by horses and the parasite completes its development to +maturity in the stomach of its definitive host. + +_Gigantorhynchus hirudinaceus_ (= _Echinorhynchus gigas_) is a common +parasite of the pig and has been reported as occurring in man. The adult +female is 20-35 cm. long and 4-9 mm. in diameter. It lacks an alimentary +canal and is provided with a strongly spined protractile rostrum, by +means of which it attaches to the intestinal mucosa of its host. + +The eggs are scattered with the feces of the host and are taken up by +certain beetle larvæ. In Europe the usual intermediate hosts are the +larvæ of the cockchafer, _Melolontha vulgaris_, or of the flower beetle, +_Cetonia aurata_. Stiles has shown that in the United States the +intermediate host is the larva of the June bug, _Lachnosterna_ (fig. +124). It is probable that several of the native species serve in this +capacity. + +A number of other nematode parasites of birds and mammals have been +reported as developing in arthropods but here, as in the case of the +cestodes, experimental proof is scant. The cases above cited are the +better established and will serve as illustrations. + + + + +CHAPTER VIII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA + + +MOSQUITOES AND MALARIA + +Under the name of malaria is included a group of morbid symptoms +formerly supposed to be due to a miasm or bad air, but now known to be +caused by protozoan parasites of the genus _Plasmodium_, which attack +the red blood corpuscles. It occurs in paroxysms, each marked by a +chill, followed by high fever and sweating. The fever is either +intermittent or remittent. + +There are three principal types of the disease, due to different species +of the parasite. They are: + +1. The benign-tertian, caused by _Plasmodium vivax_, which undergoes its +schizogony or asexual cycle in the blood in forty-eight hours or even +less. This type of the disease,--characterized by fever every two days, +is the most wide-spread and common. + +2. The quartan fever is due to the presence of _Plasmodium malariæ_, +which has an asexual cycle of seventy-two hours, and therefore the fever +recurs every three days. This type is more prevalent in temperate and +sub-tropical regions, but appears to be rare everywhere. + +3. The sub-tertian "æstivo-autumnal," or "pernicious" fever is caused by +_Plasmodium falciparum_. Schizogony usually occurs in the internal +organs, particularly in the spleen, instead of in the peripheral +circulation, as is the case of the tertian and quartan forms. The fever +produced is of an irregular type and the period of schizogony has not +been definitely determined. It is claimed by some that the variations +are due to different species of malignant parasites. + +It is one of the most wide-spread of human diseases, occurring in almost +all parts of the world, except in the polar regions and in waterless +deserts. It is most prevalent in marshy regions. + +So commonplace is malaria that it causes little of the dread inspired by +most of the epidemic diseases, and yet, as Ross says, it is perhaps the +most important of human diseases. Figures regarding its ravages are +astounding. Celli estimated that in Italy it caused an average annual +mortality of fifteen thousand, representing about two million cases. In +India alone, according to Ross (1910) "it has been officially estimated +to cause a mean annual death-rate of five per thousand; that is, to kill +every year, on the average, one million one hundred and thirty +thousand." In the United States it is widespread and though being +restricted as the country develops, it still causes enormous losses. +During the year 1911, "in Alabama alone there were seventy thousand +cases and seven hundred and seventy deaths." The weakening effects of +the disease, the invasion of other diseases due to the attacks of +malaria, are among the very serious results, but they cannot be +estimated. + +Not only is there direct effect on man, but the disease has been one of +the greatest factors in retarding the development of certain regions. +Everywhere pioneers have had to face it, and the most fertile regions +have, in many instances been those most fully dominated by it. Herrick +(1903) has presented an interesting study of its effects on the +development of the southern United States and has shown that some parts, +which are among the most fertile in the world, are rendered practically +uninhabitable by the ravages of malaria. Howard (1909) estimates that +the annual money loss from the disease in the United States is not less +than $100,000,000. + +It was formerly supposed that the disease was due to a miasm, to a +noxious effluvia, or infectious matter rising in the air from swamps. In +other words its cause was, as the name indicated "mal aria," and the +deep seated fear of night air is based largely on the belief that this +miasm was given off at night. Its production was thought to be favored +by stirring of the soil, dredging operations and the like. + +The idea of some intimate connection between malaria and mosquitoes is +not a new one. According to Manson, Lancisi noted that in some parts of +Italy the peasants for centuries have believed that malaria is produced +by the bite of mosquitoes. Celli states that one not rarely hears from +such peasants the statement that "In such a place, there is much fever, +because it is full of mosquitoes." Koch points out that in German East +Africa the natives call malaria and the mosquito by the same name, +_Mbù_. The opinion was not lacking support from medical men. Celli +quotes passages from the writings of the Italian physician, Lancisi, +which indicate that he favored the view in 1717. + +Dr. Josiah Nott is almost universally credited with having supported the +theory, in 1848, but as we have already pointed out his work has been +misinterpreted. The statements of Beauperthuy, (1853) were more +explicit. + +The clearest early presentation of the circumstantial evidence in favor +of the theory of mosquito transmission was that of A. F. A. King, an +American physician, in 1883. He presented a series of epidemiological +data and showed "how they may be explicable by the supposition that the +mosquito is the real source of the disease, rather than the inhalation +or cutaneous absorption of a marsh vapor." We may well give the space to +summarizing his argument here for it has been so remarkably +substantiated by subsequent work: + +1. Malaria, like mosquitoes, affects by preference low and moist +localities, such as swamps, fens, jungles, marshes, etc. + +2. Malaria is hardly ever developed at a lower temperature than 60° +Fahr., and such a temperature is necessary for the development of the +mosquito. + +3. Mosquitoes, like malaria, may both accumulate in and be obstructed by +forests lying in the course of winds blowing from malarious localities. + +4. By atmospheric currents malaria and mosquitoes are alike capable of +being transported for considerable distances. + +5. Malaria may be developed in previously healthy places by turning up +the soil, as in making excavations for the foundation of houses, tracks +for railroads, and beds for canals, because these operations afford +breeding places for mosquitoes. + +6. In proportion as countries, previously malarious, are cleared up and +thickly settled, periodical fevers disappear, because swamps and pools +are drained so that the mosquito cannot readily find a place suitable to +deposit her eggs. + +7. Malaria is most dangerous when the sun is down and the danger of +exposure after sunset is greatly increased by the person exposed +sleeping in the night air. Both facts are readily explicable by the +mosquito malaria theory. + +8. In malarial districts the use of fire, both indoors and to those who +sleep out, affords a comparative security against malaria, because of +the destruction of mosquitoes. + +9. It is claimed that the air of cities in some way renders the poison +innocuous, for, though a malarial disease may be raging outside, it does +not penetrate far into the interior. We may easily conceive that +mosquitoes, while invading cities during their nocturnal pilgrimages +will be so far arrested by walls and houses, as well as attracted by +lights in the suburbs, that many of them will in this way be prevented +from penetrating "far into the interior." + +10. Malarial diseases and likewise mosquitoes are most prevalent toward +the latter part of summer and in the autumn. + +11. Various writers have maintained that malaria is arrested by canvas +curtains, gauze veils and mosquito nets and have recommended the rise of +mosquito curtains, "through which malaria can seldom or never pass." It +can hardly be conceived that these intercept marsh-air but they +certainly do protect from mosquitoes. + +12. Malaria spares no age, but it affects infants much less frequently +than adults, because young infants are usually carefully housed and +protected from mosquito inoculation. + +Correlated with the miasmatic theory was the belief that some animal or +vegetable organism which lived in marshes, produced malaria, and +frequent searches were made for it. Salisbury (1862) thought this +causative organism to be an alga, of the genus _Palmella_; others +attributed it to certain fungi or bacteria. + +In 1880, the French physician, Laveran, working in Algeria, discovered +an amoeboid organism in the blood of malarial patients and definitely +established the parasitic nature of this disease. Pigmented granules had +been noted by Meckel as long ago as 1847, in the spleen and blood of a +patient who had died of malaria, and his observations had been +repeatedly verified, but the granules had been regarded as degeneration +products, and the fact that they occurred in the body of a foreign +organism had been overlooked. + +Soon after the discovery of the parasites in the blood, Gerhardt (1884) +succeeded in transferring the disease to healthy individuals by +inoculation of malarious blood, and thus proved that it is a true +infection. This was verified by numerous experimenters and it was found +that inoculation with a very minute quantity of the diseased blood would +not only produce malaria but the particular type of disease. + +Laveran traced out the life cycle of the malarial parasite as it occurs +in man. The details as we now know them and as they are illustrated by +the accompanying figure 125, are as follows: + +The infecting organism or _sporozoite_, is introduced into the +circulation, penetrates a red blood corpuscle, and forms the amoeboid +_schizont_. This lives at the expense of the corpuscle and as it +develops there are deposited in its body scattered black or reddish +black particles. These are generally called melanin granules, but are +much better referred to as hæmozoin, as they are not related to +melanin. The hæmozoin is the most conspicuous part of the parasite, a +feature of advantage in diagnosing from unstained preparations. + +[Illustration: 125. Life cycle of the malaria parasite. Adapted from +Leuckart's chart, by Miss Anna Stryke.] + +As the schizont matures, its nucleus breaks up into a number of daughter +nuclei, each with a rounded mass of protoplasm about it, and finally the +corpuscles are broken down and these rounded bodies are liberated in the +plasma as _merozoites_. These merozoites infect new corpuscles and thus +the asexual cycle is continued. The malarial paroxysm is coincident with +sporulation. + +As early as Laveran's time it was known that under conditions not yet +determined there are to be found in the blood of malarious patients +another phase of the parasite, differing in form according to the type +of the disease. In the pernicious type these appear as large, +crescent-shaped organisms which have commonly been called "crescents." +We now know that these are sexual forms. + +When the parasite became known there immediately arose speculations as +to the way in which it was transferred from man to man. It was thought +by some that in nature it occurred as a free-living amoeba, and that +it gained access to man through being taken up with impure water. +However, numerous attempts to infect healthy persons by having them +drink or inhale marsh water, or by injecting it into their circulation +resulted in failure, and influenced by Leuckart's and Melnikoff's work +on _Dipylidium_, that of Fedtschenko on _Dracunculus_, and more +especially by that of Manson on _Filaria_, search was made for some +insect which might transfer the parasite. + +Laveran had early suggested that the rôle of carrier might be played by +the mosquito, but Manson first clearly formulated the hypothesis, and it +was largely due to his suggestions that Ross in India, undertook to +solve the problem. With no knowledge of the form or of the appearance in +this stage, or of the species of mosquito concerned, Ross spent almost +two and a half years of the most arduous work in the search and finally +in August, 1897, seventeen years after the discovery of the parasite in +man, he obtained his first definite clue. In dissecting a +"dappled-winged mosquito," "every cell was searched and to my intense +disappointment nothing whatever was found, until I came to the insect's +stomach. Here, however, just as I was about to abandon the examination, +I saw a very delicate circular cell, apparently lying amongst the +ordinary cells of the organ and scarcely distinguishable from them. On +looking further, another and another similar object presented itself. I +now focused the lens carefully on one of these, and found that it +contained a few minute granules of some black substance, exactly like +the pigment of the parasite of malaria. I counted altogether twelve of +these cells in the insect." + +Further search showed that "the contents of the mature pigment cells did +not consist of clear fluid but of a multitude of delicate, thread-like +bodies which on the rupture of the parent cell, were poured into the +body cavity of the insect. They were evidently spores." + +With these facts established, confirmation and extension of Ross's +results quickly followed, from many different sources. We cannot trace +this work in detail but will only point out that much of the credit is +due to the Italian workers, Grassi, Bignami, and Bastianelli, and to +Koch and Daniels. + +It had already been found that when fresh blood was mounted and properly +protected against evaporation, a peculiar change occurred in these +crescents after about half an hour's time. From certain of them there +were pushed out long whip-like processes which moved with a very active, +lashing movement. The parasite at this stage is known as the +"flagellated body." Others, differing somewhat in details of structure, +become rounded but do not give off "flagella." + +The American worker, MacCallum (1897), in studying bird malaria as found +in crows, first recognized the true nature of these bodies. He regarded +them as sexual forms and believed that the so-called flagella played the +part of spermatozoa. Thus, the "flagellated body" is in reality a +_microgametoblast_, producing _microgametes_, or the male sexual +element, while the others constitute the _macrogametes_, or female +elements. + +It was found that when blood containing these sexual forms was sucked up +by an Anopheline mosquito and taken into its stomach, a microgamete +penetrated and fertilized a macrogamete in a way analogous to what takes +place in the fertilization of the egg in higher forms. The resultant, +mobile organism is known as the _migratory ookinete_. In this stage the +parasite bores through the epithelial lining of the "stomach" +(mid-intestine) of the mosquito and becomes encysted under the muscle +layers. Here the _oocyst_, as it is now known, matures and breaks up +into the body cavity and finally its products come to lie in the +salivary glands of the mosquito. Ten to twelve days are required for +these changes, after which the mosquito is infective, capable of +introducing the parasite with its saliva, when feeding upon a healthy +person. + +Thus the malarial parasite is known to have a double cycle, an +alternation of generations, of which the asexual stage is undergone in +man, the sexual in certain species of mosquitoes. The mosquito is +therefore the definitive host rather than the _intermediate_, as usually +stated. + +The complicated cycle may be made clearer by the diagram of Miss Stryke +(1912) which, by means of a double-headed mosquito (fig. 126) endeavors +to show how infection takes place through the biting of the human +victim, (at A), in whom asexual multiplication then takes place, and how +the sexual stages, taken up at B in the diagram, are passed in the body +of the mosquito. + +[Illustration: 126. Life cycle of the malarial parasite. After Miss Anna +Stryke.] + +The experimental proof that mosquitoes of the Anopheline group are +necessary agents in the transmission of malaria was afforded in 1900 +when two English physicians, Drs. Sambon and Low lived for the three +most malarial months in the midst of the Roman Campagna, a region +famous for centuries as a hot-bed of malaria. The two experimenters +moved about freely throughout the day, exposed themselves to rains and +all kinds of weather, drank marsh water, slept exposed to the marsh air, +and, in short, did everything which was supposed to cause malaria, +except that they protected themselves thoroughly from mosquito bites, +retiring at sunset to a mosquito-proof hut. Though they took no quinine +and all of their neighbors suffered from malaria, they were absolutely +free from the disease. + +To complete the proof, mosquitoes which had fed in Rome on malarious +patients were sent to England and allowed to bite two volunteers, one of +them Dr. Manson's own son, who had not been otherwise exposed to the +disease. Both of these gentlemen contracted typical cases of malaria and +the parasites were to be found in abundance in their blood. + +[Illustration: 127. Eggs of Anopheles. After Howard.] + +Since that time there have been many practical demonstrations of the +fact that malaria is transmitted exclusively by the bite of mosquitoes +and that the destruction of the mosquitoes means the elimination of the +disease. + +We have said that the malarial parasite is able to undergo its +development only in certain species of mosquitoes belonging to the +Anopheline group. It is by no means certain that all of this group even, +are capable of acting as the definitive host of the parasites, and much +careful experiment work is still needed along this line. In the United +States, several species have been found to be implicated, _Anopheles +quadrimaculatus_ and _Anopheles crucians_ being the most common. The +characteristics of these species and the distinctions between them and +other mosquitoes will be discussed in Chapter XII. + +In antimalarial work it is desirable to distinguish the anopheline +mosquitoes from the culicine species in all stages. The following +tabulation presents the more striking distinctions between the groups as +represented in the United States. + +_Anopheles_ _Culex, Aedes, etc_. + +_Eggs_: Laid singly in small Deposited in clumps in the +numbers upon the surface of the form of a raft (Culex group) or +water. Eggs lie upon their sides deposited singly in the water or +and float by means of lateral on the ground in places which +expansions (fig. 127). may later be submerged. + +_Larva_: When at rest floats in When at rest (with few exceptions) +a horizontal position beneath the floats suspended in an +surface film. No respiratory oblique or vertical position, or +tube but instead a flattened more rarely nearly horizontal, +area on the eighth abdominal with the respiratory tube in +segment into which the two contact with the surface film +spiracles open (fig. 128). (fig. 128). + +_Adults_: Palpi in both sexes Palpi short in the female, in +nearly or quite as long as the the male usually elongate. +proboscis. Proboscis projecting Proboscis projects forward at an +forward nearly on line with the angle with the axis of the body. +axis of the body. When at rest When at rest on a vertical wall +on a vertical wall the body is the body is usually held parallel +usually held at an angle with the or the tip of the abdomen inclined +vertical (fig. 128). Wings frequently towards the wall (fig. 128). +spotted (fig. 130). Wings usually not spotted. + +[Illustration: 128. (_a_) Normal position of the larvæ of Culex and +Anopheles in the water. Culex, left; Anopheles, middle; Culex pupa, +right hand figure.] + +These malarial-bearing species are essentially domesticated mosquitoes. +They develop in any accumulation of water which stands for a week or +more. Ponds, puddles, rain barrels, horse troughs, cess-pools, cans, +even the foot-prints of animals in marshy ground may afford them +breeding places. + +[Illustration: 128. (_b_) Normal position of Culex and Anopheles on the +wall.] + +It is clear from what has been said regarding the life cycle of the +malarial parasite that the mosquito is harmless if not itself diseased. +Hence malarial-bearing species may abound in the neighborhood where +there is no malaria, the disease being absent simply because the +mosquitoes are uninfected. Such a locality is potentially malarious and +needs only the introduction of a malarial patient who is exposed to the +mosquitoes. It is found that such patients may harbor the parasites in +their blood long after they are apparently well and thus may serve as a +menace, just as do the so-called typhoid carriers. In some malarious +regions as high as 80-90 per cent of the natives are such +malaria-carriers and must be reckoned with in antimalaria measures. + +Based upon our present day knowledge of the life cycle of the malarial +parasite the fight against the disease becomes primarily a problem in +economic entomology,--it is a question of insect control, in its +broadest interpretation. + +[Illustration: 129. Larva of Anopheles. After Howard.] + +The lines of defence and offence against the disease as outlined by +Boyce (1909) are: + + 1. Measures to avoid the reservoir (man): + + Segregation. + + Screening of patients. + + 2. Measures to avoid Anopheles: + + Choice of suitable locality, when possible. + + Screening of houses and porches. + + Sleeping under mosquito nets. + + 3. Measures to exterminate the Anopheles: + + Use of natural enemies. + + Use of culicides, oiling ponds, etc. + + Drainage and scavenging to destroy breeding places. + + Enforcement of penalties for harboring larvæ or keeping stagnant + water. + + Educational methods. + + 4. Systematic treatment with quinine to exterminate the parasites. + + +MOSQUITOES AND YELLOW FEVER + +Yellow fever was until recently one of the most dreaded of epidemic +diseases. It is an acute, specific and infectious disease, +non-contagious in character but occurring in epidemics, or endemics, +within a peculiarly limited geographical area. It is highly fatal, but +those who recover are generally immune from subsequent attacks. + +It is generally regarded as an American disease, having been found by +Cortez, in Mexico, and being confined principally to the American +continents and islands. It also occurs in Africa and attempts have been +made to show that it was originally an African disease but there is not +sufficient evidence to establish this view. + +There have been many noted outbreaks in the United States. Boston +suffered from it in 1691 and again in 1693; New York in 1668 and as late +as 1856; Baltimore in 1819. In 1793 occurred the great epidemic in +Philadelphia, with a death rate of one in ten of the population. In the +past century it was present almost every year in some locality of our +Southern States, New Orleans being the greatest sufferer. In the latter +city there were 7848 deaths from the disease in 1853, 4854 in 1858, and +4046 in 1878. The last notable outbreak was in 1905. Reed and Carroll +(1901) estimated that during the period from 1793 to 1900 there had not +been less than 500,000 cases in the United States. + +[Illustration: 130. Anopheles quadrimaculatus, male and female, +(×3½). After Howard.] + +As in the case of the plague, the most stringent methods of control +proved ineffective and helplessness, almost hopelessness marked the +great epidemics. A vivid picture of conditions is that given by Mathew +Cary, 1793 (quoted by Kelly, 1906) in "A Short Account of the Malignant +Fever Lately Prevalent in Philadelphia." + +[Illustration: 131. Anopheles punctipennis. Female, (×4). After Howard.] + +"The consternation of the people of Philadelphia at this period was +carried beyond all bounds. Dismay and affright were visible in the +countenance of almost every person. Of those who remained, many shut +themselves in their houses and were afraid to walk the streets. * * * +The corpses of the most respectable citizens, even those who did not die +of the epidemic, were carried to the grave on the shafts of a chair +(chaise), the horse driven by a negro, unattended by friends or +relative, and without any sort of ceremony. People hastily shifted their +course at the sight of a hearse coming toward them. Many never walked on +the footpath, but went into the middle of the streets to avoid being +infected by passing by houses wherein people had died. Acquaintances and +friends avoided each other in the streets and only signified their +regard by a cold nod. The old custom of shaking hands fell into such +disuse that many shrunk back with affright at even the offer of the +hand. A person with a crape, or any appearance of mourning was shunned +like a viper. And many valued themselves highly on the skill and address +with which they got to the windward of every person they met. Indeed, it +is not probable that London, at the last stage of the plague, exhibited +stronger marks of terror than were to be seen in Philadelphia from the +24th or 25th of August until pretty late in September." + +[Illustration: 132. Anopheles crucians. Female (×4). After Howard.] + +Such was the condition in Philadelphia in 1793 and, as far as methods of +control of the disease were concerned, there was practically no advance +during the last century. The dominant theory was that yellow fever was +spread by _fomites_, that is, exposed bedding, clothing, baggage, and +the like. As late as 1898 a bulletin of the United States Marine +Hospital Service stated: + +"While yellow fever is a communicable disease, it is not contagious in +the ordinary acceptance of the term, but is spread by the infection of +places and articles of bedding, clothing, and furniture." + +Based upon this theory, houses, baggage, freight, even mail, were +disinfected, and the most rigid quarantine regulations were enforced. +The hardships to which people of the stricken regions were subjected and +the financial losses are incalculable. And withal, the only efficient +check upon the disease seemed to be the heavy frosts. It was found that +for some reason, the epidemic abated with cold weather,--a measure +beyond human control. + +[Illustration: 133. Culex sollicitans. Female (×4). After Howard.] + +It is not strange that among the multitude of theories advanced to +explain the cause and method of dissemination of the disease there +should be suggestions that yellow fever was transmitted by the mosquito. +We have seen that Beauperthuy (1855) clearly urged this theory. + +More detailed, and of the greatest influence in the final solution of +the problem were the arguments of Dr. Cárlos Finlay, of Havana. In 1881, +in a paper presented before the "Real Academia de Ciencias Médicas, +Físicas y Naturales de la Habana," he said: + +"I feel convinced that any theory which attributes the origin and the +propagation of yellow fever to atmospheric influences, to miasmatic or +meteorological conditions, to filth, or to the neglect of general +hygienic precautions, must be considered as utterly indefensible." + +He postulated the existence of a material transportable substance +causing yellow fever,--"something tangible which requires to be conveyed +from the sick to the healthy before the disease can be propagated" and +after discussing the peculiarities of the spread of the disease and the +influence of meteorological conditions, he decides that the carriers of +the disease must be sought among insects. He continues: + +"On the other hand, the fact of yellow fever being characterized both +clinically and (according to recent findings) histologically, by lesions +of the blood vessels and by alterations of the physical and chemical +conditions of the blood, suggested that the insect which should convey +the infectious particles from the patient to the healthy should be +looked for among those which drive their sting into blood vessels in +order to suck human blood. Finally, by reason of other considerations +which need not be stated here, I came to think that the mosquito might +be the transmitter of yellow fever." + +"Assimilating the disease to small-pox and to vaccination, it occurred +to me that in order to inoculate yellow fever it would be necessary to +pick out the inoculable material from within the blood vessels of a +yellow fever patient and to carry it likewise into the interior of a +blood vessel of a person who was to be inoculated. All of which +conditions the mosquito satisfies most admirably through its bite." + +In the course of his study of the problem, Finlay made detailed studies +of the life history and habits of the common mosquitoes at Havana, and +arrived at the conclusion that the carrier of the yellow fever was the +_Culex mosquito_ or _Aëdes calopus_, as it is now known. With this +species he undertook direct experimental tests, and believed that he +succeeded in transmitting the disease by the bite of infected mosquitoes +in three cases. Unfortunately, possibility of other exposure was not +absolutely excluded, and the experiments attracted little attention. + +Throughout the next twenty years Finlay continued his work on yellow +fever, modifying his original theory somewhat as time went on. Among his +later suggestions was that in the light of Smith's work on Texas fever, +his theory must be "somewhat modified so as to include the important +circumstance that the faculty of transmitting the yellow fever germ need +not be limited to the parent insect, directly contaminated by stinging a +yellow fever patient (or perhaps by contact with or feeding from his +discharges), but may be likewise inherited by the next generation of +mosquitoes issued from the contaminated parent." He believed that the +bite of a single mosquito produced a light attack of the disease and was +thus effective in immunizing the patient. Throughout the period, many +apparently successful attempts to transmit the disease by mosquitoes +were made. In the light of present day knowledge we must regard these as +defective not only because possibility of other infection was not +absolutely excluded but because no account was taken of the incubation +period within the body of the mosquito. + +In 1900, while the American army was stationed in Cuba there occurred an +epidemic of yellow fever and an army medical board was appointed for +"the purpose of pursuing scientific investigations with reference to the +acute infectious diseases prevalent on the island." This was headed by +Walter Reed and associated with him were James Carroll, Jesse W. Lazear +and Aristides Agramonte, the latter a Cuban immune. For a detailed +summary of this work the lay reader cannot do better than read Dr. +Kelly's fascinating biography "Walter Reed and Yellow Fever." + +Arriving at the army barracks near Havana the Commission first took up +the study of _Bacillus icteroides_, the organism which Sanarelli, an +Italian physician, had declared the causative agent in yellow fever. +They were unable to isolate this bacillus either from the blood during +life or from the blood and organs of cadavers and therefore turned their +attention to Finlay's theory of the propagation of yellow fever by means +of the mosquito. In this work they had the unselfish and enthusiastic +support of Dr. Finlay himself, who not only consulted with them and +placed his publications at their disposal, but furnished eggs from which +their experimental mosquitoes were obtained. Inoculations of eleven +non-immunes through the bite of infected mosquitoes were made, and of +these, two gave positive results. The first of the two was Dr. Carroll +who allowed himself to be bitten by a mosquito which had been caused to +feed upon four cases of yellow fever, two of them severe and two mild. +The first patient had been bitten twelve days before. + +Three days after being bitten, Dr. Carroll came down with a typical case +of yellow fever. So severe was the attack that for three days his life +hung in the balance. During his convalescence an incident occurred which +showed how the theory of mosquito transmission of the disease was +generally regarded. We quote from Dr. Kelly: "One of his nurses who came +from Tennessee had had considerable experience with yellow fever, having +indeed, lost her husband and several children from it. One day early in +his illness Dr. Carroll mentioned to her that he had contracted the +disease through the bite of a mosquito, and noticed that she looked +surprised. Some time later, when well enough to look over the daily +records of his condition, he found this entry: 'Says he got his illness +through the bite of a mosquito,--delirious'." + +The second case was that of an American who was bitten by four +mosquitoes, two of which had bitten severe (fatal) cases of yellow fever +twelve days previously, one of which had bitten a severe case (second +day) sixteen days before and one which had bitten a severe case eight +days before. Five days later, the subject developed a well pronounced +but mild case of the disease. + +In the meantime, another member of the Commission, Dr. Lazear, was +accidentally bitten by a mosquito while collecting blood from yellow +fever patients. Five days later he contracted a typical case which +resulted fatally. + +So clear was the evidence from these preliminary experiments that the +commission felt warranted in announcing, October 27, 1900, that, "The +mosquito serves as the intermediate host for the parasite of yellow +fever, and it is highly probable that the disease is only propagated +through the bite of this insect." + +In order to extend the experimental evidence under conditions which +could leave no possibility of infection from other sources, a special +experimental sanitary station, named in honor of the deceased member of +the Commission, was established in an open field near the town of +Quemados, Cuba. Here there were constructed two small buildings known +respectively as the "infected clothing building" and the "infected +mosquito building." + +The infected clothing building, 14 × 20 feet in size, was purposely so +constructed as to exclude anything like efficient ventilation, but was +thoroughly screened to prevent the entrance of mosquitoes. Into this +building were brought sheets, pillow-slips, blankets, etc., contaminated +by contact with cases of yellow fever and their discharges,--many of +them purposely soiled with a liberal quantity of black vomit, urine, and +fecal matter from patients sick with yellow fever. Nothing could better +serve as the fomites which were supposed to convey the dread disease. + +Three non-immunes unpacked these articles, giving each a thorough +handling and shaking in order to disseminate through the air of the room +the specific agent of the disease. They were then used in making up the +beds which the volunteers occupied each night for a period of twenty +days. The experiment was repeated three times, volunteers even sleeping +in the soiled garments of yellow fever victims but in not a single case +was there the slightest symptom of disease. The theory of the spread of +yellow fever by fomites was completely demolished. + +The infected mosquito building, equal in size to its companion, was the +antithesis as far as other features were concerned. It was so +constructed as to give the best possible ventilation, and bedding which +was brought into it was thoroughly sterilized. Like the infected +clothing building it was carefully screened, but in this case it was in +order to keep mosquitoes in it as well as to prevent entrance of others. +Through the middle of the room ran a mosquito-proof screen. + +On December 5, 1900, a non-immune volunteer who had been in the +quarantine camp for fifteen days and had had no other possible exposure, +allowed himself to be bitten by five mosquitoes which had fed on yellow +fever patients fifteen or more days previously. The results were fully +confirmatory of the earlier experiments of the Commission--at the end of +three days, nine and a half hours, the subject came down with a well +marked case of yellow fever. + +In all, ten cases of experimental yellow fever, caused by the bite of +infected mosquitoes were developed in Camp Lazear. Throughout the period +of the disease, other non-immunes slept in the little building, +separated from the patient only by the mosquito-proof screen, but in no +circumstances did they suffer any ill effects. + +It was found that a yellow fever patient was capable of infecting +mosquitoes only during the first three or four days after coming down +with the disease. Moreover, after the mosquito has bitten such a +patient, a period of at least twelve days must elapse before the insect +is capable of transmitting the disease. + +Once the organism has undergone its twelve day development, the mosquito +may remain infective for weeks. In experiments of the Commission, two of +the mosquitoes transmitted the disease to a volunteer fifty-seven days +after their contamination. No other volunteers presenting themselves, +one of these mosquitoes died the sixty-ninth and one the seventy-first +day after their original contamination, without it being determined +whether they were still capable of transmitting the disease. + +So carefully carried out was this work and so conclusive were the +results that Dr. Reed was justified in writing: + +"Six months ago, when we landed on this island, absolutely nothing was +known concerning the propagation and spread of yellow fever--it was all +an unfathomable mystery--but today the curtain has been drawn--its mode +of propagation is established and we know that a case minus mosquitoes +is no more dangerous than one of chills and fever." + +The conclusions of the Commission were fully substantiated by numerous +workers, notably Dr. Guiteras of the Havana Board of Health, who had +taken a lively interest in the work and whose results were made known in +1901, and by the Brazilian and French Commission at Sao Paulo, Brazil, +in 1903. + +Throughout the work of the Army Commission and down to the present time +many fruitless efforts have been made to discover the specific organism +of yellow fever. It was clearly established that the claims of Sanarelli +for _Bacillus icteroides_ were without foundation. It was found, too, +that whatever the infective agent might be it was capable of passing +through a Berkefeld filter and thus belongs to the puzzling group of +"filterable viruses." It was further found that the virus was destroyed +by heating up to 55° C for ten minutes. It is generally believed that +the organism is a Protozoan. + +The question of the hereditary transmission of the yellow fever organism +within the mosquito was left unsettled by the Army Commission, though, +as we have seen, it was raised by Finlay. Marchoux and Simond, of the +French Commission devoted much attention to this phase of the problem +and basing their conclusions on one apparently positive case, they +decided that the disease could be transmitted through the egg of an +infected _Aëdes calopus_ to the second generation and thence to man. The +conclusion, which is of very great importance in the control of yellow +fever, has not been verified by other workers. + +Once clearly established that yellow fever was transmitted solely by +mosquitoes, the question of the characteristics, habits, and +geographical distribution of the insect carrier became of vital +importance. + +_Aëdes calopus_, more commonly known as _Stegomyia fasciata_ or +_Stegomyia calopus_ (fig. 134) is a moderate sized, rather strikingly +marked mosquito. The general color is dark-brown or reddish-brown, but +the thorax has a conspicuous broad, silvery-white curved line on each +side, with two parallel median silvery lines. Between the latter there +is a slender, broken line. The whole gives a lyre-shaped pattern to the +thorax. The abdomen is dark with silvery-white basal bands and silvery +white spots on each side of the abdominal segments. Legs black with +rings of pure white at the base of the segments. + +Size of the female 3.3 to 5 mm.; male 3 to 4.5 mm. + +[Illustration: 134. The yellow fever mosquito (Aëdes calopus). (×7). +After Howard.] + +It is preeminently a domesticated species, being found almost +exclusively about the habitation of man. "Its long association with man +is shown by many of its habits. It approaches stealthily from behind. It +retreats upon the slightest alarm. The ankles and, when one is sitting +at a table or desk, the underside of the hands and wrists are favorable +points of attack. It attacks silently, whereas other mosquitoes have a +piping or humming note. The warning sound has doubtless been suppressed +in the evolutionary process of its adaptation to man. It is extremely +wary. It hides whenever it can, concealing itself in garments, working +into the pockets, and under the lapels of coats, and crawling up under +the clothes to bite the legs. In houses, it will hide in dark corners, +under picture moldings and behind the heads of old-fashioned bedsteads. +It will enter closets and hide in the folds of garments."--Howard. + +It was claimed by the French Commission, and subsequently often stated +in discussions of the relation of the mosquito to yellow fever that the +mature _Aëdes calopus_ will bite only at night. If this were true it +would be of the greatest importance in measures to avoid the disease. +Unfortunately, the claim was illy founded and numerous workers have +clearly established that the exact converse is more nearly true, this +mosquito being pre-eminently a day species, feeding most actively in +early morning, about sunrise, and late in the afternoon. On cloudy days +it attacks at any time during the day. Thus there is peril in the +doctrine that infected regions may be visited with perfect safety during +the daytime and that measures to avoid the mosquito attack need be taken +only at night. + +[Illustration: 135_a_. Aëdes calopus. Pupa. After Howard.] + +Dr. Finlay maintained that the adult, even when starved, would not bite +when the temperature was below 23° C, but subsequent studies have shown +that this statement needs modification. The French Commission, working +at Rio de Janeiro, found that _Aëdes calopus_ would bite regularly at +temperatures between 22° and 25° and that the optimum temperature was +between 27° and 30° C, but their experiments led them to believe that it +would bite in nature at a temperature as low as 17° C. + +The yellow fever mosquito breeds in cisterns, water barrels, pitchers +and in the various water receptacles about the house. In our own +Southern States it very commonly breeds in the above-ground cisterns +which are in general use. Often the larvæ (fig. 135b) are found in +flower vases, or even in the little cups of water which are placed under +the legs of tables to prevent their being overrun by ants. They have +been repeatedly found breeding in the holy water font in churches. In +short, they breed in any collection of water in close proximity to the +dwellings or gathering places of man. + +The life cycle under favorable conditions is completed in from twelve to +fifteen days. These figures are of course very dependent upon the +temperature. The Army Commission in Cuba found that the cycle might be +completed in as brief a period as nine and a half days. Under less +favorable conditions it may be greatly lengthened. + +The adults are long lived. We have seen that during the experimental +work in Cuba specimens were kept in captivity for sixty-nine and +seventy-one days, respectively, and that they were proved to retain +their infectivity for at least fifty-seven days. Dr. Guiteras +subsequently kept an infected adult for one hundred and fifty-four days. + +Low temperatures have a very great effect not only on development, but +on the activity and even life of the adults. Long before the method of +transmission of yellow fever was discovered it was well known that the +epidemics were brought to a close by heavy frosts, and it is now known +that this is due to the killing of the mosquitoes which alone could +spread the disease. + +[Illustration: 135_b_. Aëdes calopus; larva. (×7). After Howard.] + +_Aëdes calopus_ has a very wide distribution since, as Howard says, +being a domestic mosquito, having a fairly long life in the adult stage, +and having the custom of hiding itself in the most ingenious ways, it is +particularly subject to carriage for long distances on board vessels, in +railway trains, even packed in baggage. In general, its permanent +distribution is from 40 degrees north latitude to 40 degrees south +latitude (Brumpt), in a belt extending around the world. In the United +States it breeds in most of our Southern States. + +Thus, as in the case of malaria, there are many places where the insect +carrier is abundant but where yellow fever does not occur. Such, for +instance, are Hawaii, Australia and Asia. An outbreak may occur at any +time that a patient suffering from the disease is allowed to enter and +become a source of infection for the mosquitoes. In this connection +various writers have called attention to the menace from the Panama +Canal. When it is completed, it will allow of direct passage from +regions where yellow fever is endemic and this will greatly increase the +possibility of its introduction into these places where it is now +unknown. The result, with a wholly non-immune population, would be +appalling. + +On the other hand, there are places wholly outside of the normal range +of _Aëdes calopus_ where the disease has raged. Such are New York, +Boston, and even Philadelphia, which have suffered notable epidemics. +These outbreaks have been due to the introduction of infected mosquitoes +during the heat of summer, when they have not only conveyed the disease +but have found conditions favorable for their multiplication. Or, +uninfected mosquitoes have been thus accidentally brought in and +developed in large numbers, needing then only the accidental +introduction of cases of the disease to start an epidemic. + +Methods of control of various diseases have been revolutionized by the +discovery that they were insect-borne, but in no other case has the +change been as radical or the results as spectacular as in the case of +yellow fever. The "shot-gun quarantine," the sufferings and horrors, the +hopelessness of fighting absolutely blindly have given way to an +efficient, clear-cut method of control, based upon the knowledge that +the disease is carried from man to man solely by the mosquito, _Aëdes +calopus_. The lines of defense and offense are essentially as follows: + +In the first place, when a case of yellow fever occurs, stringent +precautions must be adopted to prevent the infection of mosquitoes and +the escape of any already infected. This means that the patient must be +removed to a mosquito-proof room, or ward beyond reach of the insects, +and that the infected room must be thoroughly fumigated at once, to kill +the mosquitoes hiding within it. All cracks and openings should be +closed with strips of paper and fumigation with burning sulphur or +pyrethrum carefully carried out. + +It should be remembered that if the first case noted is that of a +resident rather than imported, it means that the mosquito carriers +became infected more than two weeks before the case was diagnosed, for +as we have seen, the germ must undergo a twelve-day period of +development within its insect host. Therefore a careful search must be +made for mild cases which, though unrecognized, may serve as foci for +the spread of the disease. + +In face of a threatened epidemic one of the most essential measures is +to educate the citizens and to gain their complete coöperation in the +fight along modern lines. This may be done through the schools, the +pulpit, places of amusement, newspapers and even bulletin boards. + +Emphasis should be placed on the necessity of both non-immunes and +immunes using mosquito curtains, and in all possible ways avoiding +exposure to the mosquitoes. + +Then the backbone of the fight must be the anti-mosquito measures. In +general, these involve screening and fumigating against adults, and +control of water supply, oiling, and drainage against the larvæ. The +region involved must be districted and a thorough survey undertaken to +locate breeding places, which must, if possible, be eradicated. If they +are necessary for water supplies, such as casks, or cisterns, they +should be carefully screened to prevent access of egg-laying adults. + +The practical results of anti-mosquito measures in the fight against +yellow fever are well illustrated by the classic examples of the work in +Havana, immediately following the discoveries of the Army Commission and +by the stamping out of the New Orleans epidemic in 1905. + +The opportunities for an immediate practical application of the theories +of the Army Commission in Havana were ideal. The city had always been a +hotbed of yellow fever and was the principal source from which the +disease was introduced year after year into our Southern States. It was +under martial law and with a military governor who was himself a +physician and thoroughly in sympathy with the views of the Commission, +the rigid enforcement of the necessary regulations was possible. The +story of the first campaign has been often told, but nowhere more +clearly than in Dr. Reed's own account, published in the _Journal of +Hygiene_ for 1902. + +Closer home was the demonstration of the efficacy of these measures in +controlling the yellow fever outbreak in New Orleans in 1905. During the +spring and early summer of the year the disease had, unperceived, gained +a firm foothold in that city and when, in early July the local Board of +Health took cognizance of its existence, it was estimated that there had +been in the neighborhood of one hundred cases. + +Conditions were not as favorable as they had been under martial law in +Havana for carrying on a rigid fight along anti-mosquito lines. The +densely populated city was unprepared, the public had to be educated, +and an efficient organization built up. The local authorities actively +began a general fight against the mosquito but in spite of their best +efforts the disease continued to spread. It was recognized that more +rigid organization was needed and on August 12th the United States +Public Health and Marine Hospital Service was put in absolute charge of +the fight. Up to this time there had been one hundred and forty-two +deaths from a total of nine hundred and thirteen cases and all of the +conditions seemed to threaten an outbreak to exceed the memorable one of +1878 when, as we have seen there were four thousand and forty-six +deaths. + +With the hearty coöperation of the citizens,--physicians and laymen +alike,--the fight was waged and long before frost or any near approach +thereto the disease was stamped out,--a thing unheard of in previous +epidemics. The total loss of life was four hundred and sixty--about 11 +per cent as great as that from the comparable epidemic of 1878. If the +disease had been promptly recognized and combated with the energy which +marked the fight later in the summer, the outbreak would have made +little headway and the great proportion of these lives would have been +saved. + + + + +CHAPTER IX + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA (Continued) + + +INSECTS AND TRYPANOSOMIASES + +By trypanosomiasis is meant a condition of animal parasitism, common to +man and the lower animals, in which trypanosomes, peculiar flagellate +protozoa, infest the blood. Depending upon the species, they may be +harmless, producing no appreciable ill-effect, or pathogenic, giving +rise to conditions of disease. A number of these are known to be +transferred by insects. + +In order that we may consider more fully the developmental stage of +these parasites within their insect host, it is necessary that we +describe briefly the structure of the blood-inhabiting stage. + +[Illustration: 136. Trypanosoma brucei. After Bruce.] + +The trypanosomes are elongated, usually pointed, flagellated protozoa +(fig. 136) in which the single flagellum, bent under the body, forms the +outer limit of a delicate undulating membrane. It arises near one end of +the organism from a minute centrosome-like body which is known as the +blepheroplast, and at the opposite end extends for a greater or less +distance as a free flagellum. Enclosing, or close beside the +blepheroplast is the small kinetonucleus. The principal nucleus, round +or oval in form, is situated near the center of the body. Asexual +reproductions occurs in this stage, by longitudinal fission, the nucleus +and the blepheroplast dividing independently of one another. From the +blepheroplast of one of the daughter cells a new flagellum is formed. + +Among the pathogenic species are to be found the causative organisms of +some of the most serious diseases of domestic animals and even of man. +It is probable that these pathogenic species secrete a specific poison. +The majority of them are tropical in distribution. + +Though we are concerned especially with the species which infest man, we +shall first consider two of the trypanosomes of lower animals, known +long before any of those of man had been found. + + +FLEAS AND LICE AS CARRIERS OF TRYPANOSOMA LEWISI.--_Trypanosoma lewisi_, +the first mammalian trypanosome known, is to be found in the blood of +wild rats. Like its host, it appears to be cosmopolitan in distribution, +having been reported from several localities in the United States, +Brazil, Argentine, England, Germany, France, Italy, Russia, Asia and +Africa. + +This species is usually regarded as non-pathogenic, but in experimental +work, especially with white rats, heavy infestations often result +fatally, and naturally infested specimens sometimes show evidence of +injury. Rats which have been infested exhibit at least temporary +immunity against new infection. + +_Trypanosoma lewisi_ is transmitted from rat to rat by fleas and by +lice. Rabinowitsch and Kempner (1899) first found that healthy rats +which were kept with infested rats, showed trypanosomes in their blood +after about two weeks. They found the trypanosomes in the alimentary +canal of fleas which had fed on the diseased rats. On teasing such fleas +in physiological salt solution and inoculating them into fresh rats they +were able to produce the infection. Finally, they showed that the fleas +which had fed upon infested rats were able to carry the parasites to +healthy rats. Corresponding experiments with lice were not successful. +Prowazek (1905) found in the rat louse (_Hæmatopinus spinulosus_) +organisms which he regarded as developmental stages of the _Trypanosoma +lewisi_. He believed that the sexual cycle was undergone in this insect. + +Nuttall (1908) readily transmitted the trypanosomes through the agency +of fleas, (_Ceratophyllus fasciatus_ and _Ctenopthalmus agyrtes_). He +believes that these insects are probably the chief transmitters of the +parasite. He was also able to transmit it from diseased to healthy rats +through the agency of the rat louse. He was unable to trace any +developmental stages in the louse and inclined to the opinion that +Prowazek was deceived by the presence of extraneous flagellates such as +are known to exist in a number of blood-sucking arthropods. + +Nuttall concludes that since three distinct kinds of blood-sucking +insects are capable of transmitting _Trypanosoma lewisi_ it appears +doubtful that this flagellate is a parasite of the invertebrate "host" +in the sense claimed by Prowazek and other investigators. + + +TSETSE-FLIES AND NAGANA--One of the greatest factors in retarding the +development of certain regions of Africa has been the presence of a +small fly, little larger than the common house-fly. This is the +tsetse-fly, _Glossina morsitans_ (fig. 165) renowned on account of the +supposed virulence of its bite for cattle, horses and other domestic +mammals. + +The technical characteristics of the tsetse-flies, or Glossinas, and +their several species, will be found in a later chapter. We need +emphasize only that they are blood-sucking Muscidæ and that, unlike the +mosquitoes, the sexes resemble each other closely in structure of the +mouth-parts, and in feeding habits. + +In 1894, Colonel David Bruce discovered that the fly was not in itself +poisonous but that the deadly effect of its bite was due to the fact +that it transmitted a highly pathogenic blood parasite, _Trypanosoma +brucei_. This trypanosome Bruce had discovered in the blood of South +African cattle suffering from a highly fatal disease known as "nagana". +On inoculating the blood of infected cattle into horses and dogs he +produced the disease and found the blood teeming with the causative +organism. In the course of his work he established beyond question that +the "nagana" and the tsetse-fly disease were identical. + +Tsetse-flies of the species _Glossina morsitans_, which fed upon +diseased animals, were found capable of giving rise to the disease in +healthy animals up to forty-eight hours after feeding. Wild tsetse-flies +taken from an infected region to a region where they did not normally +occur were able to transmit the disease to healthy animals. It was found +that many of the wild animals in the tsetse-fly regions harbored +_Trypanosoma brucei_ in their blood, though they showed no evidence of +disease. As in the case of natives of malarial districts, these animals +acted as reservoirs of the parasite. Non-immune animals subjected to the +attacks of the insect carrier, quickly succumbed to the disease. + +A question of prime importance is as to whether the insect serves as an +essential host of the pathogenic protozoan or whether it is a mere +mechanical carrier. Bruce inclined to the latter view. He was unable to +find living trypanosomes in the intestines or excrements of the fly or +to produce the disease on the many occasions when he injected the +excrement into healthy animals. Moreover, he had found that the +experimental flies were infective only during the first forty-eight +hours and that if wild flies were taken from the infected region, "kept +without food for three days and then fed on a healthy dog, they never +gave rise to the disease." + +Koch had early described what he regarded as sexual forms from the +intestine of the fly but it remained for Kleine (1909) to experimentally +demonstrate that a part of the life cycle of the parasite was undergone +in the fly. Working with _Glossina palpalis_, he found that for a period +of ten days or longer after feeding on an animal suffering from nagana +it was non-infective, but that then it became infective and was able to +transmit the disease for weeks thereafter. He discovered and described +developmental stages of the parasite within the intestine of the insect. +In other words, the tsetse-fly (in nature, _Glossina morsitans_), serves +as an essential host, within which an important part of the life cycle +of the parasite is undergone. These conclusions were quickly verified by +Bruce and numerous other workers and are no longer open to question. +Klein and Taute are even inclined to think that mechanical transmission +plays practically no rôle in nature, unless the fly is interrupted while +feeding and passes immediately to a new animal. + + +TSETSE-FLIES AND SLEEPING SICKNESS OF MAN--About the beginning of the +present century a hitherto little known disease of man began to attract +great attention on account of its ravages in Uganda and the region of +Victoria Nyanza in South Africa. It was slow, insiduous and absolutely +fatal, characterized in its later stages by dullness, apathy, and +finally absolute lethargy all day long, symptoms which gave it the name +of "sleeping sickness." + +It was soon found that the disease was not a new one but that it had +been known for over a hundred years on the west coast of Africa. Its +introduction into Central and East Africa and its rapid spread have been +attributed primarily to the development of the country, the formation of +new trade routes and the free mingling of native tribes formerly +isolated. It is estimated that in the first ten years of the present +century there were approximately two hundred thousand deaths from the +disease in the Uganda protectorate. In the British province Bugosa, on +the Victoria Nyanza there were thirty thousand deaths in the period from +1902-1905. + +While the disease is peculiarly African there are a number of instances +of its accidental introduction into temperate regions. Slaves suffering +from it were occasionally brought to America in the early part of the +last century and cases have sometimes been imported into England. In +none of the cases did the disease gain a foothold or spread at all to +other individuals. + +In 1902 Dutton described a trypanosome, _T. gambiense_, which he and +Forde had found the year before in the blood of a patient suffering from +a peculiar type of fever in Gambia. In 1902-1903 Castellani found the +same parasite in the cerebro-spinal fluid of sleeping-sickness patients +and definitely reported it as the causative organism of the disease. His +work soon found abundant confirmation, and it was discovered that the +sleeping sickness was but the ultimate phase of the fever discovered by +Dutton and Forde. + +When Castellani made known his discovery of the trypanosome of sleeping +sickness, Brumpt, in France, and Sambon, in England, independently +advanced the theory that the disease was transmitted by the tsetse-fly, +_Glossina palpalis_. This theory was based upon the geographical +distribution and epidemiology of the disease. Since then it has been +abundantly verified by experimental evidence. + +Fortunately for the elucidation of problems relating to the methods of +transfer of sleeping sickness, _Trypanosoma gambiense_ is pathogenic for +many species of animals. In monkeys it produces symptoms very similar to +those caused in man. Bruce early showed that _Glossina palpalis_ "fed on +healthy monkeys eight, twelve, twenty-four and forty-eight hours after +having fed on a native suffering from trypanosomiasis, invariably +transmitted the disease. After three days the flies failed to transmit +it." In his summary in Osler's Modern Medicine, he continues "But this +is not the only proof that these flies can carry the infective agent. On +the lake shore there was a large native population among whom we had +found about one-third to be harboring trypanosomes in their blood. The +tsetse-flies caught on this lake shore, brought to the laboratory in +cages, and placed straightway on healthy monkeys, gave them the disease +in every instance, and furnished a startling proof of the danger of +loitering along the lake shore among those infected flies." + +As in the case of nagana, Bruce and most of the earlier investigators +supposed the transmission of the sleeping sickness trypanosome by +_Glossina palpalis_ to be purely mechanical. The work of Kleine (1909) +clearly showed that for _Trypanosoma gambiense_ as well as for +_Trypanosoma brucei_ the fly served as an essential host. Indeed, Kleine +and many subsequent investigators are inclined to think that there is +practically no mechanical transmission of trypanosomes from animal to +animal by _Glossina_ in nature, and that the many successful experiments +of the earlier investigators were due to the fact that they used wild +flies which already harbored the transformed parasite rather than +directly inoculated it from the blood of the diseased experimental +animals. While the criticism is applicable to some of the work, this +extreme view is not fully justified by the evidence at hand. + +Kleine states (1912) that _Glossina palpalis_ can no longer be regarded +as the sole transmitter of sleeping sickness. Taute (1911) had shown +that under experimental conditions _Glossina morsitans_ was capable of +transferring the disease and Kleine calls attention to the fact that in +German East Africa, in the district of the Rovuma River, at least a +dozen cases of the disease have occurred recently, though only _Glossina +morsitans_ exists in the district. It appears, however, that these cases +are due to a different parasite, _Trypanosoma rhodesiense_. This +species, found especially in north-east Rhodesia and in Nyassaland, is +transferred by _Glossina morsitans_. + +Other workers maintain that the disease may be transmitted by various +blood-sucking flies, or even bugs and lice which attack man. Fülleborn +and Mayer (1907) have shown by conclusive experiments that _Aedes +(Stegomyia) calopus_ may transmit it from one animal to another if the +two bites immediately succeed each other. + +It is not possible that insects other than the tsetse-flies (and only +certain species of these), play an important rôle in the transmission of +the disease, else it would be much more wide-spread. Sambon (1908) +pointed out that the hypothesis that is spread by _Aedes calopus_ is +opposed by the fact that the disease never spread in the Antilles, +though frequently imported there by West African slaves. The same +observation would apply also to conditions in our own Southern States in +the early part of the past century. + +Since _Glossina palpalis_ acts as an essential host of the parasite and +the chief, if not the only, transmitter, the fight against sleeping +sickness, like that against malaria and yellow fever, becomes primarily +a problem in economic entomology. The minutest detail of the +life-history, biology, and habits of the fly, and of its parasites and +other natural enemies becomes of importance in attempts to eradicate the +disease. Here we can consider only the general features of the subject. + +_Glossina palpalis_ lives in limited areas, where the forest and +undergrowth is dense, along the lake shore or river banks. According to +Hodges, the natural range from shore is under thirty yards, though the +distance to which the flies may follow man greatly exceed this. + +It is a day feeder, a fact which may be taken advantage of in avoiding +exposure to its attacks. The young are brought forth alive and +full-grown, one every nine or ten days. Without feeding, they enter the +ground and under favorable conditions, complete their development in a +month or more. + +[Illustration: 137. Sleeping sickness concentration camp in German East +Africa. Report of German Commission.] + +Methods of control of the disease must look to the prevention of +infection of the flies, and to their avoidance and destruction. Along +the first line, much was hoped from temporary segregation of the sick in +regions where the fly was not found. On the assumption that the flies +acted as carriers only during the first two or three days, it was +supposed that even the "fly belts" would become safe within a few days +after the sick were removed. The problem was found to be a much more +difficult one when it was learned that after a given brief period the +fly again became infective and remained so for an indeterminate period. +Nevertheless, isolation of the sick is one of the most important +measures in preventing the spread of the disease into new districts. +Much, too, is being accomplished by moving native villages from the fly +belts. (c.f. fig. 137.) + +All measures to avoid the flies should be adopted. This means locating +and avoiding the fly belts as far as possible, careful screening of +houses, and protection of the body against bites. + +Clearing the jungle along the water courses for some yards beyond the +natural range of the fly has proved a very important measure. Castellani +recommends that the area be one hundred yards and around a village three +hundred yards at least. + +Detailed studies of the parasites and the natural enemies of the +tsetse-fly are being undertaken and may ultimately yield valuable +results. + + +SOUTH AMERICAN TRYPANOSOMIASIS--The tsetse-flies are distinctively +African in distribution and until recently there were no trypanosomes +known to infest man in America. In 1909 Dr. Chagas, of Rio de Janeiro +described a new species, _Trypanosoma cruzi_, pathogenic to man. + +_Trypanosoma cruzi_ is the causative organism of a disease common in +some regions of Brazil, where it is known as "opilacao." It is +especially to be met with in children and is characterized by extreme +anemia, wasting, and stunted development associated with fever, and +enlargement of the thyroid glands. The disease is transmitted by the +bites of several species of assassin-bugs, or Reduviidæ, notably by +_Conorhinus megistus_. The evolution of the parasite within the bug has +been studied especially by Chagas and by Brumpt. From the latter's text +we take the following summary. + +The adult trypanosomes, ingested by a _Conorhinus megistus_, of any +stage, first change into Crithidia-like forms and then those which +remain in the stomach become ovoid and non-motile. Brumpt found these +forms in immense numbers, in a _Conorhinus_ which had been infested +fourteen months before. The forms which pass into the intestine quickly +assume the _Crithidia_ form and continue to develop rapidly under this +form. Some weeks later they evolve into the trypanosome forms, +pathogenic for man. They then pass out with the excrement of the bug and +infect the vertebrate host as soon as they come in contact with any +mucous layer (buccal, ocular or rectal). More rarely they enter through +the epidermis. + +Brumpt showed that the development could take place in three species; +bed-bugs (_Cimex lectularius_, _C. hemipterus_) and in the tick +_Ornithodoros moubata_. The evolution proceeds in the first two species +of bed-bugs as rapidly as in Conorhinus, or even more rapidly, but they +remain infective for a much shorter time and hence Brumpt considers that +they play a much less important rôle in the spread of the disease. + +_Conorhinus megistus_, like related forms in our Southern States, very +commonly frequents houses and attacks man with avidity. Chagas states +that the bites are painless and do not leave any traces. They are +usually inflicted on the lips, or the cheeks and thus the buccal mucosa +of a sleeper may be soiled by the dejections of the insect and the bite +serving as a port of entry of the virus, remain unnoticed. + +The possibility of some of our North American Reduviidæ playing a +similar rôle in the transmission of disease should not be overlooked. + + +LEISHMANIOSES AND INSECTS--Closely related to the trypanosomes is a +group of intracellular parasites which have recently been grouped by +Ross under the genus _Leishmania_. Five species are known to affect man. +Three of these produce local skin infestations, but two of them, +_Leishmania donovani_ and _L. infantum_, produce serious and often fatal +systemic diseases. + +The first of these, that produced by _L. donovani_, is an exceedingly +virulent disease common in certain regions of India and China. It is +commonly known as "Kala-azar," or "dum-dum" fever, and more technically +as tropical leishmaniasis. Patton (1907) believes that the parasite is +transmitted by the bed-bug _Cimex hemipterus_, and has described a +developmental cycle similar to that which can be found in artificial +cultures. On the other hand, Donovan was unable to confirm Patton's work +and believes that the true intermediate host is a Reduviid bug, +_Conorhinus rubrofasciatus_. + +_Leishmania infantum_ is the cause of the so-called infantile splenic +leishmaniasis, occurring in northern Africa, Spain, Portugal, Italy, and +possibly other parts of Europe. The parasite occurs habitually in the +dog and is only accidentally transferred to children. Alvares and da +Silva, in Portugal (according to Brumpt, 1913) have found that the +excrement of a flea from a diseased dog contains flagellates, and they +suggest that the infection may be transmitted by the accidental +inoculation of this excrement by means of the proboscis of the flea, as +has been thought to occur in the case of the plague. To this Brumpt +objects that they and other workers who thought to trace the development +of _Leishmania infantum_ were apparently misled by the presence of a +harmless _Herpetomonas_ which infests dog fleas in all countries, even +where the leishmaniasis is unknown. + +Basile (1910 and 1911) however, carried on numerous experiments +indicating that the disease was transferred from children to dogs and +from dog to dog by the dog flea, and was able to find in the tissues of +the insects forms perfectly identical with those found in children and +in dogs suffering from leishmaniasis. He also found that _Pulex +irritans_ was capable of acting as the carrier. + +Of the cutaneous type of leishmaniasis, the best known is the so-called +"Oriental sore," an ulcerative disease of the skin which is epidemic in +many tropical and subtropical regions. The causative organism is +_Leishmania tropica_, which occurs in the diseased tissues as bodies +very similar to those found in the spleen in cases of Kala-azar. The +disease is readily inoculable and there is no doubt that it may be +transferred from the open sores to abraded surfaces of a healthy +individual by house-flies. It is also believed by a number of +investigators that it may be transferred and directly inoculated by +various blood-sucking insects. + + +TICKS AND DISEASES OF MAN AND ANIMALS + +We have seen that the way to the discoveries of the relations of +arthropods to disease was pointed out by the work of Leuckart and +Melnikoff on the life cycle of _Dipylidium_, and of Fedtschenko and +Manson on that of _Filaria_. They dealt with grosser forms, belonging to +well-recognized parasitic groups. + +This was long before the rôle of any insect as a carrier of pathogenic +micro-organisms had been established, and before the Protozoa were +generally regarded as of importance in the causation of disease. The +next important step was taken in 1889 when Smith and Kilbourne +conclusively showed that the so-called Texas fever of cattle, in the +United States, is due to an intracorpuscular blood parasite transmitted +exclusively by a tick. This discovery, antedating by eight years the +work on the relation of the mosquito to malaria, had a very great +influence on subsequent studies along these lines. + +While much of the recent work has dealt with the true insects, or +hexapods, it is now known that several of the most serious diseases of +animals, and at least two important diseases of man are tick borne. +These belong to the types known collectively as _babesioses_ (or +"_piroplasmoses_"), and _spirochætoses_. + +The term _babesiosis_ is applied to a disease of man or animals which is +caused by minute protozoan parasites of the genus _Babesia_, living in +the red blood corpuscles. These parasites have usually been given the +generic name _Piroplasma_ and hence the type of disease which they cause +is often referred to as "_piroplasmosis_." The best known illustration +is the disease known in this country as Texas fever of cattle. + + +CATTLE TICKS AND TEXAS FEVER--The cattle disease, which in the United +States is known as Texas fever, is a widely distributed, exceedingly +acute disease. In Australia it is known as _redwater fever_ and in +Europe as hæmoglobinuria, due to the fact that the urine of the diseased +animals is discolored by the breaking down of the red blood corpuscles +infested by the parasite. + +In their historical discussion, Smith and Kilbourne, point out that as +far back as 1796 it was noted that Southern cattle, in a state of +apparent health, might spread a fatal disease among Northern herds. As +observations accumulated, it was learned that this infection was carried +only during the warm season of the year and in the depth of winter +Southern cattle were harmless. Moreover, Southern cattle after remaining +for a short time in the North lost their power to transmit the disease, +and the same was true of cattle which had been driven for a considerable +distance. + +Very significant was the fact that the infection was not communicated +directly from the Southern to Northern cattle but that the ground over +which the former passed was infected by them, and that the infection was +transmitted thence to susceptible cattle _after a period of not less +than thirty days had elapsed_. + +Of course a disease as striking as this, and which caused such enormous +losses of cattle in the region invaded was fruitful in theories +concerning its causation. The most widespread was the belief that +pastures were infected by the saliva, urine, or manure of Southern +cattle. There were not wanting keen observers who suggested that the +disease was caused by ticks, but little weight was given to their view. + +Various workers had described bacteria which they had isolated from the +organs of the diseased animals, but their findings could not be +verified. In 1889, Smith and Kilbourne discovered a minute, pear-shaped +organism (fig. 138) in the red blood corpuscles of a cow which had +succumbed to Texas fever. On account of their shape they were given the +generic name _Pyrososma_ and because they were usually found two in a +corpuscle, the specific name, _bigeminum_. It is now generally accepted +that the parasite is the same which Babes had observed the year before +in Roumanian cattle suffering from hæmoglobinuria, and should be known +as _Babesia bovis_ (Babes). + +[Illustration: 138. Babesia bovis in blood corpuscles. After Calli.] + +[Illustration: 139. The cattle tick (Boophilus annulatus). (_a_) Female; +(_b_) male. After Comstock.] + +By a series of perfectly conclusive experiments carried on near +Washington, D.C., Smith and Kilbourne showed that this organism was +carried from Southern cattle to non-immune animals by the so-called +Southern cattle tick, _Boophilus annulatus_ (= _Margaropus annulatus_) +(fig. 139). + +Of fourteen head of native cattle placed in a field with tick-infested +Northern cattle all but two contracted the disease. This experiment was +repeated with similar results. Four head of native cattle kept in a plot +with three North Carolina cattle which had been carefully freed from +ticks remained healthy. A second experiment the same year gave similar +results. + +Still more conclusive was the experiment showing that fields which had +not been entered by Southern cattle but which had been infected by +mature ticks taken from such animals would produce Texas fever in native +cattle. On September 13, 1889, several thousand ticks collected from +cattle in North Carolina three and four days before, were scattered in a +small field near Washington. Three out of four native animals placed in +this field contracted the disease. The fourth animal was not examined as +to its blood but it showed no external symptoms of the disease. + +[Illustration: 140. Hyalomma ægypticum. After Nuttall and Warburton.] + +In these earlier experiments it was believed that the cattle tick acted +as a carrier of the disease between the Southern cattle and the _soil_ +of the Northern pastures. "It was believed that the tick obtained the +parasite from the blood of its host and in its dissolution on the +pasture a certain resistant spore form was set free which produced the +disease when taken in with the food." The feeding of one animal for some +time with grass from the most abundantly infected field, without any +appearance of the disease, made this hypothesis untenable. + +In the experimental work in 1890 the astonishing fact was brought out +that the disease was conveyed neither by infected ticks disintegrating +nor by their directly transferring the parasite, but that it was +conveyed by the young hatched from eggs of infected ticks. In other +words, the disease was hereditarily transferred to ticks of the second +generation and they alone were capable of conveying it. + +Thus was explained the fact that Texas fever did not appear immediately +along the route of Southern cattle being driven to Northern markets but +that after a certain definite period it manifested itself. It was +conveyed by the progeny of ticks which had dropped from the Southern +cattle and deposited their eggs on the ground. + +These results have been fully confirmed by workers in different parts of +the world,--notably by Koch, in Africa, and by Pound, in Australia. + +The disease is apparently transmitted by _Boophilus annulatus_ alone, in +the United States, but it, or an almost identical disease, is conveyed +by _Ixodes hexagonus_ in Norway, _Ixodes ricinus_ in Finland and France +and by the three species, _Boophilus decoloratus_, _Hyalomma ægypticum_ +(fig. 140 and 141), and _Hæmaphysalis punctata_ in Africa. + +[Illustration: 141. Hyalomma ægypticum. Capitulum of female; (_a_) +dorsal, (_b_) ventral aspect.] + +In spite of the detailed study which it has received, the life cycle of +_Babesia bovis_ has not been satisfactorily worked out. The asexual +reproduction in the blood of the vertebrate host has been described but +the cycle in the tick is practically unknown. + +More successful attempts have been made to work out the life cycle of a +related species, _Babesia canis_, which causes malignant jaundice in +dogs in Africa and parts of Southern Europe. In this instance, also, the +disease is transmitted by heredity to the ticks of the second +generation. Yet the larval, or "seed ticks," from an infected female are +not capable of conveying the disease, but only the nymphs and adults. +Still more complicated is the condition in the case of _Babesia ovis_ of +sheep, which Motas has shown can be conveyed solely by the adult, +sexually mature ticks of the second generation. + +In _Babesia canis_, Christopher (1907) observed developmental stages in +the tick. He found in the stomach of adult ticks, large motile +club-shaped bodies which he considered as oökinetes. These bodies pass +to the ovaries of the tick and enter the eggs where they become globular +in form and probably represent an oocyst. This breaks up into a number +of sporoblasts which enter the tissues of the developing tick and give +rise to numerous sporozoites, which collect in the salivary glands and +thence are transferred to the vertebrate host. A number of other species +of _Babesia_ are known to infest vertebrates and in all the cases where +the method has been worked out it has been found that the conveyal was +by ticks. We shall not consider the cases more fully here, as we are +concerned especially with the method of transfer of human diseases. + + +TICKS AND ROCKY MOUNTAIN SPOTTED FEVER OF MAN--Ever since 1873 there has +been known in Montana and Idaho a peculiar febrile disease of man, which +has gained the name of "Rocky Mountain spotted fever." Its onset is +marked by chills and fever which rapidly become acute. In about four to +seven days there appears a characteristic eruption on the wrists, ankles +or back, which quickly covers the body. + +McClintic (1912) states that the disease has now been reported from +practically all of the Rocky Mountain States, including Arizona, +California, Colorado, Idaho, Montana, Nevada, Oregon, Utah, Washington, +and Wyoming. "Although the disease is far more prevalent in Montana and +Idaho than in any of the other States, its spread has assumed such +proportions in the last decade as to call for the gravest consideration +on the part of both the state and national health authorities. In fact, +the disease has so spread from state to state that it has undoubtedly +become a very serious interstate problem demanding the institution of +measures for its control and suppression." + +A peculiar feature of the Rocky Mountain spotted fever is a marked +variation in its severity in different localities. In Montana, and +especially in the famous Bitter Root Valley, from 33 per cent to 75 per +cent of the cases result fatally. On the other hand, the fatality does +not exceed four per cent in Idaho. + +In 1902, Wilson and Chowning reported the causative organism of spotted +fever to be a blood parasite akin to the _Babesia_ of Texas fever, and +made the suggestion that the disease was tick-borne. The careful studies +of Stiles (1905) failed to confirm the supposed discovery of the +organism, and the disease is now generally classed as due to an +invisible virus. On the other hand, the accumulated evidence has fully +substantiated the hypothesis that it is tick-borne. + +According to Ricketts (1907) the experimental evidence in support of +this hypothesis was first afforded by Dr. L. P. McCalla and Dr. H. A. +Brereton, in 1905. These investigators transmitted the disease from man +to man in two experiments. "The tick was obtained 'from the chest of a +man very ill with spotted fever' and 'applied to the arm of a man who +had been in the hospital for two months and a half, and had lost both +feet from gangrene due to freezing.' On the eighth day the patient +became very ill and passed through a mild course of spotted fever, +leaving a characteristic eruption. The experiment was repeated by +placing the tick on a woman's leg and she likewise was infected with +spotted fever." + +The most detailed studies were those of the late Dr. H. T. Ricketts, and +it was he who clearly established the tick hypothesis. In the summer of +1906 he found that guinea pigs and monkeys are very susceptible to +spotted fever and can readily be infected by inoculation of blood from +patients suffering from the disease. This opened the way to experimental +work on tick transmission. A female tick was fed upon an infected guinea +pig for two days, removed and isolated for two days and then placed upon +a healthy guinea pig. After an incubation period of three and a half +days the experimental animal contracted a well-marked case of the +disease. + +A similar result was obtained at the same time by King, and later in the +season Ricketts proved that the male tick was also capable of +transmitting the disease. He found that there was a very intimate +relation of the virus to the tick and that the transmission must be +regarded as biological throughout. Ticks remained infective as long as +they lived and would feed for a period of several months. If they +acquired the disease in the larval or nymphal stage they retained it +during molting and were infective in the subsequent stages. In a few +cases the larvæ from an infected female were infective. + +The evidence indicated that the tick suffers from a relatively harmless, +generalized infection and the virus proliferates in its body. The +disease probably is transferred through the salivary secretion of the +tick since inoculation experiments show that the salivary glands of the +infected adult contain the virus. + +It is probable that in nature the reservoir of the virus of spotted +fever is some one or more of the native small animals. Infected ticks +have been found in nature, and as various wild animals are susceptible +to the disease, it is obvious that it may exist among them unnoticed. +Wilson and Chowning suggested that the ground squirrel plays the +principal rôle. + +Unfortunately, much confusion exists regarding the correct name of the +tick which normally conveys the disease. In the medical literature it is +usually referred to as _Dermacentor occidentalis_, but students of the +group now agree that it is specifically distinct. Banks has designated +it as _Dermacentor venustus_ and this name is used in the publications +of the Bureau of Entomology. On the other hand, Stiles maintains that +the common tick of the Bitter Root Valley, and the form which has been +collected by the authors who have worked on Rocky Mountain spotted fever +in that region, is separable from _D. venustus_, and he has described it +under the name of _Dermacentor andersoni_. + +Mayer (1911) has shown experimentally that spotted fever may be +transmitted by several different species of ticks, notably _Dermacentor +marginatus_, _Dermacentor variabilis_ and _Amblyomma americanum_. This +being the case, the question of the exact systematic status of the +species experimented upon in the Bitter Root Valley becomes less +important, for since _Dermacentor occidentalis_, _Dermacentor venustus_ +and _Dermacentor andersoni_ all readily attack man, it is probable that +either species would readily disseminate the disease if it should spread +into their range. + +Hunter and Bishop (1911) have emphasized the fact that in the eastern +and southern United States there occur several species which attack man, +and any one of which might transmit the disease from animal to animal +and from animal to man. The following species, they state, would +probably be of principal importance in the Southern and Eastern States: +the lone star tick (_Amblyomma americanum_); the American dog tick +(_Dermacentor variabilis_); and the gulf-coast tick (_Amblyomma +maculatum_). In the extreme southern portions of Texas, _Amblyomma +cajennense_, is a common pest of man. + +Since the evidence all indicates that Rocky Mountain spotted fever is +transmitted solely by the tick, and that some of the wild animals serve +as reservoirs of the virus, it is obvious that personal prophylaxis +consists in avoiding the ticks as fully as possible, and in quickly +removing those which do attack. General measures along the line of tick +eradication must be carried out if the disease is to be controlled. That +such measures are feasible has been shown by the work which has been +done in controlling the tick-borne Texas fever of cattle, and by such +work as has already been done against the spotted fever tick, which +occurs on both wild and domestic animals. Detailed consideration of +these measures is to be found in the publications of the Public Health +and Marine Hospital Service, and the Bureau of Entomology. Hunter and +Bishopp give the following summarized recommendations for control or +eradication measures in the Bitter Root Valley. + +(1) A campaign of education, whereby all the residents of the valley +will be made thoroughly familiar with the feasibility of the plan of +eradication, and with what it will mean in the development of the +valley. + +(2) The obtaining of legislation to make it possible to dip or oil all +live stock in the Bitter Root Valley. + +(3) The obtaining of an accurate census of the horses, cattle, sheep, +mules, and dogs in the valley. + +(4) The construction of ten or more dipping vats. + +(5) The providing of materials to be used in the dipping mixture. + +(6) The organization of a corps of workers to carry on the operations. + +(7) The systematic dipping of the horses, cattle, sheep, and dogs of the +valley on a definite weekly schedule from approximately March 10 to June +9. + +(8) The treatment by hand of the animals in localities remote from vats, +on the same schedule. + +They estimate that after three seasons' operations a very small annual +expenditure would provide against reinfestation of the valley by the +incoming of cattle from other places. + +Supplementary measures consist in the killing of wild mammals which may +harbor the tick; systematic burning of the brush and debris on the +mountain side; and in clearing, since the tick is seldom found on land +under cultivation. + + + + +CHAPTER X + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA [_Continued_] + + +ARTHROPODS AND SPIROCHÆTOSES OF MAN AND ANIMALS + +The term spirochætoses is applied to diseases of man or animals which +are due to protistan parasites belonging to the group of slender, spiral +organisms known as spirochætes. + +There has been much discussion concerning the relationship Of the +spirochætes. Formerly, they were regarded as bacteria closely related to +the forms grouped in the genus _Spirillum_. The results of the detailed +study which the spirochætes have received in recent years, have led most +of the workers to consider them as belonging to the protozoa. The merits +of the discussion we are not concerned with here, but rather with the +fact that a number of diseases caused by spirochætes are +arthropod-borne. The better known of these we shall discuss. + + +AFRICAN RELAPSING FEVER OF MAN--It has long been known to the natives of +Africa and to travelers in that country, that the bite of a certain +tick, _Ornithodoros moubata_, may be followed by severe or even fatal +fever of the relapsing type. Until recent years, it was supposed that +the effect was due to some special virulence of the tick, just as nagana +of cattle was attributed to the direct effect of the bite of the +tsetse-fly. The disease is commonly known as "tick-fever" or by the +various native names of the tick. + +In 1904, Ross and Milne, in Uganda, and Dutton and Todd on the Congo, +discovered that the cause of the disease is a spirochæte which is +transmitted by the tick. This organism has been designated by Novy and +Knapp as _Spirochæta duttoni_. + +_Ornithodoros moubata_ (fig. 142), the carrier of African relapsing +fever, or "tick-fever," is widely distributed in tropical Africa, and +occurs in great numbers in the huts of natives, in the dust, cracks and +crevices of the dirt floors, or the walls. It feeds voraciously on man +as well as upon birds and mammals. Like others of the _Argasidæ_, it +resembles the bed-bug in its habit of feeding primarily at night. Dutton +and Todd observed that the larval stage is undergone in the egg and that +the first free stage is that of the octopod nymph. + +[Illustration: 142. Ornithodoros moubata. (_a_) Anterior part of venter; +(_b_) second stage nymph; (_c_) capitulum; (_d_) dorsal and (_e_) +ventral aspect of female; (_f_) ventral aspect of nymph; (_g_) capitulum +of nymph. After Nuttall and Warburton.] + +The evidence that the fever is transmitted by this tick is conclusive. +Koch found that from five per cent to fifteen per cent, and in some +places, fifty per cent of the ticks captured, harbored the spirochæte. +The disease is readily transmitted to monkeys, rats, mice and other +animals and the earlier experiments along these lines have been +confirmed by many workers. + +Not only are the ticks which have fed on infected individuals capable of +conveying the disease to healthy animals but they transmit the causative +organism to their progeny. Thus Möllers (1907), working in Berlin, +repeatedly infected monkeys through the bites of nymphs which had been +bred in the laboratory from infected ticks. Still more astonishing was +his discovery that ticks of the third generation were infective. In +other words, if the progeny of infected ticks were fed throughout life +on healthy animals, and on maturity deposited eggs, the nymphs which +hatched from these eggs would still be capable of carrying the +infection. + +The developmental cycle of the spirochæte within the tick has not been +fully worked out, though the general conclusions of Leishman (1910) have +been supported by the recent works of Balfour (1911 and 1912), and +Hindle (1912), on the life cycle of spirochætes affecting fowls. + +_Spirochæta duttoni_ ingested by _Ornithodoros moubata_ apparently +disappear within a few days, but Leishman believed that in reality they +break up into minute granules which are to be found in the alimentary +canal, the salivary glands and the Malpighian tubes of the tick. These +granules, or "coccoid bodies," as Hindle calls them, are supposed to be +the form in which the spirochætes infect the new host. We shall see +later that Marchoux and Couvy (1913) dissent wholly from this +interpretation. + +According to Leishman, and Hindle, the coccoid bodies are not injected +into the vertebrate host with the saliva of the tick, as are the +sporozoites of malaria with that of the mosquito. Instead, they pass out +with the excrement and secondarily gain access to the wound inflicted by +the tick. + +Nuttall (1912) calls attention to the fact that the geographical +distribution of _Ornithodoros moubata_ is far wider than our present +records show for the distribution of the relapsing fever in man and that +there is every reason to fear the extension of the disease. Huts where +the ticks occur should be avoided and it should be remembered that in +infected localities there is special danger in sleeping on the ground. + + +EUROPEAN RELAPSING FEVER--There is widely distributed in Europe a type +of relapsing fever which is caused by _Spirochæta recurrentis_. It has +long been supposed that this disease is spread by the bed-bug and there +is some experimental evidence to show that it may be conveyed by these +insects. + +In 1897, Tictin found that he could infect monkeys by inoculating the +contents of bed-bugs which had fed upon a patient within forty-eight +hours. Nuttall, in 1907, in one experiment succeeded in transmitting +_Spirochæta recurrentis_ from mouse to mouse by bites of bed-bugs. The +bugs, thirty-five in number, were transferred at short intervals from +one mouse to another, not being allowed to take a full meal on the +first, or infected mouse. + +On the other hand, there is much clinical evidence to show that the +European relapsing fever like various other types of the disease is +transmitted from man to man by head and body lice (_Pediculus humanus_ +and _Pediculus corporis_). + +Interesting supplementary evidence is that of Bayon's observations +(1912), in Moscow. "Having visited the big municipal night hospitals at +Moscow I soon noticed that they were kept with such scrupulous +cleanliness, disinfected so lavishly, the beds of iron, the floor +cemented, that it was not possible for bed-bugs to thrive to any extent +on the premises. The people sleeping there were allowed, however, to +sleep in their own clothes. The introduction of these model homes had +not had any effect on the incidence of relapsing fever, for the places +were still hot-beds of the fever during winter. On the other hand, +though I changed my rooms several times, I found bugs in every +successive lodging, and I was told in Moscow, this can hardly be +avoided. Yet no foreigner, or Russian of the better class, ever catches +relapsing fever. To this may be added the fact that when I asked for +clothes-lice and promised to pay a kopec for two, the attendants from +the night hostel brought me next morning a small ounce bottle crammed +with _Pediculus capitis_ (= _P. humanus_), and _Pediculus vestimentorum_ +(= _P. corporis_) collected off the sleepers. If relapsing fever were +transmitted by bed-bugs, it would be much more disseminated than it is +at present in Moscow." + +Direct experimental evidence of the agency of lice in transmitting +relapsing fever is especially clear in the case of a type of the disease +prevalent in parts of North Africa. We shall consider this evidence +later. + + +OTHER TYPES OF RELAPSING FEVER OF MAN--In addition to the three types of +human relapsing fever already referred to, several others have been +distinguished and have been attributed to distinct species of +spirochætes. The various spirochætoses of man are: + +African, caused by _S. duttoni_; European, caused by _S. recurrentis_; +North African, caused by _S. berbera_; East African, caused by _S. +rossi_; East Indian, caused by _S. carteri_; North American, caused by +_S. novyi_; South American, caused by _S. duttoni_ (?). + +Nuttall (1912) in his valuable résumé of the subject, has emphasized +that "in view of the morphological similarity of the supposedly +different species of spirochætes and their individual variations in +virulence, we may well doubt if any of the 'species' are valid. As I +pointed out four years ago, the various specific names given to the +spirochætes causing relapsing fever in man may be used merely for +convenience _to distinguish strains or races_ of different origin. They +cannot be regarded as valid names, in the sense of scientific +nomenclature, for virulence and immunity reactions are not adequate +tests of specificity." + + +NORTH AFRICAN RELAPSING FEVER OF MAN--The type of human relapsing fever +to be met with in Algeria, Tunis, and Tripoli, is due to a _Spirochæta_ +to which does not differ morphologically from _Spirochæta duttoni_, but +which has been separated on biological grounds as _Spirochæta berberi_. + +Experimenting with this type of disease in Algeria, Sergent and Foly +(1910), twice succeeded in transmitting it from man to monkeys by +inoculation of crushed body lice and in two cases obtained infection of +human subjects who had received infected lice under their clothing and +who slept under coverings harboring many of the lice which had fed upon +a patient. Their results were negative with _Argas persicus_, _Cimex +lectularius_, _Musca domestica_, _Hæmatopinus spinulosus_ and +_Ceratophyllus fasciatus_. They found body lice associated with every +case of relapsing fever which they found in Algeria. + +Nicolle, Blaizot, and Conseil (1912) showed that the louse did not +transmit the parasite by its bite. Two or three hours after it has fed +on a patient, the spirochætes begin to break up and finally they +disappear, so that after a day, repeated examinations fail to reveal +them. They persist, nevertheless, in some unknown form, for if the +observations are continued they reappear in eight to twelve days. These +new forms are virulent, for a monkey was infected by inoculating a +single crushed louse which had fed on infected blood fifteen days +before. + +Natural infection is indirect. Those attacked by the insect scratch, and +in this act they excoriate the skin, crush the lice and contaminate +their fingers. The least abrasion of the skin serves for the entrance of +the spirochætes. Even the contact of the soiled fingers on the various +mucosa, such as the conjunctive of the eye, is sufficient. + +As in the case of _Spirochæta duttoni_, the organism is transmitted +hereditarily in the arthropod vector. The progeny of lice which have fed +on infected blood may themselves be infective. + + +SPIROCHÆTOSIS OF FOWLS--One of the best known of the spirochætes +transmitted by arthropods is _Spirochæta gallinarum_, the cause of a +very fatal disease of domestic fowls in widely separated regions of the +world. According to Nuttall, it occurs in Southeastern Europe, Asia, +Africa, South America and Australia. + +In 1903, Marchoux and Salimbeni, working in Brazil, made the first +detailed study of the disease, and showed that the causative organism is +transmitted from fowl to fowl by the tick _Argas persicus_. They found +that the ticks remained infective for at least five months. Specimens +which had fed upon diseased birds in Brazil were sent to Nuttall and he +promptly confirmed the experiments. Since that date many investigators, +notably Balfour and Hindle, have contributed to the elucidating of the +life-cycle of the parasite. Since it has been worked out more fully than +has that of any of the human spirochætes, we present Hindle's diagram +(fig. 143) and quote the brief summary from his preliminary paper +(1911_b_). + +"Commencing with the ordinary parasite in the blood of the fowl, the +spirochæte grows until it reaches a certain length (16-19µ) and then +divides by transverse division. This process is repeated, and is +probably the only method of multiplication of the parasite within the +blood. When the spirochætes disappear from the circulation, some of them +break up into the coccoid bodies which, however, do not usually develop +in the fowl. When the spirochætes are ingested by _Argas persicus_, some +of them pass through the gut wall into the coelomic fluid. From this +medium they bore their way into the cells of the various organs of the +tick and there break up into a number of coccoid bodies. These +intracellular forms multiply by ordinary fission in the cells of the +Malpighian tubules and gonads. Some of the coccoid bodies are formed in +the lumen of the gut and Malpighian tubules. The result is that some of +the coccoid bodies may be present in the Malpighian secretion and +excrement of an infected tick and when mixed with the coxal fluid may +gain entry into another fowl by the open wound caused by the tick's +bite. They then elongate and redevelop into ordinary spirochætes in the +blood of the fowl, and the cycle may be repeated." + +[Illustration: 143. Spirochæta gallinarum. After Hindle.] + +Hindle's account is clear cut and circumstantial, and is quite in line +with the work of Balfour, and of Leishman. Radically different is the +interpretation of Marchoux and Couvy (1913). These investigators +maintain that the granules localized in the Malpighian tubules in the +larvæ and, in the adult, also in the ovules and the genital ducts of the +male and female, are not derived from spirochætes but that they exist +normally in many acariens. They interpret the supposed disassociation +of the spirochæte into granules as simply the first phase, not of a +process of multiplication, but of a degeneration ending in the death of +the parasite. The fragmented chromatin has lost its affinity for stains, +remaining always paler than that of the normal spirochætes. On the other +hand, the granules of Leishman stain energetically with all the basic +stains. + +Further, according to Marchoux and Couvy, infection takes place without +the emission of the coxal fluid and indeed, soiling of the host by the +coxal fluid diluting the excrement is exceptional. All of the organs of +the Argasid are invaded by the parasites, but they pass from the +coelom into the acini of the salivary glands and collect in its +efferent canal. The saliva serves as the vehicle of infection. + +Thus, the question of the life cycle of _Spirochæta gallinarum_, and of +spirochætes in general, is an open one. + +It should be noted that _Argas persicus_, the carrier of _Spirochæta +gallinarum_, is a common pest of poultry in the southwestern United +States. Though the disease has not been reported from this country, +conditions are such that if accidentally introduced, it might do great +damage. + + +OTHER SPIROCHÆTE DISEASES OF ANIMALS--About a score of other blood +inhabiting spirochætes have been reported as occurring in mammals, but +little is known concerning their life-histories. One of the most +important is _Spirochæta theileri_ which produces a spirochætosis of +cattle in the Transvaal. Theiler has determined that it is transmitted +by an Ixodid tick, _Margaropus decoloratus_. + + +TYPHUS FEVER AND PEDICULIDÆ + +Typhus is an acute, and continued fever, formerly epidemically prevalent +in camps, hospitals, jails, and similar places where persons were +crowded together under insanitary conditions. It is accompanied by a +characteristic rash, which gives the disease the common name of +"spotted" or "lenticular" fever. The causative organism is unknown. + +Typhus fever has not generally been supposed to occur in the United +States, but there have been a few outbreaks and sporadic cases +recognized. According to Anderson and Goldberger (1912_a_), it has been +a subject of speculation among health authorities why, in spite of the +arrival of occasional cases in this country and of many persons from +endemic foci of the disease, typhus fever apparently does not gain a +foothold in the United States. These same workers showed that the +so-called Brill's disease, studied especially in New York City, is +identical with the typhus fever of Mexico and of Europe. + +The conditions under which the disease occurs and under which it spreads +most rapidly are such as to suggest that it is carried by some parasitic +insect. On epidemiological grounds the insects most open to suspicion +are the lice, bed-bugs and fleas. + +In 1909, Nicolle, Comte and Conseil, succeeded in transmitting typhus +fever from infected to healthy monkeys by means of the body louse +(_Pediculus corporis_). Independently of this work, Anderson and +Goldberger had undertaken work along this line in Mexico, and in 1910 +reported two attempts to transmit the disease to monkeys by means of +body lice. The first experiment resulted negatively, but the second +resulted in a slight rise in temperature, and in view of later results +it seems that this was due to infection with typhus. + +Shortly after, Ricketts and Wilder (1910) succeeded in transmitting the +disease to the monkey by the bite of body lice in two experiments, the +lice in one instance deriving their infection from a man and in another +from the monkey. Another monkey was infected by typhus through the +introduction of the feces and abdominal contents of infested lice into +small incisions. Experiments with fleas and bed-bugs resulted +negatively. + +Subsequently, Goldberger and Anderson (1912_b_) indicated that the head +louse (_Pediculus humanus_) as well, may become infected with typhus. In +an attempt to transmit typhus fever (Mexican virus) from man to monkey +by subcutaneous injection of a saline suspension of crushed head lice, +the monkeys developed a typical febrile reaction with subsequent +resistance to an inoculation of virulent typhus (Mexican) blood. In one +of the three experiments to transmit the disease from man to monkey by +means of the bite of the head louse, the animal bitten by the presumably +infected head lice proved resistant to two successive immunity tests +with virulent typhus blood. + +In 1910, Ricketts and Wilder reported an experiment undertaken with a +view to determining whether the young of infected lice were themselves +infected. Young lice were reared to maturity on the bodies of typhus +patients, so that if the eggs were susceptible to infection at any stage +of their development, they would have every opportunity of being +infected within the ovary. The eggs of these infected lice were +obtained, they were incubated, and the young lice of the second +generation were placed on a normal rhesus monkey. The experimenters were +unable to keep the monkey under very close observation during the +following three or four weeks, but from the fact that he proved +resistant to a subsequent immunity test they concluded that he probably +owed this immunity to infection by these lice of the second generation. + +Anderson and Goldberger (1912_b_) object that due consideration was not +given to the possibility of a variable susceptibility of the monkey to +typhus. Their similar experiment was "frankly negative." + +Prophylaxis against typhus fever is, therefore, primarily a question of +vermin extermination. A brief article by Dr. Goldberger (1914) so +clearly shows the practical application of his work and that of the +other investigators of the subject, that we abstract from it the +following account: + +"In general terms it may be stated that association with a case of +typhus fever in the absence of the transmitting insect is no more +dangerous than is association with a case of yellow fever in the absence +of the yellow fever mosquito. Danger threatens only when the insect +appears on the scene." + +"We may say, therefore, that to prevent infection of the individual it +is necessary for him only to avoid being bitten by the louse. In theory +this may readily be done, for we know that the body louse infests and +attaches itself almost entirely to the body linen, and that boiling +kills this insect and its eggs. Individual prophylaxis is based +essentially, therefore, on the avoidance of contact with individuals +likely to harbor lice. Practically, however, this is not always as easy +as it may seem, especially under the conditions of such intimate +association as is imposed by urban life. Particularly is this the case +in places such as some of the large Mexican cities, where a large +proportion of the population harbors this vermin. Under such +circumstances it is well to avoid crowds or crowded places, such as +public markets, crowded streets, or public assemblies at which the +'peon' gathers." + +"Community prophylaxis efficiently and intelligently carried out is, +from a certain point of view, probably easier and more effective in +protecting the individual than is the individual's own effort to guard +himself. Typhus emphasizes, perhaps better than any other disease, the +fact that fundamentally, sanitation and health are economic problems. In +proportion as the economic condition of the masses has improved--that +is, in proportion as they could afford to keep clean--the notorious +filth disease has decreased or disappeared. In localities where it still +prevails, its further reduction or complete eradication waits on a +further improvement in, or extension of, the improved economic status of +those afflicted. Economic evolution is very slow process, and, while +doing what we can to hasten it, we must take such precautions as +existing conditions permit, looking to a reduction in or complete +eradication of the disease." + +"When possible, public bath houses and public wash houses, where the +poor may bathe and do their washings at a minimum or without cost, +should be provided. Similar provision should be made in military and +construction camps. Troops in the field should be given the opportunity +as frequently as possible to wash and _scald_ or _boil_ their body +linen." + +"Lodging houses, cheap boarding houses, night shelters, hospitals, jails +and prisons, are important factors in the spread and frequently +constitute foci of the disease. They should receive rigid sanitary +supervision, including the enforcement of measures to free all inmates +of such institutions of lice on admission." + +"So far as individual foci of the disease are concerned these should be +dealt with by segregating and keeping under observation all exposed +individuals for 14 days--the period of incubation--from the last +exposure, by disinfecting (boiling or steaming) the suspected bedding, +body linen, and clothes, for the destruction of any possible vermin that +they may harbor, and by fumigating (with sulphur) the quarters that they +may have occupied." + +"It will be noted that nothing has been said as to the disposition of +the patient. So far as the patient is concerned, he should be removed to +'clean' surroundings, making sure that he does not take with him any +vermin. This may be done by bathing, treating the hair with an +insecticide (coal oil, tincture of larkspur), and a complete change of +body linen. Aside from this, the patient may be treated or cared for in +a general hospital ward or in a private house, provided the sanitary +officer is satisfied that the new surroundings to which the patient has +been removed are 'clean,' that is, free from vermin. Indeed, it is +reasonably safe to permit a 'clean' patient to remain in his own home if +this is 'clean,' for, as has already been emphasized, there can be no +spread in the absence of lice. This is a common experience in native +families of the better class and of Europeans in Mexico City." + +"Similarly the sulphur fumigation above prescribed may be dispensed with +as unnecessary in this class of cases." + + + + +CHAPTER XI + +SOME POSSIBLE, BUT IMPERFECTLY ESTABLISHED CASES OF ARTHROPOD +TRANSMISSION OF DISEASE + + +INFANTILE PARALYSIS OR ACUTE ANTERIOR POLIOMYELITIS + +The disease usually known in this country as infantile paralysis or, +more technically, as acute anterior poliomyelitis, is one which has +aroused much attention in recent years. + +The causative organism of infantile paralysis is unknown, but it has +been demonstrated that it belongs to the group of filterable viruses. It +gives rise to a general infection, producing characteristic lesions in +the central nervous system. The result of the injury to the motor nerves +is a more or less complete paralysis of the corresponding muscle. This +usually manifests itself in the legs and arms. The fatal cases are +usually the result of paralysis of the muscles of respiration. Of the +non-fatal cases about 60 per cent remain permanently crippled in varying +degrees. + +Though long known, it was not until about 1890 that it was emphasized +that the disease occurs in epidemic form. At this time Medin reported +his observations on an epidemic of forty-three cases which occurred in +and around Stockholm in 1887. Since then, according to Frost (1911), +epidemics have been observed with increasing frequency in various parts +of the world. The largest recorded epidemics have been those in Vermont, +1894, 126 cases; Norway and Sweden, 1905, about 1,500 cases; New York +City, 1907, about 2,500 cases. Since 1907 many epidemics have been +reported in the United States, and especially in the Northern States +east of the Dakotas. In 1912 there were over 300 cases of the disease in +Buffalo, N. Y., with a mortality of somewhat over 11 per cent. + +In view of the sudden prominence and the alarming spread of infantile +paralysis, there have been many attempts to determine the cause, and the +manner in which the disease spreads and develops in epidemic form. In +the course of these studies, the question of possible transmission by +insects was naturally suggested. + +C. W. Howard and Clark (1912) presented the results of studies in this +phase of the subject. They dealt especially with the house-fly, bedbug, +head, and body lice, and mosquitoes. It was found that the house-fly +(_Musca domestica_) can carry the virus of poliomyelitis in an active +state for several days upon the surface of the body and for several +hours within the gastro-intestinal tract. Mosquitoes and lice were found +not to take up or maintain the virus. On the other hand, the bedbug +(_Cimex lectularius_) was found to take the virus from the infected +monkeys and to maintain it in a living state within the body for a +period of seven days. This was demonstrated by grinding up in salt +solution, insects which had fed on poliomyelitic animals and injecting +the filtrate into a healthy monkey. The experimenters doubted that the +bedbug is a carrier of the virus in nature. + +Earlier in the same year, Brues and Sheppard published the results of an +intensive epidemiological study of the outbreak of 1911, in +Massachusetts. Special attention had been paid to the possibility of +insect transfer and the following conclusion was reached: + +"Field work during the past summer together with a consideration of the +epidemiology of the disease so far as known, points strongly toward +biting flies as possible carriers of the virus. It seems probable that +the common stable-fly (_Stomoxys calcitrans_ L.) may be responsible to a +certain extent for the spread of acute epidemic poliomyelitis, possibly +aided by other biting flies, such as _Tabanus lineola_. No facts which +disprove such a hypothesis have as yet been adduced, and experiments +based upon it are now in progress." + +As stated by Brues (1913), especial suspicion fell upon the stable-fly +because: + +1. The blood-sucking habits of the adult fly suit it for the transfer of +virus present in the blood. + +2. The seasonal abundance of the fly is very closely correlated with the +incidence of the disease, rising rapidly during the summer and reaching +a maximum in July and August, then slowly declining in September and +October. + +3. The geographical distribution of the fly is, so far as can be +ascertained, wider, or at least co-extensive with that of poliomyelitis. + +4. _Stomoxys_ is distinctly more abundant under rural conditions, than +in cities and thickly populated areas. + +5. While the disease spreads over districts quickly and in a rather +erratic way, it often appears to follow along lines of travel, and it is +known that _Stomoxys_ flies will often follow horses for long distances +along highways. + +6. In a surprisingly large number of cases, it appeared probable that +the children affected had been in the habit of frequenting places where +_Stomoxys_ is particularly abundant, i.e., about stables, barnyards, +etc. + +The experiments referred to were carried on during the summer of 1912 +and in September Dr. Rosenau announced that the disease was transferred +by the bite of the stable-fly. + +A monkey infected by inoculation was exposed to the bites of upwards of +a thousand of the _Stomoxys_ flies daily, by stretching it at full +length and rolling it in a piece of chicken wire, and then placing it on +the floor of the cage in which the flies were confined. The flies fed +freely from the first, as well as later, after paralysis had set in. +Alternating with the inoculated monkey, healthy monkeys were similarly +introduced into the cage at intervals. New monkeys were inoculated to +keep a supply of such infected animals and additional healthy ones were +exposed to the flies, which fed willingly and in considerable numbers on +each occasion. "Thus the flies were given every opportunity to obtain +infection from the monkeys, since the animals were bitten during +practically every stage of the disease from the time of the inoculation +of the virus till their death following the appearance of paralysis. By +the same arrangement the healthy monkeys were likely to be bitten by +flies that had previously fed during the various stages of the disease +on the infected monkeys. The flies had meanwhile enjoyed the opportunity +of incubating the virus for periods varying from the day or two which +usually elapses between consecutive feedings, to the two or three-week +period for which at least some (although a very small percentage) of the +flies lived in the cage." + +"In all, twelve apparently healthy monkeys of a small Japan species were +exposed to the flies in the manner described for the infected monkeys. +Some were placed in the cage only once or twice and others a number of +times after varying intervals. These exposures usually lasted for about +half an hour, but were sometimes more protracted. No results were +apparent until two or three weeks after the experiment was well under +way, and then in rather rapid succession six of the animals developed +symptoms of poliomyelitis. In three, the disease appeared in a virulent +form, resulting in death, while the other three experienced transient +tremblings, diarrhoea, partial paralysis and recovery."--Brues, 1913. + +Very soon after the announcement of the results of experiments by +Rosenau and Brues, they were apparently conclusively confirmed by +Anderson and Frost (1912), who repeated the experiments, at Washington. +They announced that through the bites of the _Stomoxys_ flies that had +previously fed on infected monkeys, they had succeeded in experimentally +infecting three healthy monkeys. + +The results of these experiments gained much publicity and in spite of +the conservative manner in which they had been announced, it was widely +proclaimed that infantile paralysis was conveyed in nature by the +stable-fly and by it alone. + +Serious doubt was cast on this theory by the results of further +experiments by Anderson and Frost, reported in May of 1913. Contrary to +the expectations justified by their first experience, the results of all +the later, and more extended, experiments were wholly negative. Not once +were these investigators again able to transmit the infection of +poliomyelitis through _Stomoxys_. They concluded that it was extremely +doubtful that the insect was an important factor in the natural +transmission of the disease, not only because of their series of +negative results, "but also because recent experiments have afforded +additional evidence of the direct transmissibility or contagiousness of +poliomyelitis, and because epidemiological studies appear to us to +indicate that the disease is more likely transmitted largely through +passive human virus carriers." + +Soon after this, Kling and Levaditi (1913) published their detailed +studies on acute anterior poliomyelitis. They considered that the +experiments of Flexner and Clark (and Howard and Clark), who fed +house-flies on emulsion of infected spinal cord, were under conditions +so different from what could occur in nature that one could not draw +precise conclusions from them regarding the epidemiology of the disease. +They cited the experiments of Josefson (1912), as being under more +reasonable conditions. He sought to determine whether the inoculation of +monkeys with flies caught in the wards of the Hospital for Contagious +Diseases at Stockholm, where they had been in contact with cases of +poliomyelitis, would produce the disease. The results were completely +negative. + +Kling and Lavaditi made four attempts of this kind. The flies were +collected in places where poliomyelitics had dwelt, three, four and +twenty-four after the beginning of the disease in the family and one, +three, and fifteen days after the patient had left the house. These +insects were for the greater part living and had certainly been in +contact with the infected person. In addition, flies were used which had +been caught in the wards of the Hospital for Contagious Diseases at +Söderkoping, when numbers of poliomyelitics were confined there. +Finally, to make the conditions as favorable as possible, the emulsions +prepared from these flies were injected without previous filtering, +since filtration often causes a weakening of the virus. In spite of +these precautions, all their results were negative, none of the +inoculated animals having contracted poliomyelitis. They also +experimented with bedbugs which had fed upon infected patients at +various stages of the disease, but the results in these cases also were +wholly negative. + +Kling and Levaditi considered at length the possibility of transmission +of the disease by _Stomoxys_. As a result of their epidemiological +studies, they found that infantile paralysis continued to spread in +epidemic form in the dead of winter, when these flies were very rare and +torpid, or were even completely absent. Numerous cases developed in the +northern part of Sweden late in October and November, long after snow +had fallen. On account of the rarity of the Stomoxys flies during the +period of their investigations they were unable to conduct satisfactory +experiments. In one instance, during a severe epidemic, they found a +number of the flies in a stable near a house inhabited by an infected +family, though none was found in the house itself. These flies were used +in preparing an emulsion which, after filtering, was injected into the +peritoneal cavity of a monkey. The result was wholly negative. + +As for the earlier experiments, Kling and Levaditi believe if the flies +were responsible for the transmission of the disease in the cases +reported by Rosenau and Brues, and the first experiments of Anderson and +Frost, it was because the virus of infantile paralysis is eliminated +with the nasal secretions of paralyzed monkeys and the flies, becoming +contaminated, had merely acted as accidental carriers. + +Still further evidence against the hypothesis of the transmission of +acute anterior poliomyelitis by _Stomoxys calcitrans_ was brought +forward by Sawyer and Herms (1913). Special precautions were used to +prevent the transference of saliva or other possibly infectious material +from the surface of one monkey to that of another, and to avoid the +possibility of complicating the experiments by introducing other +pathogenic organisms from wild flies, only laboratory-bred flies were +used. In a series of seven carefully performed experiments, in which the +conditions were varied, Sawyer and Herms were unable to transmit +poliomyelitis from monkey to monkey through the agency of _Stomoxys_, or +to obtain any indication that the fly is the usual agent for spreading +the disease in nature. + +The evidence at hand to date indicates that acute anterior +poliomyelitis, or infantile paralysis, is transmitted by contact with +infected persons. Under certain conditions insects may be agents in +spreading the disease, but their rôle is a subordinate one. + + +Pellagra + +PELLAGRA is an endemic and epidemic disease characterized by a peculiar +eruption or erythema of the skin (figs 144 and 145), digestive +disturbances and nervous trouble. Insanity is a common result, rather +than a precursor of the disease. The manifestations of pellagra are +periodic and its duration indeterminate. + +[Illustration: 144. Pellagrous eruption on the face. After Watson.] + +The disease is one the very name of which was almost unknown in the +United States until within the past decade. It has usually been regarded +as tropical, though it occurs commonly in Italy and in various parts of +Europe. Now it is known that it not only occurs quite generally in the +United States but that it is spreading. Lavinder (1911) says that "There +are certainly many thousand cases of the disease in this country, and +the present situation must be looked upon with grave concern." + +It is not within the scope of this book to undertake a general +discussion of pellagra. The subject is of such importance to every +medical man that we cannot do better than refer to Lavinder's valuable +précis. We can only touch briefly upon the entomological phases of the +problems presented. + +The most commonly accepted theories regarding the etiology of the +disease have attributed it to the use of Indian corn as an article of +diet. This supposed relationship was explained either on the basis of, +(a) insufficiency of nutriment and inappropriateness of corn as a prime +article of food; (b) toxicity of corn or, (c) parasitism of certain +organisms--fungi or bacteria--ingested with either sound or deteriorated +corn. + +In 1905, Sambon proposed the theory of the protozoal origin of pellagra +and in 1910 he marshalled an imposing array of objections to the theory +that there existed any relationship between corn and the disease. He +presented clear evidence that pellagra existed in Europe before the +introduction of Indian corn from America, as an article of diet, and +that its spread was not _pari passu_ with that of the use of corn. Cases +were found in which the patients had apparently never used corn, though +that is obviously difficult to establish. He showed that preventive +measures based on the theory had been a failure. Finally, he believed +that the recurrence of symptoms of the disease for successive springs, +in patients who abstained absolutely from the use of corn, militated +against the theory. + +[Illustration: 145. Pellagrous eruption on the hand. After Watson.] + +On the other hand, Sambon believed that the periodicity of the symptoms, +peculiarities of distribution and seasonal incidence, and analogies of +the symptoms to those of other parasitic diseases indicated that +pellagra was of protozoal origin, and that it was insect-borne. + +The insect carriers, he believed to be one or more species of Simuliidæ, +or black-flies. In support of this he stated that _Simulium_ appears to +effect the same topographical conditions as pellagra, that in its imago +stage it seems to present the same seasonal incidence, that it has a +wide geographical distribution which seems to cover that of pellagra, +and that species of the genus are known to cause severe epizootics. +Concluding from his studies in Italy, that pellagra was limited almost +wholly to agricultural laborers, he pointed out that the Simulium flies +are found only in rural districts, and as a rule do not enter towns, +villages, or houses. + +[Illustration: 146. A favorite breeding place of Simulium. Ithaca, N. +Y.] + +When Sambon's detailed report was published in 1910, his theory was +seized upon everywhere by workers who were anxious to test it and who, +in most cases, were favorably disposed towards it because of the +wonderful progress which had been made in the understanding of other +insect-borne diseases. In this country, the entomological aspects of the +subject have been dealt with especially by Forbes (1912), and by King +and Jennings, under the direction of W. D. Hunter, of the Bureau of +Entomology, and in coöperation with the Thompson-McFadden Pellagra +Commission of the Department of Tropical Medicine of the New York +Post-Graduate Medical School. An important series of experiments with +monkeys has been undertaken by S. J. Hunter, of Kansas, but +unfortunately we have as yet no satisfactory evidence that these animals +are susceptible to the disease--a fact which renders the whole problem +difficult. + +The accumulated evidence is increasingly opposed to Sambon's hypothesis +of the transmission of pellagra by _Simulium_. This has been so clearly +manifested in the work of the Thompson-McFadden Commission that we quote +here from the report by Jennings (1914): + +"Our studies in 1912 convinced us that there was little evidence to +support the incrimination of any species of _Simulium_ in South Carolina +in the transmission of pellagra. Reviewing the group as a whole, we find +that its species are essentially "wild" and lack those habits of +intimate association with man which would be expected in the vector of +such a disease as pellagra. Although these flies are excessively +abundant in some parts of their range and are moderately so in +Spartanburg County, man is merely an incidental host, and no disposition +whatever to seek him out or to invade his domicile seems to be +manifested. Critically considered, it is nearer the fact that usually +man is attacked only when he invades their habitat." + +"As our knowledge of pellagra accumulates, it is more and more evident +that its origin is in some way closely associated with the domicile. The +possibility that an insect whose association with man and his immediate +environment is, at the best, casual and desultory, can be active in the +causation of the disease becomes increasingly remote." + +"Our knowledge of the biting habits of _Simulium_ is not complete, but +it is evident, as regards American species at least, that these are +sometimes not constant for the same species in different localities. +Certain species will bite man freely when opportunity offers, while +others have never been known to attack him. To assume that the proximity +of a _Simulium_-breeding stream necessarily implies that persons in its +vicinity must be attacked and bitten is highly fallacious. In +Spartanburg County attacks by _Simulium_ seems to be confined to the +immediate vicinity of the breeding-places. Our records and observations, +exceedingly few in number, refer almost exclusively to such locations. +Statements regarding such attacks, secured with much care and +discrimination from a large number of persons, including many +pellagrins, indicate conclusively that these insects are seldom a pest +of man in this county. A certain number of the persons questioned were +familiar with the gnats in other localities, but the majority were +seemingly ignorant of the existence of such flies with biting habits. +This is especially striking, in view of the fact that the average +distance of streams from the homes of the pellagra cases studied was +about 200 yards, many being at a distance of less than 200 yards, and +that 78 per cent of these streams were found to be infested by larval +_Simulium_. Such ignorance in a large number of persons cannot be +overlooked and indicates strongly that our belief in the negligible +character of local attacks by _Simulium_ is well founded." + +"In localities infested by 'sand-flies,' mosquitoes, etc., these pests +are always well known and the ignorance described above is very +significant." + +"Such positive reports as we received nearly always referred to bites +received in the open, along streams, etc., and observations made of +their attack were of those on field laborers in similar situations. +Males engaged in agricultural pursuits are almost exempt from pellagra +in Spartanburg County. During the season of 1913, in some two or three +instances, observations were made of the biting of _Simulium_ and some +additional and entirely creditable reports were received. These +observations and reports were under conditions identical with those +referred to in the reports of 1912 and confirm the conclusions based on +the observations of that year. I would repeat with emphasis that it is +inconceivable that a fly of the appearance and habits of the prevalent +species of _Simulium_ could be present in such a region, especially +about the haunts of man and attack him with sufficient frequency and +regularity to satisfactorily account for so active and prevalent a +disease as pellagra without being a well-known and recognized pest." + +"In connection with the conditions in the Piedmont region of South +Carolina, it may be well to cite the results of a study of those in the +arid region of western Texas." + +"In May, 1913, in company with Capt. J. F. Siler of the +Thompson-McFadden Pellagra Commission, I visited the region of which +Midland in Midland County is the center. This region is very dry and +totally devoid of running water for a long distance in every direction. +The only natural source of water-supply, a few water holes and ponds, +were visited and found to be of such a nature that the survival of +_Simulium_, far less its propagation in them, is absolutely impossible. +The nearest stream affording possibilities as a source of _Simulium_ is +60 miles away, while the average distance of such possibility is not +less than 100 miles." + +"Artificial sources of water-supply were also investigated carefully and +were found to offer no opportunity for the breeding of _Simulium_." + +"At Midland the histories of five cases of pellagra were obtained, which +gave clear evidence that this place or its immediate vicinity was the +point of origin. Persons of long residence in the country were +questioned as to the occurrence of such flies as _Simulium_ and returned +negative answers. These included a retired cattle owner, who is a man of +education and a keen observer, an expert veterinarian stationed in the +country who has the cattle of the country under constant observation, +and a practical cattle man, manager of a ranch and of wide experience. +The latter had had experience with 'Buffalo gnats' in other localities +(in the East) and is well acquainted with them. His close personal +supervision of the cattle under his charge, makes it practically certain +that he would have discovered these gnats had they been present in the +country." + +"At the time the study was made, _Simulium_ was breeding and active in +the adult state in the vicinity of Dallas, Texas, in the eastern part of +the state. We have here a region in which cases of pellagra have +originated, yet in which _Simulium_ does not and cannot breed." + +Other possible insect vectors of pellagra have been studied in great +detail and the available evidence indicates that if _any_ insect plays a +rôle in the spread of the disease, _Stomoxys calcitrans_ most nearly +fills the conditions. This conclusion was announced by Jennings and King +in 1912, and has been supported by their subsequent work. + +Yet, after all the studies of the past decade, the old belief that +pellagra is essentially of dietary origin is gaining ground. Goldberger, +Waring and Willets (1914) of the United States Public Health Service +summarize their conclusions in the statement, (1) that it is dependent +on some yet undetermined fault in a diet in which the animal or +leguminous protein component is disproportionately large and (2) that no +pellagra develops in those who _consume_ a mixed, well-balanced, and +varied diet, such, for example, as that furnished by the Government to +the enlisted men of the Army, Navy, and Marine Corps. + + +Leprosy + +LEPROSY is a specific, infectious disease due to _Bacillus lepræ_, and +characterized by the formation of tubercular nodules, ulcerations, and +disturbances of sensation. In spite of the long time that the disease +has been known and the dread with which it is regarded, little is known +concerning the method of transfer of the causative organism or the means +by which it gains access to the human body. + +It is known that the bacilli are to be found in the tubercles, the scurf +of the skin, nasal secretions, the sputum and, in fact in practically +all the discharges of the leper. Under such conditions it is quite +conceivable that they may be transferred in some instances from diseased +to healthy individuals through the agency of insects and other +arthropods. Many attempts have been made to demonstrate this method of +spread of the disease, but with little success. + +Of the suggested insect carriers none seem to meet the conditions better +than mosquitoes, and there are many suggestions in literature that these +insects play an important rôle in the transmission of leprosy. The +literature has been reviewed and important experimental evidence +presented by Currie (1910). He found that mosquitoes feeding, under +natural conditions, upon cases of nodular leprosy so rarely, if ever, +imbibe the lepra bacillus that they cannot be regarded as one of the +ordinary means of transference of this bacillus from lepers to the skin +of healthy persons. He believes that the reason that mosquitoes that +have fed on lepers do not contain the lepra bacillus is that when these +insects feed they insert their proboscis directly into a blood vessel +and thus obtain bacilli-free blood, unmixed with lymph. + +The same worker undertook to determine whether flies are able to +transmit leprosy. He experimented with five species found in +Honolulu,--_Musca domestica_, _Sarcophaga pallinervis_, _Sarcophaga +barbata_, _Volucella obesa_ and an undetermined species of _Lucilia_. +The experiments with _Musca domestica_ were the most detailed. From +these experiments he concluded, first, that all of the above-named +flies, when given an opportunity to feed upon leprous fluids, will +contain the bacilli in their intestinal tracts and feces for several +days after such feeding. Second, that considering the habits of these +flies, and especially those of _Musca domestica_, it is certain that, +given an exposed leprous ulcer, these insects will frequently convey +immense numbers of lepra bacilli, directly or indirectly, to the skins, +nasal mucosa, and digestive tracts of healthy persons. Additional +evidence along this line has recently been brought forward by Honeij and +Parker (1914), who incriminate both _Musca domestica_ and _Stomoxys +calcitrans_. Whether or not such insect-borne bacilli are capable of +infecting persons whose skin and mucosa are thus contaminated, Currie +was unwilling to maintain, but he concludes that until we have more +accurate knowledge on this point, we are justified in regarding these +insects with grave suspicion of being one of the means of disseminating +leprous infection. + +Various students of the subject have suggested that bed-bugs may be the +carriers of leprosy and have determined the presence of acid-fast +bacilli in the intestines of bed-bugs which had fed on leprous patients. +Opposed to this, the careful experiments of Thompson (1913) and of +Skelton and Parkham (1913) have been wholly negative. + +Borrel has recently suggested that _Demodex_, may play a rôle in +spreading the infection in families. Many other insects and acariens +have been suggested as possible vectors, but the experimental data are +few and in no wise conclusive. The most that can be said is that it is +quite possible that under favorable conditions the infection might be +spread by any of the several blood-sucking forms or by house-flies. + + +Verruga peruviana + +VERRUGA PERUVIANA is defined by Castellani and Chalmers as "a chronic, +endemic, specific, general disorder of unknown origin, not contagious, +but apparently inoculable, and characterized by an irregular fever +associated with rheumatoid pains, anemia, followed by granulomatous +swellings in the skin, mucous membranes, and organs of the body." It has +been generally believed by medical men interested that the comparatively +benign eruptive verruga is identical with the so-called Oroya, or +Carrion's fever, a malignant type. This view is not supported by the +work of Strong, Tyzzer and Brues, (1913). + +The disease is confined to South America and to definitely limited areas +of those countries in which it does occur. It is especially prevalent in +some parts of Peru. + +The causative organism and the method of transfer of verruga are +unknown. Castellani and Chalmers pointed out in 1910 that the study of +the distribution of the disease in Peru would impress one with the +similarity to the distribution of the Rocky Mountain fever and would +lead to the conclusion that the ætiological cause must in some way be +associated with some blood-sucking animal, perhaps an arachnid, and that +this is supported by the fact that the persons most prone to the +infection are those who work in the fields. + +More recently, Townsend (1913), in a series of papers, has maintained +that verruga and Carrion's disease are identical, and that they are +transmitted to man by the bites of the Psychodid fly, _Phlebotomus +verrucarum_. He succeeded in producing the eruptive type of the disease +in experimental animals by injecting a physiological salt trituration of +wild Phlebotomus flies. A cebus monkey was exposed from October so to +November 6, by chaining him to a tree in the verruga zone, next to a +stone wall from which the flies emerged in large numbers every night. +Miliar eruption began to appear on the orbits November 13 and by +November 21, there were a number of typical eruptions, with exudation on +various parts of the body exactly like miliar eruptive sores commonly +seen on legs of human cases. + +An assistant in the verruga work, George E. Nicholson, contracted the +eruptive type of the disease, apparently as a result of being bitten by +the Phlebotomus flies. He had slept in a verruga zone, under a tight +net. During the night he evidently put his hands in contact with the +net, for in the morning there were fifty-five unmistakable Phlebotomus +bites on the backs of his hands and wrists. + +Townsend believes that in nature, lizards constitute the reservoir of +the disease and that it is from them that the Phlebotomus flies receive +the infection. + + +Cancer + +There are not wanting suggestions that this dread disease is carried, or +even caused, by arthropods. Borrel (1909) stated that he had found mites +of the genus _Demodex_ in carcinoma of the face and of the mammæ. He +believed that they acted as carriers of the virus. + +Saul (1910) and Dahl (1910) go much further, since they attribute the +production of the malignant growth to the presence of mites which Saul +had found in cancers. These Dahl described as belonging to a new +species, which he designated _Tarsonemus hominis_. These findings have +since been confirmed by several workers. Nevertheless, the presence of +the mite is so rare that it cannot be regarded as an important factor in +the causation of the disease. The theory that cancer is caused by an +external parasite is given little credence by investigators in this +field. + + +IN CONCLUSION, it should be noted that the medical and entomological +literature of the past few years abounds in suggestions, and in +unsupported direct statements that various other diseases are +insect-borne. Knab (1912) has well said "Since the discovery that +certain blood-sucking insects are the secondary hosts of pathogenic +parasites, nearly every insect that sucks blood, whether habitually or +occasionally, has been suspected or considered a possible transmitter of +disease. No thought seems to have been given to the conditions and the +characteristics of the individual species of blood-sucking insects, +which make disease transmission possible." + +He points out that "in order to be a potential transmitter of human +blood-parasites, an insect must be closely associated with man and +normally have opportunity to suck his blood repeatedly. It is not +sufficient that occasional specimens bite man, as, for example, is the +case with forest mosquitoes. Although a person may be bitten by a large +number of such mosquitoes, the chances that any of these mosquitoes +survive to develop the parasites in question, (assuming such development +to be possible), and then find opportunity to bite and infect another +person, are altogether too remote. Applying this criterion, not only the +majority of mosquitoes but many other blood-sucking insects, such as +Tabanidæ and Simuliidæ, may be confidently eliminated. Moreover, these +insects are mostly in evidence only during a brief season, so that we +have an additional difficulty of a very long interval during which there +could be no propagation of the disease in question." He makes an +exception of tick-borne diseases, where the parasites are directly +transmitted from the tick host to its offspring and where, for this +reason, the insect remains a potential transmitter for a very long +period. He also cites the trypanosome diseases as possible exceptions, +since the causative organisms apparently thrive in a number of different +vertebrate hosts and may be transmitted from cattle, or wild animals, to +man. + +Knab's article should serve a valuable end in checking irresponsible +theorizing on the subject of insect transmission of disease. +Nevertheless, the principles which he laid down cannot be applied to the +cases of accidental carriage of bacterial diseases, or to those of +direct inoculation of pyogenic organisms, or of blood parasites such as +the bacillus of anthrax, or of bubonic plague. Accumulated evidence has +justified the conclusion that certain trypanosomes pathogenic to man are +harbored by wild mammals, and so form an exception. Townsend believes +that lizards constitute the natural reservoir of verruga; and it seems +probable that field mice harbor the organism of tsutsugamushi disease. +Such instances are likely to accumulate as our knowledge of the relation +of arthropods to disease broadens. + + + + +CHAPTER XII + +HOMINOXIOUS ARTHROPODS + + +The following synoptic tables are presented in the hope that they may be +of service in giving the reader a perspective of the relationships of +the Arthropoda in general and enabling him to identify the more +important species which have been found noxious to man. Though +applicable chiefly to the arthropods found in the United States, exotic +genera and species which are concerned in the transmission of disease +are also included. For this reason the keys to the genera of the Muscids +of the world are given. As will be seen, the tables embrace a number of +groups of species which are not injurious. This was found necessary in +order that the student might not be lead to an erroneous determination +which would result were he to attempt to identify a species which +heretofore had not been considered noxious, by means of a key containing +only the noxious forms. The names printed in BOLD FACED TYPE indicate +the hominoxious arthropods which have been most commonly mentioned in +literature. + + +CRUSTACEA + +Arthropods having two pairs of antennæ which are sometimes modified for +grasping, and usually with more than five pairs of legs. With but few +exceptions they are aquatic creatures. Representatives are: Crabs, +lobsters, shrimps, crayfish, water-fleas, and woodlice. To this class +belongs the CYCLOPS (fig. 122) a genus of minute aquatic crustaceans of +which at least one species harbors _Dracunculus medinensis_, the Guinea +worm (fig. 121). + + +MYRIAPODA + +Elongate, usually vermiform, wingless, terrestrial creatures having one +pair of antennæ, legs attached to each of the many intermediate body +segments. This group is divided into two sections, now usually given +class rank: the DIPLOPODA or millipedes (fig. 13), commonly known as +thousand legs, characterized by having two pairs of legs attached to +each intermediate body segment, and the CHILOPODA or centipedes (fig. +14) having only one pair of legs to each body segment. + + +ARACHNIDA + +In this class the antennæ are apparently wanting, wings are never +present, and the adults are usually provided with four pairs of legs. +Scorpions, harvest-men, spiders, mites, etc. + + +HEXAPODA (Insects) + +True insects have a single pair of antennæ, which is rarely vestigial, +and usually one or two pairs of wings in the adult stage. Familiar +examples are cockroaches, crickets, grasshoppers, bugs, dragon-flies, +butterflies, moths, mosquitoes, flies, beetles, ants, bees and wasps. + + +ORDERS OF THE ARACHNIDA + +a. Abdomen distinctly segmented. A group of orders including scorpions, + (fig. 11), whip-scorpions (fig. 10), pseudo-scorpions, solpugids + (fig. 12) harvest-men (daddy-long-legs or harvestmen), etc. + ARTHROGASTRA + +aa. Abdomen unsegmented, though sometimes with numerous annulations + SPHÆROGASTRA + + b. A constriction between cephalothorax and abdomen (fig. 7). True + Spiders ARANEIDA + + bb. No deep constriction between these parts. + + c. Legs usually well developed, body more or less depressed (fig. + 49). Mites ACARINA + + cc. Legs stumpy or absent, body more or less elongate or vermiform, + or if shorter, the species is aquatic or semi-aquatic in habit. + + d. Four pairs of short legs; species inhabiting moss or water. + Water-bears. TARDIGRADA + + dd. Two pairs of clasping organs near the mouth, instead of legs, + in the adult; worm-like creatures parasitic within the nasal + passages, lungs, etc. of mammals and reptiles (fig. 148). + Tongue worms. LINGUATULINA + +[Illustration: 148. Linguatula. (_a_) larva; (enlarged). (_b_) adult; +(natural size).] + + +ACARINA[E] + +a. Abdomen annulate, elongate; very minute forms, often with but four + legs (fig. 62). DEMODICOIDEA + + b. With but four legs of five segments each. Living on plants, often + forming galls. ERIOPHYIDÆ + + bb. With eight legs, of three segments each. Living in the skin of + mammals. DEMODICIDÆ + + To this family belongs the genus DEMODEX found in the sebaceous + glands and hair follicles of various mammals, including man. _D. + phylloides_ Csokor has been found in Canada on swine, causing + white tubercles on the skin. _D. bovis_ Stiles has been reported + from the United States on cattle, upon the skin of which they + form swellings. D. FOLLICULORUM Simon is the species found on + man. See page 78. + +aa. Abdomen not annulate nor prolonged behind; eight legs in the adult + stage. + + b. With a distinct spiracle upon a stigmal plate on each side of the + body (usually ventral) above the third or fourth coxæ or a little + behind (fig. 50); palpi free; skin often coriaceous or leathery; + tarsi often with a sucker. + + c. Hypostome large (fig. 50), furnished below with many recurved + teeth; venter with furrows, skin leathery; large forms, usually + parasitic. IXODOIDEA + + d. Without scutum but covered by a more or less uniform leathery + integument; festoons absent; coxæ unarmed, tarsi without + ventral spurs; pulvilli absent or vestigial in the adults; + palpi cylindrical; sexual dimorphism slight. ARGASIDÆ + + e. Body flattened, oval or rounded, with a distinct flattened + margin differing in structure from the general integument; + this margin gives the body a sharp edge which is not + entirely obliterated even when the tick is full fed. + Capitulum (in adults and nymphs) entirely invisible + dorsally, distant in the adult by about its own length from + the anterior border. Eyes absent. ARGUS Latr. + + f. Body oblong; margin with quadrangular cells; anterior tibiæ + and metatarsi each about three times as long as broad. On + poultry, southwest United States. A. PERSICUS MINIATUS + + _A. brevipes_ Banks, a species with proportionately shorter + legs has been recorded from Arizona. + + ff. With another combination of characters. About six other + species of _Argas_ from various parts of the world, + parasitic on birds and mammals. + + ee. Body flattened when unfed, but usually becoming very convex + on distention; anterior end more or less pointed and + hoodlike; margin thick and not clearly defined, similar in + structure to the rest of the integument and generally + disappearing on distention; capitulum subterminal, its + anterior portions often visible dorsally in the adult; eyes + present in some species. + + f. Integument pitted, without rounded tubercles; body provided + with many short stiff bristles; eyes absent. On horses, + cattle and man (fig. 48). OTIOBIUS Banks. + + O. MEGNINI, a widely distributed species, is the type of + this genus. + + ff. Integument with rounded tubercles or granules; body + without stiff bristles. ORNITHODOROS Koch. + + g. Two pairs of eyes; tarsi IV with a prominent subterminal + spur above; leg I strongly roughened. On cattle and man. + O. CORIACEUS + + gg. No eyes; no such spur on the hind tarsi. + + h. Tarsi I without humps above. _O. talaje._ + + hh. Tarsi I with humps above. + + i. Tarsi IV without distinct humps above. On hogs, + cattle and man. O. TURICATA + + ii. Tarsi IV with humps nearly equidistant (fig. 142). + Africa. O. MOUBATA + +[Illustration: 149. Hæmaphysalis wellingtoni. Note short palpi. After +Nuttall and Warburton.] + + dd. With scutum or shield (fig. 50); festoons usually present; + coxæ usually armed with spurs, tarsi generally with one or two + ventral spurs; pulvilli present in the adults; sexual + dimorphism pronounced. IXODIDÆ + + e. With anal grooves surrounding anus in front; inornate; + without eyes; no posterior marginal festoons; venter of the + male with non-salient plates. Numerous species, 14 from the + United States, among them I. RICINUS (fig. 49 and 50), + SCAPULARIS, COOKEI, _hexagonus_, _bicornis_. IXODES Latr. + (including Ceratixodes). + + ee. With anal groove contouring anus behind, or groove faint or + obsolete. + + f. With short palpi (fig. 149). + + g. Without eyes, inornate, with posterior marginal festoons; + male without ventral plates. Numerous species. _H. + chordeilis_ and _leporis-palustris_ from the United + States. _Hæmaphysalis_ Koch. + +[Illustration: 150. Stigmal plate of Dermacentor andersoni; (_a_) of +male, (_b_) of female. After Stiles. (_c_) Dermacentor variabilis, male; +(_d_) Glyciphagus obesus; (_e_) Otodectes cynotis; (_f_) Tyroglyphus +lintneri; (_g_) Tarsonemus pallidus; (_h_) anal plate and mandible of +Liponyssus; (_c_) to (_h_) after Banks.] + + gg. With eyes. + + h. Anal groove distinct; posterior marginal festoons + present. + + i. Base of the capitulum (fig. 150c) rectangular + dorsally; usually ornate. DERMACENTOR Koch. + + j. Adults with four longitudinal rows of large + denticles on each half of hypostome; stigmal plate + nearly circular, without dorso-lateral + prolongation, goblets very large, attaining 43µ to + 115µ in diameter; not over 40 per plate, each + plate surrounded by an elevated row of regularly + arranged supporting cells; white rust wanting; + base of capitulum distinctly broader than long, + its postero-lateral angles prolonged slightly, if + at all; coxæ T with short spurs; trochanter I with + small dorso-terminal blade. Texas, Arizona, etc. + _D. nitens_ + +[Illustration: 151. Rhipicephalus bursa, male. After Nuttall and +Warburton.] + + jj. Adults with three longitudinal rows of large + denticles on each half of hypostome; goblet cells + always more than 40 per plate; whitish rust + usually present. + + k. Dorso-lateral prolongation of stigmal plate small + or absent; plates of the adults distinctly + longer than broad; goblet cells large, usually + 30µ to 85µ in diameter, appearing as very coarse + punctations on untreated specimens, but on + specimens treated with caustic potash they + appear very distinct in outline; base of + capitulum distinctly (usually about twice) + broader than long, the postero-lateral angles + distinctly produced caudad; spurs of coxæ I + long, lateral spur slightly longer than median; + trochanter I with dorso-terminal spur. _D. + albipictus_, (= _variegatus_), _salmoni_, + _nigrolineatus_. + + kk. Dorso-lateral prolongation of stigmal plate + distinct. + + l. Body of plate distinctly longer than broad; + goblet cells of medium size, usually 17.5µ to + 35µ or 40µ in diameter, appearing as medium + sized punctuations on untreated specimens, but + on the specimens treated with caustic potash + they appear very distinct in outline, which is + not circular; base of capitulum usually less + than twice as broad as long, the + postero-lateral angles always distinctly + prolonged caudad. + + m. Trochanter I with distinct dorso-subterminal + retrograde sharp, digitate spur; + postero-lateral angles of capitulum + pronouncedly prolonged caudal, 112µ to 160µ + long; goblet cells attain 13µ to 40µ in + diameter; type locality California. D. + OCCIDENTALIS + + mm. Trochanter I with dorso-terminal blade; + postero-lateral angles of capitulum with + rather short prolongations. + + n. Stigmal plate small, goblet cells not + exceeding 45 in the male or 100 in the + female; scutum with little rust, coxa I + with short spurs, the inner distinctly + shorter than the outer. _D. + parumapertus-marginatus_ + + nn. Stigmal plate larger; goblet cells over 70 + in the male and over 100 in the female; + coxa I with longer spurs, inner slightly + shorter than the outer; scutum with + considerable rust. D. VENUSTUS[F] + + ll. Goblet cells small, rarely exceeding 17.6µ, + occasionally reaching 19µ in diameter; on + untreated specimens they appear as very fine + granulations, and on specimens treated with + caustic potash they may be difficult to see, + but their large number can be determined from + the prominent stems of the goblets; surface of + outline of the goblets distinctly circular; + base of the capitulum usually less than twice + as broad as long, the postero-lateral angle + distinctly prolonged caudad; spurs of coxæ I + long. _D. reticulatus_ and _electus_ (= + _variabilis_?) + + ii. Base of the capitulum (fig. 151) usually hexagonal + (except in the male of _puchellus_); and usually + inornate. + + j. No ventral plate or shield in either sex (fig. + 153). R. BICORNIS from the United States. + RHIPICENTOR Nuttall + + jj. Males with a pair of adanal shields, and usually a + pair of accessory adanal shields. Numerous + species, among them _R. sanguineus_ (fig. 154) and + _texanus_, the latter from the United States. + _Rhipicephalus_ Koch + + hh. Anal grooves faint or obsolete; no marginal festoons. + + i. Short palpi; highly chitinized; unfed adults of large + size; coxæ conical; male with a median plate + prolonged in two long spines projecting caudad; + segments of leg pair IV greatly swollen (fig. 155, + 156). _M. winthemi_ _Margaropus_ Karsch + +[Illustration: 152. Monieziella (Histiogaster) emtomophaga-spermatica, +ventral aspect, male and female. After Trouessart.] + + ii. Very short palpi, ridged dorsally and laterally; + slightly chitinized; unfed adults of smaller size; + coxæ I bifid; male with adanal and accessory adanal + shields (fig. 139). B. ANNULATUS. BOOPHILUS Curtis + + ff. Palpi longer than broad (fig. 157). + + g. Male with pair of adanal shields, and two posterior + abdominal protrusions capped by chitinized points; + festoons present or absent. Several species, among them + H. ÆGYPTICUM (fig. 140) from the old world. HYALOMMA + Koch + + gg. Male without adanal shields but small ventral plaques + are occasionally present close to the festoons. Many + species, a few from the United States (fig. 157). + AMBLYOMMA Koch + + h. Coxa I with but one spine; metatarsi (except I) with + two thickened spurs at tips. _A. maculatum_ + + hh. Coxa I with two spines; metatarsi without stout spurs + at tips, only slender hairs. + + i. Projections of coxa I blunt and short. Large species + on the gopher tortoise in Florida. _A. tuberculatum_ + + ii. Projections of coxa I longer, and at least one of + them sharp pointed; second segment of palpus twice + as long as the third; coxa IV of the male with a + long spine. + + j. Porose areas nearly circular; shield of both sexes + pale yellowish, with some silvery streaks and + marks, and some reddish spots; shield of female as + broad as long. A. CAJENNENSE (= MIXTUM). + + jj. Porose areas elongate, shield brown, in the female + with an apical silvery mark, in the male with two + small and two or four other silvery spots; shield + of the female longer than broad (fig 158 e). A. + AMERICANUM. + +[Illustration: 153. Rhipicentor bicornis, ventral aspect, male. After +Nuttall and Warburton.] + + cc. Hypostome small, without teeth, venter without furrows; body + often with coriaceous shields, posterior margin of the body + never crenulate (i.e. without festoons); no eyes. GAMASOIDEA. + + d. Parasitic on vertebrates; mandibles fitted for piercing; body + sometimes constricted. DERMANYSSIDÆ. + + e. Anal plate present. DERMANYSSINÆ. + + f. Body short; legs stout, hind pair reaching much beyond the + tip of the body. On bats. _Pteroptus_ Dufour. + + ff. Body long; hind legs not reaching beyond the tip of the + body. + + g. Peritreme on the dorsum, very short; body distinctly + constricted. _Ptilonyssus_ Berl. + + gg. Peritreme on the venter, longer; body not distinctly + constricted. + + h. Mandibles in both sexes chelate. Parasitic on bats, + mice and birds (fig. 150, h). LIPONYSSUS Kol. + + The species L. (= LEIOGNATHUS) SYLVIARUM frequents the + nests of warblers. An instance is on record of + these mites attacking man, causing a pruritis. + + hh. Mandibles in the male chelate (fig. 158 j), in the + female long, styliform. Parasitic on birds. + DERMANYSSUS Dug. Two species of importance may be + noted, _D. hirundinus_ and D. GALLINÆ. The latter + (fig. 51) is a serious pest of poultry, sometimes + attacking man, causing itching and soreness. + + ee. Anal plate absent. In lungs and air passages of some + mammals. HALARACHNINÆ. + + dd. Free or attached to insects, rarely on vertebrates. + + e. First pair of legs inserted within the same body opening as + the oral tube; genital apertures surrounded by the sternum. + On insects. UROPODIDÆ. + +[Illustration: 154. Rhicephalus sanguineus, male. After Nuttall and +Warburton.] + + ee. First pair of legs inserted at one side of the mouth + opening; male genital aperture usually on the anterior + margin of the sternal plate. GAMASIDÆ. + + This family contains a number of genera, some of which are + found upon mammals, though the majority affect only other + arthropods. One species, LÆLAPS STABULARIS, frequents the + bedding in stables, and in one instance at least, has + occasioned irritation and itching, in man. + + bb. No distinct spiracle in the stigmal plate on each side of the + body. + + c. Body usually coriaceous, with few hairs, with a specialized seta + arising from a pore near each posterior corner of the + cephalothorax; no eyes; mouth parts and palpi very small; + ventral openings of the abdomen large; tarsi without sucker. Not + parasitic. ORIBATOIDEA. + + cc. Body softer; without such specialized seta. + + d. Aquatic species. HYDRACHNOIDEA. + + dd. Not aquatic. + + e. Palpi small, three segmented, adhering for some distance to + the lip; ventral suckers at genital opening or near anal + opening usually present; no eyes; tarsi often end in + suckers; beneath the skin on the venter are seen rod-like + epimera that support the legs; body often entire. Adults + frequently parasitic. SARCOPTOIDEA. + + f. With tracheæ; no ventral suckers; legs ending in claws; + body divided into cephalothorax and abdomen; the female + with a clavate hair between legs I and II. Usually not + parasitic on birds and mammals. TARSONEMIDÆ + + g. Hind legs of female ending in claw and sucker as in the + other pairs. PEDICULOIDINÆ + + To this sub-family belongs the genus PEDICULOIDES + + P. ventricosus is described on page 69. + +[Illustration: 155. Margaropus winthemi, male. After Nuttall and +Warburton.] + +[Illustration: 156. Margaropus winthemi, capitulum and scutum. +After Nuttall and Warburton.] + + gg. Hind legs of the female end in long hairs. TARSONEMINÆ + + TARSONEMUS INTECTUS Karpelles, normally found upon grain, + is said to attack man in Hungary and Russia. Other + species of the genus affect various plants (c.f. fig. + 150, g). + + ff. Without tracheæ; no such clavate hair. + + g. Genital suckers usually present; integument usually + without fine parallel lines. + + h. Legs short, without clavate hair on tarsi I and II. On + insects. CANESTRINIDÆ. + + hh. Legs longer, with a clavate hair on tarsi I and II. + Not normally parasitic except on bees. TYROGLYPHIDÆ + + i. Dorsal integument more or less granulate; claws very + weak, almost invisible; some hairs of the body + plainly feathered; ventral apertures large. + GLYCIPHAGUS Her. + + This genus occurs in the United States. In Europe the + mites have been found feeding on all sorts of + substances. They are known as sugar mites and + cause the disease known as grocer's itch. G. + DOMESTICUS and G. PRUNORUM are old world species + (fig. 150, d). + + ii. Dorsal integument not granulate; claws distinct; no + prominent feathered hairs; ventral aperture small. + + j. Mandibles not chelate; elongate, and toothed below; + body without long hairs; palpi enlarged at tip and + provided with two divergent bristles. Species feed + on decaying substances. _Histiostoma_ Kram. + + jj. Mandibles chelate; palpi not enlarged at the tip, + nor with two bristles. + + k. No clavate hair on the base of tarsi I and II; no + suture between cephalothorax and abdomen. Live + on bees or in their nests. _Trichotarsus_ Can. + + kk. A clavate or thickened hair at the base of tarsi + I and II. + + l. The bristle on the penultimate segment of the + legs arises from near the middle; no suture + between the cephalothorax and abdomen. The + species, some of which occur in the United + States, feed on dried fruit, etc. + _Carpoglyphus_ Robin. + + ll. The bristle on the penultimate segment of the + legs arise from near the tip; a suture between + cephalothorax and abdomen. + + m. Cephalothorax with four distinct and long + bristles in a transverse row; tarsi I and II + about twice as long as the preceding segment + (fig. 150 f). TYROGLYPHUS Latr. + + n. Some bristles on tarsi I and II near + middle, distinctly spine-like; the sense + hair about its length from the base of the + segment. Several species in the United + States belong to this group. + + nn. No spine-like bristles near the middle of + the tarsi; sense hair not its length from + the base of the segment. + + o. Of the terminal abdominal bristles, only + two are about as long as the abdomen; + leg I of the male greatly thickened and + with a spine at apex of the femur below. + T. FARINÆ. + + oo. Of the terminal abdominal bristles at + least six or more are very long, nearly + as long as the body. + + p. Bristles of the body distinctly plumose + or pectinate; tarsi very long. T. + LONGIOR. + + pp. Bristles of the body not pectinate. + + q. In mills, stored foods, grains, etc. + Third and fourth joints of hind legs + scarcely twice as long as broad; + abdominal bristles not unusually + long; legs I and II of the male not + unusually stout. T. AMERICANUS. + + qq. With other characters and habits. + _T. lintneri_ (fig. 150 f) the + mushroom mite, and several other + species. + + mm. Cephalothorax with but two long distinct + bristles (besides the frontal pair), but + sometimes a very minute intermediate pair; + tarsi I and II unusually short and not twice + as long as the preceding segment. + + n. Tarsi with some stout spines. RHIZOGLYPHUS + Clap. + + The species of this genus are vegetable + feeders. Several occur in the United + States. R. PARASITICUS and R. + SPINITARSUS have been recorded from the + old world, attacking human beings who + handle affected plants. + + nn. Tarsi with only fine hairs. MONIEZIELLA + Berl. + + The species of this genus, as far as known, + are predaceous or feed on recently + killed animal matter. Several species + occur in the United States. M. (= + HISTIOGASTER) ENTOMOPHAGA (fig. 152) + from the old world has been recorded as + injurious to man. + + gg. Genital suckers absent; integument with fine parallel + lines. Parasitic on birds and mammals. + + h. Possessing a specially developed apparatus for clinging + to hairs of mammals. LISTROPHORIDÆ. + + hh. Without such apparatus. + + i. Living on the plumage of birds. ANALGESIDÆ. + + ii. In the living tissues of birds and mammals. + + j. Vulva longitudinal. In the skin and cellular + tissues of birds. CYTOLEICHIDÆ. + + This family contains two species, both occurring in + the United States on the common fowl. + _Laminosioptes cysticola_ occurs on the skin and + also bores into the subcutaneous tissue where it + gives rise to a calcareous cyst. _Cytoleichus + nudus_ is most commonly found in the air + passages and air cells. + + jj. Vulva transverse. In the skin of mammals and + birds. SARCOPTIDÆ + + k. Anal opening on the dorsum. + + l. Third pair of legs in the male without apical + suckers. On cats and rabbits. NOTOEDRES + Rail. + + The itch mite of the cat, N. CATI (fig. 61) has + been recorded on man. + + ll. Third leg in the male with suckers. On bats + _Prosopodectes_ Can. + + kk. Anal opening below. + + l. Pedicel of the suckers jointed; mandibles + styliform and serrate near the tip. PSOROPTES + Gerv. P. COMMUNIS OVIS is the cause of sheep + scab. + + ll. Pedicel of the suckers not jointed; mandibles + chelate. + + m. No suckers on the legs of the females; + parasitic on birds, including chickens. _C. + mutans_ is itch mite of chickens. + _Cnemidocoptes_ Fürst. + + mm. Suckers at least on legs I and II; parasitic + on mammals. + + n. Legs very short; in the male the hind pairs + equal in size; body usually short. + SARCOPTES Latr. + + S. SCABIEI is the itch mite of man (fig. + 56). + +[Illustration: 157. Amblyomma, female. After Nuttall and Warburton.] + + nn. Legs more slender; in the male the third + pair is much larger than the fourth; body + more elongate. + + o. Female with suckers on the fourth pair of + legs. Species do not burrow in the skin, + but produce a scab similar to sheep + scab. They occur in the ox, horse, sheep + and goat. CHORIOPTES Gerv. C. SYMBIOTES + BOVIS of the ox has been recorded a few + times on man. + + oo. Female without suckers to the fourth + legs. + + p. Hind part of the male abdomen with two + lobes. On a few wild animals. + _Caparinia_ Can. + + pp. Hind part of the male abdomen without + lobes. Live in ears of dogs and cats. + _Otodectes_ Canestr. + + _O. cynotis_ Hering (fig. 150 e) has + been taken in the United States. + + ee. Palpi usually of four or five segments, free; rarely with + ventral suckers near genital or anal openings; eyes often + present; tarsi never end in suckers; body usually divided + into cephalothorax and abdomen; rod-like epimera rarely + visible; adults rarely parasitic. + + f. Last segment of the palpi never forms a thumb to the + preceding segment; palpi simple, or rarely formed to hold + prey; body with but few hairs. EUPODOIDEA. + + g. Palpi often geniculate, or else fitted for grasping prey; + mandibles large and snout like; cephalothorax with four + long bristles above, two in front, two behind; last + segment of leg I longer than the preceding segment, + often twice as long. BDELLIDÆ. + + gg. Palpi never geniculate (fig. 158a), nor fitted for + grasping prey: beak small; cephalothorax with bristles + in different arrangement; last segment of leg I shorter + or but little longer than the preceding joint; eyes when + present near posterior border. EUPODIDÆ + + Moniez has described a species from Belgium (TYDEUS + MOLESTUS) which attacks man. It is rose colored; + eyeless; its legs are scarcely as long as its body, + the hind femur is not thickened; the mandibles are + small and the anal opening is on the venter. The + female attains a length of about 0.3 mm. + + ff. Last segment of the palpus forms a thumb to the preceding, + which ends in a claw (with few exceptions); body often + with many hairs (fig. 158 k). TROMBIDOIDEA. + + g. Legs I and II with processes bearing spines; skin with + several shields; coxæ contiguous. CÆCULIDÆ. + + gg. Legs I and II without such processes; few if any + shields. + + h. Palpi much thickened on the base, moving laterally, + last joint often with two pectinate bristles; no eyes; + legs I ending in several long hairs; adult sometimes + parasitic. CHEYLETIDÆ + + CHEYLETUS ERUDITUS, which frequents old books, has once + been found in pus discharged from the ear of man. + + hh. Palpi less thickened, moving vertically; eyes usually + present; leg I not ending in long hairs. + + i. Coxæ contiguous, radiate; legs slender, bristly; body + with few hairs; no dorsal groove; tarsi not swollen. + ERYTHRÆIDÆ. + + ii. Coxæ more or less in two groups; legs less bristly. + +[Illustration: 158. (_a_) Tydeus, beak and leg from below; (_b_) +Cheyletus pyriformis, beak and palpus; (_c_) beak and claw of +Pediculoides; (_d_) leg of Sarcoptes; (_e_) scutum of female of +Amblyomma americana; (_f_) leg I and tip of mandible of Histiostoma +americana; (_g_) Histiogaster malus, mandible and venter; (_h_) +Aleurobius farinæ, palpus and leg I of male; (_i_) Otodectes cynotis, tip of +abdomen of male, (_j_) beak and anal plate of Dermanyssus gallinæ female; (_k_) +palpus of Allothrombium. (_a_) to (_j_) after Banks.] + + j. Body with fewer, longer hairs; often spinning + threads; no dorsal groove; tarsi never swollen; + mandibles styliform (for piercing). TETRANYCHIDÆ + + The genus TETRANYCHUS may be distinguished from the + other genera occurring in the United States by + the following characters: No scale-like + projections on the front of the cephalothorax; + legs I as long or longer than the body; palp + ends in a distinct thumb; the body is about 1.5 + times as long as broad. T. MOLESTISSIMUS Weyenb. + from South America, and T. TELARIUS from Europe + and America ordinarily infesting plants, are + said also to molest man. + + jj. Body with many fine hairs or short spines; not + spinning threads; often with dorsal groove; tarsi + often swollen. + + k. Mandibles styliform for piercing. RHYCHOLOPHIDÆ. + + kk. Mandibles chelate, for biting. TROMBIDIDÆ + + The genus TROMBIDIUM has recently been subdivided + by Berlese into a number of smaller ones, of + which some five or six occur in the United + States. The mature mite is not parasitic but + the larvæ which are very numerous in certain + localities will cause intense itching, + soreness, and even more serious complications. + They burrow beneath the skin and produce + inflamed spots. They have received the + popular name of "RED BUG." The names LEPTUS + AMERICANUS and L. IRRITANS have been applied + to them, although they are now known to be + immature stages. (Fig. 44.) + + +HEXAPODA (Insecta) + +The Thysanura (springtails and bristletails), the Neuropteroids +(may-flies, stone-flies, dragon-flies, caddis-flies, etc.), Mallophaga +(bird lice), Physopoda (thrips), Orthoptera (grasshoppers, crickets, +roaches), are of no special interest from our viewpoint. The remaining +orders are briefly characterized below. + + +SIPHUNCULATA (page 275) + +Mouth parts suctorial; beak fleshy, not jointed; insect wingless; +parasitic upon mammals. Metamorphosis incomplete. Lice. + + +HEMIPTERA (page 275) + +Mouth parts suctorial; beak or the sheath of the beak jointed; in the +mature state usually with four wings. In external appearance the +immature insect resembles the adult except that the immature form (i.e. +nymph) never has wings, the successive instars during the process of +growth, therefore, are quite similar; and the metamorphosis is thus +incomplete. To this order belong the true bugs, the plant lice, leaf +hoppers, frog hoppers, cicadas, etc. + + +LEPIDOPTERA + +The adult insect has the body covered with scales and (with the rare +exception of the females of a few species) with four wings also covered +with scales. Proboscis, when present, coiled, not segmented, adapted for +sucking. Metamorphosis complete, i.e. the young which hatches from the +egg is quite unlike the adult, and after undergoing several molts +transforms into a quiescent pupa which is frequently enclosed in a +cocoon from which the adult later emerges. The larvæ are known as +caterpillars. Butterflies and moths. + + +DIPTERA (page 285) + +The adult insect is provided with two, usually transparent, wings, the +second pair of wings of other insects being replaced by a pair of +halteres or balancers. In a few rare species the wings, or halteres, or +both, are wanting. The mouth parts, which are not segmented, are adapted +for sucking. The tarsi are five-segmented. Metamorphosis complete. The +larvæ, which are never provided with jointed legs, are variously known +as maggots, or grubs, or wrigglers. Flies, midges, mosquitoes. + + +SIPHONAPTERA (page 316) + +Mouth parts adapted for sucking; body naked or with bristles and spines; +prothorax well developed; body compressed; tarsi with five segments; +wings absent. Metamorphosis complete. The larva is a wormlike creature. +Fleas. + + +COLEOPTERA + +Adult with four wings (rarely wanting), the first pair horny or +leathery, veinless, forming wing covers which meet in a line along the +middle of the back. Mouth parts of both immature stages and adults +adapted for biting and chewing. Metamorphosis complete. The larvæ of +many species are known as grubs. Beetles. + + +HYMENOPTERA + +Adult insect with four, usually transparent, wings, wanting in some +species. Mouth parts adapted for biting and sucking; palpi small; tarsi +four or five-segmented. Metamorphosis complete. Parasitic four-winged +flies, ants, bees, and wasps. + + +SIPHUNCULATA AND HEMIPTERA + +a. Legs with claws fitted for clinging to hairs; wings wanting; + spiracles of the abdomen on the dorsal surface. (= ANOPLURA = + PARASITICA) SIPHUNCULATA. + + b. Legs not modified into clinging hooks; tibia and tarsus very long + and slender; tibia without thumb-like process; antennæ + five-segmented HÆMATOMYZIDÆ Endr. + + _Hæmatomyzus elephantis_ on the elephant. + + bb. Legs modified into clinging hooks; tibia and tarsus usually short + and stout; tibia with a thumb-like process; head not anteriorly + prolonged, tube-like. + + c. Body depressed; a pair of stigmata on the mesothorax, and + abdominal segments three to eight; antennæ three to + five-segmented. + + d. Eyes large, projecting, distinctly pigmented; pharynx short and + broad; fulturæ (inner skeleton of head) very strong and broad, + with broad arms; proboscis short, scarcely attaining the + thorax. PEDICULIDÆ + + e. Antennæ three-segmented. A few species occurring upon old + world monkeys. _Pedicinis_ Gerv. + + ee. Antennæ five-segmented. + + f. All legs stout; thumb-like process of the tibia very long + and slender, beset with strong spines, fore legs stouter + than the others; abdomen elongate, segments without + lateral processes; the divided telson with a conical + process posteriorly upon the ventral side. PEDICULUS L. + + g. Upon man. + + h. Each abdominal segment dorsally with from one to three + more or less regular transverse rows of small setæ; + antenna about as long as the width of the head. Head + louse (fig. 65). P. HUMANUS. + + hh. "No transverse rows of abdominal setæ; antenna longer + than the width of the head; species larger." Piaget. + Body louse of man. P. CORPORIS. + + gg. Upon apes and other mammals. _P. pusitatus_ (?). + + ff. Fore legs delicate, with very long and slender claws; + other legs very stout with short and stout claws; + thumb-like process of the tibia short and stout; abdomen + very short and broad; segment one to five closely crowded, + thus the stigmata of segments three to five apparently + lying in one segment; segments five to eight with lateral + processes; telson without lateral conical appendages (fig. + 69). Crab louse of man. PHTHIRUS PUBIS. + + dd. Eyes indistinct or wanting; pharynx long and slender, fulturæ + very slender and closely applied to the pharynx; proboscis + very long. Several genera found upon various mammals. + HÆMATOPINIDÆ. + + cc. Body swollen; meso- and metathorax, and abdominal segments two + to eight each with a pair of stigmata; eyes wanting; antennæ + four or five-segmented; body covered with stout spines. Three + genera found upon marine mammals. ECHINOPHTHIRIIDÆ + +aa. Legs fitted for walking or jumping; spiracles of abdomen usually + ventral; beak segmented. + + b. Apex of head usually directed anteriorly; beak arising from its + apex; sides of the face remote from the front coxæ; first pair of + wings when present thickened at base, with thinner margins. + HETEROPTERA + +[Illustration: 159. Taxonomic details of Hemiptera-Heteroptera. (_a_) +Dorsal aspect; (_b_) seta from bedbug; (_c_) wing of Heteropteron; (_d_) +leg; (_e_) wing of Sinea.] + + c. Front tarsi of one segment, spade-form (palæformes); beak short, + at most two-segmented; intermediate legs long, slender; + posterior pair adapted for swimming. CORIXIDÆ + + cc. Front tarsi rarely one-segmented, never spade-form; beak free, + at least three-segmented. + + d. Pulvilli wanting. + + e. Hemelytra usually with a distinct clavus (fig. 159), clavus + always ends behind the apex of the scutellum, forming the + commissure. (Species having the wings much reduced or + wanting should be sought for in both sections.) + + f. Antennæ very short; meso- and metasternum composite; eyes + always present. + + g. Ocelli present; beak four-segmented. OCHTERIDÆ and + NERTHRIDÆ. + + gg. Ocelli wanting; antennæ more or less hidden in a groove. + + h. Anterior coxæ inserted at or near anterior margin of + the prosternum; front legs raptorial; beak + three-segmented BELOSTOMIDÆ (with swimming legs), + NEPIDÆ, NAUCORIDÆ. + + i. Metasternum without a median longitudinal keel; + antennæ always four-segmented. + + j. Beak short, robust, conical; the hairy fleck on the + corium elongate, large, lying in the middle + between the inner angle of the membrane and the + outer vein parallel to the membrane margin; + membrane margin S-shaped. + + k. The thick fore femur with a relatively deep + longitudinal furrow to receive the tibia. + Several American species (fig. 19f.). BELOSTOMA + (= Lethocerus Mayer) + + kk. The less thickened fore femur without such a + furrow. B. GRISEUS. BENACUS Stäl. + + jj. Beak slender, cylindrical; the hairy spot on the + corium rounded lying next to the inner angle of + the membrane. + + k. Membrane large, furrow of the embolium broadened. + _Z. aurantiacum_, _fluminea_, etc. _Zaitha_ + + kk. Membrane very short; furrow of embolium not + broadened. Western genus. _Pedinocoris_ + + ii. Metasternum with a long median longitudinal keel. + Southwestern forms. _Abedus ovatus_ and _Deniostoma + dilatato_ + + hh. Anterior coxæ inserted at the posterior margin of the + prosternum; legs natatorial. Back swimmers (fig. 19 b.). + NOTONECTIDÆ + + i. Apices of the hemelytra entire; the three pairs of legs + similar in shape; beak three-segmented; abdomen not + keeled or hairy. _Plea_ Leach + + ii. Apices of hemelytra notched; legs dissimilar; beak + four-segmented; abdomen keeled and hairy. + + j. Hemelytra usually much longer than the abdomen; + fourth segment of the antenna longer than the third + segment; hind tarsi with claws. _Bueno_ Kirk. + + jj. Hemelytra but little longer than the abdomen; fourth + segment of the antenna shorter than the third + segment; hind tarsi without claws (fig. 19b). + NOTONECTA L. + + ff. Antennæ longer than the head; or if shorter, then the eyes + and ocelli absent. + + g. Eyes, ocelli, and scutellum wanting; beak + three-segmented; head short; hemelytra always short; + membrane wanting. Insects parasitic on bats. POLYCTENIDÆ + + gg. Eyes present. + + h. First two antennal segments very short, last two long, + pilose, third thickened at the base; ocelli present, + veins of the hemelytra forming cells. DIPSOCORIDÆ (= + CERATOCOMBIDÆ) including SCHIZOPTERIDÆ. + + hh. Third segment of the antenna not thickened at the + base, second as long or longer than the third, rarely + shorter. + + i. Posterior coxæ hinged (cardinate), if rarely + rotating, the cuneus is severed, the membrane is one + or two-celled, and the meso- and metasternum are + composite. + + j. Ocelli absent, clypeus dilated toward the apex; + hemelytra always short, membrane wanting. Species + parasitic. Bed bugs, etc. CIMICIDÆ + + k. Beak short, reaching to about the anterior coxæ; + scutellum acuminate at the apex; lateral margin + of the elytra but little reflexed, apical margin + more or less rounded; intermediate and posterior + coxæ very remote. + + l. Body covered with short hairs, only the sides + of the pronotum and the hemelytra fringed with + longer hairs; antennæ with the third and + fourth segments very much more slender than + the first and second; pronotum with the + anterior margin very _deeply sinuate_. CIMEX + L. + + m. Sides of the pronotum widely dilated, broader + than the breadth of one eye, and densely + fringed with backward curved hairs; apical + margin of the hemelytra nearly straight, + rounded toward the interior or exterior + angles. + + n. Body covered with very short hairs; second + segment of the antenna shorter than the + third; sides of the pronotum feebly + reflexed, fringed with shorter hairs than + the breadth of one eye; hemelytra with the + commissural (inner) margin rounded and + shorter than the scutellum, apical margin + rounded towards the interior angle. The + common bed bug (fig. 19h). C. LECTULARIUS + Linn + + nn. Body covered with longer hairs; second and + third segments of the antenna of equal + length; side of the pronotum narrowly, but + distinctly, reflexed, fringed with longer + hairs than the breadth of one eye; + hemelytra with the commissural margin + straight and longer than the scutellum, + apical margin rounded towards the exterior + angle. Species found on bats in various + parts of the United States. _C. + pillosellus_ Hov. + + mm. Sides of the pronotum neither dilated, nor + reflexed, fringed with less dense and nearly + straight hairs; hemelytra with the apical + margin distinctly rounded. Parasitic on man, + birds and bats. Occurs in the old world, + Brazil and the West Indies. C. HEMIPTERUS + Fabr. (= rotundatus) + + ll. Body clothed with rather longer silky hairs; + third and fourth segments of the antenna + somewhat more slender than the first and + second; anterior margin of the pronotum _very + slightly sinuate_ or nearly straight in the + middle, produced at the lateral angles. This + is the species which in American collections + is known as _C. hirundinis_, the latter being + an old world form. It is found in swallows + nests. O. VICARIUS. OECIACUS Stäl + + kk. Beak long, reaching to the posterior coxæ; + scutellum rounded at the apex; lateral margins + of the elytra strongly reflexed, apical margin + slightly sinuate toward the middle; intermediate + and posterior coxæ sub-contiguous. This species + infests poultry in southwest United States and + in Mexico. H. INODORUS. HÆMATOSIPHON Champ. + +[Illustration: 160. Pselliopsis (Milyas) cinctus (×2). After C. V. +Riley.] + + jj. Ocelli present, if rarely absent in the female, + then the tarsus has two segments; or if with three + tarsal segments, the wing membrane with one or two + cells. + + k. Beak four-segmented, or with two-segmented tarsi. + ISOMETOPIDÆ, MICROPHYSIDÆ, and some CAPSIDÆ. + + kk. Beak three-segmented. + + l. Hemelytra with embolium; head horizontal, more + or less conical; membrane with one to four + veins, rarely wanting. ANTHOCORIDÆ + + Several species of this family affecting man + have been noted, ANTHOCORIS KINGI and + CONGOLENSE, from Africa and LYCTOCORIS + CAMPESTRIS from various parts of the world. + LYCTOCORIS FITCHII Reuter (fig. 19 j), later + considered by Reuter as a variety of L. + CAMPESTRIS, occurs in the United States. + + ll. Hemelytra without embolium. Superfamily + ACANTHIOIDEA (= SALDÆ Fieber and LEPTOPODÆ + Fieber) + + ii. Posterior coxæ rotating. + + j. Claws preapical; aquatic forms. GERRIDÆ and VELIADÆ + + jj. Claws apical. + + k. Prosternum without stridulatory sulcus (notch for + beak). + + l. Tarsus with three segments; membrane with two + or three longitudinal cells from which veins + radiate; rarely with free longitudinal veins + (Arachnocoris) or veins nearly obsolete + (Arbela); clavus and corium coriaceous; ocelli + rarely absent. NABIDÆ + + REDUVIOLUS (= CORISCUS) SUBCOLEOPTRATUS (fig. 19 + g), a species belonging to this family, + occurring in the United States, has been + accused of biting man. This insect is flat, + of a jet black color, bordered with yellow + on the sides of the abdomen, and with + yellowish legs. It is predaceous, feeding on + other insects. + + ll. With other combinations of characters. + HYDROMETRIDÆ, HENICOCEPHALIDÆ, NÆOGEIDÆ, + MESOVELIADÆ, JOPPEICIDÆ + + kk. Prosternum with stridulatory sulcus (notch for + beak); with three segments, short, strong. + + l. Antennæ filiform or sometimes more slender + apically, geniculate; wing membrane with two + or three large basal cells; scutellum small or + moderate REDUVIIDÆ + + For a key to the genera and species see next + page. + + ll. Last antennal segment clavate or fusiform; win + membrane with the veins often forked and + anastomosing; scutellum large; tarsi each with + two segments; fore legs strong. (= PHYMATIDÆ) + MACROCEPHALIDÆ + + ee. Clavus noticeably narrowed towards the apex, never extending + beyond the scutellum, the two not meeting to form a + commissure; head horizontal, much prolonged between the + antennæ, on each side with an antennal tubercle, sometimes + acute; ocelli absent; meso- and metasternum simple; tarsi + each with two segments; body flattened (fig. 19c). ARADIDÆ, + including DYSODIIDÆ. + + dd. Pulvilli present (absent in one Australian family + THAUMATOCORIDÆ in which case there is a membranous appendage + at the tip of the tibia). CAPSIDÆ (= MIRIDÆ),[G] _Eotrechus_ + (in family GERRIDÆ), NÆOGAIDÆ, TINGITIDÆ, PIESMIDÆ, + MYODOCHIDÆ, CORIZIDÆ, COREIDÆ, ALYDIDÆ, PENTATOMIDÆ, + SCUTELLERIDÆ, etc. + + bb. Apex of head directed ventrally, beak arising from the hinder part + of the lower side of the head; sides of face contiguous to the + front coxæ; first pair of wings, when present, of uniform + thickness. Cicadas, scale insects, plant lice (Aphids), + spittle-insects, leaf hoppers, etc. HOMOPTERA + + +REDUVIIDÆ OF THE UNITED STATES + +(Adapted from a key given by Fracker). + +a. Ocelli none; wings and hemelytra always present in the adults; no + discoidal areole in the corium near the apex of the clavus. + _Orthometrops decorata_, _Oncerotrachelus acuminatus_, etc., + Pennsylvania and south. _Sarcinæ_ + +aa. Ocelli present in the winged individuals; anterior coxæ not as long + as the femora. + + b. Hemelytra without a quadrangular or discoidal areole in the corium + near the apex of the clavus. + + c. Ocelli not farther cephalad than the caudal margins of the eyes; + segment two of the antenna single. + + d. Thorax usually constricted caudad of the middle; anterior coxæ + externally flat or concave. PIRATINÆ + + e. Middle tibiæ without spongy fossa, head long, no lateral + tubercle on neck. _S. stria_, Carolina, Ill., Cal. + _Sirthenia_ Spinola + + ee. Middle tibiæ with spongy fossa; fore tibiæ convex above; + neck with a small tubercle on each side. + + f. Apical portion of anterior tibiæ angularly dilated beneath, + the spongy fossa being preceded by a small prominence. + MELANOLESTES Stäl + + g. Black, with piceous legs and antennæ. N. E. States (fig. + 19a) M. PICIPES + + gg. Sides, and sometimes the whole dorsal surface of the + abdomen red. Ill., and southward. M. ABDOMINALIS + + ff. Tibiæ not dilated as in "f"; spongy fossa elongate; + metapleural sulci close to the margin. R. BIGUTTATUS (fig. + 22). South RASAHUS A. and S. + + dd. Thorax constricted in the middle or cephalad of the middle; + anterior tarsi each three-segmented. + + e. Apex of the scutellum narrow, without spines or with a single + spine REDUVIINÆ + + f. Antennæ inserted in the lateral or dorso-lateral margins of + the head; antenniferous tubercles slightly projecting from + the sides of the head; head produced strongly cephalad; + ocelli at least as far apart as the eyes. + + g. Antennæ inserted very near the apex of the head; segments + one and three of the beak short, segment two nearly four + times as long as segment one. R. PROLIXUS. W. I. + RHODNIUS Stäl + + gg. Antennæ inserted remote from the vertex of the head. + + h. Body slightly hairy; pronotum distinctly constricted; + angles distinct; anterior lobe four-tuberculate, with + the middle tubercles large and conical. _M. + phyllosoma_, large species from the southwest. + _Meccus_ Stäl + + hh. Body smooth, margin of the pronotum sinuous, scarcely + constricted; anterior lobe lined with little + tubercles. CONORHINUS Lap. + + i. Surface of the pronotum and prosternum more or less + granular. + + j. Eyes small, head black; body very narrow, a fifth + as wide as long; beak reaches the middle of the + prosternum. California. C. PROTRACTUS + + jj. Eyes large, head fuscous; body at least a fourth + as wide as long. Southern species. _C. + rubrofasciatus_ + + ii. Pronotum and prosternum destitute of granules. + + j. Border of abdomen entirely black except for a + narrow yellowish spot at the apex of one segment. + Texas. _C. gerstaeckeri_ + + jj. Border of abdomen otherwise marked. + + k. Beak slender, joints one and two slightly pilose, + two more than twice as long as one; tubercles at + the apical angles of the pronotum slightly + acute, conical. Md. to Ill. and south. The + masked bed bug hunter (fig. 71). C. SANGUISUGUS + + kk. Beak entirely pilose, joint two a third longer + than joint one; joint one much longer than + three; tubercles at the apical angles of + pronotum slightly elevated, obtuse. Ga., Ill., + Tex., Cal. _C. variegatus_ + + ff. Antenna inserted on top of the head between margins, close + to the eyes; antenniferous tubercles not projecting from + the side of the head. + + g. Anterior lobe of the pronotum with a bispinous or + bituberculate disc; femora unarmed. _S. arizonica_, _S. + bicolor_. Southwestern species. _Spiniger_ Burm. + + gg. Disc of pronotum unarmed; apex of scutellum produced + into a spine; ocelli close to the eyes; eyes large and + close together. REDUVIUS Lamarck + + h. Color piceous. Widely distributed in the United States. + (Fig. 20). R. PERSONATUS + + hh. More or less testaceous in color. Southwestern states. + R. SENILIS + + ee. Apex of scutellum broad, with two or three spines. + ECTRICHODIINÆ + + f. First segment of the antenna about as long as the head. _E. + cruciata_ Pa. and south; _E. cinctiventris_, Tex. and Mex. + _Ectrichodia_ L. et S. + + ff. First segment of the antennæ short. _P. æneo-nitens_. + South. _Pothea_ A. et S. + + cc. Ocelli cephalad of the hind margins of the eyes; first segment + of the antennæ stout, second segment divided into many smaller + segments. South and west. _Homalocoris maculicollis_, and + _Hammatocerus purcis_. HAMMATOCERINÆ + + bb. Hemelytra with a quadrangular or discoidal areole in the corium + near the apex of the clavus (fig. 159e). + + c. Anal areole of the membrane not extending as far proximad as the + costal areole; basal segment of the antenna thickened, porrect; + the other segments slender, folding back beneath the head and + the first segment STENOPODINÆ + + d. Head armed with a ramous or furcate spine below each side, + caudad of the eyes. + + e. First segment of the antenna thickened, apex produced in a + spine beyond the insertion of the second segment. Species + from Va., Ill. and south. _Pnirontis_ Stäl. + + ee. First segment of the antenna not produced beyond the + insertion of the second segment. _Pygolampis_, N. E. states + and south; _Gnathobleda_, S. W. and Mex. + + dd. Head unarmed below or armed with a simple spine; rarely with a + subfurcate spine at the side of the base. Carolina, Missouri + and south. _Stenopoda_, _Schumannia_, _Diaditus_, _Narvesus_, + _Oncocephalus_ + + cc. Anal areole of membrane extending farther proximad than the + costal areole. + + d. Ocelli farther apart than the eyes. _A. crassipes_, widely + distributed in the United States; other species occur in the + southwest. _Apiomerus_ Hahn. + + dd. Ocelli not so far apart as the eyes. ZELINÆ + + e. Sides of mesosternum without a tubercle or fold in front. + + f. Fore femur as long as or longer than the hind femur; first + segment of the beak much shorter than the second. _Z. + audax_, in the north eastern states; other species south + and west. _Zelus_ Fabr. + + ff. Fore femur shorter than the hind femur, rarely of equal + length, in this case the first segment of the beak as long + or longer than the second. + + g. First segment of the beak shorter than the second; fore + femur a little shorter than the hind femur; the first + segment of the beak distinctly longer than the head + before the eyes. _P. cinctus_ a widely distributed + species (fig. 160). _P. punctipes_, _P. spinicollis_, + Cal., Mex. (= _Milyas_) _Pselliopus_ Berg. + + gg. First segment of the beak as long or longer than the + second. + + h. Pronotum armed with spines on the disc. + + i. Juga distinctly prominent at the apex and often acute + or subacute; fore femur distinctly thickened; + hemelytra usually not reaching the apex of the + abdomen. _Fitchia aptera_, N. Y., south and west; + _F. spinosula_, South; _Rocconata annulicornis_, + Texas, etc. + + ii. Juga when prominent, obtuse at apex; eyes full width + of the head; fore femur not thickened; pronotum with + four spines on posterior lobe. _R. taurus_, Pa., + south and west. _Repipta_ Stäl. + + hh. Pronotum unarmed on the disc. + + i. Spines on each apical angle of the penultimate + abdominal segment. _A. cinereus_, Pa., and south. + _Atrachelus_ A. et S. + + ii. Apical angle of the penultimate abdominal segment + unarmed. _Fitchia_ (in part); _Castolus ferox_, + Arizona. + + ee. Sides of the mesosternum with a tubercle or fold in front at + the hind angles of the prosternum; first segment of the beak + longer than the part of the head cephalad of the eyes. + + f. Fore femur thickened, densely granulated; hind femur + unarmed. + +[Illustration: 161. Taxonomic details of Diptera. (_a_) Ventral aspect +of abdomen of Cynomyia; (_b_) antenna of Tabanus; (_c_) ventral aspect +of abdomen of Chortophila; (_d_) ventral aspect of abdomen of Stomoxys; +(_e_) claw of Aedes (Culex) sylvestris, male; (_f_) claw of Hippoboscid; +(_g_) foot of dipterous insect showing empodium developed pulvilliform; +(_h_) hind tarsal segment of Simulium vittatum, female; (_i_) foot of +dipterous insect showing bristle-like empodium.] + + g. Fore tibiæ each with three long spines on the ventral + side. _S. diadema_ (fig. 159e), a widely distributed + species; and several southwestern species. _Sinea_ A. et + S. + + gg. Fore tibiæ unarmed. _A. multispinosa_, widely + distributed; _A. tabida_, Cal. _Acholla_ Stäl. + + ff. Fore femur unarmed, rarely a little thickened, a little + granulated. + + g. Pronotum produced caudad over the scutellum, with a high + mesal tuberculate ridge (fig. 19e). A. CRISTATUS. N. Y. + to Cal. and south. ARILUS Hahn. + + gg. Caudal lobe of the pronotum six sided, neither elevated + nor produced caudad. _H. americanus_, Southwest; also + several W. I. and Mexican genera. HARPACTOR Lap. + + +DIPTERA (Mosquitoes, Midges, Flies) + +a. Integument leathery, abdominal segments indistinct; wings often + wanting; parasitic forms. PUPIPARA + + b. Head folding back on the dorsum of the thorax; wingless flies + parasitic on bats. Genus _Nycteribia_. NYCTERIBIIDÆ + + bb. Head not folding back upon the dorsum of the thorax; flies either + winged or wingless; parasitic on birds and on bats and other + mammals. + + c. Antennæ reduced, wings when present, with distinct parallel veins + and outer crossveins; claws simple; palpi leaf-like, projecting + in front of the head. Flies chiefly found on bats. Several + genera occur in North America. STREBLIDÆ + +[Illustration: 162. Hippobosca equina, ×4. After Osborn.] + + cc. Antennæ more elongate, segments more or less distinctly + separated; head sunk into an emargination of the thorax; wings + when present with the veins crowded toward the anterior margin; + palpi not leaf-like. HIPPOBOSCIDÆ + + d. Wings absent or reduced and not adapted for flight. + + e. Wings and halteres (balancers) absent. _M. ovinus_, the sheep + tick. _Melophagus_ Latr. + + ee. Wing reduced (or cast off), halteres present. + + f. Claw bidentate; ocelli present. On deer after the wings are + cast off. _L. depressa_. _Lipoptena_ Nitsch + + ff. Claw tridentate (fig. 161 f). On _Macropis_. _B. + femorata._ _Brachypteromyia_ Will. + + dd. Wings present and adapted for flight. + + e. Claws bidentate. + + f. Ocelli present; head flat; wings frequently cast off. On + birds before casting of the wing. _Lipoptena_ Nitsch. + + ff. Ocelli absent; head round; wings present. The horse tick + H. EQUINA may attack man (fig. 162). HIPPOBOSCA L. + + ee. Claws tridentate (fig. 161 f.). + + f. Anal cell closed at apical margin by the anal crossvein. + + g. Ocelli absent. _Stilbometopa_ Coq. + + gg. Ocelli present. + + h. R_{4+5} does not form an angle at the crossvein. On + birds. There is a record of one species of this genus + attacking man. ORNITHOMYIA Latr. + + hh. R_{4+5} makes an angle at the crossvein. _O. + confluens_. _Ornithoica_ Rdi. + + ff. Anal cell not closed by an anal crossvein. _Lynchia_, + _Pseudolfersia_, and _Olfersia_ are chiefly bird + parasites. The first mentioned genus is said to be the + intermediate host of _Hæmoproteus columbæ_. + +aa. Abdominal segments chitinous; not parasitic in the adult stage. + + b. Antennæ with six or more segments and empodium not developed + pulvilliform; palpi often with four segments. + + c. Ocelli present. BLEPHAROCERIDÆ, RHYPHIDÆ, BIBIONIDÆ, + MYCETOPHILIDÆ, besides some isolated genera of other families. + + cc. Ocelli absent. + + d. Dorsum of the thorax with a V-shaped suture; wings usually with + numerous veins; legs often very long and slender. Crane flies. + TIPULIDÆ + + dd. Dorsum of the thorax without a V-shaped suture. + + e. Not more than four longitudinal veins ending in the wing + margin; wing usually hairy: antennæ slender; coxæ not long; + tibiæ: without spurs, legs long and slender. Small, delicate + flies often called Gall gnats. CECIDOMYIIDÆ. + + ee. More than four longitudinal veins ending in the wing margin. + + f. The costal vein is not produced beyond the tip of the wing; + radius with not more than three branches. + + g. Antennæ short, composed of ten or eleven closely united + segments; legs stout; body stout; abdomen oval; anterior + veins stout, posterior ones weak (fig. 163 b); eyes of + the male contiguous over the antennæ. Black flies, + buffalo flies, turkey gnats. Many North American + species, several of them notorious for their blood + sucking propensities. SIMULIIDÆ + + h. Second joint of the hind tarsus with basal scale-like + process and dorsal excision (fig. 161 h); radial + sector not forked; no small cell at the base of the + wing. _S. forbesi_, _jenningsi_, _johannseni_, + _meridionale_, _piscicidium_, VENUSTUM, VITTATUM, etc. + Widely distributed species (= EUSIMULIUM) SIMULIUM + Latr. + + hh. No basal scale-like process on the second joint of the + hind tarsus; radial sector usually forked (fig. 163 + b). + + i. Face broad, small basal cell of the wing present. _P. + fulvum_, HIRTIPES, _mutatum_, PECUARUM, _pleurale_. + PROSIMULIUM Roub. + + ii. Face linear; small basal cell of the wing absent. + One species, _P. furcatum_, from California. + _Parasimulium_ Malloch + + gg. Flies of a different structure. + + h. Antennæ composed of apparently two segments and a + terminal arista formed of a number of closely united + segments. Rare flies with aquatic larvæ. + ORPHNEPHILIDÆ + + hh. Antennæ of six to fifteen segments, those of the male + usually plumose; legs frequently slender and wings + narrow CHIRONOMIDÆ + + i. Media forked (except in the European genus + _Brachypogon_); thorax without longitudinal fissure + and not produced over the head (except in four + exotic genera); antennæ usually fourteen-jointed in + both sexes; fore tibia with a simple comb of setulæ, + hind tibia with two unequal combs, middle tibia + without comb. CERATOPOGONINÆ + + j. Thorax produced cap-like over the head, wing narrow + and very long. _Jenkinsia_, _Macroptilum_ and + _Calyptopogon_, eastern hemisphere; + _Paryphoconus_, Brazil. + + jj. Thorax not produced over the head. + + k. Eyes pubescent, empodium well developed, or if + short then R_{2+3} distinct and crossvein-like + or the branches of R coalescent; r-m crossvein + present; fore femora not thickened; wing either + with appressed hairs or with microscopic erect + setulæ _Dasyhelea_ Kieff. + + kk. Eyes bare, or otherwise differing from the + foregoing. + + l. Empodium well developed, nearly as long as the + claws and with long hairs at the base; femora + and fifth tarsal segments unarmed, i.e. + without spines or stout setæ; fourth tarsal + segment cylindrical. + + m. Wing with erect and microscopic setulæ. + Widely distributed. (= Atrichopogon) + _Ceratopogon_ Meig. + + mm. Wing with long and depressed hairs. Widely + distributed. _Forcipomyia_ + + n. Hind metatarsus shorter or not longer than + the following (i.e. the second tarsal) + segment. Subgenus _Prohelea_ Kieff + + nn. Hind metatarsus longer than the following + segment. Subgenus _Forcipomyia_ Meig. + + ll. Empodium short, scarcely reaching the middle + of the claws, or vestigial. + + m. R-m crossvein wanting. + + n. Palpi four segmented; inferior fork of the + media obliterated at the base. Australia. + _Leptoconops_ Skuse + + nn. Palpi three-segmented. + + o. Legs spinulose, tarsal claws of the + female with a basal tooth or strong + bristle, those of the male unequal, the + anterior with a long sinuous tooth, the + posterior with a short arcuate tooth. + Italy. MYCTEROTYPUS Noé + + oo. Legs unarmed; no crossvein between the + branches of the radius (fig. 163e). New + Mexico. TERSESTHES Townsend + + mm. R-m crossvein present. + + n. Fore femora very much swollen, armed with + spines below, fore tibia arcuate and + applied closely to the inferior margin of + the femur. + + o. R_{2+3} present, therefore cell R_1 and + R_2 both present; wing usually fasciate. + United States _Heteromyia_ Say. + + oo. R_{2+3} not distinct from R_{4+5}, hence + cell R_3 obliterated. South America + _Pachyleptus_ Arrib. (Walker) + + nn. Fore femur not distinctly swollen. + + o. R_{2+3} present therefore cells R_1 and + R_3 both present, or if not, then the + branches of the radius more or less + coalescent, obliterating the cells. + + p. At least the tip of the wing with erect + setulæ; tip of R_{4+5} scarcely + attaining the middle of the wing, + empodium rather indistinct, not + reaching the middle of the claws, the + claws not toothed, equal, with long + basal bristle; legs without stout + setæ. Widely distributed. CULICOIDES + Latr. + + HÆMATOMYIDIUM and OECACTA are probable + synonyms of this. + + pp. Wings bare, if rarely with hair, then + the radius reaches beyond two-thirds + the length of the wing, or the femur + or fifth tarsal segment with stout + black spines. + + q. Media unbranched. Europe. + _Brachypogon_ Kieff + + qq. Media branched. + + r. Hind femur much swollen and spined. + America and Europe. _Serromyia_ + Meg. + + rr. Hind femur not distinctly swollen. + + s. Cell R_1 not longer than high; + fork of the media distad of the + crossvein; wing with microscopic + setulæ _Stilobezzia_ Kieff + + ss. Cell R_1 elongate. + + t. Femora unarmed. Widely + distributed. (= Sphaeromias + Kieff. 1913 not Curtis?). + JOHANNSENIELLA Will. + + tt. Femora, at least in part, with + strong black spines. Widely + distributed. _Palpomyia_ + Megerle + + oo. R_{2+3} coalescent with R_{4+5} hence + cell R_3 is obliterated. + + p. In the female the lower branch of the + media with an elbow near its base + projecting proximad, the petiole of + the media coalescent with the basal + section of the radius, wing long and + narrow, radial sector ending near the + tip of the wing; venation of the male + as in _Bezzia_; front concave. United + States. _Stenoxenus_ Coq. + + pp. Venation otherwise, front not concave. + + q. Subcosta and R_1 more or less + coalescent with the costa; wing + pointed at the apex, much longer + than the body; antennæ fourteen + segmented, not plumose. India. + _Haasiella_ Kieff. + + qq. Subcosta and radius distinct from + the costa. + + r. Abdomen petiolate. _Dibezzia_ + Kieff. + + rr. Abdomen not petiolate. + + s. Head semi-globose; hind tarsi + unusually elongate in the + female; antennæ of the male not + plumose. Europe. _Macropeza_ + Meigen. + + ss. Head not globose, more or less + flattened in front; antennæ of + the male plumose. Widely + distributed. _Bezzia_ Kieff. + + t. Fore femora, at least, armed + with stout spines below. + Subgenus _Bezzia_ Kieff. + + tt. Femora unarmed. Subgenus + _Probezzia_ Kieff. + + ii. Media of the wing simple, and otherwise not as in + "i". To this group belong numerous Chironomid + genera, none of which are known to be noxious to + man. + + ff. The costal vein apparently is continued around the hind + margin of the wing; radius with at least four branches. + + g. Wing ovate pointed, with numerous veins; crossveins, if + evident, before the basal third of the wing; veins very + hairy; very small moth-like flies. PSYCHODIDÆ + + h. With elongate biting proboscis; the petiole of the + anterior forked cell of the wing (R_2) arises at or + beyond the middle of the wing (fig. 163d). PHLEBOTOMUS + Rdi. + +[Illustration: 163. Wings of Diptera. (_a_) Anopheles; (_b_) +Prosimulium; (_c_) Johannseniella; (_d_) Phlebotomus (After Doerr and +Russ); (_e_) Tersesthes (after Townsend); (_f_) Tabanus; (_g_) +Symphoromyia; (_h_) Aphiochæta; (_i_) Eristalis; (_j_) Gastrophilus; +(_k_) Fannia; (_l_) Musca.] + + hh. With shorter proboscis; the petiole of the anterior + forked cell arises near the base of the wing. + _Psychoda_, _Pericoma_, etc. + + gg. The r-m crossvein placed at or beyond the center of the + wing; wings not folded roof-like over the abdomen. + + h. Proboscis short, not adapted for piercing; wings bare + (DIXIDÆ); or wings scaled (CULICIDÆ, Subf. + CORETHRINÆ). + + hh. Proboscis elongate, adapted for piercing; wings + scaled, fringed on the hind margin; antennæ of the + male bushy plumose. Mosquitoes. CULICIDÆ (exclusive of + CORETHRINÆ) + + i. Metanotum without setæ. + + j. Proboscis strongly decurved; body with broad, + appressed, metalescent scales; cell R_2 less than + half as long as its petiole; claws of female + simple, some of the claws of the male toothed. + Several large southern species believed to feed + only on nectar of flowers. _Megarhinus_ R. D. + + jj. Proboscis straight or nearly so, or otherwise + different. + + k. Scutellum evenly rounded, not lobed; claws simple + in both sexes. ANOPHELES Meig. + + l. Abdomen with clusters of broad outstanding + scales along the sides; outstanding scales on + the veins of the wing rather narrow, + lanceolate; upper side of the thorax and + scutellum bearing many appressed lanceolate + scales. Florida and southward (CELLIA). + + m. Hind feet from the middle of the second + segment largely or wholly snow white. + + n. With a black band at the base of the last + segment of each hind foot. A. + ALBIMANUS*[H] and TARSIMACULATA* + + nn. Without such a band A. ARGYRITARSIS* + + mm. Hind feet black, mottled with whitish and + with bands of the same color at the sutures + of the segments. W. I. A. MACULIPES + + ll. Abdomen without such a cluster of scales; + outstanding scales of the wing veins rather + narrow, lanceolate; tarsi wholly black. + + m. Deep black, thorax obscurely lined with + violaceous, especially posteriorly; head, + abdomen and legs black; no markings on the + pleura; abdomen without trace of lighter + bindings; wing scales outstanding, uniform, + not forming spots, though little thicker at + the usual points indicating the spottings. + Florida. A. ATROPUS + + mm. Otherwise marked when the wings are + unspotted. + + n. Wings unspotted. + + o. Petiole of the first forked cell (R_2) + more than a third the length of the + cell. Mississippi valley. A. WALKERI + + oo. Petiole of the first forked cell a third + the length of the cell. Md. A. BARBERI + + nn. Wings spotted. + + o. Front margin of the wings with a patch of + whitish and yellow scales at a point + about two-thirds or three-fourths of the + way from base to apex of wing. + + p. Veins of the wing with many broad + obovate outstanding scales; thorax + with a black dot near the middle of + each side. W. I. A. GRABHAMI* + + pp. The outstanding scales of the wings + rather narrow, lanceolate. + + q. Scales of the last vein of the wings + white, those at each end black; + R_{4+5} black scaled, the extreme + apex white scaled. Widely + distributed north and south (fig. + 131). A. PUNCTIPENNIS + + A dark variety from Pennsylvania has + been named A. PERPLEXENS. + + qq. Scales of the last vein of the wing + white, those at its apex black; + R_{4+5} white scaled and with two + patches of black scales. South and + the tropics. A. FRANCISCANUS and + PSEUDOPUNCTIPENNIS* + + oo. Front margin of the wings wholly black + scaled. + + p. Last (anal) vein of the wings white + scaled with three patches of black + scales (fig. 132). New Jersey to + Texas. A. CRUCIANS* + + pp. Last vein of the wings wholly black + scaled. + + q. Widely distributed north and south + (fig. 130), (= MACULIPENNIS). A. + QUADRIMACULATUS* + + qq. Distributed from Rocky Mountains + westward. A. OCCIDENTALIS + + kk. Scutellum distinctly trilobed. + + l. Cell R_2 less than half as long as its petiole; + thorax with metallic blue scales; median lobe + of the scutellum not tuberculate; few small + species which are not common. URANOTÆNIA + Arrib. + + ll. Cell R_2 nearly or quite as long as its + petiole, or otherwise distinct. + + m. Femora with erect outstanding scales; occiput + broad and exposed. Large species. P. + CILIATA. P. HOWARDI PSOROPHORA R. D. + + mm. Femora without erect scales. + + n. Clypeus bearing several scales or hairs, + scutellum with broad scales only; back of + head with broad scales; scales along the + sides of the mesonotum narrow; some or the + claws toothed; thorax marked with a pair + of silvery scaled curved stripes; legs + black with white bands at the bases of + some of the segments (fig. 134). Yellow + Fever mosquito. AEDES (= STEGOMYIA) + CALOPUS. + + nn. With another combination of characters. + Numerous species of mosquitoes belonging + to several closely related genera, widely + distributed over the country. (_Culex_, + _Aedes_, _Ochlerotatus_, etc.). CULEX in + the wide sense. + + ii. Metanotum with setæ. _Wyeomyia_ (found in the United + States); and related tropic genera. + + bb. Antennæ composed of three segments with a differentiated style or + bristle; third segment sometimes complex or annulate, in which + case the empodium is usually developed like the pulvilli, i.e., + pad-like (fig. 161 g). + + c. Empodium developed pad-like (pulvilliform) i.e., three nearly + equal membranous appendages on the underside of the claw (fig. + 161g). + + d. Squamæ, head, and eyes large; occiput flattened or concave; + third segment of the antennæ with four to eight annuli or + segments, proboscis adapted for piercing; body with fine + hairs, never with bristles; middle tibia with two spurs; wing + venation as figured (fig. 163f); marginal vein encompasses the + entire wing. Horse flies, greenheads, deer flies, gad flies + TABANIDÆ[I] + + e. Hind tibia with spurs at tip; ocelli usually present + (PANGONINÆ) + + f. Third joint of the antennæ with seven or eight segments; + proboscis usually prolonged. + + g. Each section the third antennal segment branched. Central + American species, _P. festæ_. _Pityocera_ G. T. + + gg. Sections of the third antennal segment not branched. + + h. Upper corner of the eyes in the female terminating in + an acute angle; wings of both sexes dark anteriorly. + _G. chrysocoma_, a species from the eastern states. + _Goniops_ Ald. + + hh. Upper corner of the eye in the female not so + terminating; wings nearly uniform in color, or + hyaline. + + i. Proboscis scarcely extending beyond the palpi; front + of the female wide; much wider below than above. S. + W. States. _Apatolestes_ Will. + + ii. Proboscis extending beyond the palpi. + + j. Wing with cell M_3 closed. Tropic America. (= + _Diclisa_) _Scione_ Wlk. + + jj. Cell M_3 open; ocelli present or absent. Two or + three eastern species; many south and west + PANGONIA Rdi. + + ff. Third segment of the antenna with five divisions; ocelli + present. + + g. First and second segments of the antenna short, the + second only half as long as the first, three western + species. SILVIUS Rdi. + + gg. First and second segments of the antenna long, the + second distinctly over half as long as the first. Deer + flies. Many species, widely distributed. CHRYSOPS Meig. + + ee. Hind tibia without spurs; ocelli absent. + + f. Third segment of antenna with four divisions, no tooth or + angulation; wings marked with rings and circles of darker + coloring; front of the female very wide. Widely + distributed. _H. americana_, _H. punctulata_. HÆMATOPOTA + Meig. + + ff. Third segment of the antenna with five divisions (fig. + 161b). + + g. Third segment of the antenna not furnished with a tooth + or distinct angular projection. + + h. Body covered with metallic scales; front of female of + normal width; front and middle tibiæ greatly dilated. + _L. lepidota_. _Lepidoselaga_ Macq. + + hh. Body without metallic scales; antennæ not very long, + the third segment not cylindrical, not situated on a + projecting tubercle; front of the female narrow. + South. _D. ferrugatus._ (= _Diabasis_) _Diachlorus_ O. + S. + + gg. Third segment of the antenna furnished with a tooth or a + distinct angular projection. + + h. Hind tibiæ ciliate with long hairs. S. W. and tropics. + _Snowiella_ and _Stibasoma_. + + hh. Hind tibiæ not ciliate. + + i. Species of slender build, usually with a banded + thorax and abdomen; third segment of the antenna + slender, the basal prominence long; wings mostly + with brownish markings. Tropic America. + _Dichelacera_ Macq. + + ii. Species of a stouter build; third segment of the + antenna stout, its basal process short (fig. 161b). + Many species, widely distributed TABANUS L. + + dd. With another group of characters. + + e. Squamæ small, antennæ variable, thinly pilose or nearly bare + species, without distinct bristles; wing veins not crowded + anteriorly, R_4 and R_5 both present, basal cells large; + middle tibiæ at least with spurs. LEPTIDÆ + + f. Flagellum of the antenna more or less elongated, composed + of numerous more or less distinct divisions. XYLOPHAGINÆ + and ARTHROCERATINÆ. + + ff. Antennæ short, third segment simple, with arista or style; + face small, proboscis short LEPTINÆ + + g. Front tibiæ each with one or two spurs, or if absent, + then no discal cell. _Triptotricha_, _Pheneus_, + _Dialysis_, _Hilarimorpha_. + + gg. Front tibiæ without terminal spurs, discal cell present. + + h. Hind tibiæ each with a single spur. + + i. Anal cell open (fig. 163g); third antennal segment + kidney-shaped with dorsal or subdorsal arista; first + antennal segment elongate and thickened. About a + dozen species have been described from the United + States, of which at least one (S. PACHYCERAS) is + known to be a vicious blood sucker. SYMPHOROMYIA + Frauenf. + + ii. Anal cell closed; third antennal segment not + kidney-shaped. _Chrysopila_, _Ptiolina_, _Spania_. + + hh. Hind tibiæ each with two spurs. + + i. Third segment kidney-shaped, the arista subdorsal; + anal cell closed. _Atherix_ Meig. + + ii. Third segment of the antenna short and with terminal + arista; anal cell open _Leptis_ Fabr. Two European + species of this genus have been accused of blood + sucking habits, but the record seems to have been + based upon error in observation. + + ee. With another combination of characters. STRATIOMYIIDÆ, + CYRTIDÆ, etc. + + cc. Empodium bristlelike or absent. + + d. Antennæ apparently two segmented, with three-segmented arista, + wings (rarely wanting) with several stout veins anteriorly, + the weaker ones running obliquely across the wing (fig. 163h); + small, quick running, bristly, humpbacked flies. Several + genera; APHIOCHÆTA, PHORA, TRINEURA, etc. PHORIDÆ + + dd. Flies with other characters. + + e. No frontal lunule above the base of the antennæ; both R_4 and + R_5 often present; third segment of the antenna often with a + terminal bristle. ASILIDÆ, MYDAIDÆ, APIOCERIDÆ, THEREVIDÆ, + SCENOPINIDÆ, BOMBYLIIDÆ, EMPIDIDÆ, DOLICHOPODIDÆ, + LONCHOPTERIDÆ. + + ee. A frontal lunule above the base of the antennæ; third + segment of the antenna always simple, i.e., not ringed, + usually with a dorsal arista; R_4 and R_5 coalesced into a + simple vein. + + f. A spurious vein or fold between the radius and the media, + rarely absent; the cell R_{4+5} closed at the apex by vein + M_1; few or no bristles on the body, none on the head; + flies frequently with yellow markings. ERISTALIS (fig. + 163i), HELOPHILUS, and many other genera. SYRPHIDÆ + + ff. No spurious vein present. + + g. Body without bristles; proboscis elongate and slender, + often folding; front of both male and female broad. + CONOPIDÆ + + gg. Bristles almost always present on head, thorax, abdomen + and legs. + + h. Arista terminal; hind metatarsus enlarged, sometimes + ornamented, hind tarsus more or less flattened beneath + PLATYPEZIDÆ + + hh. Flies having a different combination of characters. + + i. Head large, eyes occupying nearly the entire head; + cell R_{4+5} narrowed in the margin; small flies. + PIPUNCULIDÆ + + ii. Head and eyes not unusually large. + + j. Squamæ (tegulæ, or calyptræ, or alulæ) not large, + often quite small, the lower one lacking, or at + most barely projecting from below the upper one + (antisquama); front of both male and female broad, + the eyes therefore widely separated; posthumeral + and intraalar macrochæta not simultaneously + present; thorax usually without a complete + transverse suture; postalar callus usually absent; + the connectiva adjoining the ventral sclerites + always visible; hypopleural macrochætæ absent; + last section of R_{4+5} and M_{1+2} with but few + exceptions nearly parallel; subcostal vein often + wanting or vestigial or closely approximated to + R_1; the latter often short, basal cells small, + the posterior ones often indistinct or wanting; + vibrissæ present or absent. ACALYPTRATE MUSCOIDEA + + k. Subcosta present, distinctly separated from R_1 + at the tip; R_1 usually ends distad of the + middle of the wing; the small basal cells of the + wing distinct. + + l. A bristle (vibrissa) on each side of the face + near the margin of the mouth. CORDYLURIDÆ, + SEPSIDÆ, PHYCODROMIDÆ, HETERONEURIDÆ, + HELOMYZIDÆ. + + ll. No vibrissæ present. + + m. Head nearly spherical, cheeks broad and + retreating; proboscis short; the cell R_5 + closed or narrowed in the margin; legs very + long; tarsi shorter than the tibiæ. CALOBATA + and other genera. MICROPEZIDÆ + + mm. Flies with another combination of + characters. RHOPALOMERIDÆ, TRYPETIDÆ, + ORTALIDÆ, SCIOMYZIDÆ. + + kk. Subcosta absent or vestigial, or if present, + then apparently ending in the costa at the point + where R_1 joins it; R_1 usually ends in the + costa at or before the middle of the wing. + + l. Arista long plumose, or pectinate above; oral + vibrissæ present; anal cell complete; costa + broken at the apex of R_1. DROSOPHILA, + PHORTICA, and other genera. DROSOPHILIDÆ + + ll. With another combination of characters. + + m. The cell M and first M_2 not separated by a + crossvein; anal cell absent; front bare or + only bristly above; usually light colored + flies. HIPPELATES, OSCINUS, and other + genera. (See also m m m.) OSCINIDÆ + + mm. Cell M and cell first M_2 often separated by + a crossvein; anal cell present, complete, + though frequently small; scutellum without + spines or protuberances; oral vibrissæ + present; arista bare or short plumose; front + bristly at vertex only; small dark flies. + PIOPHILA (fig. 99), SEPSIS and other genera. + SEPSIDÆ + + mmm. The GEOMYZIDÆ, AGROMYZIDÆ, PSILIDÆ, + TRYPETIDÆ, RHOPALOMERIDÆ, BORBORIDÆ and + DIOPSIDÆ differ in various particulars from + either the OSCINIDÆ and the SEPSIDÆ noted + above. + + jj. Squamæ well developed, usually large, the lower + one frequently projecting from below the upper + one; both posthumeral and intraalar macrochætæ + present; thorax with a complete transverse suture; + postalar callus present and separated by a + distinct suture from the dorsum of the thorax; + front of the female broad, of the male frequently + narrow, the eyes then nearly or quite contiguous; + the connectiva adjoining the ventral sclerites + either visible or not; hypopleural macrochætæ + present or absent; subcosta always distinct in its + whole course, R_1 never short. CALYPTRATE + MUSCOIDEA[J] + + k. Oral opening small, mouth parts usually much + reduced or vestigial. This family is undoubtedly + of polyphyletic origin but for convenience it is + here considered as a single family. OESTRIDÆ. + + l. The costal vein ends at the tip of R_{4+5}, + M_{1+2} straight, not reaching the wing + margin, hence cell R_5 wide open (fig. 163j); + squamæ small; arista bare; ovipositor of the + female elongate. Larvæ in the alimentary canal + of horses, etc. GASTROPHILUS + + m. Posterior crossvein (m-cu) wanting; wings + smoky or with clouds. Europe. G. PECORUM + + mm. Posterior crossvein (m-cu) present, at least + in part. + + n. Wing hyaline with smoky median cross band, + and two or three spots; posterior + trochanters with hook in the male and a + prominence in the female. World wide + distribution. G. EQUI. + + nn. Wings without spots. + + o. Posterior crossvein (m-cu) distad of the + anterior crossvein (r-m); legs, + particularly the femora, blackish brown. + Europe and North America. G. + HÆMORRHOIDALIS + + oo. Posterior crossvein opposite or proximad + of the anterior crossvein. Europe and + North America. G. NASALIS + + ll. The costal vein ends at the tip of M_{1+2}, + M_{1+2} with a bend, the cell R_5 hence much + narrowed in the margin, or closed. + + m. Proboscis geniculate, inserted in a deep + slit; female without extricate ovipositor; + arista either bare or plumose; squamæ large; + facial grooves approximated below. + + n. Arista bare, short. Larvæ in rodents. + Tropic America. _B. princeps_. _Bogeria_ + Austen + + nn. Arista pectinate above. + + o. Tarsi broadened and flattened, hairy, + anal lobe of the wing large. Larvæ in + rodents. A number of American species. + _Cuterebra_. + + oo. Tarsi slender, not hairy; anal lobe of + the wing moderate. Larvæ in man and + other mammals. Tropic America. D. + CYANIVENTRIS. DERMATOBIA Br. + + mm. Mouth parts very small, vestigial; arista + bare. + + n. Facial grooves approximated below, leaving + a narrow median depression or groove. + + o. Cell R_5 closed and petiolate, body + nearly bare. Larvæ in the nasal cavities + of the smaller Ungulates. The sheep bot + fly. O. OVIS. Widely distributed OESTRUS + L. + + oo. Cell R_5 narrowly open, body hairy. + Larvæ parasitic on deer. Europe and + America. _Cephenomyia_ Latr. + + nn. Facial grooves far apart, enclosing + between them a broad shield-shaped + surface; squamæ large; female with + elongate ovipositor. Larvæ hypodermatic on + Ungulates HYPODERMA Clark + + o. Palpi wanting; tibiæ thickened in the + middle. + + p. Hair at apex of the abdomen yellow; + legs including femora yellowish brown. + H. DIANA + + pp. Hair at the apex of the abdomen + reddish yellow. Europe and America. + + q. Tibiæ and tarsi yellow; femora black. + H. LINEATA + + qq. Legs black with black hair; tips of + hind tibia and tarsi yellowish + brown. H. BOVIS + + oo. Palpi small, globular; tibiæ + cylindrical, straight. On reindeer. _O. + tarandi_ _Oedemagena_ Latr. + + kk. Oral opening of the usual size; mouth parts not + vestigial. + + l. Hypopleurals wanting; if three sternopleurals + are present the arrangement is 1:2; + conjunctiva (fig. 161c) of the venter usually + present; if the terminal section of M_{1+2} is + bent it has neither fold nor appendage + (ANTHOMYIIDÆ of Girschner). + + m. Sternopleurals wanting; M_{1+2} straight + toward the apex, costa ends at or slightly + beyond the tip of R_{4+5}; mouth parts + vestigial. GASTROPHILINÆ. See OESTRIDÆ + + mm. Sternopleurals present, if rarely absent + then differing in other characters. + + n. Caudal margin of the fifth ventral + abdominal sclerite of the male deeply + notched on the median line usually to + beyond the middle; abdomen often + cylindrical or linear; abdomen often with + four to eight spots; eyes of the male + usually widely separated; sternopleurals + three, arranged in an equilateral + triangle; subapical seta of the hind tibia + placed very low; M_{1+2} straight; anal + vein abbreviated; wings not rilled. + _Cænosia_, _Caricea_, _Dexiopsis_, + _Hoplogaster_, _Schoenomyia_, etc. + (COENOSINÆ)[K]. ANTHOMYIIDÆ in part + + nn. Caudal margin of the fifth ventral + abdominal sclerite of the male incurved, + rarely deeply cleft, rarely entire, in a + few genera deeply two or three notched; + M_{1+2} straight or curved; abdomen + usually short or elongate oval; + sternopleurals, if three are present, + arranged in the order 1:2 in a right + triangle. (MUSCINÆ-ANTHOMYIINÆ of + Girschner) + + o. M_{1+2} straight, hence cell R_5 not + narrowed in the margin. ANTHOMYIIDÆ in + part + + p. Underside of the scutellum more or less + sparsely covered with fine hairs; anal + vein nearly always reaches the hind + margin of the wing; extensor surface + of the hind tibiæ with a number of + stout setæ; squamæ often small and + equal. ANTHOMYIA, _Chortophila_, + _Eustalomyia_, _Hammomyia_, + _Hylemyia_, _Prosalpia_, _Pegomyia_, + etc. HYLEMYINÆ-PEGOMYINÆ + + pp. Underside of the scutellum bare; anal + vein does not reach the wing margin. + + q. First anal vein short, second anal + suddenly flexed upwards; hind tibiæ + each with one or two strong setæ on + the extensor surface. FANNIA (= + HOMALOMYIA), _Coelomyia_, + _Choristoma_, _Euryomma_, _Azelia_, + etc. FANNINÆ-AZELINÆ + + qq. Anal veins parallel or divergent. + + r. Setæ on the exterior surface of the + hind tibiæ wanting (except in + _Limnaricia_ and _Coenosites_), + lower squama not broadened to the + margin of the scutellum. + _Leucomelina_, _Limnophora_, + _Limnospila_, _Lispa_, _Mydæa_, + _Spilogaster_, etc. + MYDÆINÆ-LIMNOPHORINÆ + + rr. One (rarely more) seta on the + extensor surface of the hind + tibia; squamæ usually large and + unequal. HYDROTAEA, _Aricia_, + _Drymeia_, _Ophyra_, _Phaonia_ (= + _Hyetodesia_), _Pogonomyia_, + _Trichophthicus_, etc. ARICINÆ + + oo. M_{1+2} curved or bent, hence the cell + R_5 more or less narrowed in the margin. + (MUSCINÆ). MUSCIDÆ in part. See page 303 + for generic synopsis. + + ll. Hypopleurals present; when three + sternopleurals are present the arrangement is + 2:1 or 1:1:1. (TACHINIDÆ of Girschner) + + m. Conjunctiva of the ventral sclerites of the + abdomen present, frequently well developed, + surrounding the sclerites. + + n. Mouth parts vestigial. OESTRIDÆ. See page + 297 for generic synopsis. + + nn. Mouth parts well developed. + + o. M_{1+2} straight, hence cell R_5 wide + open in the margin; costa ending at the + tip of R_5; three sternopleurals + present; antennal arista plumose. + _Syllegoptera_. Europe. + (SYLLEGOPTERINÆ). DEXIIDÆ in part + + oo. M_{1+2} bent, hence cell R_5 narrowed in + the margin; sternopleurals rarely + wanting, usually 1:1 or 0:1; facial + plate strongly produced below vibrissal + angle like the bridge of the nose; + antennal arista bare. Parasitic on + Hemiptera and Coleoptera. _Allophora_, + _Cistogaster_, _Clytia_, _Phasia_, etc. + (PHASIINÆ) TACHINIDÆ in part. + + mm. Conjunctiva of the ventral sclerites + invisible (fig. 161a). + + n. Second ventral sclerite of the abdomen + lying with its edges either upon or in + contact with the ventral edges of the + corresponding dorsal sclerite. + + o. Outermost posthumeral almost always lower + (more ventrad) in position than the + presutural macrochæta; fifth ventral + abdominal sclerite of the male cleft + beyond the middle, often strongly + developed; body color very frequently + metallic green or blue, or yellow; + arista plumose. (CALLIPHORINÆ) MUSCIDÆ + in part. + + See page 303 for generic synopsis. + + oo. Outermost posthumeral macrochæta on + level or higher (more dorsad) than the + presutural macrochæta; arista bare, + pubescent, or plumose only on the basal + two-thirds; body coloring usually + grayish (fig. 106). SARCOPHAGIDÆ + + p. Fifth ventral sclerite of the male + either wanting or with the caudal + margin straight; presutural intraalar + rarely present. (SARCOPHAGINÆ) + + q. Fifth ventral abdominal sclerite of + the male much reduced, the remaining + segments with straight posterior + margin, overlapping scale-like; in + the female only segment one and two + scale-like, the others wholly or in + part covered; sternopleurals usually + three or more. SARCOPHAGA and + related genera. + + qq. Fifth ventral sclerite of the male + plainly visible; sternopleurals + usually two. SARCOPHILA, + WOHLFAHRTIA, _Brachycoma_, + _Hilarella_, _Miltogramma_, + _Metopia_, _Macronychia_, _Nyctia_, + _Paramacronychia_, _Pachyphthalmus_, + etc. + + pp. Fifth ventral abdominal sclerite of + the male cleft to beyond the middle; + ventral sclerites usually visible, + shield-like. _Rhinophora_, _Phyto_, + _Melanophora_ RHINOPHORINÆ + +[Illustration: 164. Glossina palpalis. (×4.) After Austen.] + + nn. Second ventral abdominal sclerite as well + as the others more or less covered, + sometimes wholly, by the edges of the + dorsal sclerite. + + o. The presutural intraalar wanting; ventral + sclerites two to five nearly or quite + covered by the edges of the + corresponding dorsal sclerites; base of + the antennæ usually at or below the + middle of the eye; arista usually + plumose; legs usually elongate; + abdominal segments with marginal and + often discal macrochætæ. DEXIIDÆ + + oo. Presutural intraalar present, if absent, + then the ventral sclerites broadly + exposed or the fifth ventral sclerite + vestigial; base of the antennæ usually + above the middle of the eye; arista + bare; at least two posthumerals and + three posterior intraalars present. + Parasitic on caterpillars, etc. + TACHINIDÆ + + +SYNOPSIS OF THE PRINCIPAL GENERA OF THE MUSCIDÆ OF THE WORLD + +a. Proboscis long, directed forward, adapted for piercing, or oral + margin much produced, snout-like. + + b. Oral margin produced snout-like; vibrissa placed high above the + oral margin; antennal arista either pectinate or more or less + plumose. + + c. Antennal arista short or long-plumose; neither sex with distinct + orbital bristles. + + d. No facial carina between the antennæ. RHYNCHOMYIINÆ + + e. Arista short-plumose. _R. speciosa._ Europe. _Rhynchomyia_ R. + D. + + ee. Arista long-plumose. _I. phasina_. Europe and Egypt. + _Idiopsis_. B. B. + + dd. With flattened carina, the bases of the antennæ separated; no + abdominal macrochætæ. COSMININÆ _C. fuscipennis_. South + Africa. _Cosmina_ + + cc. Antennal arista pectinate; bases of the antennæ separated by a + flattened carina. RHINIINÆ R. D. + + d. Cell R_5 open, or closed at the margin. + + e. Third segment of the antenna twice as long as the second; + claws of both sexes short; cell R_5 open. _I. lunata._ + Eastern Hemisphere. _Idia_ Meigen + + ee. Third segment of the antenna three times as long as the + second; cell R_5 open or closed; claws of the male long and + slender, of the female shorter than the last tarsal joint. + _I. mandarina_, China. _Idiella_ B. B. + + dd. Cell R_5 petiolate. _Rhinia_; and _Beccarimyia_ Rdi. + + bb. Proboscis long, directed forward, adapted for piercing. STOMOXINÆ + + c. Arista flat, pectinate above with plumose rays; sternopleurals + 1:2; bases of the veins R_1 and R_{4+5} without setæ; base of + the media bowed down; apical cell opens before the apex of the + wing. African species. GLOSSINA Wied. + + d. Species measuring over twelve mm. in length. _G. longipennis_ + and FUSCA. + + dd. Species less than twelve mm. in length. + + e. All segments of the hind tarsi black. + + f. The fourth and fifth segments of the fore tarsi black; + antennæ black (fig. 164). G. PALPALIS R. D. + + ff. Otherwise marked. _G. bocagei_, _tachinoides_, + _pallicera._ + + ee. First three segments of the hind tarsi are yellow, the + fourth and fifth segments are black. + + f. Fourth and fifth segments of the first and second pair of + tarsi are black. + + g. The yellow bands of the abdominal segments occupy a third + of the segment (fig. 165). G. MORSITANS Westw. + + gg. The yellow band on each segment of the abdomen occupies + a sixth of the segment. G. LONGIPALPIS Wied. + + ff. Tarsi of the first and second pairs of legs wholly yellow. + _G. pallidipes_ Austen + + cc. Rays of the arista not plumose; only one or two sternopleurals; + base of the media not strongly bowed down; apical cell opens at + or very near the apex of the wing. + + d. Vein R_{4+5} without setæ at the base; palpi about as long as + the proboscis. + + e. Arista pectinate (i.e. rays on one side only), the rays often + undulate; two yellow sternopleurals often difficult to + detect; vein M_{1+2} only slightly bent, the apical cell + hence wide open. The horn fly, H. IRRITANS (= _Lyperosia + serrata_) and related species. Widely distributed (figs. + 167, 168). HÆMATOBIA R. D. not B. B. + +[Illustration: 165. Glossina morsitans. (×4.) After Austen.] + + ee. Arista also with rays below; vein M_{1+2} more strongly + bent, the apical cell hence less widely open. + + f. Palpi strongly spatulate at the tips, lower rays of the + arista about six in number, B. SANGUINOLENTUS. South Asia. + BDELLOLARYNX Austen + + ff. Palpi feebly spatulate; apical cell of the wing narrowly + open slightly before the tip; sternopleurals black, + anterior bristle sometimes absent. H. ATRIPALPIS. Europe. + HÆMATOBOSCA Bezzi + + dd. Vein R_{4+5} with setæ at the base.[L] + + e. Veins R_1 and R_{4+5} with setæ at the base; two equally + prominent sternopleural macrochætæ; arista with rays both + above and below; palpi as long as the proboscis; apical cell + of the wing wide open. L. TIBIALIS. (_Hæmatobia_ B. B. not + R. D.). LYPEROSIOPS Town. + + ee. Only vein R_{4+5} with basal setæ; anterior sternopleural + macrochæta wanting; arista pectinate. + + f. Palpi as long as the proboscis, the latter stout, with + fleshy terminal labellæ; apical cell narrowly open; + sternopleural macrochætæ black. S. MACULOSA from Africa + and related species from Asia. STYGEROMYIA Austen + + ff. Palpi much shorter than the proboscis, the latter pointed + at the apex, without fleshy labellæ; apical cell of the + wing wide open. S. CALCITRANS, the stable fly and related + species. Widely distributed in both hemispheres (fig. + 110). STOMOXYS Geof. + +aa. Proboscis neither slender nor elongate, the labellæ fleshy and not + adapted for piercing. + + b. Hypopleuræ without a vertical row of macrochætæ. MUSCINÆ + + c. Arista bare; distal portion of M_{1+2} broadly curved at the end; + hypopleuræ with a sparse cluster of fine hairs. _S. + braziliana_, Southern States and southward. _Synthesiomyia_ B. + B. + + cc. Arista pectinate or plumose. + + d. Arista pectinate. _H. vittigera_, with the posterior half of + the abdomen metallic blue. Mexico. _Hemichlora_ V. d. W. + + dd. Arista plumose. + + e. Middle tibia with one or more prominent setæ on the inner + (flexor) surface beyond the middle, or inner surface very + hairy. + + f. R_1 ends distad of the m-cu crossvein; M_{1+2} with a broad + curve near its apical end. (= _Neomesembrina_ Schnabl. = + _Metamesembrina_ Town). _M. meridiana._ Europe. + _Mesembrina_ Meigen + + ff. R_1 ends proximad of the m-cu crossvein. + + g. Eyes pilose, sometimes sparsely in the female. + + h. Female with two or three stout orbital setæ; the hind + metatarsus of the male thickened below at the base and + penicillate. _D. pratorum._ Europe. _Dasyphora_ R. + D.[M] + + hh. Neither sex with orbital setæ. + + i. Abdomen without macrochætæ; arista plumose. _C. + asiatica._ Eastern Hemisphere. _Cryptolucilia_ B. B. + + ii. Abdomen with strong macrochætæ; arista very + short-plumose, nearly bare. _B. tachinina._ Brazil. + _Reinwardtia_ B. B. + + gg. Eyes bare. + + h. Body densely pilose; thoracic macrochætæ wanting; + middle tibiæ much elongate and bent; last section of + M_{1+2} with a gentle curve. H. (_Mesembrina_) + _mystacea, et al._, Europe and _H. solitaria_, N. + America. _Hypodermodes_ Town. + + hh. Body not densely pilose. + + i. Dorsocentrals six; last section of M_{1+2} with a + gentle curve. + + j. Inner dorsocentrals ("acrostichals") wanting; + sternopleurals arranged 1:3. _P. cyanicolor_, + _cadaverina_, etc. Europe and America _Pyrellia_ + R. D. + + jj. Inner dorsocentrals ("acrostichals") present; + sternopleurals arranged 1:2. E. _latreillii._ + North America. _Eumesembrina_ Town. + + ii. Dorsocentrals five; inner dorsocentrals present; + last section of M_{1+2} with a rounded angle; + sternopleurals arranged 1:2. _P. cornicina_ Europe + and America. (_Pseudopyrellia_ Girsch.) _Orthellia_ + R. D. + + ee. Middle tibia without a prominent bristle on the inner + surface beyond the middle. + +[Illustration: 166. Pycnosoma marginale. (×4.) After Graham-Smith.] + + f. Squamula thoracalis broadened mesad and caudad as far as + the scutellum. + + g. Sternopleural macrochætæ arranged in an equilateral + triangle; front of both sexes broad; genæ bare; + dorsocentrals six, small; wing not rilled. (To + COENOSINÆ). _Atherigona_ Rdi. + + gg. Sternopleural macrochætæ when three are present, + arranged in a right triangle. + + h. Last section of M_{1+2} with a more or less rounded + angle (fig. 163l). + + i. Eyes of the male pilose or pubescent, of the female + nearly bare; m-cu crossvein usually at or proximad + of the mid-distance between the r-m crossvein and + the bend of M_{1+2}. P. (= _Placomyia_ R. D.) + _vitripennis_. _Plaxemyia_ R. D. + + ii. Eyes bare; the m-cu crossvein always nearer to the + bend of M_{1+2} than to the r-m crossvein. + + j. Apex of the proboscis when extended reveals a + circlet of stout chitinous teeth. P. INSIGNIS + Austen, of India, bites both man and animals. (= + _Pristirhynchomyia_.) PHILÆMATOMYIA Austen + + jj. Apex of the proboscis without black teeth. + + k. Eyes of male separated by a distance equal to a + fourth the width of the head. House or typhoid + fly. M. DOMESTICA L. Widely distributed. MUSCA + L. + + kk. Eyes of the male contiguous. E. CORVINA. Europe. + EUMUSCA Town + + hh. Last section of M_{1+2} with a gentle curve (fig. + 102). + + i. Eyes pilose. + + j. Claws in the male somewhat elongated; no orbitals + in either sex; antennæ separated at the base by a + flat carina; abdomen marked with red or yellow. + _G. maculata._ Europe and America. _Graphomyia_ R. + D. + + jj. Claws short and equal in the two sexes; two or + three stout orbital macrochætæ in the female; R_1 + scarcely produced beyond the r-m crossvein; eyes + contiguous in the male. _P. obsoleta._ Brazil. + _Phasiophana_ Br. + + ii. Eyes bare; fronto-orbital macrochætæ in a double + row, antennæ contiguous at the base. + + j. One or more pairs of well developed anterior inner + dorsocentral (acrostichal) macrochætæ; seta on + extensor surface of hind tibia. M. ASSIMILIS, + STABULANS, etc. Europe and America. MUSCINA R. D. + + jj. Anterior inner dorsocentrals and the setæ; on the + extensor surface of the hind tibia wanting. _M. + micans_, etc. Europe and North America. _Morellia_ + R. D. + + ff. Squamula thoracalis not broadened mesad and caudad, not + reaching the margin of the scutellum; macrochætæ on + extensor surface of the hind tibia wanting. + + g. Eyes pubescent. _M. meditabunda._ Europe and America. + _Myiospila_ Rdi. + + gg. Eyes bare; R_1 ends before the middle of the wing. A + number of species from the tropics of both hemispheres. + _Clinopera_ V. d. W. + + bb. Hypopleuræ with a vertical row of macrochætæ. + + c. Eyes pubescent. + + d. R_1 ends about opposite the r-m crossvein; basal section of + R_{4+5} bristly nearly to the crossvein; _S. enigmatica_. + Africa. _Somalia_ Hough + + dd. R_1 ends distad of the r-m crossvein. + + e. Eastern hemisphere. Australasia. _N. ochracea_, + _dasypthalma._ _Neocalliphora_ Br. + + ee. Western Hemisphere. _T. muscinum._ Mexico. _Tyreomma_ V. d. + W. + + cc. Eyes bare. + + d. The vibrissal angle situated at a noticeable distance above the + level of the margin of the mouth. + + e. Sternopleural macrochætæ arranged in the order 1:1. + + f. Genæ with microchætæ. + + g. Body grayish, with depressed yellow woolly hair among the + macrochætæ; wings folded longitudinally over the body + when at rest. Cluster flies. _P. rudis_ and related + species, widely distributed. _Pollenia_ R. D.[N] + + gg. Body metallic blue or green. Eastern Hemisphere. + + h. Vibrissal angle placed very high above the oral margin; + a carina between the antennæ; outer posthumeral + wanting; anterior intraalar present. _T. viridaurea_. + Java. _Thelychæta_ Br. + +[Illustration: 167. Horn fly. (_a_) egg; (_b_) larva; (_c_) puparium; +(_d_) adult. (×4). Bureau of Entomology] + + hh. Vibrissal angle moderately high above the oral margin; + carina small or wanting; no post humeral macrochæta; + lower squamæ hairy above. (= _Paracompsomyia_ Hough) + (fig. 166). _Pycnosoma_ Br. + + ff. Genæ bare. _S. terminata_. Eastern Hemisphere + _Strongyloneura_ Bigot + + ee. Sternopleurals arranged 2:1. + + f. Body metallic green or blue, with gray stripes; genæ hairy + to the lower margin; post humerals often wanting; lower + squamæ bare above. (= _Compsomyia_ Rdi.). CHRYSOMYIA R. D. + + g. With one or two orbitals; height of bucca less than half + the height of the eye. South and east U. S. (fig. 107). + C. MARCELLARIA + + gg. No orbitals; height of bucca about a third less than + height of eye. West U. S. _C. wheeleri_ Hough + + ff. Body black or sordidly metallic greenish gray, usually + yellow pollinose or variegate; genæ at most hairy above. + _N. stygia_. Eastern Hemisphere. _Neopollenia_ Br. + + dd. Vibrissal angle situated nearly on a level of the oral margin. + + e. Species wholly blackish, bluish, or greenish metallic in + color. + + f. First section of R_{4+5} with at most three or four small + bristles at the immediate base. + + g. The bend of M_{1+2} a gentle curve; costal spine present; + cell R_5 closed, ending before the apex of the wing. _S. + cuprina._ Java. _Synamphoneura_ Bigot + + gg. Bend of M_{1+2} angular; or the insect differs in other + characters; dorsal surface of the squamula thoracalis + hairy (except in _Melinda_); arista plumose only on the + basal two-thirds (except usually in _Calliphora_ and + _Eucalliphora_). + +[Illustration: 168. Head of horn-fly (Lyperosia irritans); (_a_) female; +(_b_) male; (_c_) lateral aspect of female. Bureau of Entomology] + + h. Arista plumose only on the basal two-thirds. + + i. Base of the antennæ ventrad of the middle of the eye; + eyes of the male nearly contiguous; genæ hairy; + second abdominal segment with median marginal + macrochætæ; two, rarely three, postsutural intraalar + macrochætæ. + + j. Squamula thoracalis dorsally with long black hairs; + male hypopygium two-segmented, large, projecting; + claws and pulvilli of the male elongate; three + strong sternopleural macrochætæ; genæ at least + half the width of the eye; buccæ (cheeks) half the + height of the eyes; oviviparous. _O. sepulcralis._ + Europe. _Onesia_ R. D. + + jj. Dorsal surface of the squamula thoracalis bare; + male hypopygium small, scarcely projecting below; + claws and pulvilli not elongate; two stout + sternopleural macrochætæ, sometimes with a + delicate one below the anterior; genæ nearly + linear in the male; buccæ about a third of the eye + height; oviparous. _M. cærulea_. Europe. + _Melinda_. R. D. + +[Illustration: 169. Lateral and dorsal aspects of the thorax, and +frontal aspect of the head of a muscoidean fly, with designations of the +parts commonly used in taxonomic work.] + + ii. Base of the antennæ dorsad of the middle of the eye; + eyes of both sexes distinctly separated; dorsal + surface of the squamula thoracalis with black hairs; + two postsutural intraalar macrochætæ. + + j. Hypopygium of the male large, with a pair of + slightly curved forceps whose ends are concealed + in a longitudinal slit in the fifth ventral + sclerite; third posterior inner dorso-central + (acrostichal) macrochætæ absent; anterior + intraalar rarely present; abdomen usually not + pollinose; the second segment without median + marginal macrochætæ; face yellow. _C. mortuorum_, + _cadaverina_, and related species. Both + hemispheres. _Cynomyia_ R. D.[O] + +[Illustration: 170. Sepsis violacea; puparium and adult. (See page 297.) +After Howard.] + + jj. Three pairs of posterior inner dorsocentrals + (acrostichals) present; second abdominal segment + with a row of marginal macrochætæ; genæ hairy, at + least above. + + k. Hypopygium of the male with a projecting style. + _S. stylifera_. Europe. _Steringomyia_ Pok. + + + kk. Hypopygium of the male without style. _A. + stelviana_ B. B. _Acrophaga_ B. B. + + hh. Arista usually plumose nearly to the tip; posterior + dorsocentrals and inner dorsocentrals (acrostichals) + well developed; dorsal surface of the squamula + thoracalis hairy; abdomen metallic and usually + pollinose; genæ hairy. + + i. With one pair of ocellar macrochætæ. C. VOMITORIA, + ERYTHROCEPHALA, VIRIDESCENS, and related species. + Both hemispheres. CALLIPHORA R. D. + + ii. With two strong pairs of ocellar macrochætæ. _E. + latifrons._ Pacific slope of the U. S. + _Eucalliphora_ Town. + + ff. First section of R_{4+5} bristly near or quite half way to + the small crossvein; dorsal surface of the squamula + thoracalis is bare; the hypopygium of the male is + inconspicuous. + + g. Genæ bare; posterior inner and outer dorsocentrals + distinct and well developed. _L. cæsar_, _sericata_, + _sylvarum_, and related species. Widely distributed in + both hemispheres (fig. 103). LUCILIA R. D. + + gg. Genæ with microchætæ, at least down to the level of the + base of the arista. + + h. Mesonotum flattened behind the transverse suture. + + i. Posterior dorsocentrals inconstant and unequally + developed; one pair of posterior inner + dorsocentrals. _P. terrænovæ._ North America. + _Protophormia_ Town. + + ii. Posterior dorsocentrals well developed, the inner + dorsocentrals (acrostichals) unequally developed. + _P. azurea_, _chrysorrhoea_, etc. Europe and America. + _Protocalliphora_ Hough + + hh. Mesonotum not flattened behind the transverse suture; + posterior inner and outer dorsocentrals inconstant and + unequally developed. _P. regina._ Europe and America. + _Phormia_ R. D. + + ee. Species more or less rufous or yellow in color. + + f. Anterior dorsocentrals wanting; first section of the + R_{4+5} at most only bristly at the base, bend near apex + of M_{1+2} rectangular, R_1 ends over the crossvein; + fronto-orbital macrochæta absent; eyes of the male + contiguous. _C. semiviridis._ Mexico. _Chloroprocta_ + V. d. W + + ff. With another combination of characters. + + g. Body robust, of large size, abdomen elongate, not round; + genæ with several ranges of microchætæ; vibrissal ridges + strongly convergent; abdomen with well developed + macrochætæ; costal spine usually absent; eyes of the + male widely separated. + +[Illustration: 171. Stigmata of the larvæ of Muscoidea. Third instar. +(_a_) Cynomyia cadaverina; (_b_) Phormia regina; (_c_) Chrysomyia +macellaria; (_d_) Musca domestica; (_e_) Sarcophaga sp.; (_f_) Oestris +ovis; (_g_) Gastrophilus equi; (_h_) Sarcophaga sp.; (_i_) Pegomyia +vicina; (_j_) Protocalliphora azurea; (_k_) Hypoderma lineata; (_l_) +Muscina stabulans. Magnification for f, g, and k, ×25; all others, × +50.] + + h. Peristome broad, pteropleural macrochætæ distinct; one + or two sternopleurals; in the female a single orbital + macrochæta; last abdominal segment without discal + macrochætæ; hypopygial processes of the male with a + long stylet; second abdominal segment of the female + sometimes much elongate. A. LUTEOLA (fig. 86). + Africa. The sub-genus _Choeromyia_ Roub. is included + here. AUCHMEROMYIA B. B. + + hh. Peristome narrow; no pteropleurals, two + sternopleurals; two orbitals in the female; second + segment not elongate; the fourth with two well + developed discal macrochætæ. B. DEPRESSA. Africa. + BENGALIA R. D + + gg. With another combination of characters. + + h. Costal spine present; body in part black; antennæ + noticeably shorter than the epistome, inserted above + the middle of the eye and separated from each other by + a carina; abdominal segments with marginal macrochætæ; + sternopleurals 2:1 or 1:1. _Paratricyclea_ Villen. + + hh. Costal spine not distinct, or if present, insect + otherwise different. + + i. Genæ with several ranges of microchætæ; vibrissal + ridges strongly converging; peristome broad; arista + moderately plumose; sternopleurals usually 1:1; + color entirely testaceous. C. ANTHROPOPHAGA (fig. + 87) and GRUNBERGI. Africa. CORDYLOBIA Grünb. + + ii. Genæ bare or with but one range of setæ; vibrissal + ridges less converging; peristome narrow; arista + long plumose. + + j. Genæ with a single row of microchætæ. + + k. Sternopleurals 2:1; color entirely testaceous. + _Ochromyia_ Macq.[P] + + kk. Sternopleurals 1:1. _P. varia_ Hough. Africa. + _Parochromyia_ Hough + + jj. Genæ bare. + + k. Basal section of R_{4+5} bristly only at the + immediate base, distally M_{1+2} with a broad + curve; distal portion of the abdomen metallic; + sternopleurals usually 1:1, rarely 2:1. _M. + æneiventris_ Wd. Tropic America. + _Mesembrinella._ G. T. + + kk. R_{4+5} bristly at least nearly half way to the + small crossvein; sternopleurals 1:1. + + l. Macrochætæ of the abdomen marginal; neither sex + with orbitals; no carina between the base of + the antennæ; three pairs of presutural inner + dorsocentrals. Eastern hemisphere. _T. + ferruginea._ _Tricyclea V. d. W_. (= + _Zonochroa_ B. B. according to Villeneuve + 1914). + + ll. Abdomen without macrochætæ; wing usually with + a marginal streak and gray markings. Brazil. + Hemilucilia B. B. + +[Illustration: 172. Left hand stigmata of the larvæ of muscoidea. Third +instar. (_a_) Lucilia cæsar; (_b_) Calliphora vomitoria; (_c_) Stomoxys +calcitrans; (_d_) Orthellia cornicina; (_e_) Pyrellia cadavarina; +(_f_) Hæmatobia irritans; (_g_) Mesembrina mystacea; (_h_) Mesembrina +meridiana; (_i_) Myospila meditabunda; (_j_) Mydæa urbana; (_k_) +Polietes albolineata; (_l_) Polietes lardaria; (_m_) Morellia hortorum; +(_n_) Hydrotæa dentipes; (_o_) Hebecnema umbratica; (_p_) H. vespertina; +(_q_) Limnophora septemnotata; (_r_) Muscina stabulans. (_a_ and _b_) +after MacGregor; (_d_) after Banks; all others after Portchinsky. +Magnification varies. The relative distance to the median line is +indicated in each figure.] + + +SIPHONAPTERA. Fleas + +Adapted from a table published by Oudemans. + +a. Elongated fleas, with jointed (articulated) head, with combs + (ctenidia) on head and thorax; with long, oval, free-jointed + flagellum of the antenna (fig. 92d). Suborder FRACTICIPATA + + b. With ctenidia in front of the antennæ and on the genæ (cheeks); + maxillæ with acute apices; labial palpi five-segmented, + symmetrical; eyes poorly developed or wanting. On rodents. + HYSTRICHOPSYLLIDÆ + + c. Abdominal segments without ctenidia. + + d. Post-tibial spines in pairs and not in a very close set row; + head with ctenidia. _Ctenophthalmus_ Kol. + + dd. Post-tibial spines mostly single and in a close set row. + _Ctenopsyllus_ and _Leptopsyllus_. The last genus has recently + been erected for _L. musculi_, a widely distributed species + occurring on rats and mice. + + cc. Abdominal segments with one or more ctenidia; post-tibial + spines in numerous, short, close-set transverse rows on + posterior border with about four spines in each row. _H. + americana._ _Hystrichopsylla_ Taschenb. + + bb. With only two pairs of subfrontal ctenidia; labial palpi + five-segmented, symmetrical; eyes vestigial or wanting. On bats. + (= ISCHNOPSYLLIDÆ). NYCTERIDIPSYLLIDÆ + + With more or less blunt maxilla; all tibiæ with notch; a single + antepygidial bristle; metepimeron without ctenidium. _N. + crosbyi_ from Missouri was found on bats. Rothschild suggests + that this is probably the same as _N. insignis_. (= + _Ischnopsyllus_ = _Ceratopsyllus_), _Nycteridiphilus_ + +aa. Head not jointed, i.e. the segments coalescent, traces of the + segmentation still being visible in the presence of the vertex + tubercle, the falx (sickle-shaped process), and a suture. Suborder + INTEGRICIPITA + + b. Flagellum of the antennæ long and oval. + + c. Usually elongate fleas, with a free-segmented flagellum of the + antenna; thorax not shorter than the head, longer than the first + tergite. + + d. Genæ of the head and the pronotum with ctenidia. NEOPSYLLIDÆ + + e. Labial palpi four or five-segmented; symmetrical; hind coxæ + with patch of spines inside; row of six spatulate spines on + each side in front of the antennæ. _C. ornate_ found on a + California mole. _Corypsylla_ + + ee. Labial palpi two-segmented, transparent, membranous. On + hares. _Spilopsyllus_ Baker + + dd. No ctenidium on the head. + + e. Pronotum with ctenidium. DOLICHOPSYLLIDÆ + + f. Labial palpi five-segmented, symmetrical. + + g. Antepygidial bristles one to three; eyes present. + + h. Inner side of hind coxæ distally with a comb of minute + teeth; falx present. On rodents and carnivores. + _Odontopsyllus_ Baker + + hh. Inner side of hind coxæ without comb or teeth. Many + North American species on rodents. CERATOPHYLLUS + Curtis + + gg. Antepygidial bristles five on each side; eyes absent; + suture white. _D. stylosus_ on rodents. _Dolichopsyllus_ + Baker + + ff. Labial palpi four or five-segmented; asymmetrical + (membranous behind), apex acute. _Hoplopsyllus anomalus_ + found on Spermophiles in Colorado. HOPLOPSYLLIDÆ + + ee. Pronotum without ctenidium. _Anomiopsyllus californicus_ and + _nudatus_ on rodents. ANOMIOPSYLLIDÆ + + cc. Very short fleas; flagellum of the antenna with pseudo-segments + coalescent; thorax much shorter than the head and than the first + tergite. HECTOPSYLLIDÆ + + Flagellum of the antenna with six coalescent pseudo-segments; + maxilla blunt. The chigger on man (fig. 93). D. PENETRANS. (= + RHYNCHOPRION = SARCOPSYLLA) DERMATOPHILUS Guérin + + bb. Flagellum short, round, free portion of the first segment shaped + like a mandolin. + + c. Thorax not shorter than the head, longer than the first tergite; + flagellum either with free segments or in part with the segments + coalescent. + + d. Head and pronotum with ctenidium; labial palpi asymmetrical. + ARCHÆOPSYLLIDÆ + + With four subfrontal, four genal, and one angular ctenidia. + Widely distributed. CTENOCEPHALUS Kol. + + e. Head rounded in front (fig. 92a). Dog flea. C. CANIS + + ee. Head long and flat (fig. 92b). Cat flea. C. FELIS + + dd. Neither head nor pronotum with ctenidium. Labial palpi + asymmetrical, membranous behind. PULICIDÆ + + e. Mesosternite narrow, without internal rod-like thickening + from the insertion of the coxæ upwards. Human flea, etc. + PULEX L. + + ee. Mesosternite broad with a rod-like internal thickening from + the insertion of the coxæ upwards (fig. 89). X. + (LOEMOPSYLLA) CHEOPIS, plague or rat flea. XENOPSYLLA + + cc. Thorax much shorter than the head and than the first tergite. + ECHIDNOPHAGIDÆ. E. GALLINACEA, the hen flea also attacks man + (fig. 96). (= ARGOPSYLLA = XESTOPSYLLA) ECHIDNOPHAGA Olliff. + + +FOOTNOTES: + +[E] Adapted from Banks, Nuttall, Warburton, Stiles, _et al._ + +[F] Dr. C. W. Stiles considers the species which is responsible for +spotted fever distinct from the _venustus_ of Banks, separating it as +follows: + +Goblet cells about 75 in the male or 105 in the female. Texas. _D. +venustus._ + +Goblet cells 157 in the male, or 120 in the female; stigmal plate shaped +as shown in the figure (figs. 150 a, b). Montana, etc. D. ANDERSONI. + +[G] Professor C. R. Crosby who has been working upon certain capsids +states that he and his assistant have been bitten by LYGUS PRATENSIS, +the tarnished plant bug, by CHLAMYDATUS ASSOCIATUS and by ORTHOTYLUS +FLAVOSPARSUS, though without serious results. + +[H] Species marked with an * are known to transmit malaria. Species +found only in tropical North America and not known to carry malaria have +been omitted from this table, but all found in the United States are +included. + +[I] This table to the North American genera of the Tabanidæ is adapted +from one given by Miss Ricardo. + +[J] The classification of the Muscoidea as set forth by Schiner and +other earlier writers has long been followed, although it is not +satisfactory, being admittedly more or less artificial. Within the last +two or three decades several schemes have been advanced, that of Brauer +and Bergenstamm and of Girschner, with the modifications of Schnabl and +Dziedzicki having obtained most favor in Europe. Townsend, in 1908, +proposed a system which differs from Girschner's in some respects, but +unfortunately it has not yet been published in sufficient detail to +permit us to adopt it. From considerations of expediency we use here the +arrangement given in Aldrich's Catalogue of North American Diptera, +though we have drawn very freely upon Girschner's most excellent paper +for taxonomic characters to separate the various groups. + +It may sometimes be found that a species does not agree in all the +characters with the synopsis; in this case it must be placed in the +group with which it has the most characters in common. + +[K] There are several genera of flies of the family _Cordyluridæ_; (i.e. +_Acalyptratæ_) which might be placed with the _Anthomyiidæ_ (i.e. +_Calyptratæ_), owing to the relatively large size of their squamæ. As +there is no single character which will satisfactorily separate all +doubtful genera of these two groups we must arbitrarily fix the limits. +In general those forms on the border line having a costal spine, or +lower squama larger than the upper, or the lower surface of the +scutellum more or less pubescent, or the eyes of the male nearly or +quite contiguous, or the eyes hairy, or the frontal setæ decussate in +the female; or any combination of these characters may at once be placed +with the _Anthomyiidæ_. Those forms which lack these characteristics and +have at least six abdominal segments (the first and second segments +usually being more or less coalescent) are placed with the Acalyptrates. +There are other acalyptrates with squamæ of moderate size which have +either no vibrissæ, or have the subcosta either wholly lacking or +coalescent in large part with R_1 or have spotted wings; they, therefore +will not be confused with the calyptrates. + +[L] _Pachymyia_ Macq. is closely related to _Stomoxys_. It differs in +having the arista rayed both above and below. _P. vexans_, Brazil. + +[M] The genus _Eudasyphora_ Town. has recently been erected to contain +_D. lasiophthalma_. + +[N] _Nitellia_, usually included in this genus has the apical cell +petiolate. _Apollenia_ Bezzi, has recently been separated from +_Pollenia_ to contain the species _P. nudiuscula_. Both genera belong to +the Eastern hemisphere. + +[O] The following three genera are not sufficiently well defined to +place in this synopsis. In color and structural characters they are +closely related to _Cynomyia_ from which they may be distinguished as +follows. _Catapicephala_ Macq., represented by the species _C. +splendens_ from Java, has the setæ on the facial ridges rising to the +base of the antennæ and has median marginal macrochætæ on the abdominal +segments two to four: _Blepharicnema_ Macq., represented by _B. +splendens_ from Venezuela has bare genæ, oral setæ not ascending; tibiæ +villose; claws short in both sexes; _Sarconesia_ Bigot with the species +_S. chlorogaster_ from Chile, setose genæ; legs slender, not villose; +claws of the male elongate. + +[P] _Plinthomyia_ Rdi. and _Hemigymnochæta_ Corti are related to +_Ochromyia_, though too briefly described to place in the key. + + + + +APPENDIX + +HYDROCYANIC ACID GAS AGAINST HOUSEHOLD INSECTS + + +The following directions for fumigating with hydrocyanic acid +gas are taken from Professor Herrick's circular published by the +Cornell Reading Course: + +Hydrocyanic acid gas has been used successfully against household +insects and will probably be used more and more in the future. +It is particularly effective against bed-bugs, and cockroaches, but +because _it is such a deadly poison it must be used very carefully_. + +The gas is generated from the salt potassium cyanid, by treating +it with sulfuric acid diluted with water. Potassium cyanid is a +most poisonous substance and the gas emanating from it is also +deadly to most, if not all, forms of animal life. The greatest care +must always be exercised in fumigating houses or rooms in buildings +that are occupied. Before fumigation a house should be vacated. +It is not necessary to move furniture or belongings except brass or +nickel objects, which may be somewhat tarnished, and butter, milk, +and other larder supplies that are likely to absorb gas. If the nickel +and brass fixtures or objects are carefully covered with blankets +they will usually be sufficiently protected. + +There may be danger in fumigating one house in a solid row of +houses if there is a crack in the walls through which the gas may find +its way. It also follows that the fumigation of one room in a house +may endanger the occupants of an adjoining room if the walls between +the two rooms are not perfectly tight. It is necessary to keep +all these points in mind and to do the work deliberately and thoughtfully. +The writer has fumigated a large college dormitory of 253 +rooms, once a year for several years, without the slightest accident +of any kind. In order to fumigate this building about 340 pounds +of cyanid and the same amount of sulfuric acid were used each time. +In addition to this, the writer has fumigated single rooms and smaller +houses with the gas. In one instance the generating jars were too +small; the liquid boiled over and injured the floors and the rugs. +Such an accident should be avoided by the use of large jars and by +placing old rugs or a quantity of newspapers beneath the jars. + + +THE PROPORTIONS OF INGREDIENTS + +Experiments and experience have shown that the potassium +cyanid should be ninety-eight per cent pure in order to give satisfactory +results. The purchaser should insist on the cyanid being of +at least that purity, and it should be procurable at not more than +forty cents per pound. The crude form of sulfuric acid may be used. +It is a thickish, brown liquid and should not cost more than four or +five cents a pound. If a room is made tight, one ounce of cyanid for +every one hundred cubic feet of space has been shown to be sufficient. +It is combined with the acid and water in the following proportions: + + Potassium cyanid 1 ounce + Commercial sulfuric acid 1 fluid ounce + Water 3 fluid ounces + + +A SINGLE ROOM AS AN EXAMPLE + +Suppose a room to be 12 by 15 by 8 feet. It will contain 12 × 15 × 8, or +1440 cubic feet. For convenience the writer always works on the basis of +complete hundreds; in this case he would work on the basis of 1500 cubic +feet, and thus be sure to have enough. The foregoing room, then, would +require 15 ounces of cyanid, 15 ounces of sulfuric acid, and 45 ounces +of water. The room should be made as tight as possible by stopping all +the larger openings, such as fireplaces and chimney flues, with old rags +or blankets. Cracks about windows or in other places should be sealed +with narrow strips of newspaper well soaked in water. Strips of +newspaper two or three inches wide that have been thoroughly soaked in +water may be applied quickly and effectively over the cracks around the +window sash and elsewhere. Such strips will stick closely for several +hours and may be easily removed at the conclusion of the work. + +While the room is being made tight, the ingredients should be measured +according to the formula already given. The water should be measured and +_poured first_ into a stone jar for holding at least two gallons. The +jar should be placed in the middle of the room, with an old rug or +several newspapers under it in order to protect the floor. + +The required amount of sulfuric acid should then be poured rather slowly +into the water. _This process must never be reversed; that is, the acid +must never be poured into the jar first._ The cyanid should be weighed +and put into a paper bag beside the jar. All hats, coats, or other +articles that will be needed before the work is over should be removed +from the room. When everything is ready the operator should drop the bag +of cyanid gently into the jar, holding his breath, and should walk +quickly out of the room. The steam-like gas does not rise immediately +under these conditions, and ample time is given for the operator to walk +out and shut the door. If preferred, however, the paper bag may be +suspended by a string passing through a screw eye in the ceiling and +then through the keyhole of the door. In this case the bag may be +lowered from the outside after the operator has left the room and closed +the door. + +The writer has most often started the fumigation toward evening and left +it going all night, opening the doors in the morning. The work can be +done, however, at any time during the day and should extend over a +period of five or six hours at least. It is said that better results +will be obtained in a temperature of 70° F., or above, than at a lower +degree. + +At the close of the operation the windows and doors may be opened from +the outside. In the course of two or three hours the gas should be +dissipated enough to allow a person to enter the room without danger. +The odor of the gas is like that of peach kernels and is easily +recognized. The room should not be occupied until the odor has +disappeared. + + +FUMIGATING A LARGE HOUSE + +The fumigation of a large house is merely a repetition, in each room and +hall, of the operations already described for a single room. All the +rooms should be made tight, and the proper quantities of water and +sulfuric acid should be measured and poured into jars placed in each +room with the cyanid in bags besides the jars. When all is ready, the +operator should _go to the top floor and work downward_ because the gas +is lighter than air and tends to rise. + + +PRECAUTIONS + +The cyanid should be broken up into small pieces not larger than small +eggs. This can best be done on a cement or brick pavement. It would be +advantageous to wear gloves in order to protect the hands, although the +writer has broken many pounds of cyanid without any protection on the +hands. Wash the hands thoroughly at frequent intervals in order to +remove the cyanid. + +The operations of the work must be carried out according to directions. + +The work should be done by a calm, thoughtful and careful person--best +by one who has had some experience. + +Conspicuous notices of what has been done should be placed on the doors, +and the doors should be locked so that no one can stray into the rooms. + +The gas is lighter than air, therefore one should always begin in the +rooms at the top of the house and work down. + +After fumigation is over the contents of the jar should be emptied into +the sewer or some other safe place. The jars should be washed thoroughly +before they are used again. + +_It must be remembered that cyanid is a deadly poison_; but it is very +efficient against household insects, if carefully used, and is not +particularly dangerous when properly handled. + + +LESIONS PRODUCED BY THE BITE OF THE BLACK-FLY + +While this text was in press there came to hand an important paper +presenting a phase of the subject of black fly injury so different from +others heretofore given that we deem it expedient to reproduce here the +author's summary. The paper was published in _The Journal of Cutaneous +Diseases_, for November and December, 1914, under the title of "A +Clinical, Pathological and Experimental Study of the Lesions Produced by +the Bite of the Black Fly (_Simulium venustum_)," by Dr. John Hinchman +Stokes, of the University of Michigan. + + +RESUME AND DISCUSSION OF EXPERIMENTAL FINDINGS + +The principal positive result of the work has been the experimental +reproduction of the lesion produced by the black-fly in characteristic +histological detail by the use of preserved flies. The experimental +lesions not only reproduced the pathological pictures, but followed a +clinical course, which in local symptomatology especially, tallied +closely with that of the bite. This the writer interprets as +satisfactory evidence that the lesion is not produced by any living +infective agent. The experiments performed do not identify the nature of +the toxic agent. Tentatively they seem to bring out, however, the +following characteristics. + +1. The product of alcoholic extraction of flies do not contain the toxic +agent. + +2. The toxic agent is not inactivated by alcohol. + +3. The toxic agent is not destroyed by drying fixed flies. + +4. The toxic agent is not affected by glycerin, but is, if anything, +more active in pastes made from the ground fly and glycerin, than in the +ground flies as such. + +5. The toxic agent is rendered inactive or destroyed by hydrochloric +acid in a concentration of 0.25%. + +6. The toxic agent is most abundant in the region of the anatomical +structures connected with the biting and salivary apparatus (head and +thorax). + +7. The toxic agent is not affected by a 0.5% solution of sodium +bicarbonate. + +8. The toxic agent is not affected by exposure to dry heat at 100° C. +for two hours. + +9. The toxic agent is destroyed or rendered inactive in alkaline +solution by a typical hydrolytic ferment, pancreatin. + +10. Incomplete experimental evidence suggests that the activity of the +toxic agent may be heightened by a possible lytic action of the blood +serum of a sensitive individual, and that the sensitive serum itself may +contain the toxic agent in solution. + +These results, as far as they go (omitting No. 10), accord with Langer's +except on the point of alcoholic solubility and the effect of acids. The +actual nature of the toxic agent in the black-fly is left a matter of +speculation. + +The following working theories have suggested themselves to the writer. +First, the toxin may be, as Langer believes in the case of the bee, an +alkaloidal base, toxic as such, and neutralized after injection by +antibodies produced for the occasion by the body. In such a case the +view that a partial local fixation of the toxin occurs, which prevents +its immediate diffusion, is acceptable. Through chemotactic action, +special cells capable of breaking up the toxin into harmless elements +are attracted to the scene. Their function may be, on the other hand, to +neutralize directly, not by lysis. This would explain the rôle of the +eosinophiles in the black-fly lesion. If their activities be essential +to the destruction or neutralization of the toxin, one would expect them +to be most numerous where there was least reaction. This would be at the +site of a bite in an immune individual. A point of special interest for +further investigation, would be the study of such a lesion. + +Second, it is conceivable that the injected saliva of the fly does not +contain an agent toxic as such. It is possible, that like many foreign +proteins, it only becomes toxic when broken down. The completeness and +rapidity of the breaking down depends on the number of eosinophiles +present. In such a case immunity should again be marked by intense +eosinophilia. + +[Illustration: 173. Fifth day mature lesion. Lower power drawing showing +papillary oedema and infiltrate in the region of the puncture. After Dr. +J .H. Stokes] + +Third, lytic agents in the blood serum may play the chief rôle in the +liberation of the toxic agent from its non-toxic combination. An immune +individual would then be one whose immunity was not the positive one of +antibody formation, but the negative immunity of failure to metabolize. +An immune lesion in such a case might be conceived as presenting no +eosinophilia, since no toxin is liberated. If the liberation of the +toxin is dependent upon lytic agents present in the serum rather than in +any cellular elements, a rational explanation would be available for the +apparent results (subject to confirmation) of the experiment with +sensitive and immune sera. In this experiment it will be recalled that +the sensitive serum seemed to bring out the toxicity of the ground +flies, and the serum itself seemed even to contain some of the dissolved +or liberated toxin. The slowness with which a lesion develops in the +case of the black-fly bite supports the view of the initial lack of +toxicity of the injected material. The entire absence of early +subjective symptoms, such as pain, burning, etc., is further evidence +for this view. It would appear as if no reaction occurred until lysis of +an originally non-toxic substance had begun. Regarding the toxin itself +as the chemotactic agent which attracts eosinophiles, its liberation in +the lytic process and diffusion through the blood stream attracts the +cells in question to the point at which it is being liberated. Arriving +upon the scene, these cells assist in its neutralization. + +The last view presented is the one to which the author inclines as the +one which most adequately explains the phenomena. + +A fourth view is that the initial injection of a foreign protein by the +fly (i.e., with the first bite) sensitizes the body to that protein. Its +subsequent injection at any point in the skin gives rise to a local +expression of systematic sensitization. Such local sensitization +reactions have been described by Arthus and Breton, by Hamburger and +Pollack and by Cowie. The description of such a lesion given by the +first named authors, in the rabbit, however, does not suggest, +histopathologically at least, a strong resemblance to that of the +black-fly. Such an explanation of many insect urticariæ deserves further +investigation, however, and may align them under cutaneous expressions +of anaphylaxis to a foreign protein injected by the insect. Depending on +the chemical nature of the protein injected, a specific chemotactic +reaction like eosinophilia may or may not occur. Viewed in this light +the development of immunity to insect bites assumes a place in the +larger problem of anaphylaxis. + +[Illustration: 174. Experimental lesion produced from alcohol-fixed +flies, dried and ground into a paste with glycerin. After Dr. J. H. +Stokes] + + +SUMMARY + +In order to bring the results of the foregoing studies together, the +author appends the following résumé of the clinical data presented in +the first paper. + +The black-fly, _Simulium venustum_, inflicts a painless bite, with +ecchymosis and hæmorrhage at the site of puncture. A papulo-vesicular +lesion upon an urticarial base slowly develops, the full course of the +lesion occupying several days to several weeks. Marked differences in +individual reaction occur, but the typical course involves four stages. +These are, in chronological order, the papular stage, the vesicular or +pseudovesicular, the mature vesico-papular or weeping papular stage and +the stage of involution terminating in a scar. The papule develops in +from 3 to 24 hours. The early pseudovesicle develops in 24 to 48 hours. +The mature vesico-papular lesion develops by the third to fifth day and +may last from a few days to three weeks. Involution is marked by +cessation of oozing, subsidence of the papule and scar-like changes at +the site of the lesion. The symptoms accompanying this cycle consist of +severe localized or diffused pruritus, with some heat and burning in the +earlier stages if the oedema is marked. The pruritus appears with the +pseudovesicular stage and exhibits extraordinary persistence and a +marked tendency to periodic spontaneous exacerbation. The flies tend to +group their bites and confluence of the developing lesions in such cases +may result in extensive oedema with the formation of oozing and +crusted plaques. A special tendency on the part of the flies to attack +the skin about the cheeks, eyes and the neck along the hair line and +behind the ears, is noted. In these sites inflammation and oedema may +be extreme. + +A distinctive satellite adenopathy of the cervical glands develops in +the majority of susceptible persons within 48 hours after being bitten +in the typical sites. This adenopathy is marked, discrete and painful, +the glands often exquisitely tender on pressure. It subsides without +suppuration. + +Immunity may be developed to all except the earliest manifestations, by +repeated exposures. 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Science, n. s. xxxviii, p. 513. + +HONEIJ, J. A. and PARKER, R. R. 1914. Leprosy: flies in relation to the + transmission of the disease. Journ. Med. Research, Boston, xxx, p. + 127-130. + +HOOKER, W. A. 1908 a. Life history, habits, and methods of study of the + Ixodoidea. Jour. Econ. Ent. i, p. 34-51. + +---- 1908 b. A review of the present knowledge of the rôle of ticks in + the transmission of disease. ibid., i, p. 65-76. + +HOPE, F. W. 1837. On insects and their larvæ occasionally found in the + human body. Trans. Ent. Soc., London, ii, p. 256-271. + +HOUGH, G. DE N. 1899 a. Synopsis of the Calliphorinæ of the United + States. Zoological Bulletin, ii, p. 283-290. + +---- 1899 b. Some Muscinæ of North America, Biological Bulletin i, p. + 19-33. + +---- 1899 c. Some North American Genera of Calliphorinæ. Entom. News, + x, p. 62-66. + +HOVARTH, G. 1912. Revision of the American Cimicidæ. Ann. Mus. Nat. + Hungarici, x, p. 257-262. + +HOWARD, C. W. 1908. A list of the ticks of South Africa, with + descriptions and keys to all the forms known. Ann. Transvaal Mus. 1, + p. 73-170. + +HOWARD, C. W. and CLARK, P. F. 1912. Experiments on insect transmission + of the virus of poliomyelitis. Journ. Exper. Med. xvi, p. 805-859. + +HOWARD, L. O. 1899. Spider bites and kissing bugs. Pop. Sci. Mo. lv, p. + 31-42. + +---- 1900. A contribution to the study of the insect fauna of human + excrement. Proc. Wash. Acad. Sci. ii, p. 541-604. + +---- 1901. Mosquitoes, how they live, how they carry disease, how they + are classified, how they may be destroyed. 8vo. New York, Doubleday, + Page & Co. (xv + 241 p.) + +---- 1909. Economic loss to the people of the United States, through + insects that carry disease. U. S. Dept. Agric. Bur. of Ent. Bul. 78, + p. 1-40. + +HOWARD, L. O., DYAR, H. G. and KNAB, F. 1913-. The mosquitoes of North + and Central America and the West Indies. Vol. I. A general + consideration of mosquitoes, their habits, and their relations to the + human species. 4^o. Carnegie Institution of Washington (vii + 520 + p.). + +HOWARD, L. O. and MARLATT, C. L. 1902. The principal household insects + of the United States. U. S. Dept. Agric., Bur. Ent. Bul. 4. + +HUEBNER, W. 1907. Ueber das Pfeilgift der Kalahari. Arch. exper. Path. + und Pharm., lvii, p. 358-366. + +HUNTER, S. J. 1913. Pellagra and the sand-fly. Jour. econ. Ent. vi, p. + 96-99. + +HUNTER, W. D. 1913. American interest in medical entomology. Jour. econ. + Ent. vi, p. 27-39. + +HUNTER, W. D. and BISHOPP, F. C. 1910. Some of the more important ticks + of the United States. U. S. Dept. Agric. Yearbook 1910, p. 219-230, + pls. xv-xvi. + +---- 1911. The Rocky Mountain spotted fever tick. With special reference + to the problem of its control in the Bitter Root Valley in Montana. + U. S. Dept. Agric., Bur. Ent. Bul. 105, p. 1-47. + +HUTCHISON, R. H. 1914. The migratory habit of housefly larvæ as + indicating a favorable remedial measure. An account of progress. U. + S. Dept. Agric., Bul. 14, p. 1-11. + +JENNINGS, A. H. 1914. Summary of two years' study of insects in relation + to pellagra. Journ. of Parasitology, i, p. 10-21. + +JENNINGS, A. H. and KING, W. V. 1913. One of the possible factors in the + causation of pellagra. Journ. Amer. Med. Assoc., lx, p. 271-274. + +JEPSON, F. P. 1909. Notes on colouring flies for purposes of + identification. Rep't to the Local Gov't Board on Publ. Health, n. s. + 16, p. 4-9. + +JOHANNSEN, O. A. 1903. Aquatic Nematocerous Diptera. N. Y. State Mus. + Bul., 68, p. 328-448, pls. 32-50. + +---- 1905. Aquatic Nematocerous Diptera II. (Chironomidæ). ibid. 86, p. + 76-330, pls. 16-37. + +---- 1908. North America Chironomidæ. ibid., 124, p. 264-285. + +---- 1911. The typhoid fly and its allies. Maine Agric. Exp. Sta. Bul., + 401, p. 1-7. + +---- 1911. Simulium and pellagra. Insect Notes for 1910. Maine Agr. + Exper. Station. Bul. 187, p. 4. + +KELLOGG, V. L. 1915. Spider poison. Jour. of Parasitology, i, p. 107+ + +KELLY, H. A. 1907. Walter Reed and yellow fever. 8vo. New York, McClure, + Phillips & Co. (xix + 310 p.). + +KEPHART, CORNELIA F. 1914. The poison glands of the larva of the + browntail moth (Euproctis chrysorrhoea Linn.). Journ. Parasit., i, + p. + +KIEFFER, J. J. 1906. Chironomidæ. Genera Insectorum. Fasc. 42, p. 1-78. + +---- 1913. Nouv. étude sur les Chironomides de l'Indien Museum de + Calcutta. Records of the Indian Mus., ix, p. 119-197. + +KING, A. F. A. 1883. Insects and disease--mosquitoes and malaria. Pop. + Sci. Mo. xxiii, p. 644-658. + +KIRKLAND, A. H. 1907. Second annual report of the Superintendent for + suppressing the gypsy and browntail moths. 8vo. Boston. 170 p. + +KLEINE, E. 1909. Postive Infektionsversuche mit _Trypanosoma brucei_ + durch _Glossina palpalis._ Deutsche med. Wochenschr., xxxv, p. + 469-470. + + Weitere wissenschaftliche Beobachtungen über die Entwicklung von + Trypanosomen in Glossinen. ibid. p. 924-925. + + Weitere Untersuchungen über die Ætiologie der Schlafkrankheit. ibid., + p. 1257-1260. + + Weitere Beobachtungen über Tsetsefliegen und Trypanosomen. ibid., p. + 1956-1958. + +KLING, C. and LEVADITI, C. 1913. Études sur la poliomyélite aiguë + épidémique. Ann. Inst. Pasteur, xxvii, p. 718-749, 739-855. + +KNAB, F. 1912. Unconsidered factors in disease-transmission by + blood-sucking insects. Journ. Econ. Ent., v, p. 196-200. + +---- 1913 a. The species of Anopheles that transmit human malaria. Amer. + Journ. Trop. Dis. and Preventive Med., i, p. 24-43. + +---- 1913 b. Anopheles and malaria. ibid., i, p. 217. + +---- 1913 c. The life history of _Dermatobia hominis_. ibid., i, p. + 464-468. + +KNAB, F. See Howard, Dyar, and Knab. + +KOBERT, R. 1893. Lehrbuch der Intoxikationen. 4^o. Stuttgart, Enke. 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Observations on the mechanism of infection in + tick fever and on the hereditary transmission of _Spirochæta duttoni_ + in the tick. Trans. Soc. Trop. Med. Hyg., iii, p. 77-95. Abstr. in + Bul. Inst. Pasteur, viii, p. 312-313. + +---- 1910 b. On the hereditary transmission and mechanism of infection + in tick fever and on the hereditary transmission of _Spirochæta + duttoni_ in the tick. Lancet., clxxvii, p. 11. + +LINNELL, R. McC. 1914. Notes on a case of death following the sting of a + scorpion. Lancet, 1914, p. 1608-1609. + +LIVINGSTONE, D. 1857. Missionary travels and researches in South Africa. + +LUCAS, H. 1843. (note) _Latrodectus malmignatus_ Bul. Soc. Ent., France, + 1843, p. viii. + +LUDLOW, C. S. 1914. Disease bearing mosquitoes of North and Central + America, the West Indies and the Philippine Islands. War Dept., + Office of Surgeon General. Bul. No. 4, 1-96. + +LUGGER, 1896. Insects injurious in 1896. Agr. Exp. Sta. Bul. 48. p. 33 + to 270. + +MacCALLUM, W. C. 1898. On the hæmatozoan infection of birds. Journ. Exp. + Med. iii, p. 117. + +MacGREGOR, M. E. 1914. The posterior stigmata of dipterous larvæ as a + diagnostic character. Parasitology, vii, p. 176-188. + +MacLOSKIE, G. 1888. The poison apparatus of the mosquito. Amer. + Naturalist, xxii, p. 884-888. + +MALLOCH, J. P. 1913. American black-flies or Buffalo gnats. U. S. Dept. + Agric. Bur. Ent. Tech. Bul. 26, p. 1-72. + +---- 1914. Notes on North American Diptera. Bul. Illinois State Lab. + Nat. Hist., x, p. 213-243. + +MANSON, P. 1911. Tropical diseases: a manual of the diseases of warm + climates. 8^o. London, Cassell & Co. (xx + 876 p.). 4 ed. (1907). + Reprinted. + +MARCHOUX, E. and COUVY, L. 1913. Argas et spirochætes (1 mémoire). Les + granules de Leishman. Ann. Inst. Pasteur, xxvii, p. 450-480. 2 + mémoire. Le virus chez l'acarien. ibid. p. 620-643. + +MARCHOUX, E. and SELIMBENI, A. 1903. La spirillose des poules. Ann. + Inst. Pasteur, xvii, p. 569-580. + +MARCHOUX, E. and SIMOND, P. 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Ueber die Jugendzustände der _Tænia cucumerina_. + Arch. f. Naturg., xxxv, p. 62-70. + +MENSE, C. 1913. Handbuch der Tropenkrankheiten. 1 Band. 4^o. Leipzig, + Barth (xv + 295 p.) Entomological parts by A. Eysell, and by Doerr + and Russ. + +MINCHIN, E. A. 1912. An introduction to the study of the Protozoa, with + special reference to the parasitic forms. 8^o. London. Arnold (xi + + 517 p.). + +MITCHELL, EVELYN G. 1907. Mosquito life. 8vo. New York, Putmans. (xxii + + 281 p.). + +MITZMAIN, M. B. 1910. General observations on the bionomics of the + rodent and human flies. U. S. Publ. Health Service. Bul., 38, p. + 1-34. + +---- 1912. The rôle of _Stomoxys calcitrans_ in the transmission of + _Trypanosoma evansi_. Philippine Journ. Sci., vii, p. 475-519, 5 pls. + +---- 1913 a. The biology of _Tabanus striatus_ Fabricius, the horsefly + of the Philippines. ibid., vii, B. p. 197-221. + +---- 1913 b. The mechanical transmission of surra. ibid., viii, sec. B., + p. 223-229. + +---- 1914 a. Experimental insect transmission of anthrax. U. S. Public + Health Repts. xxix, p. 75-77. + +---- 1914 b. I. Collected studies on the insect transmission of + _Trypanosoma evansi_. II. Summary of experiments in the transmission + of anthrax by biting flies. U. S. Pub. Health Service, Hyg. Lab. + Bul., 94, p. 1-48. + +MIYAKE, H. and SCRIBA, J. 1893. Vorläufige Mitteilung über einen neuen + Parasit des Menschen. Berl. klin. Wochenschr., xxx, p. 374. + +MOLLERS, B. 1907. Experimentelle Studien über die Uebertragung des + Rückfallfiebers durch Zecken. Zeitschr. für Hyg. u. + Infektionskrankheiten, lviii, p. 277-286. + +MOTE, D. C. 1914. The cheese-skipper (_Piophila casei_). Ohio Naturalist + xiv, p. 309-310. + +NEIVA, A. 1910. Beiträge zur Biologie der _Conorhinus megistus_ Burm. + Memorias de Institute Oswaldo Cruz., ii, p. 206-212. + +NEVEU-LEMAIRE, M. 1907. Un nouveau cas de parasitisme accidental d'un + myriapode dans le tube digestif de l'homme, C. R. 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Egyptian Gov't + School of Medicine, Cairo, ii, p. 7-44. + + + + +INDEX + + +Abscess, 178 + +Acanthia, 87 + +Acariasis, 58 + +Acarina, 23, 58, 131, 259 + +Acarus dysenteriæ, 132 + +Accidental parasites, 131, 132, 134 + +Aedes, 194, 293 + +Aedes calopus, 182, 201, 205, 206, 208 + +Aedes cantator, 101 + +Aedes sollicitans, 101 + +Aedes tæniorhynchus, 101 + +Aerobic bacteria, 152 + +Æstivo-autumnal, 186 + +African Relapsing Fever, 230 + +Akis spinosa, 177 + +Alternation of Generations, 175 + +Amblyomma, 264 + +Amblyomma americanum, 67 + +Amblyomma cajennense, 67 + +American dog tick, 228 + +Amoeboid organism, 189 + +Anisolabis annulipes, 177 + +Anterior poliomyelitis, 241 + +Anopheles, 194, 291 + +Anopheles crucians, 199 + +Anopheles maculipennis, 182 + +Anopheles punctipennis, 198 + +Anopheles quadrimaculatus, 197 + +Anopheline, 192 + +Anthocoris, 279 + +Anthomyiidæ, 300 + +Anthomyia, 138 + +Anthrax, 165 + +Antipruritic treatment, 72 + +Ants, 42 + +Aphiochæta, 295 + +Apis mellifica, 36 + +Arachnida, 258 + +Araneida, 6 + +Argas, 64 + +Argas persicus, 63, 235, 237 + +Argasidæ, 62 + +Argopsylla, 317 + +Argus, 259 + +Arilus, 284 + +Arthropods, poisonous, 6 + +Asopia farinalis, 177 + +Assassin-bugs, 31, 219 + +Auchmeromyia, 117 + +Automeris io, 47 + +Avicularoidea, 12 + + +Babesia, 226 + +Babesia bovis, 223 + +Babesia ovis, 225 + +Babesiosis, 221-222 + +Bacilli, 170 + +Bacillus icteroides, 202, 205 + +Bacillus pestis, 166 + +Bacillus typhosus, 153 + +Back swimmers, 30 + +Bdellolarynx, 304 + +Beauperthuy, Louis Daniel, 2 + +Bed-bug, 86, 88, 90, 173, 219-220 + +Bed-bug, cone-nosed, 92 + +Blister beetles, 54 + +Belostoma, 28, 277 + +Belostoma americana, 31 + +Belostomatidæ, 30 + +Bengalia, 314 + +Bird-spiders, 10 + +Black death, 1, 166 + +Black flies, 33, 104, 247 + +Black heads, 80 + +Blaps mortisaga, 134 + +Blepharoceridæ, 286 + +Boophilus, 264 + +Boophilus annulatus, 67, 223-225 + +Bot-flies, 112 + +Blue bottle flies, 140 + +Brill's disease, 238 + +Brown-tailed moth, 48 + +Bruck, 34 + +Buthus quinquestriatus, 21 + + +Cabbage butterfly, 56 + +Calliphora, 136, 140, 312 + +Calliphora erythrocephala, 141 + +Calobata, 296 + +Camponotinæ, 43 + +Cancer, 254 + +Cantharidin, 54 + +Cantharidin poison, 55 + +Canthariasis, 134 + +Capsidæ, 280 + +Carriers, simple, 4, 144 + +Carriers of disease, 144 + +Carrion's fever, 253 + +Caterpillar rash, 45 + +Cat flea, 172 + +Cattle ticks, 222 + +Causative organism, 170 + +Cellia, 291 + +Centipedes, 25, 257 + +Ceratophyllus, 120, 316 + +Ceratophyllus acutus, 123 + +Ceratophyllus fasciatus, 122, 172, 213 + +Ceratopogon, 108 + +Cheese-fly, 137 + +Cheyletus eruditus, 271 + +Chigger, 60, 70 + +Chigoes, 126 + +Chilopoda, 25, 257 + +Chiracanthium nutrix, 18 + +Chironomidæ, 107 + +Chorioptes, 270 + +Chrysomelid, 55 + +Chrysomyia, 136, 308 + +Chrysomyia macellaria, 117, 140 + +Chrysops, 294 + +Chylous dropsy, 179 + +Chyluria, 178 + +Cicadidæ, 55 + +Cimex L., 278 + +Cimex boueti, 92 + +Cimex columbarius, 92 + +Cimex hemipterus, 91, 220 + +Cimex hirundinis, 92 + +Cimex inodorus, 92 + +Cimex lectularius, 87, 219 + +Citheronia regalis, 44 + +Clinocoris, 87 + +Coleoptera, 134, 274 + +Comedons, 80 + +Complete metamorphosis, 80 + +Compressor muscle, 20 + +Compsomyia, 117 + +Cone-nosed bed-bug, 92 + +Conjunctivitis, nodular, 52 + +Conorhinus, 282 + +Conorhinus megistus, 93, 219-220 + +Conorhinus rubrofasciatus, 220 + +Conorhinus sanguisugus, 32, 92 + +Copra itch, 72 + +Cordylobia, 118 + +Coriscus, 280 + +Coriscus subcoleoptratus, 32 + +Creeping myasis, 112 + +Crustacea, 257 + +Cryptocystis, 176 + +Cryptotoxic, 54-55 + +Cteniza sauvagei, 13 + +Ctenocephalus, 120, 172, 213, 317 + +Culex, 194, 201, 293 + +Culex pipiens, 35, 98 + +Culex quinquefasciatus, 180 + +Culex sollicitans, 200 + +Culex territans, 101 + +Culicidæ, 33, 97 + +Culicin, 34 + +Culicoides, 109, 288 + +Cyclops, 183, 257 + +Cynomyia, 136, 311 + + +Dance, St. Vitus, 8 + +Dancing mania, 8 + +Deer-flies, 110 + +Definitive host, 192 + +Demodecidæ, 78 + +Demodex, 259 + +Demodex folliculorum, 78 + +Dermacentor, 262 + +Dermacentor andersoni, 67, 228 + +Dermacentor occidentalis, 227 + +Dermacentor variabilis, 67 + +Dermacentor venustus, 24, 228 + +Dermanyssidæ, 68 + +Dermanyssus, 266 + +Dermanyssus gallinæ, 68 + +Dermatitis, 72, 77, 85 + +Dermatobia, 115, 298 + +Dermatobia cyaniventris, 163 + +Dermatophilus, 317 + +Dermatophilus penetrans, 60, 126 + +Diamphidia simplex, 55 + +Dimorphism, 65 + +Direct inoculators, 4 + +Diplopoda, 25, 257 + +Diptera, 33, 94, 274 + +Dipterous Larvæ, 135 + +Dipylidium, 175, 221 + +Dipylidium caninum, 4, 175-176 + +Dog flea, 172 + +Dracunculus, 257 + +Dracunculus medinensis, 182 + +Drosophila, 296 + +Dum-dum fever, 220 + +Dysentery, 154 + + +Ear-flies, 110 + +Earwig, 177 + +Echidnophaga, 317 + +Echinorhynchus, 185 + +Elephantiasis, 178-179 + +Empoasca mali, 33 + +Empretia, 46 + +English Plague Commission, 171 + +Epeira diadema, 18 + +Epizootic, 170 + +Eristalis, 137, 295 + +Essential hosts, 4, 165 + +Eumusca, 307 + +European Relapsing Fever, 233 + +Euproctis chrysorrhoea, 48 + +Eusimulium, 286 + + +Facultative parasites, 131 + +Fannia, 136, 138, 145, 300 + +Federal Health Service, 169 + +Fever, lenticular, 237 + African Relapsing, 230, 234 + Carrion's, 253 + dum-dum, 154 + European Relapsing, 233 + pappatici, 96 + red water, 220 + Rocky Mt. Spotted, 226 + three day, 96 + Typhus, 237 + +Filaria, 178, 221 + immitis, 182 + +Filariasis, 178 + +Flannel-moth larvæ, 44 + +Fleas, 119, 166, 213 + cat, 172 + dog, 172 + human, 172, 176 + rodent, 123, 172 + rat, 171 + +Flesope, 125 + +Formaldehyde, 91 + +Fomites, 199, 204 + +Fulgoridæ, 28 + +Fumigation, 320 + + +Gamasid, 68 + +Gangrene, 129 + +Gastrophilus, 113, 297 + +Giant crab spiders, 13 + +Giant water bugs, 30 + +Gigantorhynchus, 185 + +Glossina, 117, 297, 303 + +Glossina morsitans, 214, 217 + palpalis, 215, 217, 218 + +Glyciphagus, 267 + +Grain moth, 69 + +Grocer's itch, 72 + +Guinea-worm, 182 + + +Habronema muscæ, 156, 183 + +Hæmatobia, 166, 304 + irritans, 146 + +Hæmatobosca, 304 + +Hæmatomyidium, 288 + +Hæmatopinus spinulosus, 213 + +Hæmatopota, 294 + +Hæmatosiphon, 279 + +Hæmoglobinuria, 222 + +Hæmozoin, 189 + +Harpactor, 284 + +Harvest mites, 60 + effect of, 59 + +Head-louse, 173 + +Helminthiasis, 138 + +Helophilus, 295 + +Hemiptera, 27, 86, 273-275 + +Heteropodidæ, 13 + +Heuchis sanguinea, 55 + +Hexapod larvæ, 58 + +Hexapoda, 27, 80, 258 + +Hippelates, 297 + +Hippobosca, 285 + +Histiogaster, 269 + spermaticus, 132 + +Homalomyia, 136, 138, 300 + +Honey bee, 36 + poison of, 37 + +Hornets, 43 + +Horn-fly, 137, 304, 308 + +Horse-fly, 110, 165 + +House-fly, 137-139, 144, 183 + control of, 156, 160 + +Human flea, 124 + +Host, definitive, 175 + intermediate, 175 + primary, 175 + +Hyalomma, 264 + ægypticum, 224-225 + +Hydrocyanic Acid Gas, 318 + +Hydrotæa, 300 + +Hymenolepis diminuta, 176 + +Hymenoptera, 36, 275 + +Hypoderma, 113, 298 + diana, 113 + lineata, 113 + +Hypopharynx, 80 + + +Immunity from stings, 39 + +Incomplete metamorphosis, 80 + +Infantile paralysis, 162, 241 + splenic, 220 + Direct inoculation, 164 + +Insects, 258 + blood-sucking, 170 + +Intermediate host, 192, 203 + +Intestinal infestation, 112, 133 + myasis, 137 + +Isosoma, 69 + +Itch, 73-74 + mite, 73 + Norwegian, 77 + +Ixodes, 260 + ricinus, 66, 225 + scapularis, 66 + +Ixodidæ, 64-65 + +Ixodoidea, 62 + + +Janthinosoma lutzi, 116 + +Jigger, 60 + +Johannseniella, 110, 288 + +Journal of Tropical Medicine and Hygiene, 36 + +Julus terrestris, 25 + +June bug, 185 + + +Kala-azar, 220 + +Karakurte, 14 + +Katipo, 14 + +King, A. F. A., 3 + +Kircher, Athanasius, 1, 8 + +Kissing-bug, 31 + + +Labium, 29, 80 + +Labrum, 28, 80 + +Lachnosterna, 185 + +Lælaps, 266 + +Lagoa crispata, 45 + +Lamblia intestinalis, 154 + +Langer, Josef, 37 + +Larder beetles, 135 + +Latrodectus, 12, 14, 17 + mactans, 15 + +Leishmanioses, 220 + +Lenticular fever, 237 + +Lepidoptera, 274 + +Lepidopterous larvæ, 134 + +Leprosy, 252 + +Leptidæ, 112 + +Leptis, 295 + +Leptus, 60, 273 + +Lice, 80 + +Linguatulina, 258 + +Liponyssus, 265 + +Loemopsylla, 172, 317 + +Lone star tick, 228 + +Louse, body, 84 + crab, 85 + dog, 176 + head, 82 + pubic, 85 + +Lucilia, 136, 312 + +Lycosa tarantula, 10 + +Lycosidæ, 10 + +Lyctocoris, 279 + +Lygus pratensis, 33 + +Lymphangitis, 67 + +Lymph scrotum, 178 + +Lyperosia, 304 + +Lyperosiops, 305 + + +Macloskie, 34 + +Maggots, rat-tail, 137 + +Magnes sive de Arte Magnetica, 8 + +Malaria, 186 + +Malmigniatte, 14 + +Mandibles, 28, 80 + +Mange, 73-75 + +Margaropus, 237, 264 + annulatus, 223 + +Masked bed-bug hunter, 32 + +Mastigoproctus giganteus, 19, 80 + +Maxillæ, 28 + +Meal infesting species, 135 + +Melanin granules, 189 + +Melanolestes, 280 + picipes, 32 + +Mena-vodi, 14 + +Mercurialis, 1 + +Merozoites, 190 + +Metamorphosis, 80 + +Miana bug, 63 + +Microgametoblast, 192 + +Midges, 107 + +Migratory ookinete, 192 + +Millipedes, 25, 257 + +Mites, 23, 58 + +Monieziella, 269 + +Mosquitoes, 33, 97, 178, 196, 250 + treatment for bites of, 34, 36, 102 + +Musca, 137, 307 + domestica, 139, 145, 146, 157, 162 + +Muscidæ, 117 + +Muscina, 137, 146, 307 + stabulans, 140 + +Mutualism, 57 + +Myasis, 112, 135 + intestinal, 135-140 + nasal, 141 + +Mycterotypus, 287 + +Myiospila, 146, 307 + +Myriapoda, 225, 132, 257 + + +Nagana, 165, 214 + +Nasal infestation, 114, 133 + +Necrobia, 135 + +Nematode parasite, 182 + +Nepa, 28 + +Nephrophages sanguinarius, 132 + +Nettling insects, 43 + larvæ, poison of, 53 + +Neurasthenia, 89 + +Nits, 86 + +North African Relapsing Fever, 234 + +Norwegian itch, 77 + +No-see-ums, 109 + +Notoedres, 269 + cati, 78 + +Notonecta, 28, 277 + +Notonectidæ, 30 + +Nott, Dr. Josiah, 2 + +Nuttall, 34 + + +Occipital headaches, 138 + +Oecacta, 288 + +Oeciacus, 279 + +Oesophageal diverticula, 35 + +Oestridæ, 112, 136 + +Oestris ovis, 113 + +Oestrus, 298 + oocyst, 192 + ookinete, 192 + +Opsicoetes personatus, 32 + +Opthalmia, 155 + nodosa, 52 + +Oriental sore, 221 + +Ornithodoros, 65, 260 + moubata, 220, 230 + +Orthotylus flavosparsus, 33 + +Ornithomyia, 286 + +Oroya, 253 + +Oscinus, 297 + +Otiobius, 259 + megnini, 65 + +Otodectes, 271 + + +Pangonia, 294 + +Pappatici fever, 96 + +Parasimulium, 286 + +Parasite, 3, 57, 131, 134, 182 + accidental, 3, 131, 134 + facultative, 3, 57, 131 + nematode, 182 + stationary, 57 + temporary, 57 + true, 3 + +Parasitism, accidental, 134 + +Pathogenic bacteria, 152 + organisms, 144, 164 + +Pawlowsky, 81 + +Pediculoides, 267 + ventricosus, 69, 72 + +Pediculosis, 81 + +Pediculus, 275 + corporis, 84, 233, 238 + humanus, 82, 173 + +Pellagra, 162, 246 + +Pernicious fever, 186 + +Pest, 166 + +Phidippus audax, 19 + +Philæmatomyia, 306 + +Phisalix, 13, 43 + +Phlebotomus, 289 + papatasii, 94 + verrucarum, 254 + vexator, 95 + +Phora, 295 + +Phormia, 136 + +Phormictopus carcerides, 13 + +Phthirus pubis, 85, 275 + +Phortica, 296 + +Pieris brassicæ, 56 + +Piophila, 297 + +Piophila casei, 136, 137 + +Piroplasmosis, 222 + +Plague, 166 + bubonic, 166, 169, 170 + pneumonic, 167 + +Plasmodium, 186 + +Platymetopius acutus, 33 + +Plica palonica, 83 + +Pneumonic, 166 + plague, 167, 173 + +Poisoning by nettling larvæ, 53 + +Poison of spiders, 7 + +Pollenia, 308 + rudis, 146, 147 + +Primary gland, 28 + +Prionurus citrinus, 20 + +Prosimulium, 286 + +Protocalliphora, 136, 312 + +Protozoan blood parasite, 165 + +Pseudo-tubercular, 52 + +Psorophora, 293 + +Psoroptes, 270 + +Psychodidæ, 94 + +Pulex, 120, 124, 126, 172, 317 + cheopis, 172 + irritans, 124 + penetrans, 126 + serraticeps, 120 + +Pulvillus, 150 + +Punkies, 109 + +Pycnosoma, 308 + + +Rasahus, 280 + thoracicus, 32 + +Rat fleas, 120, 124, 171 + +Rat louse, 213 + +Red bugs, 70-72 + +Reduviidæ, 31 + +Reduviolus, 280 + +Reduvius, 282 + personatus, 32 + +Redwater fever, 222 + +Relapsing fever, 230, 233 + +Rhinoestrus nasalis, 115 + +Rhipicentor, 264 + +Rhipicephalus, 264 + +Rhizoglyphus, 269 + +Rhodnius, 281 + +Rocky Mountain Spotted Fever, 226 + spotted fever tick, 67 + +Russian gad-fly, 115 + + +St. Vitus's or St. John's dance, 8 + +Salivary syringe, 28 + +Sand-flies, 109, 250 + +Sanguinetti, 11 + +Sarcophaga, 136, 142, 143 + +Sarcophila, 302 + +Sarcopsylla, 317 + penetrans, 126 + +Sarcoptes, 270 + minor, 78 + scabiei, 73 + +Sarcoptidæ, 72 + +Scabies, 72, 73, 74, 75 + +Scaurus striatus, 177 + +Schaudinn, 34 + +Schizont, 189, 190 + +Scholeciasis, 134 + +Scolopendra morsitans, 26 + +Scorpions, 20 + poison of, 21 + +Screw worm fly, 140 + +Sepsidæ, 296 + +Sepsis, 136, 297 + +Shipley, 34 + +Sibine, 46 + +Silvius, 294 + +Simple carriers, 4, 144 + +Simuliidæ, 33, 104 + +Simulium, 247, 249, 286, 321 + pictipes, 104 + +Siphonaptera, 119, 274, 316 + +Siphunculata, 80, 275 + +Sitotroga cerealella, 69 + +Skippers, 137 + +Sleeping sickness, 166, 215 + +Snipe-flies, 112 + +Solpugida, 22 + +Spanish fly, 54 + +Spermatozoa, 192 + +Spinose ear-tick, 65 + +Spirochæta, 35 + berberi, 234 + duttoni, 234 + +Spirochætosis, 235 + +Sporozoite, 189 + +Spotted fever, 67, 226 + +Squirrel flea, 123 + +Stable-fly, 137, 160, 163, 165 + +Stegomyia, 182, 293 + calopus, 206 + fasciata, 206 + +Stomoxys, 137, 305 + calcitrans, 117, 146, 160, 161, 165, 242 + +Straw-worm, 69 + +Stygeromyia, 305 + +Sucking stomach, 35 + +Sulphur ointment, 77 + +Surra, 165 + +Symbiosis, 57 + +Symphoromyia, 112, 295 + + +Tabanidæ, 110 + +Tabanus, 110, 166, 294 + striatus, 165 + +Taenia, 175 + +Tapeworm, 4, 176 + +Tarantella, 8 + +Tarantism, 8 + +Tarantula, 10 + +Tarsonemidæ, 69 + +Tarsonemus, 267 + +Tenebrionid beetles, 127 + +Tersesthes, 110, 288 + +Tetanus, 129 + +Tetranychus, 273 + +Texas fever, 220-223 + +Three-day fever, 96 + +Tick, 23, 226 + bites, Treatment of, 68 + fever, 230 + paralysis, 67 + +Treatment, + Bee stings, 36, 41 + Bites of, + Bed-bugs, 90, 93 + Blackflies, 107 + Buffalo flies, 107 + Bugs, 31, 33 + Centipedes, 26, 27 + Chiggers, 127 + Chigoes, 127 + Fleas, 127 + Harvest mites, 61 + Jiggers, 129 + Lice, 83, 85 + Mosquitoes, 34, 36, 102 + Phlebotomus flies, 97 + Sand flies, 96, 107, 109 + Scorpions, 22, 23 + Spiders, 19 + Ticks, 61, 68, 72 + Ticks, ear, 65 + Blister beetle poison, 55 + Brown-tail moth rash, 45 + Cantharidin poison, 55 + Caterpillar rash, 45 + Ear ticks, 65 + House fly control, 156, 160 + Itch, 77 + Itch, grocer's, 72 + Lice, 85 + Nasal myasis, 143 + Rocky Mt. spotted fever, 228, 229 + Rash, caterpillar, 45 + Scabies, 77 + Sleeping sickness control, 218 + Spotted fever, 228, 229 + Stings, bee, 36, 41 + Typhus fever, prophylaxis, 239 + +Trichodectes canis, 176 + +Trichoma, 82 + +Trineura, 295 + +Trochosa singoriensis, 11 + +Trombidium, 60, 273 + +True insects, 80 + +Trypanosoma, 35 + +Trypanosoma, brucei, 165 + +Trypanosoma cruzi, 219 + +Trypanosoma lewisi, 213 + +Trypanosomiases, 212 + +Trypanosomiasis, 165, 219 + +Tsetse flies, 117, 166, 214, 219 + +Tsetse flies disease, 165 + +Tuberculosis, 155 + +Tumbu-fly, 118 + +Tydeus, 271 + +Typhoid, 155 + +Typhoid fever, 154 + +Typhus, 237 + +Typhus fever, 237 + +Tyroglyphus, 72, 131, 268 + +Dr. Tyzzer, 49 + + +Uranotænia, 292 + + +Vancoho, 14 + +Varicose groin glands, 178 + +Verruga peruviana, 253 + +Vescicating insects, 54 + + +Wanzenspritze, 29 + +Warble-flies, 112 + +Wasps, 43 + +Whip-scorpions, 19 + +Wohlfahrtia, 143, 302 + +Wolf-spiders, 10 + +Wyeomyia smithii, 101, 293 + + +Xenopsylla, 172, 317 + +Xenopsylla cheopis, 171, 124 + +Xestopsylla, 317 + + +Yaws, 2 + +Yellow fever, 196, 203, 205 + + + + +TRANSCRIBER'S NOTES + +The following discrepancies in the text are as in the original: + + The inconsistent hyphenation of the following: + assassin-bugs/assassin bugs; + bedbug/bed-bug (and bedbugs); + beekeeper/bee-keeper (and beekeepers); + blackflies/black-flies; + blow-flies/blow flies; + bluebottles/blue-bottles; + bot-flies/bot flies; + bristlelike/bristle-like; + browntail/brown-tail; + coextensive/co-extensive; + deer-flies/deer flies; + dorsocentral/dorso-central; + ectoparasites/ecto-parasites; + endoparasites/endo-parasites; + flesh-fly/flesh fly (and flesh flies); + hotbed/hot-bed; + housefly/house-fly (and houseflies); + horsefly/horse-fly (and horse flies); + horse-manure/horse manure; + midsummer/mid-summer; + preeminently/pre-eminently; + sandfly/sand-fly (and sandflies); + screw-worm fly/screw worm fly; + stable-fly/stable fly; + subequal/sub-equal; + subfamily/sub-family; + subtropical/sub-tropical; + tapeworm/tape-worm (and tapeworms); + today/to-day; + tsetse-flies/tsetse flies; + widespread/wide-spread; + wormlike/worm-like. + + Inconsistent use of diaeresis in Aëdes/Aedes + + Inconsistent spelling of the following: + defence/defense; + disc/disk; + hemolysis/hæmolysis; + hemolytic/hæmolytic; + hexapod/hexopod; + Levaditi/Lavaditi; + metalescent/metallescent; + Mitzmain/Mitzman; + Neveau-Lemaire/Neveau-Lamaire; + offence/offense; + Phthirus/Phthirius + Portschinsky/Portchinsky/Portchinski; + travelled/traveled; + ventra-/ventro-; + Villot/Villet; + Wohlfartia/Wohlfahrtia; + + Inconsistent use of ligatures in Taenia/Tænia + +toxine, insiduous, efficaceous, cyanid are spelt as in the original + +In the first paragraph of chapter XII, "the student might not be lead" +is as in the original. Lead perhaps should be led + +In the bibliography, the entry for Vaughan without a text specified is +as in the original + + + + + +End of the Project Gutenberg EBook of Handbook of Medical Entomology, by +William Albert Riley and Oskar Augustus Johanssen + +*** END OF THIS PROJECT GUTENBERG EBOOK HANDBOOK OF MEDICAL ENTOMOLOGY *** + +***** This file should be named 34279-8.txt or 34279-8.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/4/2/7/34279/ + +Produced by Bryan Ness, Iris Schimandle, Brownfox and the +Online Distributed Proofreading Team at http://www.pgdp.net +(This file was produced from images generously made +available by The Internet Archive/American Libraries.) + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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A. Riley, Ph.D., and O. A. Johanssen, Ph.D. + </title> + <style type="text/css"> + +body { + margin-left: 10%; + margin-right: 10%; +} + + h1,h2,h3 { + text-align: center; /* all headings centered */ + clear: both; +} + +p { + margin-top: .75em; + text-align: justify; + margin-bottom: .75em; +} + +hr { + width: 33%; + margin-top: 2em; + margin-bottom: 2em; + margin-left: auto; + margin-right: auto; + clear: both; +} + +table { + margin-left: auto; + margin-right: auto; +} + +.pagenum { /* uncomment the next line for invisible page numbers */ + /* visibility: hidden; */ + position: absolute; + left: 93%; + font-size: smaller; + text-align: right; +} /* page numbers */ + + +.blockquot { + margin-left: 5%; + margin-right: 10%; +} + +.bbox {border: solid 2px;} + +.center {text-align: center;} + +.smcap {font-variant: small-caps;} + +.u {text-decoration: underline;} + +.caption {font-weight: bold;} + +.hanging { + margin-left: 20px; + text-indent: -20px; +} + +.rightalign { + float: right; +} + +/* Images */ +.figcenter { + margin: auto; + text-align: center; +} + +.figleft { + float: left; + clear: left; + margin-left: 0; + margin-bottom: 1em; + margin-top: 1em; + margin-right: 1em; + padding: 0; + text-align: center; +} + +.figright { + float: right; + clear: right; + margin-left: 1em; + margin-bottom: + 1em; + margin-top: 1em; + margin-right: 0; + padding: 0; + text-align: center; +} + +/* Footnotes */ +.footnotes {border: dashed 1px;} + +.footnote {margin-left: 10%; margin-right: 10%; font-size: 0.9em;} + +.footnote .label {position: absolute; right: 84%; text-align: right;} + +.fnanchor { + vertical-align: super; + font-size: .8em; + text-decoration: + none; +} + +/* Poetry */ +.poem { + margin-left:10%; + margin-right:10%; + text-align: left; +} + +.poem br {display: none;} + +.poem .stanza {margin: 1em 0em 1em 0em;} + +.poem span.i0 { + display: block; + margin-left: 0em; + padding-left: 3em; + text-indent: -3em; +} + +.poem span.i2 { + display: block; + margin-left: 2em; + padding-left: 3em; + text-indent: -3em; +} + +.poem span.i4 { + display: block; + margin-left: 4em; + padding-left: 3em; + text-indent: -3em; +} + + </style> + </head> +<body> + + +<pre> + +The Project Gutenberg EBook of Handbook of Medical Entomology, by +William Albert Riley and Oskar Augustus Johanssen + +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: Handbook of Medical Entomology + +Author: William Albert Riley + Oskar Augustus Johanssen + +Release Date: November 11, 2010 [EBook #34279] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK HANDBOOK OF MEDICAL ENTOMOLOGY *** + + + + +Produced by Bryan Ness, Iris Schimandle, Brownfox and the +Online Distributed Proofreading Team at http://www.pgdp.net +(This file was produced from images generously made +available by The Internet Archive/American Libraries.) + + + + + + +</pre> + + + +<div class="bbox" style="padding:1em;"> +<h3>TRANSCRIBERS' NOTES</h3> +<p>Barring some obvious typos, the text has been left +as printed. Discrepancies identified are listed at the end of the text. +Most images are linked to a larger image of the same picture.</p> +</div> + + +<div class="figcenter" style="width: 300px;margin-top:2em;"> +<a href="images/frontispiece-full.png"><img src="images/frontispiece.png" width="300" height="436" alt="Some early medical entomology. Athanasius Kircher's illustration of the Italian tarantula +and the music prescribed as an antidote for the poison of its bite. (1643)." title="Some early medical entomology. Athanasius Kircher's illustration of the Italian tarantula +and the music prescribed as an antidote for the poison of its bite. (1643)." /></a> +<span class="caption">Some early medical entomology. Athanasius Kircher's illustration of the Italian tarantula +and the music prescribed as an antidote for the poison of its bite. (1643).</span> +</div> + + + + +<hr style="width: 65%;" /> +<h1>HANDBOOK OF MEDICAL +ENTOMOLOGY</h1> + +<h2>WM. A. RILEY, <span class="smcap">Ph.D.</span></h2> + +<p class="center">Professor of Insect Morphology and Parasitology, Cornell University</p> + +<p class="center">and</p> + +<h2>O. A. JOHANNSEN, <span class="smcap">Ph.D.</span></h2> + +<p class="center">Professor of Biology, Cornell University</p> + +<div class="figcenter" style="width: 125px;"> +<img src="images/tpage.png" width="125" height="152" alt="" title="" /> +</div> + +<p class="center">ITHACA, NEW YORK</p> + +<p class="center">THE COMSTOCK PUBLISHING COMPANY</p> + +<p class="center">1915</p> + + +<p class="center">COPYRIGHT, 1915</p> + +<p class="center">BY THE COMSTOCK PUBLISHING COMPANY,</p> + +<p class="center">ITHACA, N. Y.</p> + + +<p class="center">Press of W. F. Humphrey<br /> +Geneva, N. Y.</p> +<p><span class="pagenum"><a name="Page_v" id="Page_v">[Pg v]</a></span></p> + + + + +<hr style="width: 65%;" /> +<h2>PREFACE</h2> + + +<p>The Handbook of Medical Entomology is the outgrowth of a +course of lectures along the lines of insect transmission and +dissemination of diseases of man given by the senior author +in the Department of Entomology of Cornell University during the +past six years. More specifically it is an illustrated revision and +elaboration of his "Notes on the Relation of Insects to Disease" +published January, 1912.</p> + +<p>Its object is to afford a general survey of the field, and primarily +to put the student of medicine and entomology in touch with the +discoveries and theories which underlie some of the most important +modern work in preventive medicine. At the same time the older +phases of the subject—the consideration of poisonous and parasitic +forms—have not been ignored.</p> + +<p>Considering the rapid shifts in viewpoint, and the development +of the subject within recent years, the authors do not indulge in any +hopes that the present text will exactly meet the needs of every +one specializing in the field,—still less do they regard it as complete +or final. The fact that the enormous literature of isolated articles is +to be found principally in foreign periodicals and is therefore difficult +of access to many American workers, has led the authors to hope +that a summary of the important advances, in the form of a reference +book may not prove unwelcome to physicians, sanitarians and +working entomologists, and to teachers as a text supplementing +lecture work in the subject.</p> + +<p>Lengthy as is the bibliography, it covers but a very small fraction +of the important contributions to the subject. It will serve only to +put those interested in touch with original sources and to open up +the field. Of the more general works, special acknowledgment +should be made to those of Banks, Brumpt, Castellani and Chalmers, +Comstock, Hewitt, Howard, Manson, Mense, Neveau-Lemaire, +Nuttall, and Stiles.</p> + +<p>To the many who have aided the authors in the years past, by +suggestions and by sending specimens and other materials, sincerest +thanks is tendered. This is especially due to their colleagues in +the Department of Entomology of Cornell University, and to Professor +Charles W. Howard, Dr. John Uri Lloyd, Mr. A. H. Ritchie, +Dr. I. M. Unger, and Dr. Luzerne Coville.<span class="pagenum"><a name="Page_vi" id="Page_vi">[Pg vi]</a></span></p> + +<p>They wish to express indebtedness to the authors and publishers +who have so willingly given permission to use certain illustrations. +Especially is this acknowledgment due to Professor John Henry +Comstock, Dr. L. O. Howard, Dr. Graham-Smith, and Professor +G. H. T. Nuttall. Professor Comstock not only authorized the use +of departmental negatives by the late Professor M. V. Slingerland +(credited as M. V. S.), but generously put at their disposal the illustrations +from the <span class="smcap">Manual for the Study of Insects</span> and from +the <span class="smcap">Spider Book</span>. Figures <a href="#Fig_5">5</a> and <a href="#Fig_111">111</a> are from Peter's "Der Arzt +und die <a name="AC_1" id="AC_1"><span title="for Heilkunft read Heilkunst">Heilkunft</span></a> in der deutschen Vergangenheit." It should be +noted that on examining the original, it is found that Gottfried's +figure relates to an event antedating the typical epidemic of dancing +mania.</p> + +<div style="margin-left: 75%;"><p><span class="smcap">Wm. A. Riley.</span><br /> +<span class="smcap">O. A. Johannsen.</span></p></div> + +<div style="margin-left: 2em;"><p><span class="smcap">Cornell University</span>,<br /> +January, 1915.</p></div> + + + + +<hr style="width: 65%;" /> +<h2>ADDITIONS AND CORRECTIONS</h2> + + +<div class="hanging"><p>vi <a href="#AC_1">line 11</a>, for Heilkunft read Heilkunst.</p> + +<p>18 <a href="#AC_2">line 2</a>, for tarsi read tarsus.</p> + +<p>32 <a href="#AC_3">line 21</a>, and legend under <a href="#Fig_23">fig. 23</a>, for C. (Conorhinus) +abdominalis read Melanolestes abdominalis.</p> + +<p>47 legend under <a href="#Fig_33c">figure</a> for 33c read 34.</p> + +<p>92 line <a href="#AC_5a">22</a> and <a href="#AC_5b">25</a>, for sangiusugus read sanguisugus.</p> + +<p>116 legend under <a href="#Fig_83">fig. 83</a>, for Graham-Smith read Manson.</p> + +<p>136 <a href="#AC_7">line 10, from bottom</a>, insert "ring" after "chitin".</p> + +<p>137 <a href="#AC_8">line 3</a>, for meditatunda read meditabunda.</p> + +<p>145 <a href="#AC_9">line 7, from bottom</a>, for Rs read R<sub>5</sub>.</p> + +<p>158 <a href="#AC_10">line 20</a>, for have read has.</p> + +<p>212 after the <a href="#CHAPTER_IX">chapter heading</a> insert "continued".</p> + +<p>219 <a href="#AC_12">line 10, from bottom</a>, for Cornohinus read Conorhinus.</p> + +<p>266 <a href="#AC_13">line 1</a>, <a href="#Fig_158">fig. 158j</a> refers to the female.</p> + +<p>272 <a href="#AC_14">line 5</a>, insert "palpus" before "and leg".</p> + +<p>281 <a href="#AC_15">line 6</a>, for discodial read discoidal.</p> + +<p>281 <a href="#AC_16">last line</a>, insert "from" before "the".</p> + +<p>284 <a href="#AC_17">line 5</a>, for "tubercle of" read "tubercle or".</p> + +<p>305 lines <a href="#AC_18a">19</a>, <a href="#AC_18b">28</a>, <a href="#AC_18c">44</a>, page 306 lines <a href="#AC_18d">1</a>, <a href="#AC_18e">9</a>, <a href="#AC_18f">22</a>, <a href="#AC_18g">27</a>, <a href="#AC_18h">30</a>, page 307 <a href="#AC_18i">line 7</a>, +page 309 lines <a href="#AC_18j">8</a>, <a href="#AC_18k">11</a>, for R<sub>4+5</sub> read M<sub>1+2</sub>.</p> + +<p>309 legend under <a href="#Fig_168">fig. 168</a> add Bureau of Entomology.</p> + +<p>312 <a href="#AC_20">line 36</a>, for "near apex" read "of M<sub>1+2</sub>".</p> + +<p>313 running head, for Muscidæ read Muscoidea.</p> + +<p>314 <a href="#AC_22">line 29</a>, for "distal section" read "distally M<sub>1+2</sub>".</p> + +<p>315 legend under <a href="#Fig_172">fig. 172</a>, for Pseudopyrellia read Orthellia, +for Lyperosia read Hæmatobia, for Umbana read urbana.</p> + +<p><a href="#Fig_173">323</a> and <a href="#Fig_174">325</a> legends under the figures, add "After Dr. J. H. +Stokes".</p> + +<p>328 <a href="#AC_25">line 7 from bottom</a> for Apiochæta read Aphiochæta.</p></div><p><span class="pagenum"><a name="Page_vii" id="Page_vii">[Pg vii]</a></span></p> + + + +<hr style="width: 65%;" /> +<h2>CONTENTS</h2> + + +<pre> +<a href="#CHAPTER_I"><b>CHAPTER I</b></a> + +<a href="#CHAPTER_I"><b>INTRODUCTION</b></a> <a href="#Page_1">1-5</a> + + <a href="#EARLY_SUGGESTIONS_REGARDING_THE_TRANSMISSION_OF_DISEASE_BY_INSECTS">Early suggestions regarding the transmission of disease by</a> + <a href="#EARLY_SUGGESTIONS_REGARDING_THE_TRANSMISSION_OF_DISEASE_BY_INSECTS">insects.</a> + <a href="#THE_WAYS_IN_WHICH_ARTHROPODS_MAY_AFFECT_THE_HEALTH_OF_MAN">The ways in which arthropods may affect the health of man.</a> + + +<a href="#CHAPTER_II"><b>CHAPTER II</b></a> + +<a href="#CHAPTER_II"><b>ARTHROPODS WHICH ARE DIRECTLY POISONOUS</b></a> <a href="#Page_6">6-56</a> + + <a href="#ARANEIDA_OR_SPIDERS">The Araneida, or Spiders.</a> + <a href="#The_Tarantulas">The tarantulas.</a> <a href="#Avicularoidea">Bird spiders.</a> <a href="#Spiders_of_the_Genus_Latrodectus">Spiders of the genus</a> + <a href="#Spiders_of_the_Genus_Latrodectus">Latrodectus.</a> <a href="#Other_Venomous_Spiders">Other venomous spiders.</a> <a href="#Summary">Summary.</a> + <a href="#THE_PEDIPALPIDA_OR_WHIP-SCORPIONS">The Pedipalpida, or whip-scorpions.</a> + <a href="#THE_SCORPIONIDA_OR_TRUE_SCORPIONS">The Scorpionida, or true scorpions.</a> + <a href="#THE_SOLPUGIDA_OR_SOLPUGIDS">The Solpugida, or solpugids.</a> + <a href="#THE_ACARINA_OR_MITES_AND_TICKS">The Acarina, or mites and ticks.</a> + <a href="#THE_MYRIAPODA_OR_CENTIPEDES_AND_MILLIPEDES">The Myriapoda, or centipedes and millipedes.</a> + <a href="#THE_HEXAPODA_OR_TRUE_INSECTS">The Hexapoda, or true insects.</a> + <a href="#PIERCING_OR_BITING_INSECTS_POISONOUS_TO_MAN">Piercing or biting insects poisonous to man.</a> + <a href="#Hemiptera">Hemiptera, or true bugs.</a> + <a href="#The_Notonectidae_or_back_swimmers">The Notonectidæ or back-swimmers.</a> <a href="#The_Belostomatidae_or_giant_water_bugs">Belostomidæ or giant</a> + <a href="#The_Belostomatidae_or_giant_water_bugs">water-bugs.</a> <a href="#The_Reduviidae_or_assassin-bugs">Reduviidæ, or assassin bugs.</a> <a href="#Other_Hemiptera_Reported_as_Poisonous_to_Man">Other</a> + <a href="#Other_Hemiptera_Reported_as_Poisonous_to_Man">Hemiptera reported as poisonous to man.</a> + <a href="#DIPTERA">Diptera; the midges, mosquitoes and flies.</a> + <a href="#STINGING_INSECTS">Stinging insects.</a> + <a href="#Apis_mellifica_the_honey_bee">Apis mellifica, the honey bee.</a> <a href="#Other_Stinging_Forms">Other stinging forms.</a> + <a href="#NETTLING_INSECTS">Nettling insects.</a> + <a href="#LEPIDOPTERA">Lepidoptera, or butterflies and moths.</a> <a href="#Relief_from_Poisoning_by_Nettling_Larvae">Relief from</a> + <a href="#Relief_from_Poisoning_by_Nettling_Larvae">poisoning by nettling larvæ.</a> + <a href="#Vescicating_Insects_and_those_Possessing_Other_Poisons_in_their_Blood_Plasma">Vescicating insects and those possessing other poisons</a> + <a href="#Vescicating_Insects_and_those_Possessing_Other_Poisons_in_their_Blood_Plasma">in their blood plasma.</a> <a href="#The_Blister_Beetles">The blister beetles.</a> <a href="#Other_Cryptotoxic_Insects">Other</a> + <a href="#Other_Cryptotoxic_Insects">cryptotoxic insects.</a> + + +<a href="#CHAPTER_III"><b>CHAPTER III</b></a> + +<a href="#CHAPTER_III"><b>PARASITIC ARTHROPODS AFFECTING MAN</b></a> <a href="#Page_57">57-130</a> + + <a href="#Acarina_or_Mites">Acarina, or mites.</a> + <a href="#The_Trombidiidae_or_Harvest_Mites">The Trombidiidæ, or harvest mites.</a> + <a href="#Ixodoidea_or_Ticks">The Ixodoidea, or ticks.</a> + <a href="#Argasidae">Argasidæ.</a> <a href="#Ixodidae">Ixodidæ.</a> <a href="#Treatment_of_Tick_Bites">Treatment of tick bites.</a> + <a href="#Dermanyssidae">The mites.</a> + <a href="#Dermanyssidae">Dermanyssidæ.</a> <a href="#Tarsonemidae">Tarsonemidæ.</a> <a href="#Sarcoptidae">Sarcoptidæ, the itch mites.</a> + <a href="#Demodecidae">Demodecidæ, the follicle mites.</a> + <a href="#Hexapoda_or_True_Insects">Hexapoda, or true insects.</a> + <a href="#Siphunculata">Siphunculata, or sucking lice.</a> + <a href="#Hemiptera">Hemiptera.</a><span class="pagenum"><a name="Page_viii" id="Page_viii">[Pg viii]</a></span> + <a href="#The_Bed-bugs">The bed-bug.</a> <a href="#Other_Bed-bugs">Other bed-bugs.</a> + <a href="#Parasitic_Diptera_or_Flies">Parasitic Diptera, or flies.</a> + <a href="#Psychodidae_or_Moth-Flies">Psychodidæ, or moth flies.</a> <a href="#The_Phlebotominae">Phlebotominæ.</a> <a href="#Culicidae_or_Mosquitoes">Culicidæ, or</a> + <a href="#Culicidae_or_Mosquitoes">mosquitoes.</a> <a href="#The_Simuliidae_or_Black_Flies">Simuliidæ, or black-flies.</a> <a href="#Chironomidae_or_Midges">Chironomidæ, or</a> + <a href="#Chironomidae_or_Midges">midges.</a> <a href="#Tabanidae_or_Horse-Flies">Tabanidæ, or horse-flies.</a> <a href="#Leptidae_or_Snipe-Flies">Leptidæ or</a> + <a href="#Leptidae_or_Snipe-Flies">snipe-flies.</a> <a href="#Oestridae_or_Bot-flies">Oestridæ, or bot-flies.</a> <a href="#Muscidae">Muscidæ, the</a> + <a href="#Muscidae">stable-fly and others.</a> + <a href="#The_Siphonaptera_or_Fleas">Siphonaptera, or fleas.</a> + <a href="#The_Siphonaptera_or_Fleas">The fleas affecting man, the dog, cat, and rat.</a> + <a href="#The_true_chiggers">The true chiggers, or chigoes.</a> + +<a href="#CHAPTER_IV"><b>CHAPTER IV</b></a> + +<a href="#CHAPTER_IV"><b>ACCIDENTAL OR FACULTATIVE PARASITES</b></a> <a href="#Page_131">131-143</a> + + <a href="#Acarina">Acarina, or mites.</a> + <a href="#Myriapoda">Myriapoda, or centipedes and millipedes.</a> + <a href="#Lepidopterous_Larvae">Lepidopterous larvæ.</a> + <a href="#Coleoptera">Coleoptera, or beetles.</a> + <a href="#Dipterous_Larvae">Dipterous larvæ causing myiasis.</a> + <a href="#Piophila_casei">Piophila casei, the cheese skipper.</a> <a href="#Chrysomyia_macellaria">Chrysomyia macellaria,</a> + <a href="#Chrysomyia_macellaria">the screw-worm fly.</a> <a href="#Calliphorinae">Calliphorinæ, the bluebottles.</a> + <a href="#Muscinae">Muscinæ, the house or typhoid fly, and others.</a> + <a href="#Anthyomyiidae">Anthomyiidæ, the lesser house-fly and others.</a> + <a href="#Sarcophagidae">Sarcophagidæ, the flesh-flies.</a> + + +<a href="#CHAPTER_V"><b>CHAPTER V</b></a> + +<a href="#CHAPTER_V"><b>ARTHROPODS AS SIMPLE CARRIERS OF DISEASE</b></a> <a href="#Page_144">144-163</a> + + <a href="#The_House-fly_As_a_Carrier_of_Disease">The house or typhoid fly as a carrier of disease.</a> + <a href="#Stomoxys_calcitrans_the_stable-fly">Stomoxys calcitrans, the stable-fly.</a> + <a href="#Other_arthropods_which_may_serve_as_simple_carriers_of_pathogenic_organisms">Other arthropods which may serve as simple carriers of</a> + <a href="#Other_arthropods_which_may_serve_as_simple_carriers_of_pathogenic_organisms">pathogenic organisms.</a> + + +<a href="#CHAPTER_VI"><b>CHAPTER VI</b></a> + +<a href="#CHAPTER_VI"><b>ARTHROPODS AS DIRECT INOCULATORS OF DISEASE GERMS</b></a> <a href="#Page_164">164-174</a> + + <a href="#Some_Illustrations_of_Direct_Inoculation_of_Disease_Germs_by_Arthropods">Some illustrations of direct inoculations of disease germs</a> + <a href="#Some_Illustrations_of_Direct_Inoculation_of_Disease_Germs_by_Arthropods">by arthropods.</a> + <a href="#The_Role_of_Fleas_in_the_Transmission_of_the_Plague">The rôle of fleas in the transmission of the plague.</a> + + +<a href="#CHAPTER_VII"><b>CHAPTER VII</b></a> + +<a href="#CHAPTER_VII"><b>ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC ORGANISMS</b></a> <a href="#Page_175">175-185</a> + + <a href="#Insects_as_Intermediate_Hosts_of_Tapeworms">Insects as intermediate hosts of tape-worms.</a> + <a href="#Arthropods_as_Intermediate_Hosts_of_Nematode_Worms">Arthropods as intermediate hosts of nematode worms.</a> + <a href="#Filariasis_and_Mosquitoes">Filariasis and mosquitoes.</a> + <a href="#Other_Nematode_Parasites_of_Man_and_Animals_Developing_in_Arthropods">Other nematode parasites of man and animals.</a> + + +<a href="#CHAPTER_VIII"><b>CHAPTER VIII</b></a> + +<a href="#CHAPTER_VIII"><b>ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA</b></a> <a href="#Page_186">186-211</a> + + <a href="#Mosquitoes_and_Malaria">Mosquitoes and malaria.</a> + <a href="#Mosquitoes_and_Yellow_Fever">Mosquitoes and yellow fever.</a><span class="pagenum"><a name="Page_ix" id="Page_ix">[Pg ix]</a></span> + + +<a href="#CHAPTER_IX"><b>CHAPTER IX</b></a> + +<a href="#CHAPTER_IX"><b>ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA</b></a> <a href="#Page_212">212-229</a> + + <a href="#Insects_and_Trypanosomiases">Insects and trypanosomiases.</a> + <a href="#Fleas_and_Lice_as_Carriers_of_Trypanosoma_lewisi">Fleas and lice as carriers of Trypanosoma lewisi.</a> + <a href="#Tsetse-flies_and_Nagana">Tsetse-flies and nagana.</a> + <a href="#Tsetse-flies_and_Sleeping_Sickness_of_Man">Tsetse-flies and sleeping sickness in man.</a> + <a href="#South_American_Trypanosomiasis">South American trypanosomiasis.</a> + <a href="#Leishmanioses_and_Insects">Leishmanioses and insects.</a> + <a href="#Ticks_and_Diseases_of_Man_and_Animals">Ticks and diseases of man and animals.</a> + <a href="#Cattle_Ticks_and_Texas_Fever">Cattle tick and Texas fever.</a> + <a href="#Ticks_and_Rocky_Mountain_Spotted_Fever_of_Man">Ticks and Rocky Mountain Spotted fever of man.</a> + + +<a href="#CHAPTER_X"><b>CHAPTER X</b></a> + +<b><a href="#CHAPTER_X">ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA</a> +<a href="#CHAPTER_X">(Continued)</a></b> <a href="#Page_230">230-240</a> + + <a href="#Arthropods_and_Spirochaetoses_of_Man_and_Animals">Arthropods and Spirochætoses of man and animals.</a> + <a href="#African_Relapsing_Fever_of_Man">African relapsing fever of man.</a> + <a href="#European_Relapsing_Fever">European relapsing fever.</a> + <a href="#North_African_Relapsing_Fever_of_Man">North African relapsing fever of man.</a> + <a href="#Other_Types_of_Relapsing_Fever_of_Man">Other types of relapsing fever of man.</a> + <a href="#Spirochaetosis_of_Fowls">Spirochætosis of fowls.</a> + <a href="#Other_Spirochaete_Diseases_of_Animals">Other spirochæte diseases of animals.</a> + <a href="#Typhus_Fever_and_Pediculidae">Typhus fever and lice.</a> + + +<a href="#CHAPTER_XI"><b>CHAPTER XI</b></a> + +<b><a href="#CHAPTER_XI">SOME POSSIBLE, BUT IMPERFECTLY KNOWN CASES OF</a> +<a href="#CHAPTER_XI">ARTHROPOD TRANSMISSION OF DISEASE</a></b> <a href="#Page_241">241-256</a> + + <a href="#Infantile_Paralysis_or_Acute_Anterior_Poliomyelitis">Infantile paralysis, or acute anterior poliomyelitis.</a> + <a href="#Pellagra">Pellagra.</a> <a href="#Leprosy">Leprosy.</a> <a href="#Verruga_peruviana">Verruga peruviana.</a> <a href="#Cancer">Cancer.</a> + + +<a href="#CHAPTER_XII"><b>CHAPTER XII</b></a> + +<a href="#CHAPTER_XII"><b>KEYS TO THE ARTHROPODS NOXIOUS TO MAN</b></a> <a href="#Page_257">257-317</a> + + <a href="#CRUSTACEA">Crustacea.</a> + <a href="#MYRIAPODA_1">Myriapoda, or centipedes and millipedes.</a> + <a href="#ORDERS_OF_THE_ARACHNIDA">Arachnida (Orders of).</a> + <a href="#ACARINA_KEY">Acarina or ticks.</a> + <a href="#HEXAPODA_Insecta">Hexapoda (Insecta).</a> + <a href="#SIPHUNCULATA_AND_HEMIPTERA">Siphunculata and Hemiptera (lice and true bugs).</a> + <a href="#DIPTERA_Mosquitoes_Midges_Flies">Diptera (mosquitoes, midges, and flies).</a> + <a href="#SIPHONAPTERA_Fleas">Siphonaptera (fleas).</a> + + +<a href="#APPENDIX"><b>APPENDIX</b></a> + +<a href="#HYDROCYANIC_ACID_GAS_AGAINST_HOUSEHOLD_INSECTS">Hydrocyanic acid gas against household insects</a> <a href="#Page_318">318-320</a> + <a href="#The_Proportions_of_Ingredients">Proportion of ingredients.</a> <a href="#A_Single_Room_As_an_Example">A single room as an example.</a> + <a href="#Fumigating_a_Large_House">Fumigating a large house.</a> <a href="#Precautions">Precautions.</a> + +<a href="#LESIONS_PRODUCED_BY_THE_BITE_OF_THE_BLACK-FLY">Lesions produced by the bite of the black-fly</a> <a href="#Page_321">321-326</a> + + +<a href="#BIBLIOGRAPHY"><b>BIBLIOGRAPHY</b></a> <a href="#Page_327">327-340</a> + + +<a href="#INDEX"><b>INDEX</b></a> <a href="#Page_341">341-348</a><span class="pagenum"><a name="Page_1" id="Page_1">[Pg 1]</a></span></pre> + + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_I" id="CHAPTER_I"></a>CHAPTER I.</h2> + +<h2>INTRODUCTION</h2> + + +<a name="EARLY_SUGGESTIONS_REGARDING_THE_TRANSMISSION_OF_DISEASE_BY_INSECTS" id="EARLY_SUGGESTIONS_REGARDING_THE_TRANSMISSION_OF_DISEASE_BY_INSECTS"></a><h3>EARLY SUGGESTIONS REGARDING THE TRANSMISSION OF DISEASE +BY INSECTS</h3> + +<p>Until very recent years insects and their allies have been considered +as of economic importance merely in so far as they are an annoyance +or direct menace to man, or his flocks and herds, or are injurious to +his crops. It is only within the past fifteen years that there has +sprung into prominence the knowledge that in another and much more +insiduous manner, they may be the enemy of mankind, that they +may be among the most important of the disseminators of disease. +In this brief period, such knowledge has completely revolutionized +our methods of control of certain diseases, and has become an important +weapon in the fight for the conservation of health.</p> + +<p>It is nowhere truer than in the case under consideration that however +abrupt may be their coming into prominence, great movements +and great discoveries do not arise suddenly. Centuries ago +there was suggested the possibility that insects were concerned with +the spread of disease, and from time to time there have appeared keen +suggestions and logical hypotheses along this line, that lead us to +marvel that the establishment of the truths should have been so long +delayed.</p> + +<p>One of the earliest of these references is by the Italian physician, +Mercurialis, who lived from 1530 to 1607, during a period when +Europe was being ravaged by the dread "black death", or plague. +Concerning its transmission he wrote: "There can be no doubt that +flies feed on the internal secretions of the diseased and dying, then, +flying away, they deposit their excretions on the food in neighboring +dwellings, and persons who eat of it are thus infected."</p> + +<p>It would be difficult to formulate more clearly this aspect of the +facts as we know them to-day, though it must always be borne in +mind that we are prone to interpret such statements in the light of +present-day knowledge. Mercurialis had no conception of the animate +nature of contagion, and his statement was little more than a lucky +guess.</p> + +<p>Much more worthy of consideration is the approval which was +given to his view by the German Jesuit, Athanasius Kircher in 1658.<span class="pagenum"><a name="Page_2" id="Page_2">[Pg 2]</a></span> +One cannot read carefully his works without believing that long +before Leeuwenhook's discovery, Kircher had seen the larger species of +bacteria. Moreover, he attributed the production of disease to these +organisms and formulated, vaguely, to be sure, a theory of the animate +nature of contagion. It has taken two and a half centuries to +accumulate the facts to prove his hypothesis.</p> + +<p>The theory of Mercurialis was not wholly lost sight of, for in the +medical literature of the eighteenth century there are scattered +references to flies as carriers of disease. Such a view seems even to +have been more or less popularly accepted, in some cases. Gudger +(1910), has pointed out that, as far back as 1769, Edward Bancroft, +in "An Essay on the Natural History of Guiana in South America," +wrote concerning the contagious skin-disease known as "Yaws": +"It is usually believed that this disorder is communicated by the flies +who have been feasting on a diseased object, to those persons who have +sores, or scratches, which are uncovered; and from many observations, +I think this is not improbable, as none ever receive this disorder +whose skins are whole."</p> + +<p>Approaching more closely the present epoch, we find that in 1848, +Dr. Josiah Nott, of Mobile, Alabama, published a remarkable +article on the cause of yellow fever, in which he presented "reasons for +supposing its specific cause to exist in some form of insect life." +As a matter of fact, the bearing of Nott's work on present day ideas +of the insect transmission of disease has been very curiously overrated. +The common interpretation of his theory has been deduced from a few +isolated sentences, but his argument appears quite differently when +the entire article is studied. It must be remembered that he wrote at +a period before the epoch-making discoveries of Pasteur and before +the recognition of micro-organisms as factors in the cause of disease. +His article is a masterly refutation of the theory of "malarial" origin +of "all the fevers of hot climates," but he uses the term "insect" as +applicable to the lower forms of life, and specific references to "mosquitoes," +"aphids," "cotton-worms," and others, are merely in the +way of similes.</p> + +<p>But, while Nott's ideas regarding the relation of insects to yellow +fever were vague and indefinite, it was almost contemporaneously +that the French physician, Louis Daniel Beauperthuy argued in the +most explicit possible manner, that yellow fever and various others +are transmitted by mosquitoes. In the light of the data which were +available when he wrote, in 1853, it is not surprising that he erred by<span class="pagenum"><a name="Page_3" id="Page_3">[Pg 3]</a></span> +thinking that the source of the virus was decomposing matter which +the mosquito took up and accidentally inoculated into man. Beauperthuy +not only discussed the rôle of mosquitoes in the transmission +of disease, but he taught, less clearly, that house-flies scatter pathogenic +organisms. It seems that Boyce (1909) who quotes extensively +from this pioneer work, does not go too far when he says "It is Dr. +Beauperthuy whom we must regard as the father of the doctrine of +insect-borne disease."</p> + +<p>In this connection, mention must be made of the scholarly article +by the American physician, A. F. A. King who, in 1883, brought +together an all but conclusive mass of argument in support of his +belief that malaria was caused by mosquitoes. At about the same +time, Finley, of Havana, was forcefully presenting his view that the +mosquito played the chief rôle in the spread of yellow fever.</p> + +<p>To enter more fully into the general historical discussion is beyond +the scope of this book. We shall have occasion to make more +explicit references in considering various insect-borne diseases. +Enough has been said here to emphasize that the recognition of +insects as factors in the spread of disease was long presaged, and that +there were not wanting keen thinkers who, with a background of +present-day conceptions of the nature of disease, might have been in +the front rank of investigators along these lines.</p> + + +<a name="THE_WAYS_IN_WHICH_ARTHROPODS_MAY_AFFECT_THE_HEALTH_OF_MAN" id="THE_WAYS_IN_WHICH_ARTHROPODS_MAY_AFFECT_THE_HEALTH_OF_MAN"></a><h3>THE WAYS IN WHICH ARTHROPODS MAY AFFECT THE HEALTH +OF MAN</h3> + +<p>When we consider the ways in which insects and their allies may +affect the health of man, we find that we may treat them under three +main groups:</p> + +<p>A. They may be directly poisonous. Such, for example, are the +scorpions, certain spiders and mites, some of the predaceous bugs, +and stinging insects. Even such forms as the mosquito deserve +some consideration from this viewpoint.</p> + +<p>B. They may be parasitic, living more or less permanently on +or in the body and deriving their sustenance from it.</p> + +<p>Of the parasitic arthropods we may distinguish, first, the <i>true +parasites</i>, those which have adopted and become confirmed in the +parasitic habit. Such are the itch mites, the lice, fleas, and the +majority of the forms to be considered as parasitic.</p> + +<p>In addition to these, we may distinguish a group of <i>accidental</i>, or +<i>facultative parasites</i>, species which are normally free-living, feeding on<span class="pagenum"><a name="Page_4" id="Page_4">[Pg 4]</a></span> +decaying substances, but which when accidentally introduced into +the alimentary canal or other cavities of man, may exist there +for a greater or less period. For example, certain fly larvæ, or maggots, +normally feeding in putrifying meat, have been known to occur +as accidental or facultative parasites in the stomach of man.</p> + +<p>C. Finally, and most important, arthropods may be transmitters +and disseminators of disease. In this capacity they may +function in one of three ways; as <i>simple carriers</i>, as <i>direct inoculators</i>, +or as <i>essential hosts</i> of disease germs.</p> + +<p>As simple carriers, they may, in a wholly incidental manner, +transport from the diseased to the healthy, or from filth to food, +pathogenic germs which cling to their bodies or appendages. Such, +for instance, is the relation of the house-fly to the dissemination of +typhoid.</p> + +<p>As direct inoculators, biting or piercing species may take up from +a diseased man or animal, germs which, clinging to the mouth parts, +are inoculated directly into the blood of the insect's next victim. It +it thus that horse-flies may occasionally transmit anthrax. Similarly, +species of spiders and other forms which are ordinarily perfectly +harmless, may accidentally convey and inoculate pyogenic bacteria.</p> + +<p>It is as essential hosts of disease germs that arthropods play their +most important rôle. In such cases an essential part of the life cycle +of the pathogenic organism is undergone in the insect. In other +words, without the arthropod host the disease-producing organism +cannot complete its development. As illustrations may be cited the +relation of the Anopheles mosquito to the malarial parasite, and the +relation of the cattle tick to Texas fever.</p> + +<p>A little consideration will show that this is the most important of +the group. Typhoid fever is carried by water or by contaminated +milk, and in various other ways, as well as by the house-fly. Kill all +the house-flies and typhoid would still exist. On the other hand, +malaria is carried only by the mosquito, because an essential part of +the development of the malarial parasite is undergone in this insect. +Exterminate all of the mosquitoes of certain species and the dissemination +of human malaria is absolutely prevented.</p> + +<p>Once an arthropod becomes an essential host for a given parasite +it may disseminate infection in three different ways:</p> + +<p>1. By infecting man or animals who ingest it. It is thus, for +example, that man, dog, or cat, becomes infected with the double-pored +dog tapeworm, <i>Dipylidium caninum</i>. The cysticercoid stage<span class="pagenum"><a name="Page_5" id="Page_5">[Pg 5]</a></span> +occurs in the dog louse, or in the dog or cat fleas, and by accidentally +ingesting the infested insect the vertebrate becomes infested. Similarly, +<i>Hymenolepis diminuta</i>, a common tapeworm of rats and mice, +and occasional in man, undergoes part of its life cycle in various meal-infesting +insects, and is accidentally taken up by its definitive host. +It is very probable that man becomes infested with <i>Dracunculus +(Filaria) medinensis</i> through swallowing in drinking water, the +crustacean, <i>Cyclops</i>, containing the larvæ of this worm.</p> + +<p>2. By infecting man or animals on whose skin or mucous membranes the +insect host may be crushed or may deposit its excrement. +The pathogenic organism may then actively penetrate, or may be +inoculated by scratching. The causative organism of typhus fever +is thus transmitted by the body louse.</p> + +<p>3. By direct inoculation by its bite, the insect host may transfer +the parasite which has undergone development within it. The +malarial parasite is thus transferred by mosquitoes; the Texas fever +parasite by cattle ticks.<span class="pagenum"><a name="Page_6" id="Page_6">[Pg 6]</a></span></p> + + + + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_II" id="CHAPTER_II"></a>CHAPTER II.</h2> + +<h2>ARTHROPODS WHICH ARE DIRECTLY POISONOUS</h2> + + +<p>Of all the myriads of insects and related forms, a very few are of +direct use to man, some few others have forced his approbation on +account of their wonderful beauty, but the great hordes of them are +loathed or regarded as directly dangerous. As a matter of fact, only +a very small number are in the slightest degree poisonous to man or +to the higher animals. The result is that entomologists and lovers +of nature, intent upon dissipating the foolish dread of insects, are +sometimes inclined to go to the extreme of discrediting all statements +of serious injury from the bites or stings of any species.</p> + +<p>Nevertheless, it must not be overlooked that poisonous forms do +exist, and they must receive attention in a consideration of the ways +in which arthropods may affect the health of man. Moreover, it +must be recognized that "what is one man's meat, is another man's +poison," and that in considering the possibilities of injury we must not +ignore individual idiosyncrasies. Just as certain individuals may be +poisoned by what, for others, are common articles of food, so some +persons may be abnormally susceptible to insect poison. Thus, the +poison of a bee sting may be of varying severity, but there are individuals +who are made seriously sick by a single sting, regardless of the +point of entry. Some individuals scarcely notice a mosquito bite, +others find it very painful, and so illustrations of this difference in +individuals might be multiplied.</p> + +<p>In considering the poisonous arthropods, we shall take them up by +groups. The reader who is unacquainted with the systematic relationship +of insects and their allies is referred to <a href="#CHAPTER_XII">Chapter XII</a>. No +attempt will be made to make the lists under the various headings +exhaustive, but typical forms will be discussed.</p> + + +<a name="ARANEIDA_OR_SPIDERS" id="ARANEIDA_OR_SPIDERS"></a><h3>ARANEIDA OR SPIDERS</h3> + +<p>Of all the arthropods there are none which are more universally +feared than are the spiders. It is commonly supposed that the +majority, if not all the species are poisonous and that they are aggressive +enemies of man and the higher animals, as well as of lower forms.</p> + +<p>That they really secrete a poison may be readily inferred from the +effect of their bite upon insects and other small forms. Moreover,<span class="pagenum"><a name="Page_7" id="Page_7">[Pg 7]</a></span> +the presence of definite and well-developed poison glands can easily +be shown. They occur as a pair of pouches (<a href="#Fig_1">fig. 1</a>) lying within the +cephalothorax and connected by a delicate +duct with a pore on the claw of the chelicera, +or so-called "mandible" on the convex surface +of the claw in such a position that it is not +plugged and closed by the flesh of the victim.</p> + +<div class="figleft" style="width: 300px;"><a name="Fig_1" id="Fig_1"></a> +<a href="images/f001-full.png"><img src="images/f001.png" width="300" height="263" alt="1. Head of a spider showing +poison gland (c) and its relation +to the chelicera (a)." title="1. Head of a spider showing +poison gland (c) and its relation +to the chelicera (a)." /></a> +<span class="caption">1. Head of a spider showing +poison gland (<i>c</i>) and its relation +to the chelicera (<i>a</i>).</span> +</div> + +<p>The glands may be demonstrated by slowly +and carefully twisting off a chelicera and +pushing aside the stumps of muscles at its +base. By exercising care, the chitinous wall +of the chelicera and its claw may be broken +away and the duct traced from the gland to its outlet. The inner +lining of the sac is constituted by a highly developed glandular +epithelium, supported by a basement membrane of connective +tissue and covered by a muscular layer, (<a href="#Fig_2">fig. 2</a>). The muscles, which +are striated, are spirally arranged (<a href="#Fig_1">fig. 1</a>), and are doubtless under +control of the spider, so that the amount of poison to be injected into +a wound may be varied.</p> + +<div class="figright" style="width: 300px;"><a name="Fig_2" id="Fig_2"></a> +<a href="images/f002-full.png"><img src="images/f002.png" width="300" height="254" alt="2. Section through a venom +gland of Latrodectus +13-guttatus showing +the peritoneal, muscular +and epithelial layers. +After Bordas." title="2. Section through a venom +gland of Latrodectus +13-guttatus showing +the peritoneal, muscular +and epithelial layers. +After Bordas." /></a> +<span class="caption">2. Section through a venom +gland of Latrodectus +13-guttatus showing +the peritoneal, muscular +and epithelial layers. +After Bordas.</span> +</div> + +<p>The poison itself, according to Kobert (1901), is a clear, colorless +fluid, of oily consistency, acid reaction, and very bitter taste. After +the spider has bitten two or three times, its supply is exhausted and +therefore, as in the case of snakes, the poison of the bite decreases +quickly with use, until it is null. To what extent the content of the +poison sacs may contain blood serum or, at least, active principles of +serum, in addition to a specific poison formed by the poison glands +themselves, Kobert regards as an open question. He believes that +the acid part of the poison, if really present, +is formed by the glands and that, +in the case of some spiders, the +ferment-like, or better, active +toxine, comes from the blood.</p> + +<div class="figleft" style="width: 225px;"> +<a href="images/f003-full.png"><img src="images/f003.png" width="225" height="342" alt="3. Chelicera of a +spider." title="3. Chelicera of a +spider." /></a> +<span class="caption">3. Chelicera of a +spider.</span> +</div> + +<p>But there is a wide difference +between a poison which may kill +an insect and one which is harmful +to men. Certain it is that +there is no lack of popular belief +and newspaper records of fatal +cases, but the evidence regarding the possibility of fatal or even very +serious results for man is most contradictory. For some years, +we have attempted to trace the more circumstantial newspaper<span class="pagenum"><a name="Page_8" id="Page_8">[Pg 8]</a></span> +accounts, which have come to our notice, of injury by North +American species. The results have served, mainly, to emphasize +the straits to which reporters are sometimes driven when +there is a dearth of news. The accounts are usually vague and lacking +in any definite clue for locating the supposed victim. In the +comparatively few cases where the patient, or his physician, could +be located, there was either no claim that the injury was due to +spider venom, or there was no evidence to support the belief. +Rarely, there was evidence that a secondary blood poisoning, such +as might be brought about by the prick of a pin, or by any mechanical +injury, had followed the bite of a spider. Such instances have +no bearing on the question of the +venomous nature of these forms.</p> + +<div class="figleft" style="width: 300px;"><a name="Fig_4" id="Fig_4"></a> +<a href="images/f004-full.png"><img src="images/f004.png" width="300" height="321" alt="4. The Italian tarantula (Lycosa tarantula). +After Kobert." title="4. The Italian tarantula (Lycosa tarantula). +After Kobert." /></a> +<span class="caption">4. The Italian tarantula (Lycosa tarantula). +After Kobert.</span> +</div> + +<p>The extreme to which unreasonable +fear of the bites of spiders +influenced the popular mind was +evidenced by the accepted explanation +of the remarkable dancing +mania, or tarantism, of Italy during +the Middle Ages. This was a nervous +disorder, supposed to be due +to the bite of a spider, the European +tarantula (<a href="#Fig_4">fig. 4</a>), though it was +also, at times, attributed to the +bite of the scorpion. In its typical +form, it was characterized by so +great a sensibility to music that under its influence the victims +indulged in the wildest and most frenzied dancing, until they sank +to the ground utterly exhausted and almost lifeless. The profuse +perspiring resulting from these exertions was supposed to be the +only efficacious remedy for the disease. Certain forms of music +were regarded as of especial value in treating this tarantism, and +hence the name of "tarantella" was applied to them. Our frontispiece, +taken from Athanasius Kircher's <i>Magnes sive de Arte Magnetica</i>, +1643 ed., represents the most commonly implicated spider and illustrates +some of what Fabre has aptly designated as "medical +choreography."</p> + +<p>The disease was, in reality, a form of hysteria, spreading by sympathy +until whole communities were involved, and was paralleled by +the outbreaks of the so-called St. Vitus's or St. John's dance, which<span class="pagenum"><a name="Page_9" id="Page_9">[Pg 9]</a></span> +swept Germany at about the same time (<a href="#Fig_5">fig. 5</a>). The evidence that +the spider was the cause of the first is about as conclusive as is that +of the demoniacal origin of the latter. The true explanation of the +outbreaks is doubtless to be found in the depleted physical and mental +condition of the people, resulting from the wars and the frightful +plagues which devastated all Europe previous to, and during these +times. An interesting discussion of these aspects of the question is to +be found in Hecker.</p> + +<div class="figcenter" style="width: 300px;"><a name="Fig_5" id="Fig_5"></a> +<a href="images/f005-full.png"><img src="images/f005.png" width="300" height="218" alt="5. Dancing mania. Illustration from Johann Ludwig Gottfried's Chronik. 1632." title="5. Dancing mania. Illustration from Johann Ludwig Gottfried's Chronik. 1632." /></a> +<span class="caption">5. Dancing mania. Illustration from Johann Ludwig Gottfried's Chronik. 1632.</span> +</div> + +<p>So gross has been the exaggeration and so baseless the popular fear +regarding spiders that entomologists have been inclined to discredit +all accounts of serious injury from their bites. Not only have the +most circumstantial of newspaper accounts proved to be without +foundation but there are on record a number of cases where the bite +of many of the commoner species have been intentionally provoked +and where the effect has been insignificant. Some years ago the +senior author personally experimented with a number of the largest of +our northern species, and with unexpected results. The first surprise +was that the spiders were very unwilling to bite and that it required a +considerable effort to get them to attempt to do so. In the second<span class="pagenum"><a name="Page_10" id="Page_10">[Pg 10]</a></span> +place, most of those experimented with were unable to pierce the skin +of the palm or the back of the hand, but had to be applied to the thin +skin between the fingers before they were able to draw blood. Unfortunately, +no special attempt was made to determine, at the time, the +species experimented with, but among them were <i>Theridion tepidariorum</i>, +<i>Miranda aurantia</i> (<i>Argiopa</i>), <i>Metargiope trifasciata</i>, <i>Marxia +stellata</i>, <i>Aranea trifolium</i>, <i>Misumena vatia</i>, and <i>Agelena nævia</i>. In +no case was the bite more severe than a pin prick and though in some +cases the sensation seemed to last longer, it was probably due to the +fact that the mind was intent upon the experiment.</p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_6" id="Fig_6"></a> +<a href="images/f006-full.png"><img src="images/f006.png" width="400" height="171" alt="6. An American tarantula (Eurypelma hentzii). Natural size. After Comstock." title="6. An American tarantula (Eurypelma hentzii). Natural size. After Comstock." /></a> +<span class="caption">6. An American tarantula (Eurypelma hentzii). Natural size. After Comstock.</span> +</div> + +<p>Similar experiments were carried out by Blackwell (1855), who +believed that in the case of insects bitten, death did not result any +more promptly than it would have from a purely mechanical injury of +equal extent. He was inclined to regard all accounts of serious +injury to man as baseless. The question cannot be so summarily +dismissed, and we shall now consider some of the groups which have +been more explicitly implicated.</p> + + +<p><a name="The_Tarantulas" id="The_Tarantulas"></a><b>The Tarantulas.</b>—In popular usage, the term "tarantula" is +loosely applied to any one of a number of large spiders. The famous +tarantulas of southern Europe, whose bites were supposed to cause the +dancing mania, were Lycosidæ, or wolf-spiders. Though various +species of this group were doubtless so designated, the one which +seems to have been most implicated was <i>Lycosa tarantula</i> (L.), +(<a href="#Fig_4">fig. 4</a>). On the other hand, in this country, though there are many +Lycosidæ, the term "tarantula" has been applied to members of the +superfamily Avicularoidea (<a href="#Fig_6">fig. 6</a>), including the bird-spiders.</p> + +<p>Of the Old World Lycosidæ there is no doubt that several species +were implicated as the supposed cause of the tarantism. In fact, as +we have already noted, the blame was sometimes attached to a scorpion.<span class="pagenum"><a name="Page_11" id="Page_11">[Pg 11]</a></span> +However, there seems to be no doubt that most of the accounts +refer to the spider known as <i>Lycosa tarantula</i>.</p> + +<p>There is no need to enter into further details here regarding the +supposed virulence of these forms, popular and the older medical +literature abound in circumstantial accounts of the terrible effects of +the bite. Fortunately, there is direct experimental evidence which +bears on the question.</p> + +<p>Fabre induced a common south European wolf-spider, <i>Lycosa +narbonensis</i>, to bite the leg of a young sparrow, ready to leave the +nest. The leg seemed paralyzed as a result of the bite, and though +the bird seemed lively and clamored for food the next day, on the +third day it died. A mole, bitten on the nose, succumbed after thirty-six +hours. From these experiments Fabre seemed justified in his +conclusion that the bite of this spider is not an accident which man +can afford to treat lightly. Unfortunately, there is nothing in the +experiments, or in the symptoms detailed, to exclude the probability +that the death of the animals was the result of secondary infection.</p> + +<p>As far back as 1693, as we learn from the valuable account of +Kobert, (1901), the Italian physician, Sanguinetti allowed himself to +be bitten on the arm by two tarantulas, in the presence of witnesses. +The sensation was equivalent to that from an ant or a mosquito bite +and there were no other phenomena the first day. On the second day +the wound was inflamed and there was slight ulceration. It is clear +that these later symptoms were due to a secondary infection. These +experiments have been repeated by various observers, among whom +may be mentioned Leon Dufour, Josef Erker and Heinzel, and with +the similar conclusion that the bite of the Italian tarantula ordinarily +causes no severe symptoms. In this conclusion, Kobert, though +firmly convinced of the poisonous nature of some spiders, coincides. +He also believes that striking symptoms may be simulated or artificially +induced by patients in order to attract interest, or because +they have been assured that the bite, under all circumstances, caused +tarantism.</p> + +<p>The so-called Russian tarantula, <i>Trochosa singoriensis</i> (<a href="#Fig_7">fig. 7</a>), is +much larger than the Italian species, and is much feared. Kobert +carried out a series of careful experiments with this species and his +results have such an important bearing on the question of the venomous +nature of the tarantula that we quote his summary. Experimenting +first on nearly a hundred living specimens of <i>Trochosa +singoriensis</i> from Crimea he says that:<span class="pagenum"><a name="Page_12" id="Page_12">[Pg 12]</a></span></p> + +<p>"The tarantulas, no matter how often they were placed on the +skin, handled, and irritated, could not be induced to bite either myself, +the janitor, or the ordinary experimental animals. The objection +that the tarantulas were weak and indifferent cannot stand, for as +soon as I placed two of them on the shaved skin of a rabbit, instead of +an attack on the animal, there began a furious battle between the +two spiders, which did not cease until one of the two was killed."</p> + +<div class="figleft" style="width: 300px;"><a name="Fig_7" id="Fig_7"></a> +<a href="images/f007-full.png"><img src="images/f007.png" width="300" height="373" alt="7. Trochosa singoriensis. After Kobert." title="7. Trochosa singoriensis. After Kobert." /></a> +<span class="caption">7. Trochosa singoriensis. After Kobert.</span> +</div> + +<p>"Since the spiders would not +bite, I carefully ground up the +fresh animals in physiological +salt solution, preparing an extract +which must have contained, in +solution, all of the poisonous +substance of their bodies. While +in the case of <i>Latrodectus</i>, as we +shall see, less than one specimen +sufficed to yield an active extract, +I have injected the filtered extract +of six fresh Russian tarantulas, +of which each one was much +heavier than an average <i>Latrodectus</i>, +subcutaneously and into +the jugular vein of various cats +without the animals dying or +showing any special symptoms. +On the basis of my experiments I can therefore only say that the +quantity of the poison soluble in physiological salt solution, even +when the spiders are perfectly fresh and well nourished, is very +insignificant. That the poison of the Russian tarantula is not +soluble in physiological salt solution, is exceedingly improbable. +Moreover, I have prepared alcoholic extracts and was unable to +find them active. Since the Russian spider exceeds the Italian in +size and in intensity of the bite, it seems very improbable to me that +the pharmacological test of the Italian tarantula would yield +essentially other results than those from the Russian species."</p> + +<p>To the <a name="Avicularoidea" id="Avicularoidea"></a><b>Avicularoidea</b> belong the largest and most formidable +appearing of the spiders and it is not strange that in the New World +they have fallen heir to the bad reputation, as well as to the name of +the tarantula of Europe. In this country they occur only in the +South or in the far West, but occasionally living specimens are brought<span class="pagenum"><a name="Page_13" id="Page_13">[Pg 13]</a></span> +to our northern ports in shipments of bananas and other tropical +produce, and are the source of much alarm. It should be mentioned, +however, that the large spider most frequently found under such circumstances +is not a tarantula at all, but one of the Heteropodidæ, or +giant crab-spiders, (<a href="#Fig_8">fig. 8</a>).</p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_8" id="Fig_8"></a> +<a href="images/f008-full.png"><img src="images/f008.png" width="400" height="341" alt="8. The giant crab-spider or banana spider (Heteropoda venatoria). +Natural size. After Comstock." title="8. The giant crab-spider or banana spider (Heteropoda venatoria). +Natural size. After Comstock." /></a> +<span class="caption">8. The giant crab-spider or banana spider (Heteropoda venatoria). +Natural size. After Comstock.</span> +</div> + +<p>In spite of their prominence and the fear which they arouse there +are few accurate data regarding these American tarantulas. It has +often been shown experimentally that they can kill small birds and +mammals, though it is doubtful if these form the normal prey of any +of the species, as has been claimed. There is no question but that +the mere mechanical injury which they may inflict, and the consequent +chances of secondary infection, justify, in part, their bad reputation. +In addition to the injury from their bite, it is claimed that the body +hairs of several of the South American species are readily detached +and are urticating.</p> + +<p>Recently, Phisalix (1912) has made a study of the physiological +effects of the venom of two Avicularoidea, <i>Phormictopus carcerides</i> +Pocock, from Haiti and <i>Cteniza sauvagei</i> Rossi, from Corsica. The +glands were removed aseptically and ground up with fine, sterilized +sand in distilled water. The resultant liquid was somewhat viscid, +colorless, and feebly alkaline. Injected into sparrows and mice the<span class="pagenum"><a name="Page_14" id="Page_14">[Pg 14]</a></span> +extract of <i>Phormictopus</i> proved very actively poisonous, that from a +single spider being sufficient to kill ten sparrows or twenty mice. It +manifested itself first and, above all, as a narcotic, slightly lowering +the temperature and paralyzing the respiration. Muscular and +cardiac weakening, loss of general sensibility, and the disappearance +of reflexes did not occur until near the end. The extract from <i>Cteniza</i> +was less active and, curiously enough, the comparative effect on +sparrows and on mice was just reversed.</p> + + +<p><a name="Spiders_of_the_Genus_Latrodectus" id="Spiders_of_the_Genus_Latrodectus"></a><b>Spiders of the Genus Latrodectus.</b>—While most of the popular +accounts of evil effects from the bites of spiders will not stand investigation, +it is a significant fact that, the world over, the best authenticated +records refer to a group of small and comparatively insignificant +spiders belonging to the genus <i>Latrodectus</i>, of the family Theridiidæ. +The dread "Malmigniatte" of Corsica and South Europe, the "Karakurte" +of southeastern Russia, the "Katipo" of New Zealand, the +"Mena-vodi" and "Vancoho" of Madagascar, and our own <i>Latrodectus +mactans</i>, all belong to this genus, and concerning all of these the most +circumstantial accounts of their venomous nature are given. These +accounts are not mere fantastic stories by uneducated natives but in +many cases are reports from thoroughly trained medical men.</p> + +<p>The symptoms produced are general, rather than local. As +summarized by Kobert (1901) from a study of twenty-two cases +treated in 1888, in the Kherson (Russia) Government Hospital and +Berislaw (Kherson) District Hospital the typical case, aside from +complications, exhibits the following symptoms. The victim suddenly +feels the bite, like the sting of a bee. Swelling of the barely +reddened spot seldom follows. The shooting pains, which quickly +set in, are not manifested at the point of injury but localized at the +joints of the lower limb and in the region of the hip. The severity +of the pain forces the victim to the hospital, in spite of the fact that +they otherwise have a great abhorrence of it. The patient is unable +to reach the hospital afoot, or, at least, not without help, for there is +usually inability to walk. The patient, even if he has ridden, reaches +the hospital covered with cold sweat and continues to perspire for a +considerable period. His expression indicates great suffering. The +respiration may be somewhat dyspnœic, and a feeling of oppression +in the region of the heart is common. There is great aversion to +solid food, but increasing thirst for milk and tea. Retention of +urine, and constipation occur. Cathartics and, at night, strong<span class="pagenum"><a name="Page_15" id="Page_15">[Pg 15]</a></span> +narcotics are desired. Warm baths give great relief. After three +days, there is marked improvement and usually the patient is dismissed +after the fifth. This summary of symptoms agrees well with +other trustworthy records.</p> + +<p>It would seem, then, that Riley and Howard (1889), who discussed +a number of accounts in the entomological literature, were fully +justified in their statement that "It must be admitted that certain +spiders of the genus <i>Latrodectus</i> have the power to inflict poisonous +bites, which may (probably exceptionally and depending upon exceptional +conditions) bring about the death of a human being."</p> + +<p>And yet, until recently the evidence bearing on the question has +been most conflicting. The eminent arachnologist, Lucas, (1843) +states that he himself, had been repeatedly bitten by the Malmigniatte +without any bad effects. Dr. Marx, in 1890, gave before the Entomological +Society of Washington, an account of a series of experiments +to determine whether the bite of <i>Latrodectus mactans</i> is poisonous or +not. He described the poison glands as remarkably small<a name="FNanchor_A_1" id="FNanchor_A_1"></a><a href="#Footnote_A_1" class="fnanchor">[A]</a> and stated +that he had introduced the poison in various ways into guinea-pigs +and rabbits without obtaining any satisfactory results. Obviously, +carefully conducted experiments with the supposed venom were +needed and fortunately they have been carried out in the greatest +detail by Kobert (1901).</p> + +<p>This investigator pointed out that there were two factors which +might account for the discrepancies in the earlier experiments. In +the first place, the poison of spiders, as of snakes, might be so exhausted +after two or three bites that further bites, following directly, +might be without visible effect. Secondly, the application of the +poison by means of the bite, is exceedingly inexact, since even after +the most careful selection of the point of application, the poison might +in one instance enter a little vein or lymph vessel, and in another case +fail to do so. Besides, there would always remain an incalculable and +very large amount externally, in the nonabsorptive epithelium. +While all of these factors enter into the question of the effect of the +bite in specific instances, they must be as nearly as possible obviated +in considering the question of whether the spiders really secrete a +venom harmful to man.</p> +<p><span class="pagenum"><a name="Page_16" id="Page_16">[Pg 16]</a></span></p> +<p>Kobert therefore sought to prepare extracts which would contain +the active principles of the poison and which could be injected in +definite quantities directly into the blood of the experimental animal. +For this purpose various parts of the spiders were rubbed up in a mortar +with distilled water, or physiological salt solution, allowed to +stand for an hour, filtered, and then carefully washed, by adding water +drop by drop for twenty-four hours. The filtrate and the wash-water +were then united, well mixed and, if necessary, cleared by centrifuging +or by exposure to cold. The mixture was again filtered, +measured, and used, in part, for injection and, in part, for the determination +of the organic materials.</p> + +<p>Such an extract was prepared from the cephalothoraces of eight +dried specimens of the Russian <i>Latrodectus</i> and three cubic centimeters +of this, containing 4.29 mg. of organic material, were injected into +the jugular vein of a cat weighing 2450 grams. The previously very +active animal was paralyzed and lay in whatever position it was +placed. The sensibility of the skin of the extremities and the rump +was so reduced that there was no reaction from cutting or sticking. +There quickly followed dyspnœa, convulsions, paralysis of the +respiratory muscles and of the heart. In twenty-eight minutes the +cat was dead, after having exhibited exactly the symptoms observed +in severe cases of poisoning of man from the bite of this spider.</p> + +<p>These experiments were continued on cats, dogs, guinea pigs and +various other animals. Not only extracts from the cephalothorax, +but from other parts of the body, from newly hatched spiders, and +from the eggs were used and all showed a similar virulence. Every +effort was made to avoid sources of error and the experiments, conducted +by such a recognized authority in the field of toxicology, must +be accepted as conclusively showing that this spider and, presumably, +other species of the genus <i>Latrodectus</i> against which the clinical evidence +is quite parallel, possess a poison which paralyzes the heart and +central nervous system, with or without preliminary stimulus of the +motor center. If the quantity of the poison which comes into direct +contact with the blood is large, there may occur hæmolysis and +thrombosis of the vessels.</p> + +<p>On the other hand, check experiments were carried out, using +similar extracts of many common European spiders of the genera +<i>Tegenaria</i>, <i>Drassus</i>, <i>Agelena</i>, <i>Eucharia</i> and <i>Argyroneta</i>, as well as +the Russian tarantula, <i>Lycosa singoriensis</i>. In no other case was the +effect on experimental animals comparable to the <i>Latrodectus</i> extract.<span class="pagenum"><a name="Page_17" id="Page_17">[Pg 17]</a></span></p> + +<p>Kobert concludes that in its chemical nature the poison is neither +an alkaloid, nor a glycoside, nor an acid, but a toxalbumen, or poisonous +enzyme which is very similar to certain other animal poisons, +notably that of the scorpion.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_9" id="Fig_9"></a> +<a href="images/f009-full.png"><img src="images/f009.png" width="350" height="396" alt="9. Latrodectus mactans; (a) female, x 3; (b) venter of female; (c) dorsum of male. +After Comstock." title="9. Latrodectus mactans; (a) female, x 3; (b) venter of female; (c) dorsum of male. +After Comstock." /></a> +<span class="caption">9. Latrodectus mactans; (<i>a</i>) female, ×3; (<i>b</i>) venter of female; (<i>c</i>) dorsum of male. +After Comstock.</span> +</div> + +<p>The genus <i>Latrodectus</i> is represented in the United States by at +least two species, <i>L. mactans</i> and <i>L. geometricus</i>. Concerning <i>L. +mactans</i> there are very circumstantial accounts of serious injury and +even death in man<a name="FNanchor_B_2" id="FNanchor_B_2"></a><a href="#Footnote_B_2" class="fnanchor">[B]</a>. <i>Latrodectus mactans</i> is coal black, marked with +red or yellow or both. It has eight eyes, which are dissimilar in<span class="pagenum"><a name="Page_18" id="Page_18">[Pg 18]</a></span> +color and are distinctly in front of the middle of the thorax, the +lateral eyes of each side widely separate. The <span title="for tarsi read tarsus"><a name="AC_2" id="AC_2"></a>tarsi</span> of the fourth +pair of legs has a number of curved setæ in a single series. It has on +the ventral side of its abdomen an hour-glass shaped spot. The full-grown +female is about half an inch in length. Its globose abdomen is +usually marked with one or more red spots dorsally along the middle +line. The male is about half as long but has in addition to the dorsal +spots, four pairs of stripes along the sides. Immature females +resemble the male in coloring (<a href="#Fig_9">fig. 9</a>).</p> + +<p>Regarding the distribution of <i>Latrodectus mactans</i>, Comstock +states that: "Although it is essentially a Southern species, it occurs +in Indiana, Ohio, Pennsylvania, New Hampshire, and doubtless other +of the Northern States." <i>L. geometricus</i> has been reported from +California.</p> + + +<p><a name="Other_Venomous_Spiders" id="Other_Venomous_Spiders"></a><b>Other Venomous Spiders</b>—While conclusive evidence regarding +the venomous nature of spiders is meager and relates almost wholly +to that of the genus <i>Latrodectus</i>, the group is a large one and we are +not justified in dismissing arbitrarily, all accounts of injury from their +bites. Several species stand out as especially needing more detailed +investigation.</p> + +<p><i>Chiracanthium nutrix</i> is a common European species of the family +Clubionidæ, concerning which there is much conflicting testimony. +Among the reports are two by distinguished scientists whose accounts +of personal experiences cannot be ignored. A. Forel allowed a spider +of this species to bite him and not only was the pain extreme, but the +general symptoms were so severe that he had to be helped to his +house. The distinguished arachnologist, Bertkau reports that he, +himself, was bitten and that an extreme, burning pain spread almost +instantaneously over the arm and into the breast. There were slight +chills the same day and throbbing pain at the wound lasted for days. +While this particular species is not found in the United States, there +are two other representatives of the genus and it is possible that they +possess the same properties. We are unaware of any direct experimental +work on the poison.</p> + +<p><i>Epeira diadema</i>, of Europe, belongs to a wholly different group, +that of the orb-weavers, but has long been reputed venomous. Kobert +was able to prepare from it an extract whose effects were very similar +to that prepared from <i>Latrodectus</i>, though feebler in its action. Under +ordinary circumstances this spider is unable to pierce the skin of man<span class="pagenum"><a name="Page_19" id="Page_19">[Pg 19]</a></span> +and though Kobert's results seem conclusive, the spider is little to +be feared.</p> + +<p><i>Phidippus audax</i> (<i>P. tripunctatus</i>) is one of our largest Attids, +or jumping spiders. The late Dr. O. Lugger describes a case of severe +poisoning from the bite of this spider and though details are lacking, +it is quite possible that this and other large species of the same group, +which stalk their prey, may possess a more active poison than that of +web-building species.</p> + + +<p><a name="Summary" id="Summary"></a><b>Summary</b>—It is clearly established that our common spiders are +not to be feared and that the stories regarding their virulence are +almost wholly without foundation. +On the other hand, the +chances of secondary infection +from the bites of some of the +more powerful species are not +to be ignored.</p> + +<p>Probably all species possess +a toxin secreted by the poison +gland, virulent for insects and +other normal prey of the +spiders, but with little or no +effect on man.</p> + +<p>There are a very few species, +notably of the genus <i>Latrodectus</i>, +and possibly including the European +<i>Chiracanthium nutrix</i> and +<i>Epeira diadema</i>, which possess, +in addition, a toxalbumen +derived from the general body +tissue, which is of great virulence +and may even cause death in man and the higher animals.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_10" id="Fig_10"></a> +<a href="images/f010-full.png"><img src="images/f010.png" width="350" height="482" alt="10. A whip-scorpion (Mastigoproctus giganteus). +Half natural size. After Comstock." title="10. A whip-scorpion (Mastigoproctus giganteus). +Half natural size. After Comstock." /></a> +<span class="caption">10. A whip-scorpion (Mastigoproctus giganteus). +Half natural size. After Comstock.</span> +</div> + + +<h3><a name="THE_PEDIPALPIDA_OR_WHIP-SCORPIONS" id="THE_PEDIPALPIDA_OR_WHIP-SCORPIONS"></a>THE PEDIPALPIDA OR WHIP-SCORPIONS</h3> + +<p>The tailed whip-scorpions, belonging to the family Thelyphonidæ, +are represented in the United States by the giant whip-scorpion +<i>Mastigoproctus giganteus</i> (<a href="#Fig_10">fig. 10</a>), which is common in Florida, Texas +and some other parts of the South. In Florida, it is locally known as +the "grampus" or "mule-killer" and is very greatly feared. There is +no evidence that these fears have any foundation, and Dr. Marx +states that there is neither a poison gland nor a pore in the claw of the +chelicera.<span class="pagenum"><a name="Page_20" id="Page_20">[Pg 20]</a></span></p> + + +<h3><a name="THE_SCORPIONIDA_OR_TRUE_SCORPIONS" id="THE_SCORPIONIDA_OR_TRUE_SCORPIONS"></a>THE SCORPIONIDA, OR TRUE SCORPIONS</h3> + +<p>The true scorpions are widely distributed throughout warm countries +and everywhere bear an evil reputation. According to Comstock +(1912), about a score of species occur in the Southern United States. +These are comparatively small forms but in the tropics members of +this group may reach a length of seven or eight inches. They are +pre-eminently predaceous forms, which lie hidden during the day and +seek their prey by night.</p> + +<div class="figleft" style="width: 300px;"><a name="Fig_11" id="Fig_11"></a> +<a href="images/f011-full.png"><img src="images/f011.png" width="300" height="591" alt="11. A true scorpion. After +Comstock." title="11. A true scorpion. After +Comstock." /></a> +<span class="caption">11. A true scorpion. After +Comstock.</span> +</div> + +<p>The scorpions (<a href="#Fig_11">fig. 11</a>) possess large pedipalpi, terminated by +strongly developed claws, or chelæ. They may be distinguished from +all other Arachnids by the fact that the distinctly +segmented abdomen is divided into a +broad basal region of seven segments and a +terminal, slender, tail-like division of five +distinct segments.</p> + +<p>The last segment of the abdomen, or +telson, terminates in a ventrally-directed, +sharp spine, and contains a pair of highly +developed poison glands. These glands open +by two small pores near the tip of the spine. +Most of the species when running carry the +tip of the abdomen bent upward over the +back, and the prey, caught and held by the +pedipalpi, is stung by inserting the spine of +the telson and allowing it to remain for a +time in the wound.</p> + +<p>The glands themselves have been studied +in <i>Prionurus citrinus</i> by Wilson (1904). +He found that each gland is covered by a sheet of muscle on its +mesal and dorsal aspects, which may be described as the <i>compressor +muscle</i>. The muscle of each side is inserted by its edge along the +ventral inner surface of the chitinous wall of the telson, close to the +middle line, and by a broader insertion laterally. A layer of fine +connective tissue completely envelops each gland and forms the +basis upon which the secreting cells rest. The secreting epithelium +is columnar; and apparently of three different types of cells.</p> + +<p>1. The most numerous have the appearance of mucous cells, +resembling the goblet cells of columnar mucous membranes. The +nucleus, surrounded by a small quantity of protoplasm staining with +hæmatoxylin, lies close to the base of the cell.<span class="pagenum"><a name="Page_21" id="Page_21">[Pg 21]</a></span></p> + +<p>2. Cells present in considerable numbers, the peripheral portions +of which are filled with very numerous fine granules, staining +with acid dyes such as methyl orange.</p> + +<p>3. Cells few in number, filled with very large granules, or irregular +masses of a substance staining with hæmatoxylin.</p> + +<p>The poison, according to Kobert (1893), is a limpid, acid-reacting +fluid, soluble in water but insoluble in absolute alcohol and ether. +There are few data relative to its chemical nature. Wilson (1901) +states that a common Egyptian species, <i>Buthus quinquestriatus</i>, has +a specific gravity of 1.092, and contains 20.3% of solids and 8.4% ash.</p> + +<p>The venom of different species appears to differ not only quantitatively +but qualitatively. The effects of the bite of the smaller species +of the Southern United States may be painful but there is no satisfactory +evidence that it is ever fatal. On the other hand, certain +tropical species are exceedingly virulent and cases of death of man +from the bite are common.</p> + +<p>In the case of <i>Buthus quinquestriatus</i>, Wilson (1904) found the +symptoms in animals to be hypersecretion, salivation and lachrymation, +especially marked, convulsions followed by prolonged muscular +spasm; death from asphyxia. The temperature shows a +slight, rarely considerable, rise. Rapid and considerable increase +of blood-pressure (observed in dogs) is followed by a gradual fall with +slowing of the heart-beat. The coagulability of the blood is not +affected.</p> + +<p>An interesting phase of Wilson's work was the experiments on +desert mammals. The condition under which these animals exist +must frequently bring them in contact with scorpions, and he found +that they possess a degree of immunity to the venom sufficient at +least to protect them from the fatal effects of the sting.</p> + +<p>As far as concerns its effect on man, Wilson found that much +depended upon the age. As high as 60 per cent of the cases of +children under five, resulted fatally. Caroroz (1865), states that in a +Mexican state of 15,000 inhabitants, the scorpions were so abundant +and so much feared that the authorities offered a bounty for their +destruction. A result was a large number of fatalities, over two +hundred per year. Most of the victims were children who had +attempted to collect the scorpions.</p> + +<p>The treatment usually employed in the case of bites by the more +poisonous forms is similar to that for the bite of venomous snakes. +First, a tight ligature is applied above the wound so as to stop the<span class="pagenum"><a name="Page_22" id="Page_22">[Pg 22]</a></span> +flow of blood and lymph from that region. The wound is then +freely excised and treated with a strong solution of permanganate +of potash, or with lead and opium lotion.</p> + +<p>In recent years there have been many attempts to prepare an +antivenom, or antiserum comparable to what has been used so +effectively in the case of snake bites. The most promising of these +is that of Todd (1909), produced by the immunization of suitable +animals. This antivenom proved capable of neutralizing the venom +when mixed <i>in vitro</i> and also acts both prophylactically and curatively +in animals. Employed curatively in man, it appears to have +a very marked effect on the intense pain following the sting, and +the evidence so far indicates that its prompt use greatly reduces +the chance of fatal results.</p> + + +<h3><a name="THE_SOLPUGIDA_OR_SOLPUGIDS" id="THE_SOLPUGIDA_OR_SOLPUGIDS"></a>THE SOLPUGIDA, OR SOLPUGIDS</h3> + +<p>The <b>Solpugida</b> are peculiar spider-like forms which are distinguished +from nearly all other +arachnids by the fact that +they possess no true cephalothorax, +the last two leg-bearing +segments being distinct, +resembling those of the abdomen +in this respect. The +first pair of legs is not used +in locomotion but seemingly +functions as a second pair of +pedipalpi. <a href="#Fig_12">Figure 12</a> illustrates +the striking peculiarities +of the group. They are +primarily desert forms and +occur in the warm zones of +all countries. Of the two +hundred or more species, +Comstock lists twelve as +occurring in our fauna. +These occur primarily in the +southwest.</p> + +<div class="figleft" style="width: 300px;"><a name="Fig_12" id="Fig_12"></a> +<a href="images/f012-full.png"><img src="images/f012.png" width="300" height="424" alt="12. A solpugid (Eremobates cinerea). After Comstock." title="12. A solpugid (Eremobates cinerea). After Comstock." /></a> +<span class="caption">12. A solpugid (Eremobates cinerea). After Comstock.</span> +</div> + +<p>The Solpugida have long +borne a bad reputation and, regarding virulence, have been classed +with the scorpions. Among the effects of their bites have been<span class="pagenum"><a name="Page_23" id="Page_23">[Pg 23]</a></span> +described painful swelling, gangrene, loss of speech, cramps, delirium, +unconsciousness and even death. Opposed to the numerous loose +accounts of poisoning, there are a number of careful records by +physicians and zoölogists which indicate clearly that the effects are +local and though they may be severe, they show not the slightest +symptom of direct poisoning.</p> + +<p>More important in the consideration of the question is the fact +that there are neither poison glands nor pores in the fangs for the +exit of any poisonous secretion. This is the testimony of a number +of prominent zoölogists, among whom is Dr. A. Walter, who wrote +to Kobert at length on the subject and whose conclusions are presented +by him.</p> + +<p>However, it should be noted that the fangs are very powerful +and are used in such a manner that they may inflict especially severe +wounds. Thus, there may be more opportunity for secondary +infection than is usual in the case of insect wounds.</p> + +<p>The treatment of the bite of the Solpugida is, therefore, a matter +of preventing infection. The wound should be allowed to bleed +freely and then washed out with a 1:3000 solution of corrosive +sublimate, and, if severe, a wet dressing of this should be applied. +If infection takes place, it should be treated in the usual manner, +regardless of its origin.</p> + + +<h3><a name="THE_ACARINA_OR_MITES_AND_TICKS" id="THE_ACARINA_OR_MITES_AND_TICKS"></a>THE ACARINA, OR MITES AND TICKS</h3> + +<p>A number of the parasitic Acarina evidently secrete a +specific poison, presumably carried by the saliva, but in most cases +its effect on man is insignificant. There is an abundant literature +dealing with the poisonous effect of the bite of these forms, especially +the ticks, but until recently it has been confused by failure to recognize +that various species may transmit diseases of man, rather than +produce injury through direct poisoning. We shall therefore +discuss the Acarina more especially in subsequent chapters, dealing +with parasitism and with disease transmission.</p> + +<p>Nevertheless, after the evidence is sifted, there can be no doubt +that the bites of certain ticks may occasionally be followed by a +direct poisoning, which may be either local or general in its effects. +Nuttall (1908) was unable to determine the cause of the toxic effect, +for, in <i>Argas persicus</i>, the species most often implicated, he failed to +get the slightest local or general effect on experimental animals, from +the injection of an emulsion prepared by crushing three of the ticks.<span class="pagenum"><a name="Page_24" id="Page_24">[Pg 24]</a></span></p> + +<p>It seems clearly established that the bite of certain ticks may +cause a temporary paralysis, or even complete paralysis, involving +the organs of respiration or the heart, and causing death. In 1912, +Dr. I. U. Temple, of Pendleton, Oregon, reported several cases of +what he called "acute ascending paralysis" associated with the occurrence +of ticks on the head or the back of the neck. A typical severe +case was that of a six year old child, who had retired in her usual +normal health. The following morning upon arising she was unable +to stand on her feet. She exhibited paralysis extending to the knees, +slight temperature, no pain, sensory nerves normal, motor nerves +completely paralyzed, reflexes absent. The following day the paralysis +had extended to the upper limbs, and before night of the third +day the nerves of the throat (hypoglossal) were affected. The thorax +and larynx were involved, breathing was labored, she was unable +to swallow liquids, phonation was impossible and she could only make +low, guttural sounds. At this stage, two ticks, fully distended with +blood, were found over the junction of the spinal column with the +occipital bones in the hollow depression. They were removed by +the application of undiluted creoline. Though the child's life was +despaired of, by the following morning she was very much improved. +By evening she was able to speak. The paralysis gradually receded, +remaining longest in the feet, and at the end of one week the patient +was able to go home.</p> + +<p>There was some doubt as to the exact species of tick implicated +in the cases which Dr. Temple reported, although the evidence +pointed strongly to <i>Dermacentor venustus</i>.<a name="FNanchor_C_3" id="FNanchor_C_3"></a><a href="#Footnote_C_3" class="fnanchor">[C]</a> Somewhat later, Hadwen +(1913) reported that "tick paralysis" occurs in British Columbia, +where it affects not only man, but sheep and probably other animals. +It is caused by the bites of <i>Dermacentor venustus</i> and was experimentally +produced in lambs and a dog (Hadwen and Nuttall, 1913). +It is only when the tick begins to engorge or feed rapidly, some days +after it has become attached, that its saliva produces pathogenic +effects.</p> + +<p>Ulceration following tick bite is not uncommon. In many of the +instances it is due to the file-like hypostome, with its recurved teeth, +being left in the wound when the tick is forcibly pulled off.</p> +<p><span class="pagenum"><a name="Page_25" id="Page_25">[Pg 25]</a></span></p> + + +<h3><a name="THE_MYRIAPODA_OR_CENTIPEDES_AND_MILLIPEDES" id="THE_MYRIAPODA_OR_CENTIPEDES_AND_MILLIPEDES"></a>THE MYRIAPODA, OR CENTIPEDES AND MILLIPEDES</h3> + +<p>The old class, Myriapoda includes the <b>Diplopoda</b>, or +millipedes, and the <b>Chilopoda</b>, or centipedes. The present +tendency is to raise these groups to the rank of +classes.</p> + + +<h3>The Diplopoda</h3> + +<div class="figleft" style="width: 125px;"><a name="Fig_13" id="Fig_13"></a> +<a href="images/f013-full.png"><img src="images/f013.png" width="125" height="527" alt="13. A millipede. +After Comstock." title="13. A millipede. +After Comstock." /></a> +<span class="caption">13. A millipede. +After Comstock.</span> +</div> + +<p>The <b>Diplopoda</b>, or millipedes (<a href="#Fig_13">fig. 13</a>), are characterized +by the presence of two pairs of legs to a segment. +The largest of our local myriapods belong to this group. +They live in moist places, feeding primarily on decaying +vegetable matter, though a few species occasionally +attack growing plants.</p> + +<p>The millipedes are inoffensive and harmless. <i>Julus +terrestris</i>, and related species, when irritated pour out +over the entire body a yellowish secretion which escapes +from cutaneous glands. It is +volatile, with a pungent odor, +and Phisalix (1900) has shown +that it is an active poison when +injected into the blood of experimental +animals. This, however, +does not entitle +them to be considered +as poisonous arthropods, +in the sense of this +chapter, any more than +the toad can be considered +poisonous to +man because it secretes +a venom from its cutaneous +glands.</p> + + +<h3>The Chilopoda</h3> + +<div class="figright" style="width: 323px;"><a name="Fig_14" id="Fig_14"></a> +<a href="images/f014-full.png"><img src="images/f014.png" width="323" height="456" alt="14. Two common centipedes. + +(a) Lithobius forficatus. +After Comstock. + +(b) Scutigera forceps. Natural +size; after Howard." title="14. Two common centipedes. + +(a) Lithobius forficatus. +After Comstock. + +(b) Scutigera forceps. Natural +size; after Howard." /></a> +<span class="caption">14. Two common centipedes. + +(<i>a</i>) Lithobius forficatus. +After Comstock. + +(<i>b</i>) Scutigera forceps. Natural +size; after Howard.</span> +</div> + +<p>The <b>Chilopoda</b>, or +centipedes (<a href="#Fig_14">fig. 14</a>), unlike +the millipedes, are +predaceous forms, and +possess well developed +poison glands for killing +their prey. These<span class="pagenum"><a name="Page_26" id="Page_26">[Pg 26]</a></span> +glands are at the base of the first pair of legs (<a href="#Fig_15">fig. 15</a>), which are +bent forward so as to be used in holding their prey. The legs +terminate in a powerful claw, at the tip of which is the +outlet of the poison glands.</p> + +<p>The poison is a limpid, homogeneous, slightly acid +fluid, which precipitates in distilled water. Briot (1904) +extracted it from the glands of <i>Scolopendra morsitans</i>, a +species common in central France, and found that it was +actively venomous for the ordinary experimental animals. +A rabbit of two kilograms weight received an +injection of three cubic centimeters in the vein of the ear +and died in a minute. A white rat, weighing forty-eight +grams, received one and a half cubic centimeters in the +hind leg. There was an almost immediate paralysis of +the leg and marked necrosis of the tissues.</p> + +<div class="figleft" style="width: 200px;"><a name="Fig_15" id="Fig_15"></a> +<a href="images/f015-full.png"><img src="images/f015.png" width="200" height="367" alt="15. Mandible of +Scolopendra +cingulata +showing +venom +gland. After +Dubosq." title="15. Mandible of +Scolopendra +cingulata +showing +venom +gland. After +Dubosq." /></a> +<span class="caption">15. Mandible of +Scolopendra +cingulata +showing +venom +gland. After +Dubosq.</span> +</div> + +<p>As for the effect on man, there is little foundation for the fear +with which centipedes are regarded. Our native species produce, +at most, local symptoms,—sometimes severe local pain and swelling,—but +there is no authentic record of fatal results. In the tropics, +some of the species attain a large size, <i>Scolopendra gigantea</i> reaching +a length of nearly a foot. These forms are justly feared, and there +is good evidence that death sometimes, though rarely, results from +their bite.</p> + +<p>One of the most careful accounts of death from the sting of the +scorpion is that of Linnell, (1914), which relates to a comparatively +small Malayan species, unfortunately undetermined. The patient, +a coolie, aged twenty, was admitted to a hospital after having been +stung two days previously on the left heel. For cure, the other +coolies had made him eat the head of the scorpion. On admission, +the patient complained of "things creeping all over the body". +Temp. 102.8°. On the fourth day he had paralysis of the legs, and +on the fifth day motor paralysis to the umbilicus, sensation being +unaltered. On the sixth day there was retention of the urine and +on the ninth day (first test after third day) sugar was present. On +the thirteenth day the patient became comatose, but could be +roused to eat and drink. The temperature on the following day fell +below 95° and the patient was still comatose. Death fifteenth day.</p> + +<p>Examination of the spinal (lumbar) cord showed acute disseminated +myelitis. In one part there was an acute destruction of the +anterior horn and an infiltration of round cells. In another portion<span class="pagenum"><a name="Page_27" id="Page_27">[Pg 27]</a></span> +Clarke's column had been destroyed. The perivascular sheaths +were crowded with small round cells and the meninges were congested. +Some of the cells of the anterior horn were swollen and the +nuclei eccentric; chromatolysis had occurred in many of them.</p> + +<p>As for treatment, Castellani and Chalmers (1910), recommend +bathing the part well with a solution of ammonia (one in five, or one +in ten). After bathing, apply a dressing of the same alkali or, if +there is much swelling and redness, an ice-bag. If necessary, hypodermic +injections of morphine may be given to relieve the pain. +At a later period fomentations may be required to reduce the local +inflammation.</p> + + +<h3><a name="THE_HEXAPODA_OR_TRUE_INSECTS" id="THE_HEXAPODA_OR_TRUE_INSECTS"></a>THE HEXAPODA OR TRUE INSECTS</h3> + +<p>There are a number of <b>Hexapoda</b>, or true insects, which are, in +one way or another, poisonous to man. These belong primarily +to the orders Hemiptera, or true bugs; Lepidoptera, or butterflies +and moths (larval forms); Diptera, or flies; Coleoptera, or beetles; +and Hymenoptera, or ants, bees, and wasps. There are various ways +in which they may be poisonous.</p> + +<p>1. <i>Piercing</i> or <i>biting</i> forms may inject an irritating or poisonous +saliva into the wound caused by their mouth-parts.</p> + +<p>2. <i>Stinging forms</i> may inject a poison, from glands at the caudal +end of the abdomen, into wounds produced by a specially modified +ovipositer, the <i>sting</i>.</p> + +<p>3. <i>Nettling</i> properties may be possessed by the hairs of the insect.</p> + +<p>4. <i>Vescicating</i>, or <i>poisonous blood plasma</i>, or <i>body fluids</i> are +known to exist in a large number of species and may, under exceptional +circumstances, affect man.</p> + +<p>For convenience of discussion, we shall consider poisonous insects +under these various headings. In this, as in the preceding discussion, +no attempt will be made to give an exhaustive list of the poisonous +forms. Typical instances will be selected and these will be chosen +largely from North American species.</p> + + +<h3><a name="PIERCING_OR_BITING_INSECTS_POISONOUS_TO_MAN" id="PIERCING_OR_BITING_INSECTS_POISONOUS_TO_MAN"></a>PIERCING OR BITING INSECTS POISONOUS TO MAN</h3> + + +<h3><span class="smcap"><a name="Hemiptera" id="Hemiptera"></a>Hemiptera</span></h3> + +<p>Several families of the true bugs include forms which, while +normally inoffensive, are capable of inflicting painful wounds on man. +In these, as in all of the Hemiptera, the mouth-parts are modified<span class="pagenum"><a name="Page_28" id="Page_28">[Pg 28]</a></span> +to form an organ for piercing and +sucking. This is well shown by the +accompanying illustration (<a href="#Fig_16">fig. 16</a>).</p> + +<p>The upper lip, or <i>labrum</i>, is much +reduced and immovable, the lower +lip, or <i>labium</i>, is elongated to form +a jointed sheath, within which the +lance-like mandibles and maxillæ +are enclosed. The mandibles are +more or less deeply serrate, depending +on the species concerned.</p> + +<div class="figleft" style="width: 400px;"><a name="Fig_16" id="Fig_16"></a> +<a href="images/f016-full.png"><img src="images/f016.png" width="400" height="339" alt="16. Beak of hemipteron." title="16. Beak of hemipteron." /></a> +<span class="caption">16. Beak of hemipteron.</span> +</div> + +<p>The poison is elaborated by the salivary glands, excepting, possibly, +in <i>Belostoma</i> where Locy is inclined to believe that it is secreted +by the maxillary glands. The salivary glands +of the Hemiptera have been the subject of +much study but the most recent, comprehensive +work has been done by Bugnion and Popoff, +(1908 and 1910) to whose text the reader is +referred for details.</p> + +<p>The Hemiptera have two +pairs of salivary glands: the +<i>primary gland</i>, of which the +efferent duct leads to the +salivary syringe, and the +<i>accessory gland</i>, of which the +very long and flexuous duct +empties into the primary duct +at its point of insertion. +Thus, when one observes the +isolated primary gland it appears as though it +had efferent ducts inserted at the same point. In +<i>Nepa</i> and the <i>Fulgoridæ</i> there are two accessory +glands and therefore apparently three ducts +at the same point on the primary gland. The +<i>ensemble</i> differs greatly in appearance in different +species but we shall show here Bugnion and +Popoff's figure of the apparatus of <i>Notonecta +maculata</i>, a species capable of inflicting a painful +bite on man (<a href="#Fig_17">fig. 17</a>).</p> + +<div class="figcenter" style="width: 200px;"><a name="Fig_17" id="Fig_17"></a> +<a href="images/f017-full.png"><img src="images/f017.png" width="200" height="361" alt="17. Salivary glands of +Notonecta maculata. +After Bugnion and +Popoff." title="17. Salivary glands of +Notonecta maculata. +After Bugnion and +Popoff." /></a> +<span class="caption">17. Salivary glands of +Notonecta maculata. +After Bugnion and +Popoff.</span> +</div> + +<div class="figcenter" style="width: 200px;"><a name="Fig_18" id="Fig_18"></a> +<a href="images/f018-full.png"><img src="images/f018.png" width="200" height="524" alt="18. Pharyngeal syringe or +salivary pump of Fulgora +maculata. After +Bugnion and Popoff." title="18. Pharyngeal syringe or +salivary pump of Fulgora +maculata. After +Bugnion and Popoff." /></a> +<span class="caption">18. Pharyngeal syringe or +salivary pump of Fulgora +maculata. After +Bugnion and Popoff.</span> +</div><p><span class="pagenum"><a name="Page_29" id="Page_29">[Pg 29]</a></span></p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_19" id="Fig_19"></a> +<a href="images/f019-full.png"><img src="images/f019.png" width="400" height="515" alt="19. Heteroptera, (a) Melanolestes picipes; (b) Notonecta undulata; (c, d) Aradus robustus +(c) adult, (d) nymph, much enlarged; (e) Arilus cristatus; (f) Belostoma americana; +(g) Nabis (Coriscus) subcoleoptratus, enlarged; (h) Cimex lectularius, (i) Oeciacus +vicarius, much enlarged; (j) Lyctocoris fitchii, much enlarged. After Lugger." title="19. Heteroptera, (a) Melanolestes picipes; (b) Notonecta undulata; (c, d) Aradus robustus +(c) adult, (d) nymph, much enlarged; (e) Arilus cristatus; (f) Belostoma americana; +(g) Nabis (Coriscus) subcoleoptratus, enlarged; (h) Cimex lectularius, (i) Oeciacus +vicarius, much enlarged; (j) Lyctocoris fitchii, much enlarged. After Lugger." /></a> +<span class="caption">19. Heteroptera, (<i>a</i>) Melanolestes picipes; (<i>b</i>) Notonecta undulata; (<i>c</i>, <i>d</i>) Aradus robustus +(<i>c</i>) adult, (<i>d</i>) nymph, much enlarged; (<i>e</i>) Arilus cristatus; (<i>f</i>) Belostoma americana; +(<i>g</i>) Nabis (Coriscus) subcoleoptratus, enlarged; (<i>h</i>) Cimex lectularius, (<i>i</i>) Oeciacus +vicarius, much enlarged; (<i>j</i>) Lyctocoris fitchii, much enlarged. After Lugger.</span> +</div> + +<p>Accessory to the salivary apparatus there is on the ventral side +of the head, underneath the pharynx, a peculiar organ which the +Germans have called the "Wanzenspritze," or syringe. The accompanying +figure of the structure in <i>Fulgora maculata</i> (<a href="#Fig_18">fig. 18</a>) shows +its relation to the ducts of the salivary glands and to the beak. It is<span class="pagenum"><a name="Page_30" id="Page_30">[Pg 30]</a></span> +made up of a dilatation forming the body of the pump, in which there +is a chitinous piston. Attached to the piston is a strong retractor +muscle. The function of the salivary pump is to suck up the saliva +from the salivary ducts and to force it out through the beak.</p> + +<p>Of the Hemiptera reported as attacking man, we shall consider +briefly the forms most frequently noted.</p> + + +<p><a name="The_Notonectidae_or_back_swimmers" id="The_Notonectidae_or_back_swimmers"></a>The <b>Notonectidæ</b>, or <i>back swimmers</i>, (<a href="#Fig_19">fig. 19<i>b</i></a>) are small, aquatic +bugs that differ from all others in that they always swim on their +backs. They are predaceous; feeding on insects and other small +forms. When handled carelessly they are able to inflict a painful +bite, which is sometimes as severe as the sting of a bee. In fact, +they are known in Germany as "Wasserbienen."</p> + + +<p><a name="The_Belostomatidae_or_giant_water_bugs" id="The_Belostomatidae_or_giant_water_bugs"></a>The <b>Belostomatidæ</b>, or <i>giant water bugs</i>, (<a href="#Fig_19">fig. 19<i>f</i></a>) include the largest +living Hemiptera. They are attracted to lights and on account of the +large numbers which swarm about the electric street lamps in some +localities they have received the popular name "electric light bugs." +Our largest representatives in the northern United States belong to +the two genera <i>Belostoma</i> and <i>Banacus</i>, distinguished from each +other by the fact that <i>Belostoma</i> has a groove on the under side of +the femur of the front leg, for the reception of the tibia.</p> + +<p>The salivary glands of Belostoma were figured by Leidy (1847) +and later were studied in more detail by Locy (1884). There are +two pairs of the glands, those of one pair being long and extending +back as far as the beginning of the abdomen, while the others are +about one-fourth as long. They lie on either side of the œsophagus. +On each side of the œsophagus there is a slender tube with a +sigmoid swelling which may serve as a poison reservoir. In addition +to this salivary system, there is a pair of very prominent glands +on the ventral side of the head, opening just above the base of the +beak. These Locy has called the "cephalic glands" and he suggests +that they are the source of the poison. They are the homologues +of the maxillary glands described for other Hemiptera, and it is by +no means clear that they are concerned with the production of +venom. It seems more probable that in <i>Belostoma</i>, as in other +Hemiptera, it is produced by the salivary glands, though the question +is an open one.</p> + +<p>The Belostomatidæ feed not only on insects, but on small frogs, +fish, salamanders and the like. Matheson (1907) has recorded the +killing of a good-sized bird by <i>Belostoma americana</i>. A woodpecker,<span class="pagenum"><a name="Page_31" id="Page_31">[Pg 31]</a></span> +or flicker, was heard to utter cries of distress, +and fluttered and fell from a tree. On examination +it was found that a bug of this species +had inserted its beak into the back part of +the skull and was apparently busily engaged +in sucking the blood or brains of the bird. +Various species of <i>Belostoma</i> have been cited +as causing painful bites in man. We can +testify from personal experience that the bite +of <i>Belostoma americana</i> may almost immediately cause severe, +shooting pains that +may extend throughout +the arm and that +they may be felt for several days.</p> + +<div class="figleft" style="width: 300px;"><a name="Fig_20" id="Fig_20"></a> +<a href="images/f020-full.png"><img src="images/f020.png" width="300" height="437" alt="20. Reduvius (Opsicœtus) +personatus. (×2)." title="20. Reduvius (Opsicœtus) +personatus. (×2)." /></a> +<span class="caption">20. Reduvius (Opsicœtus) +personatus. (×2).</span> +</div> + +<p>Relief from the pain may be obtained by +the use of dilute ammonia, or a menthol +ointment. In the not uncommon case of +secondary infection the usual treatment for +that should be adopted.</p> + +<div class="figright" style="width: 300px;"><a name="Fig_21a" id="Fig_21a"></a> +<a href="images/f021-full.jpg"><img src="images/f021.png" width="300" height="511" alt="21. (a) Reduvius personatus, +nymph. +Photograph by M. V. S." title="21. (a) Reduvius personatus, +nymph. +Photograph by M. V. S." /></a> +<span class="caption">21. (<i>a</i>) Reduvius personatus, +nymph. +Photograph by M. V. S.</span> +</div> + + +<p><a name="The_Reduviidae_or_assassin-bugs" id="The_Reduviidae_or_assassin-bugs"></a>The <b>Reduviidæ</b>, or <i>assassin-bugs</i> are capable +of inflicting very painful wounds, as +most collectors +of Hemiptera +know to +their sorrow. +Some species are frequently to be +found in houses and outhouses and +Dr. Howard suggests that many of +the stories of painful spider bites +relate to the attack of these forms.</p> + +<div class="figleft" style="width: 375px;"> +<a href="images/f021b-full.jpg"><img src="images/f021b.png" width="375" height="512" alt="21. (b) Reduvius personatus, +adult (×2) +Photograph by M. V. S." title="21. (b) Reduvius personatus, +adult (×2) +Photograph by M. V. S." /></a> +<span class="caption">21. (<i>b</i>) Reduvius personatus, +adult (×2) +Photograph by M. V. S.</span> +</div> + +<p>An interesting psychological study +was afforded in the summer of 1899, +by the "kissing-bug" scare which +swept over the country. It was +reported in the daily papers that a +new and deadly bug had made its +appearance, which had the unpleasant +habit of choosing the lips or cheeks<span class="pagenum"><a name="Page_32" id="Page_32">[Pg 32]</a></span> +for its point of attack on man. So widespread were the stories +regarding this supposedly new insect that station entomologists all +over the country began to receive suspected +specimens for identification. At +Cornell there were received, among +others, specimens of stone-flies, may-flies +and even small moths, with inquiries +as to whether they were "kissing-bugs."</p> + +<div class="figleft" style="width: 300px;"><a name="Fig_22" id="Fig_22"></a> +<a href="images/f022-full.png"><img src="images/f022.png" width="300" height="391" alt="22. Rasahus biguttatus. (×2). After Howard." title="22. Rasahus biguttatus. (×2). After Howard." /></a> +<span class="caption">22. Rasahus biguttatus. (×2). After Howard.</span> +</div> + +<p>Dr. L. O. Howard has shown that the +scare had its origin in newspaper reports +of some instances of bites by either +<i>Melanolestes picipes</i> (<a href="#Fig_19">fig. 19a</a>) or <i>Opsicoetes +personatus</i> (<a href="#Fig_20">fig. 20</a>), in the vicinity +of Washington, D. C. He then discusses +in considerable detail the more prominent +of the Reduviidæ which, with +greater or less frequency pierce the skin +of human beings. These are <i>Opsicoetes personatus</i>, <i>Melanolestes +picipes</i>, <i>Coriscus subcoleoptratus</i> (<a href="#Fig_19">fig. 19<i>g</i></a>), <i>Rasahus thoracicus</i>, +<i>Rasahus biguttatus</i> (<a href="#Fig_22">fig. 22</a>), <i>Conorhinus sanguisugus</i> (<a href="#Fig_71">fig. 71</a>), and <span title="for C. (Conorhinus) abdominalis read Melanolestes abdominalis"><a name="AC_3" id="AC_3"></a><i>C. +abdominalis</i></span> (<a href="#Fig_23">fig. 23</a>).</p> + +<div class="figright" style="width: 400px;"><a name="Fig_23" id="Fig_23"></a> +<a href="images/f023-full.png"><img src="images/f023.png" width="400" height="386" alt="23. Conorhinus abdominalis (×2). After +Marlatt." title="23. Conorhinus abdominalis (×2). After +Marlatt." /></a> +<span class="caption">23. <span title="for C. (Conorhinus) abdominalis read Melanolestes abdominalis">Conorhinus abdominalis</span> (×2). After +Marlatt.</span> +</div> + +<p>One of the most interesting of these species is <i>Reduvius personatus</i>, +(= <i>Opsicœtus personatus</i>), which is popularly known as the "masked +bed-bug hunter." It owes this +name to the fact that the immature +nymphs (<a href="#Fig_21a">fig. 21</a>) have their +bodies and legs completely covered +by dust and lint, and that they +are supposed to prey upon bed-bugs. +LeConte is quoted by Howard +as stating that "This species is +remarkable for the intense pain +caused by its bite. I do not know +whether it ever willingly plunges +its rostrum into any person, but +when caught, or unskilfully handled +it always stings. In this case the +pain is almost equal to the bite of a snake, and the swelling and +irritation which result from it will sometimes last for a week."<span class="pagenum"><a name="Page_33" id="Page_33">[Pg 33]</a></span></p> + +<p>A species which very commonly attacks man is <i>Conorhinus +sanguisugus</i>, the so-called "big bed-bug" of the south and southern +United States. It is frequently found in houses and is known to +inflict an exceedingly painful bite. As in the case of a number of +other predaceous Hemiptera, the salivary glands of these forms are +highly developed. The effect of the bite on their prey and, as Marlatt +has pointed out, the constant and uniform character of the symptoms +in nearly all cases of bites in man, clearly indicate that their saliva +contains a specific substance. No satisfactory studies of the secretions +have been made. On the other hand, Dr. Howard is doubtless +right in maintaining that the very serious results which sometimes +follow the bite are due to the introduction of extraneous poison +germs. This is borne out by the symptoms of most of the cases +cited in literature and also by the fact that treatment with corrosive +sublimate, locally applied to the wound, has yielded favorable results.</p> + + +<p><a name="Other_Hemiptera_Reported_as_Poisonous_to_Man" id="Other_Hemiptera_Reported_as_Poisonous_to_Man"></a><b>Other Hemiptera Reported as Poisonous to Man</b>—A large number +of other Hemiptera have been reported as attacking man. Of these, +there are several species of Lygæidæ, Coreidæ, and Capsidæ. Of the +latter, <i>Lygus pratensis</i>, the tarnished plant-bug, is reported by +Professor Crosby as sucking blood. <i>Orthotylus flavosparsus</i> is another +Capsid which has been implicated. <i>Empoasca mali</i> and <i>Platymetopius +acutus</i> of the Jassidæ have also been reported as having similar +habits.</p> + +<p>Whenever the periodical cicada or "seventeen-year locust" becomes +abundant, the newspapers contain accounts of serious results +from its bites. The senior author has made scores of attempts to +induce this species to bite and only once successfully. At that +time the bite was in no wise more severe than a pin-prick. A student +in our department reports a similar experience. There is no +case on record which bears evidence of being worthy of any credence, +whatsoever.</p> + +<p>Under the heading of poisonous Hemiptera we might consider the +bed-bugs and the lice. These will be discussed later, as parasites +and as carriers of disease, and therefore need only be mentioned here.</p> + + +<h3><a name="DIPTERA" id="DIPTERA"></a>DIPTERA</h3> + +<p>Several species of blood-sucking Diptera undoubtedly secrete a +saliva possessing poisonous properties. Chief among these are the +Culicidæ, or mosquitoes, and the Simuliidæ, or black-flies. As we +shall consider these forms in detail under the heading of parasitic<span class="pagenum"><a name="Page_34" id="Page_34">[Pg 34]</a></span> +species and insects transmitting disease, we shall discuss here only +the poison of the mosquitoes.</p> + +<p>It is well known that mosquitoes, when they bite, inject into the +wound a minute quantity of poison. The effect of this varies according +to the species of mosquito and also depends very much on the +susceptibility of the individual. Soon after the bite a sensation of +itching is noticed and often a wheal, or eminence, is produced on the +skin, which may increase to a considerable swelling. The scratching +which is induced may cause a secondary infection and thus lead to +serious results. Some people seem to acquire an immunity against +the poison.</p> + +<p>The purpose of this irritating fluid may be, as Reaumur suggested, +to prevent the coagulation of the blood and thus not only to cause +it to flow freely when the insect bites but to prevent its rapid coagulation +in the stomach. Obviously, it is not developed as a protective +fluid, and its presence subjects the group to the additional handicap +of the vengeance of man.</p> + +<div class="figleft" style="width: 425px;"><a name="Fig_24" id="Fig_24"></a> +<a href="images/f024-full.png"><img src="images/f024.png" width="425" height="150" alt="24. Diagram of a longitudinal section of a mosquito." title="24. Diagram of a longitudinal section of a mosquito." /></a> +<span class="caption">24. Diagram of a longitudinal section of a mosquito.</span> +</div> + +<p>As to the origin of the poison, there has been little question, +until recent years, that it was a secretion from the salivary glands. +Macloskie (1888) showed +that each gland is subdivided +into three lobes, +the middle of which differs +from the others in having +evenly granulated contents +and staining more deeply +than the others (<a href="#Fig_24">fig. 24</a>). This middle lobe he regarded as the source +of the poison. Bruck, (1911), by the use of water, glycerine, chloroform, +and other fluids, extracted from the bodies of a large number +of mosquitoes a toxine which he calls <i>culicin</i>. This he assumes +comes from the salivary glands. Animal experimentation showed +that this extract possessed hemolytic powers. Inoculated into the +experimenter's own skin it produced lesions which behaved exactly +as do those of mosquito bites.</p> + +<p>Similarly, most writers on the subject have concurred with the +view that the salivary glands are the source of the poison. However, +recent work, especially that of Nuttall and Shipley (1903), +and Schaudinn (1904), has shown that the evidence is by no means +conclusive. Nuttall dissected out six sets (thirty-six acini) of glands +from freshly killed <i>Culex pipiens</i> and placed them in a drop of salt<span class="pagenum"><a name="Page_35" id="Page_35">[Pg 35]</a></span> +solution. The drop was allowed to dry, it being thought that the +salt crystals would facilitate the grinding up of the glands with the +end of a small glass rod, this being done under microscopic control. +After grinding up, a small drop of water was added of the size of the +original drop of saline, and an equal volume of human blood taken +from the clean finger-tip was quickly mixed therewith, and the whole +drawn up into a capillary tube. Clotting was not prevented and no +hemolysis occurred. Salivary gland emulsion added to a dilute +suspension of corpuscles did not lead to hemolysis. This experiment +was repeated a number of times, with slight modification, but +with similar results. The data obtained from the series "do not +support the hypothesis that the salivary glands, at any rate in <i>Culex +pipiens</i>, contain a substance which prevents coagulation."</p> + +<p>Much more detailed, and the more important experiments made +along this line, are those of Schaudinn (1904). The results of these +experiments were published in connection with a technical paper +on the alternation of generations and of hosts in <i>Trypanosoma</i> and +<i>Spirochæta</i>, and for this reason seem to have largely escaped the notice +of entomologists. They are so suggestive that we shall refer to them +in some detail.</p> + +<p>Schaudinn observed that the three œsophageal diverticula (commonly, +but incorrectly, known as the "sucking stomach") (<a href="#Fig_24">fig. 24</a>) +usually contain large bubbles of gas and in addition, he always found +yeast cells. On the ground of numerous observations, Schaudinn +was convinced that these yeast plants are normal and constant +commensals of the insect. He regarded them as the cause of the gas +bubbles to be found in diverticula. It was found that as the insect +fed, from time to time the abdomen underwent convulsive contractions +which resulted in the emptying of the œsophageal diverticula and +the salivary glands through blood pressure.</p> + +<p>In order to test the supposed toxic action of the salivary glands, +Schaudinn repeatedly introduced them under his skin and that of +his assistant, in a drop of salt solution, and never obtained a suggestion +of the irritation following a bite of the insect, even though the +glands were carefully rubbed to fragments after their implantation. +Like Nuttall, he failed to get satisfactory evidence that the secretion +of the salivary glands retarded coagulation of the blood.</p> + +<p>He then carefully removed the œsophageal diverticula with their +content of yeast and introduced them into an opening in the skin +of the hand. Within a few seconds there was noticeable the characteristic<span class="pagenum"><a name="Page_36" id="Page_36">[Pg 36]</a></span> +itching irritation of the mosquito bite; and in a short time +there appeared reddening and typical swelling. This was usually +much more severe than after the usual mosquito bite, and the swelling +persisted and itched longer. This was because by the ordinary +bite of the mosquito most of the yeast cells are again sucked up, +while in these experiments they remained in the wound. These +experiments were repeated a number of times on himself, his assistant +and others, and always with the same result. From them Schaudinn +decided that the poisonous action of the mosquito bite is caused by +an enzyme from a commensal fungus. These conclusions have not, +as yet, been satisfactorily tested.</p> + +<p>Relief from the effect of the mosquito bite may be obtained by +bathing the swellings with weak ammonia or, according to Howard, +by using moist soap. The latter is to be rubbed gently on the puncture +and is said to speedily allay the irritation. Howard also quotes +from the <i>Journal of Tropical Medicine and Hygiene</i> to the effect that +a few drops of a solution of thirty to forty grains of iodine to an ounce +of saponated petroleum rubbed into the mosquito bite, or wasp sting, +allay the pain instantaneously.</p> + +<p>Methods of mosquito control will be discussed later, in considering +these insects as parasites and as carriers of disease.</p> + + +<h3><a name="STINGING_INSECTS" id="STINGING_INSECTS"></a>STINGING INSECTS</h3> + +<p>The stinging insects all belong to the order <b>Hymenoptera</b>. In a +number of families of this group the ovipositor is modified to form a +sting and is connected with poison-secreting glands. We shall +consider the apparatus of the honey-bee and then make briefer reference +to that of other forms.</p> + + +<p><a name="Apis_mellifica_the_honey_bee" id="Apis_mellifica_the_honey_bee"></a><b>Apis mellifica, the honey bee</b>—The sting of the worker honey-bee +is situated within a so-called sting chamber at the end of the +abdomen. This chamber is produced by the infolding of the greatly +reduced and modified eighth, ninth and tenth abdominal segments +into the seventh.<a name="FNanchor_D_4" id="FNanchor_D_4"></a><a href="#Footnote_D_4" class="fnanchor">[D]</a> From it the dart-like sting can be quickly exserted.</p> + +<div class="figright" style="width: 400px;"><a name="Fig_25" id="Fig_25"></a> +<a href="images/f025-full.png"><img src="images/f025.png" width="400" height="245" alt="25. Sting of a honey bee. Psn Sc, base of acid +poison gland; B Gl, alkaline poison gland; +Stn Plp, sting palpi; Sh B, bulb of sting; +Sh A, basal arm; Lct, lancets or darts; Sh s, +shaft of sting. Modified from Snodgrass." title="25. Sting of a honey bee. Psn Sc, base of acid +poison gland; B Gl, alkaline poison gland; +Stn Plp, sting palpi; Sh B, bulb of sting; +Sh A, basal arm; Lct, lancets or darts; Sh s, +shaft of sting. Modified from Snodgrass." /></a> +<span class="caption">25. Sting of a honey bee. <i>Psn Sc</i>, base of acid +poison gland; <i>B Gl</i>, alkaline poison gland; +<i>Stn Plp</i>, sting palpi; <i>Sh B</i>, bulb of sting; +<i>Sh A</i>, basal arm; <i>Lct</i>, lancets or darts; <i>Sh s</i>, +shaft of sting. Modified from Snodgrass.</span> +</div> + +<p>The sting (<a href="#Fig_25">fig. 25</a>) is made up of a central shaft, ventro-laterad of +which are the paired <i>lancets</i>, or darts, which are provided with sharp, +recurved teeth. Still further laterad lie the paired whitish, finger-like<span class="pagenum"><a name="Page_37" id="Page_37">[Pg 37]</a></span> +<i>sting palpi</i>. Comparative morphological as well as embryological +studies have clearly established that these three parts correspond +to the three pairs of +gonopophyses of the ovipositor +of more generalized insects.</p> + +<p>An examination of the internal +structures (<a href="#Fig_26">fig. 26</a>) reveals +two distinct types of poison +glands, the acid-secreting and +the alkaline-secreting glands, +and a prominent poison reservoir. +In addition, there is a +small pair of accessory structures +which have been called lubricating glands, on account of the +supposed function of their product. The acid-secreting gland empties +into the distal end of the poison reservoir which in turn pours the +secretion into the muscular bulb-like enlargement at the base of the +shaft. The alkaline secreting gland empties into the bulb ventrad +of the narrow neck of the reservoir.</p> + +<div class="figleft" style="width: 300px;"><a name="Fig_26" id="Fig_26"></a> +<a href="images/f026-full.png"><img src="images/f026.png" width="300" height="438" alt="26. Poison apparatus of a honey +bee. Modified from Snodgrass." title="26. Poison apparatus of a honey +bee. Modified from Snodgrass." /></a> +<span class="caption">26. Poison apparatus of a honey +bee. Modified from Snodgrass.</span> +</div> + +<p>The poison is usually referred to as formic acid. That it is not so +easily explained has been repeatedly shown and is evidenced by the +presence of the two types of glands. Carlet maintains that the product +of either gland is in itself innocent,—it +is only when they are combined that +the toxic properties appear.</p> + +<p>The most detailed study of the poison +of the honey-bee is that of Josef Langer +(1897), who in the course of his work used +some 25,000 bees. Various methods of +obtaining the active poison for experimental +purposes were used. For obtaining +the pure secretion, bees were held in the +fingers and compressed until the sting was +exserted, when a clear drop of the poison +was visible at its tip. This was then taken +up in a capillary tube or dilute solutions +obtained by dipping the tip of the sting into +a definite amount of distilled water.</p> + +<p>An aqueous solution of the poison was more readily obtained by +pulling out the sting and poison sacs by means of forceps, and grinding<span class="pagenum"><a name="Page_38" id="Page_38">[Pg 38]</a></span> +them up in water. The somewhat clouded fluid was then filtered +one or more times. For obtaining still greater quantities, advantage +was taken of the fact that while alcohol coagulates the poison, the +active principle remains soluble in water. Hence the stings with +the annexed glands where collected in 96 per cent alcohol, after +filtering off of the alcohol were dried at 40° C., then rubbed to a fine +powder and this was repeatedly extracted with water. Through +filtering of this aqueous extract there was obtained a yellowish-brown +fluid which produced the typical reactions, according to concentration +of the poison.</p> + +<p>The freshly expelled drop of poison is limpid, of distinct acid +reaction, tastes bitter and has a delicate aromatic odor. On evaporation, +it leaves a sticky residue, which at 100 degrees becomes fissured, +and suggests dried gum arabic. The poison is readily soluble in +water and possesses a specific gravity of 1.1313. On drying at room +temperature, it leaves a residue of 30 per cent, which has not lost in +poisonous action or in solubility. In spite of extended experiments, +Langer was unable to determine the nature of the active principle. +He showed that it was not, as had been supposed, an albuminous +body, but rather an organic base.</p> + +<p>The pure poison, or the two per cent aqueous solution, placed on +the uninjured skin showed absolutely no irritating effect, though it +produced a marked reaction on the mucus membrane of the nose or +eye. A single drop of one-tenth per cent aqueous solution of the +poison brought about a typical irritation in the conjunctiva of the +rabbit's eye. On the other hand, the application of a drop of the +poison, or its solution, to the slightest break in the skin, or by means +of a needle piercing the skin, produced typical effects. There is produced +a local necrosis, in the neighborhood of which there is infiltration +of lymphocytes, œdema, and hyperæmia.</p> + +<p>The effect of the sting on man (<a href="#Fig_27">fig. 27</a>) is usually transitory but +there are some individuals who are made sick for hours, by a single +sting. Much depends, too, on the place struck. It is a common +experience that an angry bee will attempt to reach the eye of its +victim and a sting on the lid may result in severe and prolonged +swelling. In the case of a man stung on the cheek, Legiehn observed +complete aphonia and a breaking out of red blotches all over the +body. A sting on the tongue has been known to cause such collateral +œdema as to endanger life through suffocation. Cases of death of +man from the attacks of bees are rare but are not unknown. Such<span class="pagenum"><a name="Page_39" id="Page_39">[Pg 39]</a></span> +results are usually from a number of stings but, rarely, death has +been known to follow a single sting, entering a blood vessel of a +particularly susceptible individual.</p> + +<div class="figcenter" style="width: 425px;"><a name="Fig_27" id="Fig_27"></a> +<a href="images/f027-full.png"><img src="images/f027.png" width="425" height="407" alt="Effect of bee stings. After Root." title="Effect of bee stings. After Root." /></a> +<span class="caption">Effect of bee stings. After Root.</span> +</div> + +<p>It is clearly established that partial immunity from the effects +of the poison may be acquired. By repeated injections of the venom, +mice have been rendered capable of bearing doses that certainly +would have killed them at first. It is a well-known fact that most +bee-keepers become gradually hardened to the stings, so that the +irritation and the swelling become less and less. Some individuals +have found this immunity a temporary one, to be reacquired each +season. A striking case of acquired immunity is related by the +Roots in their "A B C and X Y Z of Bee Culture." The evidence +in the case is so clear that it should be made more widely available +and hence we quote it here.</p> + +<p>A young man who was determined to become a bee-keeper, was so +susceptible to the poison that he was most seriously affected by a +single sting, his body breaking out with red blotches, breathing growing +difficult, and his heart action being painfully accelerated. "We +finally suggested taking a live bee and pressing it on the back of his +hand until it merely pierced his skin with the sting, then immediately +brushing off both bee and sting. This was done and since no serious +effect followed, it was repeated inside of four or five days. This +was continued for some three or four weeks, when the patient began to +have a sort of itching sensation all over his body. The hypodermic<span class="pagenum"><a name="Page_40" id="Page_40">[Pg 40]</a></span> +injections of bee-sting poison were then discontinued. At the end +of a month they were repeated at intervals of four or five days. +Again, after two or three weeks the itching sensation came on, but +it was less pronounced. The patient was given a rest of about a +month, when the doses were repeated as before." By this course +of treatment the young man became so thoroughly immunized that +neither unpleasant results nor swelling followed the attacks of the +insects and he is able to handle bees with the same freedom that any +experienced bee-keeper does.</p> + +<p>In an interesting article in the <i>Entomological News</i> for November, +1914, J. H. Lovell calls attention to the fact that "There has been a +widespread belief among apiarists that a beekeeper will receive more +stings when dressed in black than when wearing white clothing. +A large amount of evidence has been published in the various bee +journals showing beyond question that honey-bees under certain +conditions discriminate against black. A few instances may be +cited in illustration. Of a flock of twelve chickens running in a bee-yard +seven black ones were stung to death, while five light colored +ones escaped uninjured. A white dog ran among the bee-hives +without attracting much attention, while at the same time a black +dog was furiously assailed by the bees. Mr. J. D. Byer, a prominent +Canadian beekeeper, relates that a black and white cow, tethered +about forty feet from an apiary, was one afternoon attacked and +badly stung by bees. On examination it was found that the black +spots had five or six stings to one on the white. All noticed this fact, +although no one was able to offer any explanation. A white horse +is in much less danger of being stung, when driven near an apiary, +than a black one. It has, indeed, been observed repeatedly that +domestic animals of all kinds, if wholly or partially black, are much +more liable to be attacked by bees, if they wander among the hives, +than those which are entirely white."</p> + +<p>In order to test the matter experimentally, the following series +of experiments was performed. In the language of the investigator:</p> + +<p>"On a clear, warm day in August I dressed wholly in white with +the exception of a black veil. Midway on the sleeve of my right arm +there was sewed a band of black cloth ten inches wide. I then +entered the bee-yard and, removing the cover from one of the hives, +lifted a piece of comb with both hands and gently shook it. Instantly +many of the bees flew to the black band, which they continued to<span class="pagenum"><a name="Page_41" id="Page_41">[Pg 41]</a></span> +attack as long as they were disturbed. Not a single bee attempted +to sting the left sleeve, which was of course entirely white, and very +few even alighted upon it."</p> + +<p>"This experiment was repeated a second, third and fourth time; +in each instance with similar results. I estimated the number of bees +on the band of black cloth at various moments was from thirty to +forty; it was evident from their behavior that they were extremely +irritable. To the left white sleeve and other portions of my clothing +they paid very little attention; but the black veil was very frequently +attacked."</p> + +<p>"A few days later the experiments were repeated, but the band of +black cloth, ten inches wide, was sewed around my left arm instead +of around the right arm as before. When the bees were disturbed, +after the hive cover had been removed, they fiercely attacked the +band of black cloth as in the previous experiences; but the right white +sleeve and the white suit were scarcely noticed. At one time a part +of the black cloth was almost literally covered with furiously stinging +bees, and the black veil was assailed by hundreds. The bees behaved +in a similar manner when a second hive on the opposite side of the +apiary was opened."</p> + +<p>"A white veil which had been procured for this purpose, was next +substituted for the black veil. The result was most surprising, +for, whereas in the previous experiments hundreds of bees had +attacked the black veil, so few flew against the white veil as to cause +me no inconvenience. Undoubtedly beekeepers will find it greatly +to their advantage to wear white clothing when working among their +colonies of bees and manipulating the frames of the hives."</p> + +<p>When a honey-bee stings, the tip of the abdomen, with the entire +sting apparatus, is torn off and remains in the wound. Here the +muscles continue to contract, for some minutes, forcing the barbs +deeper and deeper into the skin, and forcing out additional poison +from the reservoir.</p> + +<p>Treatment, therefore, first consists in removing the sting without +squeezing out additional poison. This is accomplished by lifting +and scraping it out with a knife-blade or the fingernail instead of +grasping and pulling it out. Local application of alkalines, such as +weak ammonia, are often recommended on the assumption that the +poison is an acid to be neutralized on this manner, but these are of +little or no avail. They should certainly not be rubbed in, as that +would only accelerate the absorption of the poison. The use of<span class="pagenum"><a name="Page_42" id="Page_42">[Pg 42]</a></span> +cloths wrung out in hot water and applied as hot as can be borne, +affords much relief in the case of severe stings. The application of +wet clay, or of the end of a freshly cut potato is sometimes helpful.</p> + +<p>In extreme cases, where there is great susceptibility, or where +there may have been many stings, a physician should be called. He +may find strychnine injections or other treatment necessary, if +general symptoms develop.</p> + +<div class="figcenter" style="width: 300px;"><a name="Fig_28" id="Fig_28"></a> +<a href="images/f028-full.png"><img src="images/f028.png" width="300" height="215" alt="28. The poison apparatus of Formica. Wheeler, after Forel." title="28. The poison apparatus of Formica. Wheeler, after Forel." /></a> +<span class="caption">28. The poison apparatus of Formica. Wheeler, after Forel.</span> +</div> + + +<p><a name="Other_Stinging_Forms" id="Other_Stinging_Forms"></a><b>Other Stinging Forms</b>—Of the five thousand, or more, species +of bees, most possess a sting and poison apparatus and some of the +larger forms are capable of inflicting a much more painful sting than +that of the common honey-bee. In fact, some, like the bumble bees, +possess the advantage that they do not lose the sting from once using +it, but are capable of driving it in repeatedly. In the tropics there +are found many species of stingless bees but these are noted for their +united efforts to drive away intruders by biting. Certain species +possess a very irritating saliva which they inject into the wounds.</p> + +<p>The ants are not ordinarily regarded as worthy of consideration +under the heading of "stinging insects" but as a matter of fact, +most of them possess well developed stings and some of them, especially +in the tropics, are very justly feared. Even those which lack +the sting possess well-developed poison glands and the parts of the +entire stinging apparatus, in so far as it is developed in the various +species, may readily be homologized with those of the honey-bee.<span class="pagenum"><a name="Page_43" id="Page_43">[Pg 43]</a></span></p> + +<p>The ants lacking a sting are those of the subfamily <b>Camponotinæ</b>, +which includes the largest of our local species. It is an interesting +fact that some of these species possess the largest poison glands and +reservoir (<a href="#Fig_28">fig. 28</a>) and it is found that when they attack an enemy +they bring the tip of the abdomen forward and spray the poison in +such a way that it is introduced into the wound made by the powerful +mandibles.</p> + +<div class="figcenter" style="width: 450px;"><a name="Fig_29" id="Fig_29"></a> +<a href="images/f029-full.png"><img src="images/f029.png" width="450" height="367" alt="29. A harmless, but much feared larva, the "tomato worm." +Natural size. Photograph by M. V. S." title="29. A harmless, but much feared larva, the "tomato worm." +Natural size. Photograph by M. V. S." /></a> +<span class="caption">29. A harmless, but much feared larva, the "tomato worm." +Natural size. Photograph by M. V. S.</span> +</div> + +<p>More feared than any of the other Hymenoptera are the hornets +and wasps. Of these there are many species, some of which attain +a large size and are truly formidable. Phisalix (1897), has made a +study of the venom of the common hornet and finds that, like the +poison of the honey-bee, it is neither an albuminoid nor an alkaloid. +Its toxic properties are destroyed at 120° C. Phisalix also says that +the venom is soluble in alcohol. If this be true, it differs in this +respect from that of the bee. An interesting phase of the work of +Phisalix is that several of her experiments go to show that the venom +of hornets acts as a vaccine against that of vipers.</p> + + +<h3><a name="NETTLING_INSECTS" id="NETTLING_INSECTS"></a>NETTLING INSECTS</h3> + +<p>So far, we have considered insects which possess poison glands +connected with the mouth-parts or a special sting and which actively<span class="pagenum"><a name="Page_44" id="Page_44">[Pg 44]</a></span> +inject their poison into man. There remain to be considered those +insects which possess poisonous hairs or body fluids which, under +favorable circumstances, may act as poisons. To the first of these +belong primarily the larvæ of certain Lepidoptera.</p> + + +<h3><a name="LEPIDOPTERA" id="LEPIDOPTERA"></a>LEPIDOPTERA</h3> + +<div class="figcenter" style="width: 500px;"><a name="Fig_30" id="Fig_30"></a> +<a href="images/f030-full.png"><img src="images/f030.png" width="500" height="353" alt="30. Another innocent but much maligned caterpillar, the larva of the Regal moth. +Photograph by M. V. S." title="50030. Another innocent but much maligned caterpillar, the larva of the Regal moth. +Photograph by M. V. S." /></a> +<span class="caption">30. Another innocent but much maligned caterpillar, the larva of the Regal moth. +Photograph by M. V. S.</span> +</div> + +<p>When we consider the reputedly poisonous larvæ of moths and +butterflies, one of the first things to impress us is that we cannot +judge by mere appearance. Various species of Sphingid, or hawk-moth +larvæ, bear at the end of the body a chitinous horn, which is +often referred to as a "sting" and regarded as capable of inflicting +dangerous wounds. It would seem unnecessary to refer to this +absurd belief if it were not that each summer the newspapers contain +supposed accounts of injury from the "tomato worm" (<a href="#Fig_29">fig. 29</a>) +and others of this group. The grotesque, spiny larva (<a href="#Fig_30">fig. 30</a>) of +one of our largest moths, <i>Citheronia regalis</i> is much feared though +perfectly harmless, and similar instances could be multiplied.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_31a" id="Fig_31a"></a> +<a href="images/f031a-full.png"><img src="images/f031a.png" width="500" height="361" alt="31. The flannel moth (Lagoa crispata). (a) Poisonous larva." title="31. The flannel moth (Lagoa crispata). (a) Poisonous larva." /></a> +<span class="caption">31. The flannel moth (Lagoa crispata). (<i>a</i>) Poisonous larva.</span> +</div> + +<div class="figcenter" style="width: 400px;"> +<a href="images/f031b-full.png"><img src="images/f031b.png" width="400" height="323" alt="31. (b) Adult. Enlarged. Photographs by M. V. S." title="31. (b) Adult. Enlarged. Photographs by M. V. S." /></a> +<span class="caption">31. (<i>b</i>) Adult. Enlarged. Photographs by M. V. S.</span> +</div> + +<div class="figcenter" style="width: 396px;"><a name="Fig_32" id="Fig_32"></a> +<a href="images/f032-full.png"><img src="images/f032.png" width="396" height="372" alt="32. The poisonous saddle back caterpillar. Empretia +(Sibine) stimulea. Photograph by M. V. S." title="32. The poisonous saddle back caterpillar. Empretia +(Sibine) stimulea. Photograph by M. V. S." /></a> +<span class="caption">32. The poisonous saddle back caterpillar. Empretia +(Sibine) stimulea. Photograph by M. V. S.</span> +</div> + +<p>But if the larvæ are often misjudged on account of their ferocious +appearance, the reverse may be true. A group of most innocent +looking and attractive caterpillars is that of the flannel-moth larvæ,<span class="pagenum"><a name="Page_45" id="Page_45">[Pg 45]</a></span> +of which <i>Lagoa crispata</i> may be taken as an example. Its larva +(<a href="#Fig_31a">fig. 31</a>) has a very short and thick body, which is fleshy and completely +covered and hidden by long silken hairs of a tawny or brown +color, giving a convex form to the upper side. Interspersed among<span class="pagenum"><a name="Page_46" id="Page_46">[Pg 46]</a></span> +these long hairs are numerous +short spines connected +with underlying hypodermal +poison glands. These +hairs are capable of producing +a marked nettling +effect when they come in +contact with the skin. +This species is found in +our Atlantic and Southern +States. Satisfactory +studies of its poisonous +hairs and their glands have +not yet been made.</p> + +<div class="figcenter" style="width: 350px;"> +<a href="images/f033a-full.png"><img src="images/f033a.png" width="350" height="368" alt="33a. Io moth larvæ on willow. Photograph by M. V. S." title="33a. Io moth larvæ on willow. Photograph by M. V. S." /></a> +<span class="caption">33<i>a</i>. Io moth larvæ on willow. Photograph by M. V. S.</span> +</div> + +<p><i>Sibine stimulea (Empretia stimulea)</i>, or the +saddle-back caterpillar +(<a href="#Fig_32">fig. 32</a>), is another which possesses nettling hairs. This species +belongs to the group of Eucleidæ, or slug caterpillars. It can be +readily recognized +by its flattened +form, lateral, bristling +spines and by +the large green +patch on the back +resembling a +saddle-cloth, while +the saddle is represented +by an oval, +purplish-brown +spot. The small +spines are venomous +and affect +some persons very +painfully. The +larva feeds on the +leaves of a large +variety of forest +trees and also on +cherry, plum, and<span class="pagenum"><a name="Page_47" id="Page_47">[Pg 47]</a></span> +even corn leaves. It is to be found throughout the Eastern and +Southern United States.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_33b" id="Fig_33b"></a> +<a href="images/f033b-full.png"><img src="images/f033b.png" width="350" height="281" alt="33b. Io moth. Full grown larva. Photograph by M. V. S." title="33b. Io moth. Full grown larva. Photograph by M. V. S." /></a> +<span class="caption">33<i>b</i>. Io moth. Full grown larva. Photograph by M. V. S.</span> +</div> + +<div class="figcenter" style="width: 350px;"><a name="Fig_33c" id="Fig_33c"></a> +<a href="images/f033c-full.png"><img src="images/f033c.png" width="350" height="237" alt="33c. Io moth. Adult. Photograph by M. V. S." title="33c. Io moth. Adult. Photograph by M. V. S." /></a> +<span class="caption"><span title="for 33c read 34">33<i>c</i>.</span> Io moth. Adult. Photograph by M. V. S.</span> +</div> + +<p><i>Automeris io</i> is the best known of the nettling caterpillars. It is +the larva of the Io moth, one of the Saturniidæ. The mature caterpillar, +(<a href="#Fig_33b">fig. 33</a>), which reaches a length of two and one-half inches, is +of a beautiful pale green with sublateral stripes of cream and red color +and a few black spines among the green ones. The green radiating +spines give the body a mossy appearance. They are tipped with a<span class="pagenum"><a name="Page_48" id="Page_48">[Pg 48]</a></span> +slender chitinous hair whose tip is readily broken off in the skin and +whose poisonous content causes great irritation. Some individuals +are very susceptible to the poison, while others are able to handle +the larvæ freely without any discomfort. The larvæ feed on a wide +range of food plants. They are most commonly encountered on +corn and on willow, because of the opportunities for coming in contact +with them.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_35" id="Fig_35"></a> +<a href="images/f035-full.png"><img src="images/f035.png" width="350" height="258" alt="35. Larva of brown-tail moth. (Natural size). Photograph by M. V. S." title="35. Larva of brown-tail moth. (Natural size). Photograph by M. V. S." /></a> +<span class="caption">35. Larva of brown-tail moth. (Natural size). Photograph by M. V. S.</span> +</div> + +<p>The larvæ of the brown-tail moth (<i>Euproctis chrysorrhœa</i>) (fig. <a href="#Fig_35">35</a> +and <a href="#Fig_36">36</a>), where they occur in this country, are, on account of their +great numbers, the most serious of all poisonous caterpillars. It is +not necessary here, to go into details regarding the introduction of +this species from Europe into the New England States. This is all +available in the literature from the United States Bureau of Entomology +and from that of the various states which are fighting the species. +Suffice to say, there is every prospect that the pest will continue to +spread throughout the Eastern United States and Canada and that +wherever it goes it will prove a direct pest to man as well as to his +plants.</p> + +<p>Very soon after the introduction of the species there occurred in +the region where it had gained a foothold, a mysterious dermatitis of +man. The breaking out which usually occurred on the neck or other +exposed part of the body was always accompanied by an intense<span class="pagenum"><a name="Page_49" id="Page_49">[Pg 49]</a></span> +itching. It was soon found that this dermatitis was caused by certain +short, barbed hairs of the brown-tail caterpillars and that not only the +caterpillars but their cocoons and even the adult female moths might +harbor these nettling hairs and thus give rise to the irritation. In +many cases the hairs were wafted to clothing on the line and when this +was worn it might cause the same trouble. Still worse, it was found +that very serious internal injury was often caused by breathing or +swallowing the poisonous hairs.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_36" id="Fig_36"></a> +<a href="images/f036-full.png"><img src="images/f036.png" width="350" height="292" alt="36. Browntail moths. One male and two females. Photograph by +M. V. S." title="36. Browntail moths. One male and two females. Photograph by +M. V. S." /></a> +<span class="caption">36. Browntail moths. One male and two females. Photograph by +M. V. S.</span> +</div> + +<div class="figleft" style="width: 400px;"><a name="Fig_37a" id="Fig_37a"></a> +<a href="images/f037a-full.png"><img src="images/f037a.png" width="400" height="341" alt="37. (a) Ordinary hairs and three poison hairs of subdorsal +and lateral tubercles of the larva of the +browntail moth. Drawing by Miss Kephart." title="37. (a) Ordinary hairs and three poison hairs of subdorsal +and lateral tubercles of the larva of the +browntail moth. Drawing by Miss Kephart." /></a> +<span class="caption">37. (<i>a</i>) Ordinary hairs and three poison hairs of subdorsal +and lateral tubercles of the larva of the +browntail moth. Drawing by Miss Kephart.</span> +</div> + +<p>The earlier studies seemed to indicate that the irritation was +purely mechanical in origin, the result of the minute barbed hairs +working into the skin in large numbers. Subsequently, however, +Dr. Tyzzer (1907) demonstrated beyond question that the trouble +was due to a poison contained in the hairs. In the first place, it is +only the peculiar short barbed hairs which will produce the dermatitis +when rubbed on the skin, although most of the other hairs are sharply +barbed. Moreover, it was found that in various ways the nettling +properties could be destroyed without modifying the structure of the +hairs. This was accomplished by baking for one hour at 110° C, by +warming to 60° C in distilled water, or by soaking in one per cent. or in +one-tenth per cent. of potassium hydrate or sodium hydrate. The +most significant part of his work was the demonstration of the fact<span class="pagenum"><a name="Page_50" id="Page_50">[Pg 50]</a></span> +that if the nettling hairs are mingled with blood, they immediately +produce a change in the red corpuscles. These at once become +coarsely crenated, and the +roleaux are broken up in the +vicinity of the hair (<a href="#Fig_37b">fig. 37<i>b</i></a>). +The corpuscles decrease in +size, the coarse crenations +are transformed into slender +spines which rapidly disappear, +leaving the corpuscles +in the form of spheres, the +light refraction of which contrasts +them sharply with the +normal corpuscles. The +reaction always begins at the +basal sharp point of the hair. +It could not be produced by +purely mechanical means, such as the mingling of minute particles +of glass wool, the barbed hairs of a tussock moth, or the other +coarser hairs of the brown-tail, with the blood.</p> + +<p>The question of the source of the poison has been studied in our +laboratory by Miss Cornelia +Kephart. She first confirmed +Dr. Tyzzer's general results +and then studied carefully fixed +specimens of the larvæ to +determine the distribution of +the hairs and their relation to +the underlying tissues.</p> + +<div class="figcenter" style="width: 360px;"><a name="Fig_37b" id="Fig_37b"></a> +<a href="images/f037b-full.png"><img src="images/f037b.png" width="360" height="391" alt="37. (b) Effect of the poison on the blood corpuscles +of man. After Tyzzer." title="37. (b) Effect of the poison on the blood corpuscles +of man. After Tyzzer." /></a> +<span class="caption">37. (<i>b</i>) Effect of the poison on the blood corpuscles +of man. After Tyzzer.</span> +</div> + +<p>The poison hairs (<a href="#Fig_37a">fig. 37</a>), +are found on the subdorsal +and lateral tubercles (<a href="#Fig_38">fig. 38</a>), +in bunches of from three to +twelve on the minute papillæ +with which the tubercles are +thickly covered. The underlying +hypodermis is very +greatly thickened, the cells +being three or four times the length of the ordinary hypodermal +cells and being closely crowded together. Instead of a pore canal<span class="pagenum"><a name="Page_51" id="Page_51">[Pg 51]</a></span> +through the cuticula for each individual hair, there is a single pore +for each papillæ on a tubercle, all the hairs of the papilla being +connected with the +underlying cells +through the same +pore canal, (figs. +<a href="#Fig_39">39</a> and <a href="#Fig_40">40</a>).</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_38" id="Fig_38"></a> +<a href="images/f038-full.png"><img src="images/f038.png" width="500" height="352" alt="38. Cross section of the larva of the browntail moth showing the +tubercles bearing the poison hairs. Drawing by Miss +Kephart." title="38. Cross section of the larva of the browntail moth showing the +tubercles bearing the poison hairs. Drawing by Miss +Kephart." /></a> +<span class="caption">38. Cross section of the larva of the browntail moth showing the +tubercles bearing the poison hairs. Drawing by Miss +Kephart.</span> +</div> + +<p>The hypodermis +of this region is of +two distinct types +of cells. First, +there is a group of +slender fusiform +cells, one for each +poison hair on the +papilla, which are +the trichogen, or +hair-formative cells. They are crowded to one side and towards +the basement membrane by a series of much larger, and more prominent +cells (<a href="#Fig_40">fig. 40</a>), of which there is a single one for each papilla. +These larger cells have a granular protoplasm with large nuclei and +are obviously actively secreting. They are so characteristic in +appearance as to leave no question but that they are the true +poison glands.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_39" id="Fig_39"></a> +<a href="images/f039-full.png"><img src="images/f039.png" width="500" height="285" alt="39. Epithelium underlying poison hairs of the larva of the +browntail moth. Drawing by Miss Kephart." title="39. Epithelium underlying poison hairs of the larva of the +browntail moth. Drawing by Miss Kephart." /></a> +<span class="caption">39. Epithelium underlying poison hairs of the larva of the +browntail moth. Drawing by Miss Kephart.</span> +</div> + +<p>Poisonous larvæ of many other species have been reported from +Europe and especially from the tropics but the above-mentioned +species are the more important of those occurring in the United States +and will serve as types. It should be noted in this connection that<span class="pagenum"><a name="Page_52" id="Page_52">[Pg 52]</a></span> +through some curious misunderstanding +Gœldi (1913) +has featured the larva of +<i>Orgyia leucostigma</i>, the white-marked +tussock moth, as the +most important of the poisonous +caterpillars of this +country. Though there are +occasional reports of irritation +from its hairs such cases are +rare and there is no evidence +that there is any poison present. +Indeed, subcutaneous +implantation of the hairs +leads to no poisoning, but merely to temporary irritation.</p> + +<div class="figcenter" style="width: 450px;"><a name="Fig_40" id="Fig_40"></a> +<a href="images/f040-full.png"><img src="images/f040.png" width="450" height="410" alt="40. Same as figure 39, on larger scale." title="40. Same as figure 39, on larger scale." /></a> +<span class="caption">40. Same as <a href="#Fig_39">figure 39</a>, on larger scale.</span> +</div> + +<p>Occasionally, the hairs of certain species of caterpillars find lodgement +in the conjunctiva, cornea, or iris of the eye of man and give +rise to the condition known as <i>opthalmia nodosa</i>. The essential +feature of this trouble is a nodular conjunctivitis which simulates +tuberculosis of the conjunctiva and hence has been called <i>pseudo-tubercular</i>. +It may be distinguished microscopically by the presence +of the hairs.</p> + +<div class="figcenter" style="width: 375px;"><a name="Fig_41a" id="Fig_41a"></a> +<a href="images/f041a-full.png"><img src="images/f041a.png" width="375" height="317" alt="41. (a) Nodular conjunctivitis in the eye of a child. +De Schweinitz and Shumway." title="41. (a) Nodular conjunctivitis in the eye of a child. +De Schweinitz and Shumway." /></a> +<span class="caption">41. (<i>a</i>) Nodular conjunctivitis in the eye of a child. +De Schweinitz and Shumway.</span> +</div> + +<p>Numerous cases of opthalmia nodosa are on record. Of those +from this country, one of the most interesting is reported by de +Schweinitz and +Shumway (1904). It +is that of a child of +fifteen years whose +eye had become inflamed +owing to the +presence of some foreign +body. Downward +and inward on +the bulbar conjunctiva +were a number +of flattened, grayish-yellow +nodules, between +which was a +marked congestion of +the conjunctival and<span class="pagenum"><a name="Page_53" id="Page_53">[Pg 53]</a></span> +episcleral vessels (<a href="#Fig_41a">fig. 41<i>a</i></a>). Twenty-seven nodules could be differentiated, +those directly in the center of the collection being somewhat +confluent and +assuming a crescentic +and circular appearance. +The nodules +were excised +and, on sectioning, +were found to be +composed of a layer +of spindle cells and +round cells, outside +of which the tissue +was condensed into +a capsule. The +interior consisted of +epithelioid cells, between +which was a +considerable intercellular +substance. Directly in the center of a certain number of +nodules was found the section of a hair (<a href="#Fig_41b">fig. 41<i>b</i></a>). The evidence +indicated that the injury had resulted from playing with caterpillars +of one of the Arctiid moths, <i>Spilosoma virginica</i>. Other reported +cases have been caused by the hairs of larvæ of <i>Lasiocampa rubi</i>, +<i>L. pini</i>, <i>Porthetria dispar</i>, <i>Psilura monacha</i> and <i>Cnethocampa +processionea</i>.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_41b" id="Fig_41b"></a> +<a href="images/f041b-full.png"><img src="images/f041b.png" width="350" height="278" alt="41b. Section through one of the nodules showing the caterpillar +hair. De Schweinitz and Shumway." title="41b. Section through one of the nodules showing the caterpillar +hair. De Schweinitz and Shumway." /></a> +<span class="caption">41<i>b</i>. Section through one of the nodules showing the caterpillar +hair. De Schweinitz and Shumway.</span> +</div> + + +<p><a name="Relief_from_Poisoning_by_Nettling_Larvae" id="Relief_from_Poisoning_by_Nettling_Larvae"></a><b>Relief from Poisoning by Nettling Larvæ</b>—The irritation from +nettling larvæ is often severe and, especially in regions where the +brown-tail abounds, inquiries as to treatment arise. In general, it +may be said that cooling lotions afford relief, and that scratching, +with the possibilities of secondary infection, should be avoided, in +so far as possible.</p> + +<p>Among the remedies usually at hand, weak solutions of ammonia, +or a paste of ordinary baking soda are helpful. Castellani and Chalmers +recommend cleaning away the hairs by bathing the region with +an alkaline lotion, such as two per cent solution of bicarbonate of +soda, and then applying an ointment of ichthyol (10%).<span class="pagenum"><a name="Page_54" id="Page_54">[Pg 54]</a></span></p> + +<p>In the brown-tail district, there are many proprietary remedies of +which the best ones are essentially the following, as recommended +by Kirkland (1907):</p> + + +<div class="center"> +<table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td align="left">Carbolic acid</td><td align="left">½ drachm.</td></tr> +<tr><td align="left">Zinc oxide</td><td align="left">½ oz.</td></tr> +<tr><td align="left">Lime water</td><td align="left">8 oz.</td></tr> +</table></div> + +<p>Shake thoroughly and rub well into the affected parts.</p> + +<p>In some cases, and especially where there is danger of secondary +infection, the use of a weak solution of creoline (one teaspoonful to a +quart of water), is to be advised.</p> + + +<h3><a name="Vescicating_Insects_and_those_Possessing_Other_Poisons_in_their_Blood_Plasma" id="Vescicating_Insects_and_those_Possessing_Other_Poisons_in_their_Blood_Plasma"></a><span class="smcap">Vescicating Insects and those Possessing Other Poisons in +their Blood Plasma</span></h3> + +<div class="figleft" style="width: 150px;"><a name="Fig_42a" id="Fig_42a"></a> +<a href="images/f042a-full.png"><img src="images/f042a.png" width="150" height="185" alt="42a. Blister beetle." title="42a. Blister beetle." /></a> +<span class="caption">42<i>a</i>. Blister beetle.</span> +</div> + +<p>We have seen that certain forms, for example, the +poisonous spiders, not only secrete a toxine in their +poison glands, but that such a substance may be extracted +from other parts of their body, or even their +eggs. There are many insects which likewise possess a +poisonous blood plasma. Such forms have been well +designated by Taschenberg as <i>cryptotoxic</i> (κρυπτος = +hidden). We shall consider a few representative forms.</p> + +<div class="figright" style="width: 400px;"> +<a href="images/f042b-full.png"><img src="images/f042b.png" width="400" height="505" alt="42b. An American blister beetle. Meloe angusticollis. +Photograph by M. V. S." title="42b. An American blister beetle. Meloe angusticollis. +Photograph by M. V. S." /></a> +<span class="caption">42<i>b</i>. An American blister beetle. Meloe angusticollis. +Photograph by M. V. S.</span> +</div> + + +<p><a name="The_Blister_Beetles" id="The_Blister_Beetles"></a><b>The Blister Beetles</b>—Foremost +among the cryptotoxic +insects are the <i>Meloidæ</i> or +"blister beetles," to which the +well-known "Spanish fly" (<a href="#Fig_42a">fig. 42<i>a</i></a>), +formerly very generally +used in medical practice, belongs. +The vescicating property +is due to the presence in the +blood plasma of a peculiar, +volatile, crystalline substance +known as <i>cantharidin</i>, which is +especially abundant in the reproductive +organs of the beetle. According +to Kobert, the amount +of this varies in different species +from .4 or .5% to 2.57% of the +dry weight of the beetle.<span class="pagenum"><a name="Page_55" id="Page_55">[Pg 55]</a></span></p> + +<p>While blister beetles have been especially used for external application, +they are also at times used internally as a stimulant and a +diuretic. The powder or extract was formerly much in vogue as an +aphrodisiac, and formed the essential constituent of various philters, +or "love powders". It is now known that its effects on the reproductive +organs appear primarily after the kidneys have been affected to +such an extent as to endanger life, and that many cases of fatal poison +have been due to its ignorant use.</p> + +<p>There are many cases on record of poisoning and death due to +internal use, and in some instances from merely external application. +There are not rarely cases of poisoning of cattle from feeding on +herbage bearing a large number of the beetles and authentic cases are +known of human beings who have been poisoned by eating the flesh +of such cattle. Kobert states that the beetles are not poisonous to +birds but that the flesh of birds which have fed on them is poisonous +to man, and that if the flesh of chickens or frogs which have fed on +the cantharidin be fed to cats it causes in them the same symptoms +as does the cantharidin.</p> + +<p>Treatment of cases of cantharidin poison is a matter for a skilled +physician. Until he can be obtained, emetics should be administered +and these should be followed by white of egg in water. Oils should +be avoided, as they hasten the absorption of the poison.</p> + + +<p><a name="Other_Cryptotoxic_Insects" id="Other_Cryptotoxic_Insects"></a><b>Other Cryptotoxic Insects</b>—Though the blister beetles are the +best known of the insects with poisonous blood plasma, various +others have been reported and we shall refer to a few of the best +authenticated.</p> + +<p>One of the most famous is the Chrysomelid beetle, <i>Diamphidia +simplex</i>, the body fluids of whose larvæ are used by certain South +African bushmen as an arrow poison. Its action is due to the presence +of a toxalbumin which exerts a hæmolytic action on the blood, and +produces inflammation of the subcutaneous connective tissue and +mucous membranes. Death results from general paralysis. Krause +(1907) has surmised that the active principle may be a bacterial toxin +arising from decomposition of the tissues of the larva, but he presents +no support of this view and it is opposed by all the available evidence.</p> + +<p>In China, a bug, <i>Heuchis sanguinea</i>, belonging to the family +Cicadidæ, is used like the Meloidæ, to produce blistering, and often +causes poisoning. It has been assumed that its vescicating properties +are due to cantharidin, but the presence of this substance has not +been demonstrated.<span class="pagenum"><a name="Page_56" id="Page_56">[Pg 56]</a></span></p> + +<p>Certain Aphididæ contain a strongly irritating substance which +produces, not merely on mucous membranes but on outer skin, a +characteristic inflammation.</p> + +<p>It has been frequently reported that the larvæ of the European +cabbage butterfly, <i>Pieris brassicæ</i>, accidentally eaten by cows, horses, +ducks, and other domestic animals, cause severe colic, attempts to +vomit, paralysis of the hind legs, salivation, and stomatitis. On +<i>postmortem</i> there are to be found hæmorrhagic gastro-enteritis, +splenitis, and nephritis. Kobert has recently investigated the subject +and has found a poisonous substance in the blood of not only the +larvæ but also the pupæ.<span class="pagenum"><a name="Page_57" id="Page_57">[Pg 57]</a></span></p> + + +<div class="footnotes"><h3>FOOTNOTES:</h3> + +<div class="footnote"><p><a name="Footnote_A_1" id="Footnote_A_1"></a><a href="#FNanchor_A_1"><span class="label">[A]</span></a> This is diametrically opposed to the findings of Bordas (1905) in the case +of the European <i>Latrodectus 13-guttatus</i>, whose glands are "much larger than +those of other spiders." From a considerable comparative study, we should also +unhesitatingly make this statement regarding the glands of our American species, +<i>L. mactans</i>.</p></div> + +<div class="footnote"><p><a name="Footnote_B_2" id="Footnote_B_2"></a><a href="#FNanchor_B_2"><span class="label">[B]</span></a> Dr. E. H. Coleman (Kellogg, 1915) has demonstrated its virulence by a series +of experiments comparable with those of Kobert.</p></div> + +<div class="footnote"><p><a name="Footnote_C_3" id="Footnote_C_3"></a><a href="#FNanchor_C_3"><span class="label">[C]</span></a> According to Stiles, the species occurring in the Northwest which is commonly +identified as <i>D. venustus</i> should be called <i>D. andersoni</i> (see footnote, chapter 12).</p></div> + +<div class="footnote"><p><a name="Footnote_D_4" id="Footnote_D_4"></a><a href="#FNanchor_D_4"><span class="label">[D]</span></a> It should be remembered that in all the higher Hymenoptera the first abdominal +segment is fused with the thorax and that what is apparently the sixth +segment is, in reality, the seventh.</p></div> +</div> + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_III" id="CHAPTER_III"></a>CHAPTER III</h2> + +<h2>PARASITIC ARTHROPODA AFFECTING MAN</h2> + + +<p>The relation of insects to man as simple parasites has long been +studied, and until very recent years the bulk of the literature of medical +entomology referred to this phase of the subject. This is now +completely overshadowed by the fact that so many of these parasitic +forms are more than simple parasites, they are transmitters of other +microscopic parasites which are pathogenic to man. Yet the importance +of insects as parasites still remains and must be considered in a +discussion of the relation of insects to the health of man. In taking +up the subject we shall first consider some general features of the +phenomenon of animal parasitism.</p> + +<p>Parasitism is an adaptation which has originated very often among +living organisms and in widely separated groups. It would seem +simple to define what is meant by a "parasite" but, in reality, the +term is not easily limited. It is often stated that a parasite is "An +organism which lives at the expense of another," but this definition +is applicable to a predatory species or, in its broadest sense, to all +organisms. For our purpose we may say with Braun: "A parasite +is an organism which, for the purpose of obtaining food, takes up its +abode, temporarily or permanently, on or within another living +organism".</p> + +<p>Thus, parasitism is a phase of the broad biological phenomenon of +<i>symbiosis</i>, or living together of organisms. It is distinguished from +<i>mutualism</i>, or symbiosis in the narrow sense, by the fact that only one +party to the arrangement obtains any advantage, while the other is +to a greater or less extent injured.</p> + +<p>Of parasites we may distinguish on the basis of their location on or +in the host, <i>ecto-parasites</i>, which live outside of the body; and <i>endo-parasites</i>, +which live within the body. On account of their method +of breathing the parasitic arthropods belong almost exclusively to the +first of these groups.</p> + +<p>On the basis of relation to their host, we find <i>temporary parasites</i>, +those which seek the host only occasionally, to obtain food; and the +<i>stationary</i> or <i>permanent</i> parasites which, at least during certain stages, +do not leave their host.</p> + +<p><i>Facultative parasites</i> are forms which are not normally parasitic, +but which, when accidentally ingested, or otherwise brought into the<span class="pagenum"><a name="Page_58" id="Page_58">[Pg 58]</a></span> +body, are able to exist for a greater or less period of time in their +unusual environment. These are generally called in the medical +literature "pseudoparasites" but the term is an unfortunate one.</p> + +<p>We shall now take up the different groups of arthropods, discussing +the more important of the parasitic forms attacking man. The +systematic relationship of these forms, and key for determining +important species will be found in Chapter XII.</p> + + +<h3><a name="Acarina_or_Mites" id="Acarina_or_Mites"></a><span class="smcap">Acarina or Mites</span></h3> + +<p>The <b>Acarina</b>, or <i>mites</i>, form a fairly natural group of arachnids, +characterized, in general, by a sac-like, unsegmented body which is +generally fused with the cephalothorax. The mouth-parts have been +united to form a beak or rostrum.</p> + +<p>The representatives of this group undergo a marked metamorphosis. +Commonly, the larvæ on hatching from the egg, possess but +three pairs of legs, and hence are called <i>hexapod larvæ</i>. After a molt, +they transform into nymphs which, like the adult, have four pairs of +legs and are called <i>octopod nymphs</i>. These after a period of growth, +molt one or more times and, acquiring external sexual organs, become +adult.</p> + +<p>Most of the mites are free-living, but there are many parasitic +species and as these have originated in widely separated families, the +Acarina form an especially favorable group for study of the origin of +parasitism. Such a study has been made by Ewing (1911), who has +reached the following conclusions:</p> + +<p>"We have strong evidence indicating that the parasitic habit has +originated independently at least eleven times in the phylogeny of the +Ararina. Among the zoophagous parasites, the parasitic habit has +been developed from three different types of free-living Acarina: +(a) predaceous forms, (b) scavengers, (c) forms living upon the juices +of plants."</p> + +<p>Ewing also showed that among the living forms of Acarina we can +trace out all the stages of advancing parasitism, semiparasitism, +facultative parasitism, even to the fixed and permanent type, and +finally to endoparasitism.</p> + +<p>Of the many parasitic forms, there are several species which are +serious parasites of man and we shall consider the more important of +these. Infestation by mites is technically known as <i>acariasis</i>.<span class="pagenum"><a name="Page_59" id="Page_59">[Pg 59]</a></span></p> + +<div class="figcenter" style="width: 225px;"><a name="Fig_43" id="Fig_43"></a> +<a href="images/f043-full.png"><img src="images/f043.png" width="225" height="373" alt="43. Effect of the harvest mites on the skin of man. Photograph by +J. C. Bradley." title="43. Effect of the harvest mites on the skin of man. Photograph by +J. C. Bradley." /></a> +<span class="caption">43. Effect of the harvest mites on the skin of man. Photograph by +J. C. Bradley.</span> +</div><p><span class="pagenum"><a name="Page_60" id="Page_60">[Pg 60]</a></span></p> + + +<h3><a name="The_Trombidiidae_or_Harvest_Mites" id="The_Trombidiidae_or_Harvest_Mites"></a>The Trombidiidæ, or Harvest Mites</h3> + +<p>In many parts of this country it is impossible for a visitor to go +into the fields and, particularly, into berry patches and among tall +weeds and grass in the summer or early fall without being affected by +an intolerable itching, which is followed, later, by a breaking out of +wheals, or papules, surrounded by a bright red or violaceous aureola, +(<a href="#Fig_43">fig. 43</a>). It is often regarded as a urticaria or eczema, produced by +change of climate, an error in diet, or some condition of general health.</p> + +<p>Sooner or later, the victim finds that it is due to none of these, but +to the attacks of an almost microscopic red mite, usually called +"jigger" or "chigger" in this country. As the term "chigger" is +applied to one of the true fleas, <i>Dermatophilus penetrans</i>, of the tropics, +these forms are more +correctly known as +"harvest mites." +Natives of an infested +region may be so +immune or accustomed +to its attacks +as to be unaware of +its presence, though +such immunity is by +no means possessed +by all who have been +long exposed to the +annoyance.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_44" id="Fig_44"></a> +<a href="images/f044-full.png"><img src="images/f044.png" width="500" height="305" alt="44. Harvest mites. (Larvæ of Trombidium). After C. V. +Riley." title="44. Harvest mites. (Larvæ of Trombidium). After C. V. +Riley." /></a> +<span class="caption">44. Harvest mites. (Larvæ of Trombidium). After C. V. +Riley.</span> +</div> + +<p>The harvest mites, or chiggers, attacking man are larval forms, +possessing three pairs of legs (<a href="#Fig_44">fig. 44</a>). Their systematic position was +at first unknown and they were classed under a special genus <i>Leptus</i>, +a name which is very commonly still retained in the medical literature. +It is now known that they are the larval forms of various species of +the genus <i>Trombidium</i>, a group of predaceous forms, the adults of +which feed primarily on insects and their eggs. In this country the +species best known are those to be found late in summer, as larvæ +at the base of the wings of houseflies or grasshoppers.</p> + +<p>There is much uncertainty as to the species of the larvæ attacking +man but it is clear that several are implicated. Bruyant has shown +that in France the larvæ <i>Trombidium inapinatum</i> and <i>Trombidium +holosericeum</i> are those most frequently found. The habit of attacking +man is abnormal and the larvæ die after entering the skin. Normally +they are parasitic on various insects.<span class="pagenum"><a name="Page_61" id="Page_61">[Pg 61]</a></span></p> + +<p>Most recent writers agree that, on man, they do not bore into the +skin, as is generally supposed, but enter a hair follicle or sebaceous +gland and from the bottom of this, pierce the cutis with their elongate +hypopharynx. According to Braun, there arises about the inserted +hypopharynx a fibrous secretion—the so-called "beak" which is, in +reality, a product of the host. Dr. J. C. Bradley, however, has made +careful observations on their method of attack, and he assures us that +the mite ordinarily remains for a long time feeding on the surface of +the skin, where it produces the erythema above described. During +this time it is not buried in the skin but is able to retreat rapidly into +it through a hair follicle or sweat gland. The irritation from the +mites ceases after a few days, but not infrequently the intolerable +itching leads to so much scratching that secondary infection follows.</p> + +<p>Relief from the irritation may be afforded by taking a warm salt +bath as soon as possible after exposure or by killing the mites by +application of benzine, sulphur ointment or carbolized vaseline. +When they are few in number, they can be picked out with a sterile +needle.</p> + +<p>Much may be done in the way of warding off their attacks by +wearing gaiters or close-woven stockings extending from ankle to the +knee. Still more efficacious is the sprinkling of flowers of sulphur in +the stockings and the underclothes from a little above the knee, down. +The writers have known this to make it possible for persons who were +especially susceptible to work with perfect comfort in badly infested +regions. Powdered naphthalene is successfully used in the same way +and as Chittenden (1906) points out, is a safeguard against various +forms of man-infesting tropical insect pests.</p> + +<p>The question of the destruction of the mites in the field is sometimes +an important one, and under some conditions, is feasible. +Chittenden states that much can be accomplished by keeping the +grass, weeds, and useless herbage mowed closely, so as to expose the +mites to the sun. He believes that in some cases good may be done +by dusting the grass and other plants, after cutting, with flowers of +sulphur or by spraying with dilute kerosene emulsion in which +sulphur has been mixed. More recently (1914) he calls attention to +the value of cattle, and more especially sheep, in destroying the pests +by tramping on them and by keeping the grass and herbage closely +cropped.<span class="pagenum"><a name="Page_62" id="Page_62">[Pg 62]</a></span></p> + + +<h3><a name="Ixodoidea_or_Ticks" id="Ixodoidea_or_Ticks"></a><span class="smcap">Ixodoidea or Ticks</span></h3> + +<p>Until recently, the ticks attracted comparatively little attention +from entomologists. Since their importance as carriers of disease +has been established, interest in +the group has been enormously +stimulated and now they rank +second only to the mosquitoes +in the amount of detailed study +that has been devoted to them.</p> + +<div class="figcenter" style="width: 450px;"><a name="Fig_45a" id="Fig_45a"></a> +<a href="images/f045a-full.png"><img src="images/f045a.png" width="450" height="481" alt="45a. Argus persicus. Capitulum of male. +After Nuttall and Warburton." title="45a. Argus persicus. Capitulum of male. +After Nuttall and Warburton." /></a> +<span class="caption">45<i>a</i>. Argus persicus. Capitulum of male. +After Nuttall and Warburton.</span> +</div> + +<p>The ticks are the largest of +the Acarina. They are characterized +by the fact that the +hypostome, or "tongue" (<a href="#Fig_45a">fig. 45</a>) +is large and file-like, roughened +by sharp teeth. They possess +a breathing pore on each side +of the body, above the third +or fourth coxæ (<a href="#Fig_45b">fig. 45<i>b</i></a>).</p> + +<div class="figcenter" style="width: 450px;"><a name="Fig_45b" id="Fig_45b"></a> +<a href="images/f045b-full.png"><img src="images/f045b.png" width="450" height="398" alt="45b. Left spiracle of nymph of Argas persicus. +After Nuttall and Warburton." title="45b. Left spiracle of nymph of Argas persicus. +After Nuttall and Warburton." /></a> +<span class="caption">45<i>b</i>. Left spiracle of nymph of <i>Argas persicus</i>. +After Nuttall and Warburton.</span> +</div> + +<p>There are two distinct families +of the <b>Ixodoidea</b>, differing +greatly in structure, life-history and habits. These are the <b>Argasidæ</b> +and the <b>Ixodidæ</b>. We shall follow Nuttall (1908) in characterizing +these two families and in pointing out their biological differences, and +shall discuss briefly the more important species which attack man. +The consideration of the +ticks as carriers of disease +will be reserved for a later +chapter.</p> + + +<h3><a name="Argasidae" id="Argasidae"></a>Argasidæ</h3> + +<p>In the ticks belonging to +the family <b>Argasidæ</b>, there +is comparatively little sexual +dimorphism, while this is +very marked in the Ixodidæ. +The capitulum, or so-called +"head" is ventral, instead of +terminal; the palpi are leg-like, +with the segments subequal; the scutum, or dorsal shield, is +absent; eyes, when present, are lateral, on supracoxal folds. The<span class="pagenum"><a name="Page_63" id="Page_63">[Pg 63]</a></span> +spiracles are very small; coxæ unarmed; tarsi without ventral spurs, +and the pulvilli are absent or rudimentary.</p> + +<p>In habits and life history the Argasidæ present striking characteristics. +In the first place, they are long-lived, a factor which counts for +much in the maintenance of the species. They are intermittent +feeders, being comparable with the bed-bug in this respect. There are +two or more nymphal stages, and they may molt after attaining maturity. +The female lays comparatively few eggs in several small batches.</p> + +<p>Nuttall (1911) concludes that "The Argasidæ represent the relatively +primitive type of ticks because they are less constantly parasitic +than are the Ixodidæ. Their nymphs and adults are rapid +feeders and chiefly infest the habitat of their hosts. * * * Owing +to the Argasidæ infesting the habitats of their hosts, their resistance +to prolonged starvation and their rapid feeding habits, they do not +need to bring forth a large progeny, because there is less loss of life +in the various stages, as compared with the Ixodidæ, prior to their +attaining maturity."</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_46" id="Fig_46"></a> +<a href="images/f046-full.png"><img src="images/f046.png" width="350" height="264" alt="46. Argus persicus. Dorsal and ventral aspects. (×4). After Hassell." title="46. Argus persicus. Dorsal and ventral aspects. (×4). After Hassell." /></a> +<span class="caption">46. Argus persicus. Dorsal and ventral aspects. (×4). After Hassell.</span> +</div> + +<p>Of the Argasidæ, we have in the United States, several species +which have been reported as attacking man.</p> + +<div class="figleft" style="width: 300px;"><a name="Fig_47" id="Fig_47"></a> +<a href="images/f047-full.png"><img src="images/f047.png" width="300" height="354" alt="47. Otiobius (Ornithodoros) megnini, head +of nymph. After Stiles." title="47. Otiobius (Ornithodoros) megnini, head +of nymph. After Stiles." /></a> +<span class="caption">47. Otiobius (Ornithodoros) megnini, head +of nymph. After Stiles.</span> +</div> + +<p><i>Argas persicus</i>, the famous "Miana bug" (<a href="#Fig_46">fig. 46</a>), is a very widely +distributed species, being reported from Europe, Asia, Africa, and +Australia. It is everywhere preeminently a parasite of fowls.<span class="pagenum"><a name="Page_64" id="Page_64">[Pg 64]</a></span> +According to Nuttall it is specifically identical with <i>Argas americanus</i> +Packard or <i>Argas miniatus</i> Koch, which is commonly found on fowls +in the United States, in the South +and Southwest. Its habits are comparable +to those of the bed-bug. It +feeds intermittently, primarily at +night, and instead of remaining on its +host, it then retreats to cracks and +crevices. Hunter and Hooker (1908) +record that they have found the larva +to remain attached for five or eight +days before dropping. Unlike the +Ixodidæ, the adults oviposit frequently.</p> + +<p>The most remarkable feature of +the biology of this species is the great +longevity, especially of the adult. +Hunter and Hooker report keeping +larvæ confined in summer in pill boxes immediately after hatching +for about two months while under similar conditions those +of the Ixodid, <i>Boophilus annulatus</i> lived for but two or three days. +Many writers have recorded keeping adults for long periods without +food. We have kept specimens in a tin box for over a year and a half +and at the end of that time a number were still alive. Laboulliene +kept unfed adults for over three years. In view of the effectiveness of<span class="pagenum"><a name="Page_65" id="Page_65">[Pg 65]</a></span> +sulphur in warding off the attacks of Trombidiidæ, it is astonishing +to find that Lounsbury has kept adults of <i>Argas persicus</i> for three +months in a box nearly filled with flowers of sulphur, with no apparent +effect on them.</p> + + +<div class="figcenter" style="width: 500px;"><a name="Fig_48" id="Fig_48"></a> +<a href="images/f048-full.png"><img src="images/f048.png" width="500" height="335" alt="48. Otiobius (Ornithodoros) megnini, male. (a) dorsal, (b) ventral +aspect. After Nuttall and Warburton." title="48. Otiobius (Ornithodoros) megnini, male. (a) dorsal, (b) ventral +aspect. After Nuttall and Warburton." /></a> +<span class="caption">48. Otiobius (Ornithodoros) megnini, male. (<i>a</i>) dorsal, (<i>b</i>) ventral +aspect. After Nuttall and Warburton.</span> +</div> + +<p>We have already called attention to the occasional serious effects +of the bites of this species. While such reports have been frequently +discredited there can be no doubt that they have foundation in fact. +The readiness with which this tick attacks man, and the extent to +which old huts may be infested makes it especially troublesome.</p> + +<p><i>Otiobius (Ornithodoros) megnini</i>, the "spinose ear-tick" (figs. <a href="#Fig_47">47</a>, <a href="#Fig_48">48</a>), +first described from Mexico, as occurring in the ears of horses, is a +common species in our Southwestern States and is recorded by Banks +as occurring as far north as Iowa.</p> + +<p>The species is remarkable for the great difference between the +spiny nymph stage and the adult. The life history has been worked +out by Hooker (1908). Seed ticks, having gained entrance to the +ear, attach deeply down in the folds, engorge, and in about five days, +molt; as nymphs with their spinose body they appear entirely unlike +the larvæ. As nymphs they continue feeding sometimes for months. +Finally the nymph leaves the host, molts to form the unspined adult, +and without further feeding is fertilized and commences oviposition.</p> + +<p>The common name is due to the fact that in the young stage the +ticks occur in the ear of their hosts, usually horses or cattle. Not +uncommonly it has been reported as occurring in the ear of man and +causing very severe pain. Stiles recommends that it be removed by +pouring some bland oil into the ear.</p> + +<p>Banks (1908) reports three species of <i>Ornithodoros</i>—<i>O. turicata</i>, +<i>coriaceus</i> and <i>talaje</i>—as occurring in the United States. All of these +attack man and are capable of inflicting very painful bites.</p> + + +<h3><a name="Ixodidae" id="Ixodidae"></a>Ixodidæ</h3> + +<p>The ticks belonging to the family <b>Ixodidæ</b> (figs. <a href="#Fig_49">49</a> and <a href="#Fig_50">50</a>) exhibit +a marked sexual dimorphism. The capitulum is anterior, terminal, +instead of ventral as in the Argasidæ; the palpi are relatively rigid +(except in the subfamily Ixodinæ), with rudimentary fourth segment; +scutum present; eyes, when present, dorsal, on side of scutum. The +spiracles are generally large, situated well behind the fourth coxæ; +coxæ generally with spurs; pulvilli always present.</p> + +<p>In habits and life history the typical Ixodidæ differ greatly from +the Argasidæ. They are relatively short-lived, though some recent<span class="pagenum"><a name="Page_66" id="Page_66">[Pg 66]</a></span> +work indicates that their longevity +has been considerably +under-estimated. Typically, +they are permanent feeders, +remaining on the host, or hosts, +during the greater part of their +life. They molt twice only, +on leaving the larval and the +nymphal stages. The adult +female deposits a single, large +batch of eggs. Contrasting +the habits of the Ixodidæ to +those of the Argasidæ, Nuttall +(1911) emphasizes that the +Ixodidæ are more highly +specialized parasites. "The +majority are parasitic on hosts +having no fixed habitat and +consequently all stages, as a +rule, occur upon the host."</p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_49" id="Fig_49"></a> +<a href="images/f049-full.png"><img src="images/f049.png" width="400" height="524" alt="49. Ixodes ricinus; male, ventral aspect. After +Braun and Luehe." title="49. Ixodes ricinus; male, ventral aspect. After +Braun and Luehe." /></a> +<span class="caption">49. Ixodes ricinus; male, ventral aspect. After +Braun and Luehe.</span> +</div> + +<p>As mere parasites of man, apart from their power to transmit +disease, the Ixodidæ are much less important than the Argasidæ. +Many are reported as occasionally attacking man and of these the +following native species +may be mentioned.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_50" id="Fig_50"></a> +<a href="images/f050-full.png"><img src="images/f050.png" width="500" height="427" alt="50. Ixodes ricinus, var. scapularis, female. Capitulum and +scutum; ventral aspect of capitulum; coxæ; tarsus 4; +spiracle; genital and anal grooves. After Nuttall and +Warburton." title="50. Ixodes ricinus, var. scapularis, female. Capitulum and +scutum; ventral aspect of capitulum; coxæ; tarsus 4; +spiracle; genital and anal grooves. After Nuttall and +Warburton." /></a> +<span class="caption">50. Ixodes ricinus, var. scapularis, female. Capitulum and +scutum; ventral aspect of capitulum; coxæ; tarsus 4; +spiracle; genital and anal grooves. After Nuttall and +Warburton.</span> +</div> + +<p><i>Ixodes ricinus</i>, the +European castor bean +tick (figs. <a href="#Fig_49">49</a>, <a href="#Fig_50">50</a>), is a +species which has been +often reported from +this country but Banks +(1908) has shown that, +though it does occur, +practically all of the +records apply to <i>Ixodes +scapularis</i> or <i>Ixodes +cookei</i>. In Europe, +<i>Ixodes ricinus</i> is very +abundant and very +commonly attacks<span class="pagenum"><a name="Page_67" id="Page_67">[Pg 67]</a></span> +man. At the point of penetration of the hypostome there is more or +less inflammation but serious injury does not occur unless there have +been introduced pathogenic bacteria or, unless the tick has been +abruptly removed, leaving the capitulum in the wound. Under the +latter circumstances, there may be an abscess formed about the foreign +body and occasionally, serious results have followed. Under certain +conditions the tick, in various stages, may penetrate under the skin +and produce a tumor, within which it may survive for a considerable +period of time.</p> + +<p><i>Ixodes cookei</i> is given by Banks as "common on mammals in the +Eastern States as far west as the Rockies." It is said to affect man +severely.</p> + +<p><i>Amblyomma americanum</i>, (<a href="#Fig_158">fig. 158c</a>), the "lone star tick," is +widely distributed in the United States. Its common name is derived +from the single silvery spot on the scutum of the female. Hunter +and Hooker regard this species as, next to <i>Boophilus annulatus</i>, the +most important tick in the United States. Though more common on +cattle, it appears to attack mammals generally, and "in portions of +Louisiana and Texas it becomes a pest of considerable importance to +moss gatherers and other persons who spend much time in the forests."</p> + +<p><i>Amblyomma cajennense</i>, noted as a pest of man in central and +tropical America, is reported from various places in the south and +southwestern United States.</p> + +<p><i>Dermacentor variabilis</i> is a common dog tick of the eastern United +States. It frequently attacks man, but the direct effects of its bite +are negligible.</p> + +<p>The "Rocky Mountain spotted fever tick" (<i>Dermacentor andersoni</i> +according to Stiles, <i>D. venustus</i> according to Banks) is, from the viewpoint +of its effects on man, the most important of the ticks of the +United States. This is because, as has been clearly established, it +transmits the so-called "spotted fever" of man in our northwestern +states. This phase of the subject will be discussed later and it need +merely be mentioned here, that this species has been reported as +causing painful injuries by its bites. Dr. Stiles states that he has +seen cases of rather severe lymphangitis and various sores and swellings +developing from this cause. In one case, of an individual bitten +near the elbow, the arm became very much swollen and the patient +was confined in bed for several days. The so-called tick paralysis +produced by this species is discussed in a preceding chapter.<span class="pagenum"><a name="Page_68" id="Page_68">[Pg 68]</a></span></p> + +<p>There are many other records of various species of ticks attacking +man, but the above-mentioned will serve as typical and it is not necessary +to enter into greater detail.</p> + + +<p><a name="Treatment_of_Tick_Bites" id="Treatment_of_Tick_Bites"></a><b>Treatment of Tick Bites</b>—When a tick attaches to man the first +thing to be done is to remove it without leaving the hypostome in the +wound to fester and bring about secondary effects. This is best +accomplished by applying to the tick's body some substance which +will cause it to more readily loosen its hold. Gasoline or petroleum, +oil or vaseline will serve. +For removing the +spinose ear-tick, Stiles +recommends pouring +some bland oil into the +ear. Others have used +effectively a pledget of +cotton soaked in chloroform.</p> + +<p>In general, the treatment +recommended by +Wellman for the bites +of <i>Ornithodoros moubata</i> +will prove helpful. It +consists of prolonged +bathing in very hot +water, followed by the +application of a strong +solution of bicarbonate +of soda, which is allowed to dry upon the skin. He states that this +treatment is comforting. For severe itching he advises smearing +the bites with vaseline, which is slightly impregnated with camphor +or menthol. Medical aid should be sought when complications arise.</p> + +<p>The <a name="Dermanyssidae" id="Dermanyssidae"></a><b>Dermanyssidæ</b> are Gamasid mites which differ from others of +the group in that they are parasitic on vertebrates. None of the +species normally attack man, but certain of them, especially the +poultry mite, may be accidental annoyances.</p> + +<div class="figcenter" style="width: 475px;"><a name="Fig_51" id="Fig_51"></a> +<a href="images/f051-full.png"><img src="images/f051.png" width="475" height="473" alt="51. Dermanyssus gallinæ, female. After Delafond." title="51. Dermanyssus gallinæ, female. After Delafond." /></a> +<span class="caption">51. Dermanyssus gallinæ, female. After Delafond.</span> +</div> + +<p><i>Dermanyssus gallinæ</i> (<a href="#Fig_51">fig. 51</a>), the red mite of poultry, is an exceedingly +common and widespread parasite of fowls. During the day +it lives in cracks and crevices of poultry houses, under supports of +roosts, and in litter of the food and nests, coming out at night to feed.<span class="pagenum"><a name="Page_69" id="Page_69">[Pg 69]</a></span> +They often attack people working in poultry houses or handling and +plucking infested fowls. They may cause an intense pruritis, but they +do not produce a true dermatosis, for +they do not find conditions favorable for +multiplication on the skin of man.</p> + + +<h3><a name="Tarsonemidae" id="Tarsonemidae"></a>Tarsonemidæ</h3> + +<p>The representatives of the family <b>Tarsonemidæ</b> +are minute mites, with the body +divided into cephalothorax and abdomen. +There is marked sexual dimorphism. +The females possess stigmata at the +anterior part of the body, at the base of +the rostrum, and differ from all other mites +in having on each side, a prominent clavate +organ between the first and second legs. +The larva, when it exists, is hexapodous +and resembles the adult. A number of the +species are true parasites on insects, while others attack plants. +Several of them may be accidental parasites of man.</p> + +<div class="figcenter" style="width: 275px;"><a name="Fig_52" id="Fig_52"></a> +<a href="images/f052-full.png"><img src="images/f052.png" width="275" height="421" alt="52. Pediculoides ventricosus, female. +After Webster." title="52. Pediculoides ventricosus, female. +After Webster." /></a> +<span class="caption">52. Pediculoides ventricosus, female. +After Webster.</span> +</div> + +<div class="figcenter" style="width: 400px;"><a name="Fig_53" id="Fig_53"></a> +<a href="images/f053-full.png"><img src="images/f053.png" width="400" height="495" alt="53. Pediculoides ventricosus, gravid female. (×80). +After Webster." title="53. Pediculoides ventricosus, gravid female. (×80). +After Webster." /></a> +<span class="caption">53. Pediculoides ventricosus, gravid female. (×80). +After Webster.</span> +</div> + +<p><i>Pediculoides ventricosus</i> +(fig. <a href="#Fig_52">52</a> and <a href="#Fig_53">53</a>) is, of all the +Tarsonemidæ reported, the +one which has proved most +troublesome to man. It is a +predaceous species which +attacks a large number of +insects but which has most +commonly been met with by +man through its fondness for +certain grain-infesting insects, +notably the Angoumois grain +moth, <i>Sitotroga cerealella</i>, and +the wheat straw-worm, <i>Isosoma +grande</i>. In recent years +it has attracted much attention +in the United States and +its distribution and habits +have been the object of detailed +study by Webster (1901).<span class="pagenum"><a name="Page_70" id="Page_70">[Pg 70]</a></span></p> + +<div class="figcenter" style="width: 250px;"><a name="Fig_54" id="Fig_54"></a> +<a href="images/f054-full.png"><img src="images/f054.png" width="250" height="264" alt="54. Pediculoides ventricosus, male. After Braun." title="54. Pediculoides ventricosus, male. After Braun." /></a> +<span class="caption">54. Pediculoides ventricosus, male. After Braun.</span> +</div> + +<p>There is a very striking sexual dimorphism in this species. The +non-gravid female is elongate, about 200µ by 70µ (<a href="#Fig_52">fig. 52</a>), with the +abdomen slightly striated longitudinally. The gravid female (<a href="#Fig_53">fig. 53</a>) +has the abdomen enormously swollen, so that it is from twenty to a +hundred times greater than the rest of the body. The species is +viviparous and the larvæ undergo their entire growth in the body of +the mother. They emerge as sexually mature males and females +which soon pair. The male (<a href="#Fig_54">fig. 54</a>) is much smaller, reaching a +length of only 320µ but +is relatively broad, +80µ, and angular. Its +abdomen is very greatly +reduced.</p> + +<p>As far back as 1850 +it was noted as causing +serious outbreaks +of peculiar dermatitis +among men handling +infested grain. For +some time the true +source of the difficulty +was unknown and it +was even believed that +the grain had been +poisoned. Webster +has shown that in this +country (and probably +in Europe as well) its +attacks have been mistaken for those of the red bugs or "chiggers" +(larval Trombiidæ). More recently a number of outbreaks of a +mysterious "skin disease" were traced to the use of straw mattresses, +which were found to be swarming with these almost microscopic +forms which had turned their attentions to the occupants of the beds. +Other cases cited were those of farmers running wheat through a +fanning mill, and of thrashers engaged in feeding unthrashed grain +into the cylinder of the machine.</p> + +<div class="figcenter" style="width: 250px;"><a name="Fig_55" id="Fig_55"></a> +<a href="images/f055-full.png"><img src="images/f055.png" width="250" height="338" alt="55. Lesions produced by the attacks of Pediculoides ventricosus. +After Webster." title="55. Lesions produced by the attacks of Pediculoides ventricosus. +After Webster." /></a> +<span class="caption">55. Lesions produced by the attacks of Pediculoides ventricosus. +After Webster.</span> +</div> + +<p>The medical aspects of the question have been studied especially +by Schamberg and Goldberger and from the latter's summary (1910) +we derive the following data. Within twelve to sixteen hours after +exposure, itching appears and in severe cases, especially where exposure<span class="pagenum"><a name="Page_71" id="Page_71">[Pg 71]</a></span> +is continued night after night by sleeping on an infested bed, the +itching may become almost intolerable. Simultaneously, there +appears an eruption which characteristically consists of wheals +surrounded by a vesicle (<a href="#Fig_55">fig. 55</a>). The vesicle as a rule does not exceed +a pin head in size but may become as large as a pea. Its contents +rapidly become turbid and in a few hours it is converted into a pustule. +The eruption is most abundant on the trunk, slight on the face and +extremities and almost absent on the feet and hands. In severe cases +there may be constitutional disturbances marked, at the outset, by +chilliness, nausea, and vomiting, followed for a few days by a slight +elevation of temperature, with the appearance of albumin in the +urine. In some cases the eruption may simulate that of chicken-pox +or small-pox.<span class="pagenum"><a name="Page_72" id="Page_72">[Pg 72]</a></span></p> + +<p>Treatment for the purpose of killing the mites is hardly necessary +as they attach feebly to the surface and are readily brushed off by +friction of the clothes. "Antipruritic treatment is always called for; +warm, mildly alkaline baths or some soothing ointment, such as zinc +oxide will be found to fulfil this indication." Of course, reinfestation +must be guarded against, by discarding, or thoroughly fumigating +infested mattresses, or by avoiding other sources. Goldberger suggests +that farm laborers who must work with infested wheat or straw +might protect themselves by anointing the body freely with some +bland oil or grease, followed by a change of clothes and bath as soon +as their work is done. We are not aware of any experiments to +determine the effect of flowers of sulphur, but their efficiency in the +case of "red bugs" suggests that they are worth a trial against +<i>Pediculoides</i>.</p> + +<p>Various species of <b>Tyroglyphidæ</b> (<a href="#Fig_150">fig. 150<i>f</i></a>) may abound on dried +fruits and other products and attacking persons handling them, may +cause a severe dermatitis, comparable to that described above for +<i>Pediculoides ventricosus</i>. Many instances of their occurrence as such +temporary ectoparasites are on record. Thus, workers who handle +vanilla pods are subject to a severe dermatitis, known as vanillism, +which is due to the attacks of <i>Tyroglyphus siro</i>, or a closely related +species. The so-called "grocer's itch" is similarly caused by mites +infesting various products. Castellani has shown that in Ceylon, +workers employed in the copra mills, where dried cocoanut is ground +up for export, are much annoyed by mites, which produce the so-called +"copra itch." The skin of the hands, arms and legs, and sometimes +of the whole body, except the face, is covered by fairly numerous, very +pruriginous papules, often covered by small, bloody crusts due to +scratching. The condition is readily mistaken for scabies. It is +due to the attacks of <i>Tyroglyphus longior castellanii</i> which occur in +enormous numbers in some samples of the copra.</p> + + +<h3><a name="Sarcoptidae" id="Sarcoptidae"></a>Sarcoptidæ</h3> + +<p>The <b>Sarcoptidæ</b> are minute whitish mites, semi-globular in shape, +with a delicate transversely striated cuticula. They lack eyes and +tracheæ. The mouth-parts are fused at the base to form a cone +which is usually designated as the head. The legs are short and +stout, and composed of five segments. The tarsi may or may not +possess a claw and may terminate in a pedunculated sucker, or simple +long bristle, or both. The presence or absence of these structures<span class="pagenum"><a name="Page_73" id="Page_73">[Pg 73]</a></span> +and their distribution are much used in classification. The mites +live on or under the skin of mammals and birds, where they produce +the disease known as scabies, mange, or +itch. Several species of the Sarcoptidæ +attack man but the most important of +these, and the one pre-eminent as the +"itch mite" is <i>Sarcoptes scabiei</i>.</p> + +<p>The female of <i>Sarcoptes scabiei</i>, of man, +is oval and yellowish white; the male +more rounded and of a somewhat reddish +tinge, and much smaller. The body is +marked by transverse striæ which are +partly interrupted on the back. There +are transverse rows of scales, or pointed +spines, and scattered bristles on the +dorsum.</p> + +<div class="figleft" style="width: 300px;"><a name="Fig_56a" id="Fig_56a"></a> +<a href="images/f056a-full.png"><img src="images/f056a.png" width="300" height="427" alt="56a. Sarcoptes scabiei, male. +(×100). After Fürstenberg." title="56a. Sarcoptes scabiei, male. +(×100). After Fürstenberg." /></a> +<span class="caption">56<i>a</i>. Sarcoptes scabiei, male. +(×100). After Fürstenberg.</span> +</div> + +<p>The male (<a href="#Fig_56a">fig. 56</a>) which is from 200-240µ +in length, and 150-200µ in breadth, +possesses pedunculated suckers on each +pair of legs except the third, which bears, instead, a long bristle. +The female (<a href="#Fig_56b">fig. 56</a>) 300-450µ in length and 250-350µ in breadth, has +the pedunculated suckers on the first and second pairs of legs, only, +the third and fourth terminating in bristles.</p> + +<div class="figright" style="width: 300px;"><a name="Fig_56b" id="Fig_56b"></a> +<a href="images/f056b-full.png"><img src="images/f056b.png" width="300" height="416" alt="56b. Sarcoptes scabiei, female. +(×100.) After Fürstenberg." title="56b. Sarcoptes scabiei, female. +(×100.) After Fürstenberg." /></a> +<span class="caption">56<i>b</i>. Sarcoptes scabiei, female. +(×100.) After Fürstenberg.</span> +</div> + +<p>The mite lives in irregular galleries from +a few millimeters to several centimeters in +length, which it excavates in the epidermis +(<a href="#Fig_57">fig. 57</a>). It works especially where the +skin is thin, such as between the fingers, +in the bend of the elbows and knees, and +in the groin, but it is by no means restricted +to these localities. The female, alone, +tunnels into the skin; the males remain +under the superficial epidermal scales, and +seldom are found, as they die soon after +mating.</p> + +<p>As she burrows into the skin the female +deposits her eggs, which measure about +150 × 100µ. Fürstenberg says that each +deposits an average of twenty-two to twenty-four eggs, though +Gudden reports a single burrow as containing fifty-one. From these<span class="pagenum"><a name="Page_74" id="Page_74">[Pg 74]</a></span> +there develop after about seven days, the hexapod larvæ. These +molt on the sixteenth day to form an octopod nymph, which molts +again the twenty-first day. At the end of the fourth week the +nymphs molt to form the sexually mature males and the so-called +pubescent females. These pair, the males die, and the females again +cast their skin, and become the oviparous females. Thus the life +cycle is completed in about twenty-eight days.</p> + +<p>The external temperature exercises a great influence on the development +of the mites and thus, during the winter, the areas of infestation +not only do not spread, but they become restricted. As soon as +the temperature rises, the mites increase and the infestation becomes +much more extensive.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_57" id="Fig_57"></a> +<a href="images/f057-full.png"><img src="images/f057.png" width="500" height="259" alt="57. Sarcoptes scabiei. Diagrammatic representation of the course in +the skin of man." title="57. Sarcoptes scabiei. Diagrammatic representation of the course in +the skin of man." /></a> +<span class="caption">57. Sarcoptes scabiei. Diagrammatic representation of the course in +the skin of man.</span> +</div> + +<p>In considering the possible sources of infestation, and the chances +of reinfestation after treatment, the question of the ability of the mite +to live apart from its host is a very important one. Unfortunately +there are few reliable data on this subject. Gerlach found that, +exposed in the dry, warm air of a room they became very inactive +within twenty-four hours, that after two days they showed only +slight movement, and that after three or four days they could not +be revived by moisture and warming. The important fact was +brought out that in moist air, in folded soiled underwear, they survived +as long as ten days. Bourguignon found that under the most +favorable conditions the mites of <i>Sarcoptes scabiei equi</i> would live for +sixteen days.</p> + +<p>The disease designated the "itch" or "scabies," in man has been +known from time immemorial, but until within less than a hundred +years it was almost universally attributed to malnutrition, errors of<span class="pagenum"><a name="Page_75" id="Page_75">[Pg 75]</a></span> +diet, or "bad blood." This was in spite of the fact that the mite was +known to Mouffet and that Bonomo had figured both the adult and +the egg and had declared the mite the sole cause of the disease. In +1834 the Corsican medical student, Francis Renucci, demonstrated +the mite before a clinic in Saint Louis Hospital in Paris and soon +thereafter there followed detailed studies of the life history of the +various itch mites of +man and animals.</p> + +<div class="figcenter" style="width: 270px;"><a name="Fig_58" id="Fig_58"></a> +<a href="images/f058-full.png"><img src="images/f058.png" width="270" height="390" alt="58. Scabies on the hand. From portfolio of Dermochromes +by permission of Rebman & Co., of +New York. Publishers." title="58. Scabies on the hand. From portfolio of Dermochromes +by permission of Rebman & Co., of +New York. Publishers." /></a> +<span class="caption">58. Scabies on the hand. From portfolio of Dermochromes +by permission of Rebman & Co., of +New York. Publishers.</span> +</div> + +<p>The disease is a cosmopolitan +one, being exceedingly +abundant in +some localities. Its +spread is much favored +where large numbers of +people are crowded together +under insanitary +conditions and hence it +increases greatly during +wars and is widely disseminated +and abundant +immediately afterwards. +Though more commonly +to be met with among +the lower classes, it not +infrequently appears +among those of the most +cleanly, careful habits, +and it is such cases that +are most liable to wrong +diagnosis by the physician.</p> + +<p>Infection occurs solely through the passage, direct or indirect, +of the young fertilized females to the skin of a healthy individual. +The adult, oviparous females do not quit their galleries and hence +do not serve to spread the disease. The young females move about +more or less at night and thus the principal source of infestation is +through sleeping in the same bed with an infested person, or indirectly +through bedclothes, or even towels or clothing. Diurnal infestation +through contact or clothing is exceptional. Many cases are known +of the disease being contracted from animals suffering from scabies, +or mange.<span class="pagenum"><a name="Page_76" id="Page_76">[Pg 76]</a></span></p> + +<p>When a person is exposed to infestation, the trouble manifests +itself after eight or ten days, though there usually elapses a period of +twenty to thirty days before +there is a suspicion of +anything serious. The first +symptom is an intense +itching which increases +when the patient is in bed. +When the point of irritation +is examined the galleries +may usually be seen +as characteristic sinuous +lines, at first whitish in +color but soon becoming +blackish because of the contained +eggs and excrement. +The galleries, which may +not be very distinct in +some cases, may measure +as much as four centimeters +in length. Little +vesicles, of the size of a +pin head are produced by +the secretions of the feeding mite; they are firm, and projecting, and +contain a limpid fluid. Figures <a href="#Fig_58">58</a> +and <a href="#Fig_59">59</a> show the typical appearance +of scabies on the hands, while <a href="#Fig_60">figure 60</a> +shows a severe general infestation. +The intolerable itching induces +scratching and through this various +complications may arise. The lesions +are not normally found on the face +and scalp, and are rare on the back.</p> + +<div class="figcenter" style="width: 315px;"><a name="Fig_59" id="Fig_59"></a> +<a href="images/f059-full.png"><img src="images/f059.png" width="315" height="396" alt="59. Scabies on the hand. After Duhring." title="59. Scabies on the hand. After Duhring." /></a> +<span class="caption">59. Scabies on the hand. After Duhring.</span> +</div> + +<div class="figcenter" style="width: 315px;"><a name="Fig_60" id="Fig_60"></a> +<a href="images/f060-full.png"><img src="images/f060.png" width="315" height="391" alt="60. Generalized infection of Scabies. +After Morrow." title="60. Generalized infection of Scabies. +After Morrow." /></a> +<span class="caption">60. Generalized infection of Scabies. +After Morrow.</span> +</div> + +<p>Formerly, scabies was considered +a very serious disease, for its cause +and method of treatment were unknown, +and potentially it may continue +indefinitely. Generation after +generation of the mites may develop +and finally their number become so<span class="pagenum"><a name="Page_77" id="Page_77">[Pg 77]</a></span> +great that the general health of the individual is seriously affected. +Now that the true cause of the disease is known, it is easily controlled.</p> + +<p>Treatment usually consists in softening the skin by friction with +soap and warm water, followed by a warm bath, and then applying +some substance to kill the mites. Stiles gives the following directions, +modified from Bourguignon's, as "a rather radical guide, to +be modified according to facilities and according to the delicacy of the +skin or condition of the patient":</p> + +<p>1. The patient, stripped naked, is energetically rubbed all over +(except the head) for twenty minutes, with green soap and warm +water. 2. He is then placed in a warm bath for thirty minutes, +during which time the rubbing is continued. 3. The parasiticide +is next rubbed in for twenty minutes and is allowed to remain on the +body for four or five hours; in the meantime the patient's clothes are +sterilized, to kill the eggs or mites attached to them. 4. A final +bath is taken to remove the parasiticide.</p> + +<p>The parasiticide usually relied on is the officinal sulphur ointment +of the United States pharmacopœia. When infestation is severe it +is necessary to repeat treatment after three or four days in order +to kill mites which have hatched from the eggs.</p> + +<p>The above treatment is too severe for some individuals and may, +of itself, produce a troublesome dermatitis. We have seen cases +where the treatment was persisted in and aggravated the condition +because it was supposed to be due to the parasite. For delicate-skinned +patients the use of balsam of Peru is very satisfactory, +and usually causes no irritation whatever. Of course, sources +of reinfection should be carefully guarded against.</p> + +<p><i>Sarcoptes scabiei crustosæ</i>, which is a distinct variety, if not species, +of the human itch mite, is the cause of so-called Norwegian itch. +This disease is very contagious, and is much more resistant than the +ordinary scabies. Unlike the latter, it may occur on the face and +scalp.</p> + +<p><i>Sarcoptes scabiei</i> not only attacks man but also occurs on a large +number of mammals. Many species, based on choice of host, and +minute differences in size and secondary characters, have been +established, but most students of the subject relegate these to +varietal rank. Many of them readily attack man, but they have +become sufficiently adapted to their normal host so that they are +usually less persistent on man.<span class="pagenum"><a name="Page_78" id="Page_78">[Pg 78]</a></span></p> + +<p><i>Notoedres cati</i> (usually known as <i>Sarcoptes minor</i>) is a species +of itch mites which produce an often fatal disease of cats. The body +is rounded and it is considerably smaller than <i>Sarcoptes scabiei</i>, +the female (<a href="#Fig_61">fig. 61</a>) measuring 215-230µ long and 165-175µ wide; +the males 145-150µ by 120-125µ. The most important character +separating <i>Notoedres</i> from <i>Sarcoptes</i> is the position of the anus, +which is dorsal instead of terminal. The mite readily transfers +to man but does not persist, the infestation usually disappearing +spontaneously in about two weeks. Infested cats are +very difficult to cure, unless treatment is begun at +the very inception of the outbreak, and under ordinary +circumstances it is better to kill them promptly, +to avoid spread of the disease to children and others +who may be exposed.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_61" id="Fig_61"></a> +<a href="images/f061-full.png"><img src="images/f061.png" width="500" height="312" alt="61. Notoedres cati, male and female. After Railliet." title="61. Notœdres cati, male and female. After Railliet." /></a> +<span class="caption">61. Notœdres cati, male and female. After Railliet.</span> +</div> + + +<h3><a name="Demodecidae" id="Demodecidae"></a>Demodecidæ</h3> + +<div class="figcenter" style="width: 125px;"><a name="Fig_62" id="Fig_62"></a> +<a href="images/f062-full.png"><img src="images/f062.png" width="125" height="463" alt="62. Demodex folliculorum. +(×200). +After Blanchard." title="62. Demodex folliculorum. +(×200). +After Blanchard." /></a> +<span class="caption">62. Demodex folliculorum. +(×200). +After Blanchard.</span> +</div> + +<p>The <b>Demodecidæ</b> are small, elongate, vermiform +mites which live in the hair follicles of mammals. +The family characteristics will be brought out in the +discussion of the species infesting man, <i>Demodex +folliculorum</i>.</p> + +<p><i>Demodex folliculorum</i> (<a href="#Fig_62">fig. 62</a>) is to be found very +commonly in the hair follicles and sebaceous glands +of man. It is vermiform in appearance, and with the +elongate abdomen transversely striated so as to give +it the appearance of segmentation. The female is 380-400µ +long by 45µ; the male 300µ by 40µ. The three-jointed<span class="pagenum"><a name="Page_79" id="Page_79">[Pg 79]</a></span> +legs, eight in number, are reduced to mere stubs in the adult. +The larval form is hexopod. These mites thus show in their form a +striking adaptation to their environment. In the sebaceous glands +and hair follicles they lie with their heads down (<a href="#Fig_63">fig. 63</a>). Usually +there are only a few in a gland, but Gruby has counted as many as +two hundred.</p> + +<div class="figcenter" style="width: 250px;"><a name="Fig_63" id="Fig_63"></a> +<a href="images/f063-full.png"><img src="images/f063.png" width="250" height="378" alt="63. Demodex folliculorum. Section through skin showing the +mites in situ. Magnification of Nos. 1, 2, 6 and 7, ×150; +Nos, 3, 4, 5, ×450. After Megnin." title="63. Demodex folliculorum. Section through skin showing the +mites in situ. Magnification of Nos. 1, 2, 6 and 7, ×150; +Nos, 3, 4, 5, ×450. After Megnin." /></a> +<span class="caption">63. Demodex folliculorum. Section through skin showing the +mites in situ. Magnification of Nos. 1, 2, 6 and 7, ×150; +Nos, 3, 4, 5, ×450. After Megnin.</span> +</div> + +<p>The frequency with which they occur in man is surprising. According +to European statistics they are found in 50 per cent to 60 per +cent or even more. Gruby found them in forty out of sixty persons<span class="pagenum"><a name="Page_80" id="Page_80">[Pg 80]</a></span> +examined. These figures are very commonly quoted, but reliable +data for the United States seem to be lacking. Our studies indicate +that it is very much less common in this country than is generally +assumed.</p> + +<p>The Demodex in man does not, as a rule, cause the slightest +inconvenience to its host. It is often stated that they give rise to +comedons or "black-heads" but there is no clear evidence that they +are ever implicated. Certain it is that they are not the usual cause. +A variety of the same, or a very closely related species of <i>Demodex</i>, +on the dog gives rise to the very resistant and often fatal follicular +mange.</p> + + +<h3><a name="Hexapoda_or_True_Insects" id="Hexapoda_or_True_Insects"></a><span class="smcap">Hexapoda or True Insects</span></h3> + +<p>The <b>Hexapoda</b>, or true insects, are characterized by the fact that +the adult possesses three pairs of legs. The body is distinctly +segmented and is divided into head, thorax, and abdomen.</p> + +<p>The mouth-parts in a generalized form, consist of an upper lip, +or <i>labrum</i>, which is a part of the head capsule, and a central unpaired +<i>hypopharynx</i>, two <i>mandibles</i>, two <i>maxillæ</i> and a lower lip, or <i>labium</i>, +made up of the fused pair of second maxillæ. These parts may be +greatly modified, dependent upon whether they are used for biting, +sucking, piercing and sucking, or a combination of biting and sucking.</p> + +<p>Roughly speaking, insects may be grouped into those which +undergo <i>complete metamorphosis</i> and those which have <i>incomplete +metamorphosis</i>. They are said to undergo complete metamorphosis +when the young form, as it leaves the egg, bears no resemblance to +the adult. For example, the maggot changes to a quiescent pupa +and from this emerges the winged active fly. They undergo incomplete +metamorphosis, when the young insect, as it leaves the egg, +resembles the adult to a greater or less extent, and after undergoing +a certain number of molts becomes sexually mature.</p> + +<p>Representatives of several orders have been reported as accidental +or faculative parasites of man, but the true parasites are restricted +to four orders. These are the Siphunculata; the Hemiptera, the +Diptera and the Siphonaptera.</p> + + +<h3><a name="Siphunculata" id="Siphunculata"></a><span class="smcap">Siphunculata</span></h3> + +<p>The order <b>Siphunculata</b> was established by Meinert to include the +true sucking lice. These are small wingless insects, with reduced +mouth-parts, adapted for sucking; thorax apparently a single piece +due to indistinct separation of its three segments: the compound eyes<span class="pagenum"><a name="Page_81" id="Page_81">[Pg 81]</a></span> +reduced to a single ommatidium on each side. The short, powerful +legs are terminated by a single long claw. Metamorphosis incomplete.</p> + +<p>There has been a great deal of discussion regarding the structure +of the mouth-parts, and the relationships of the sucking lice, and the +questions cannot yet be regarded +as settled. The conflicting +views are well represented +by Cholodkovsky +(1904 and 1905) and by +Enderlein (1904).</p> + +<div class="figcenter" style="width: 450px;"><a name="Fig_64" id="Fig_64"></a> +<a href="images/f064-full.png"><img src="images/f064.png" width="450" height="252" alt="64. Pediculus showing the blind sac (b) containing the +mouth parts (a) beneath the alimentary canal +(p). After Pawlowsky." title="64. Pediculus showing the blind sac (b) containing the +mouth parts (a) beneath the alimentary canal +(p). After Pawlowsky." /></a> +<span class="caption">64. Pediculus showing the blind sac (<i>b</i>) containing the +mouth parts (<i>a</i>) beneath the alimentary canal (<i>p</i>). +After Pawlowsky.</span> +</div> + +<p>Following Graber, it is +generally stated that the +mouth-parts consist of a +short tube furnished with +hooks in front, which constitutes +the lower lip, and that within this is a delicate sucking tube +derived from the fusion of the labrum and the mandibles. Opposed +to this, Cholodkovsky and, more recently, Pawlowsky, (1906), have +shown that the piercing apparatus lies in a blind sac under the +pharynx and opening into the mouth cavity (<a href="#Fig_64">fig. 64</a>). It does not +form a true tube but a furrow with its open surface uppermost. +Eysell has shown that, in addition, there is a pair of chitinous rods +which he regards as the homologues of the maxillæ.</p> + +<p>When the louse feeds, it everts the anterior part of the mouth +cavity, with its circle of hooks. The latter serve for anchoring +the bug, and the piercing apparatus is then pushed +out.</p> + +<div class="figright" style="width: 150px;"><a name="Fig_65" id="Fig_65"></a> +<a href="images/f065-full.png"><img src="images/f065.png" width="150" height="275" alt="65. Pediculus humanus, +ventral aspect +of male. (×10)" title="65. Pediculus humanus, +ventral aspect +of male. (×10)" /></a> +<span class="caption">65. Pediculus humanus, +ventral aspect +of male. (×10)</span> +</div> + +<p>Most writers have classed the sucking lice as a +sub-order of the Hemiptera, but the more recent +anatomical and developmental studies render this +grouping untenable. An important fact, bearing on +the question, is that, as shown by Gross, (1905), +the structure of the ovaries is radically different +from that of the Hemiptera.</p> + +<p>Lice infestation and its effects are known medically +as <i>pediculosis</i>. Though their continued presence +is the result of the grossest neglect and filthiness, the original +infestation may be innocently obtained and by people of the most +careful habits.<span class="pagenum"><a name="Page_82" id="Page_82">[Pg 82]</a></span></p> + +<p>Three species commonly attack man. Strangely enough, there +are very few accurate data regarding their life history.</p> + +<p><i>Pediculus humanus</i> (<a href="#Fig_65">fig. 65</a>), the head louse, is the most widely +distributed. It is usually referred to in medical literature as <i>Pediculus +capitis</i>, but the Linnean specific name has priority. In color +it is of a pale gray, blackish on the margins. It is claimed by some +authors that the color varies according to the color of the skin of the +host. The abdomen is +composed of seven distinct +segments, bearing +spiracles laterally. +There is considerable +variation in size. The +males average 1.8 mm. +and the females 2.7 mm. +in length.</p> + +<p>The eggs, fifty to +sixty in number, stick +firmly to the hairs of +the host and are known +as nits. They are large +and conspicuous, especially +on dark hair and +are provided with an +operculum, or cap, at +the free end, where the +nymphs emerge. They +hatch in about six days +and about the eighteenth +day the young +lice are sexually mature.</p> + +<div class="figcenter" style="width: 300px;"><a name="Fig_66" id="Fig_66"></a> +<a href="images/f066-full.png"><img src="images/f066.png" width="300" height="353" alt="66. Pediculosis of the head. The illustration shows the +characteristic indications of the presence of lice, viz: +the occipital eczema gluing the hairs together, the +swollen cervical glands, and the porrigo, or eruption +of contagious pustules upon the neck. After +Fox." title="66. Pediculosis of the head. The illustration shows the +characteristic indications of the presence of lice, viz: +the occipital eczema gluing the hairs together, the +swollen cervical glands, and the porrigo, or eruption +of contagious pustules upon the neck. After +Fox." /></a> +<span class="caption">66. Pediculosis of the head. The illustration shows the +characteristic indications of the presence of lice, viz: +the occipital eczema gluing the hairs together, the +swollen cervical glands, and the porrigo, or eruption +of contagious pustules upon the neck. After +Fox.</span> +</div> + +<p>The head lice live by preference on the scalp of their host but +occasionally they are found on the eyelashes and beard, or in the +pubic region. They may also occur elsewhere on the body. The +penetration of the rostrum into the skin and the discharge of an irritating +saliva produce a severe itching, accompanied by the formation +of an eczema-like eruption (<a href="#Fig_66">fig. 66</a>). When the infestation is severe, +the discharge from the pustules mats down the hair, and scabs are +formed, under which the insects swarm. "If allowed to run, a regular +carapace may form, called <i>trichoma</i>, and the head exudes a fœtid<span class="pagenum"><a name="Page_83" id="Page_83">[Pg 83]</a></span> +odor. Various low plants may grow in the trichoma, the whole +being known as <i>plica palonica</i>."—Stiles.</p> + +<p>Sources of infestation are various. School children may obtain +the lice from seatmates, by wearing the hats or caps of infested mates, +or by the use, in common, of brushes and combs. They may be +obtained from infested beds or sleeper berths. Stiles reports an instance +in which a large number of girls in a fashionable boarding +school developed lousiness a short time after traveling in a sleeping +car.</p> + +<p>Treatment is simple, for the parasites may readily be controlled +by cleanliness and washing the head with a two per cent solution of +carbolic acid or even kerosene. The latter is better used mixed with +equal parts of olive oil, to avoid irritation. The treatment should +be applied at night and followed the next morning by a shampoo with +soap and warm water. It is necessary to repeat the operation in a +few days. Xylol, used pure, or with the addition of five per cent +of vaseline, is also very efficacious. Of course, the patient must be +cautioned to stay away from a lighted lamp or fire while using either +the kerosene or xylol. While these treatments will kill the eggs or +nits, they will not remove them from the hairs. Pusey recommends +repeated washings with vinegar or 25 per cent of acetic acid in water, +for the purpose of loosening and removing the nits.</p> + +<p>Treatment of severe infestations in females is often troublesome +on account of long hair. For such cases the following method recommended +by Whitfield (1912) is especially applicable:</p> + +<p>The patient is laid on her back on the bed with her head over the +edge, and beneath the head is placed a basin on a chair so that the +hair lies in the basin. A solution of 1 in 40 carbolic acid is then poured +over the hair into the basin and sluiced backwards and forwards +until the whole of the hair is thoroughly soaked with it. It is especially +necessary that care should be taken to secure thorough saturation +of the hair over the ears and at the nape of the neck, since these +parts are not only the sites of predilection of the parasites but they +are apt to escape the solution. This sluicing is carried out for ten +minutes by the clock. At the end of ten minutes the hair is lifted +from the basin and allowed to drain, but is not dried or even thoroughly +wrung out. The whole head is then swathed with a thick +towel or better, a large piece of common house flannel, which is +fastened up to form a sort of turban, and is allowed to remain thus +for an hour. It can then be washed or simply allowed to dry, as the<span class="pagenum"><a name="Page_84" id="Page_84">[Pg 84]</a></span> +carbolic quickly disperses. At the end of this period every pediculus +and what is better, every ovum is dead and no relapse will occur +unless there is exposure to fresh contagion. Whitfield states that +there seem to be no disadvantages in this method, which he has used +for years. He has never seen carboluria result from it, but would +advise first cutting the hair of children under five years of age.</p> + +<p><i>Pediculus corporis</i> (= <i>P. vestimenti</i>) the body louse, is larger than +the preceding species, the female measuring 3.3 mm., and the male +3 mm. in length. The color is a dirty white, or grayish. <i>P. corporis</i> +has been regarded by some authorities as merely a variety of <i>P. +humanus</i> but Piaget maintains there are good characters separating +the two species.</p> + +<p>The body louse lives in the folds and seams of the clothing of its +host, passing to the skin only when it wishes to feed. Brumpt +states that he has found enormous numbers of them in the collars +of glass-ware or grains worn by certain naked tribes in Africa.</p> + +<p>Exact data regarding the life-history of this species have been +supplied, in part, by the work of Warburton (1910), cited by Nuttall. +He found that <i>Pediculus corporis</i> lives longer than <i>P. humanus</i> under +adverse conditions. This is doubtless due to its living habitually +on the clothing, whereas <i>humanus</i> lives upon the head, where it has +more frequent opportunities of feeding. He reared a single female +upon his own person, keeping the louse enclosed in a cotton-plugged +tube with a particle of cloth to which it could cling. The tube was +kept next to his body, thus simulating the natural conditions of +warmth and moisture under which the lice thrive. The specimen +was fed twice daily, while it clung to the cloth upon which it rested. +Under these conditions she lived for one month. Copulation commenced +five days after the female had hatched and was repeated a +number of times, sexual union lasting for hours. The female laid +one hundred and twenty-four eggs within twenty-five days.</p> + +<p>The eggs hatched after eight days, under favorable conditions, +such as those under which the female was kept. They did not +hatch in the cold. Eggs kept near the person during the day and +hung in clothing by the bedside at night, during the winter, in a cold +room, did not hatch until the thirty-fifth day. When the nymphs +emerge from the eggs, they feed at once, if given a chance to do so. +They are prone to scatter about the person and abandon the fragment +of cloth to which the adult clings.<span class="pagenum"><a name="Page_85" id="Page_85">[Pg 85]</a></span></p> + +<p>The adult stage is reached on the eleventh day, after three molts, +about four days apart. Adults enter into copulation about the +fifth day and as the eggs require eight days for development, +the total cycle, under favorable conditions, is about twenty-four +days. Warburton's data differ considerably from those commonly +quoted and serve to emphasize the necessity for detailed studies +of some of the commonest of parasitic insects.</p> + +<p>Body lice are voracious feeders, producing by their bites and the +irritating saliva which they inject, rosy elevations and papules which +become covered with a brownish +crust. The intense itching provokes +scratching, and characteristic +white scars (<a href="#Fig_67">fig. 67</a>) surrounded +by brownish pigment +(<a href="#Fig_68">fig. 68</a>) are formed. The skin +may become thickened and take +on a bronze tinge. This melanoderma +is especially marked +in the region between the shoulders +but it may become generalized, +a prominent characteristic +of "vagabond's disease." According +to Dubre and Beille, +this melanoderma is due to a +toxic substance secreted by the +lice, which indirectly provokes +the formation of pigment.</p> + +<div class="figcenter" style="width: 300px;"><a name="Fig_67" id="Fig_67"></a> +<a href="images/f067-full.png"><img src="images/f067.png" width="300" height="391" alt="67. Pediculosis in man caused by the body +louse. After Morrow." title="67. Pediculosis in man caused by the body +louse. After Morrow." /></a> +<span class="caption">67. Pediculosis in man caused by the body +louse. After Morrow.</span> +</div> + +<p>Control measures, in the case +of the body louse, consist in +boiling or steaming the clothes or in some cases, sterilizing by dry heat. +The dermatitis may be relieved by the use of zinc-oxide ointment, +to which Pusey recommends that there be added, on account of their +parasiticidal properties, sulphur and balsam of Peru, equal parts, 15 +to 30 grains to the ounce.</p> + +<div class="figcenter" style="width: 245px;"><a name="Fig_68" id="Fig_68"></a> +<a href="images/f068-full.png"><img src="images/f068.png" width="245" height="463" alt="68. Melanoderma caused by the body +louse. From Portfolio of Dermochromes, +by permission of Rebman +& Co., New York, Publishers." title="68. Melanoderma caused by the body +louse. From Portfolio of Dermochromes, +by permission of Rebman +& Co., New York, Publishers." /></a> +<span class="caption">68. Melanoderma caused by the body +louse. From Portfolio of Dermochromes, +by permission of Rebman +& Co., New York, Publishers.</span> +</div> + +<div class="figright" style="width: 200px;"><a name="Fig_69" id="Fig_69"></a> +<a href="images/f069-full.png"><img src="images/f069.png" width="200" height="182" alt="69. Phthirius pubis. Ventral +aspect of female. +(×12)." title="69. Phthirius pubis. Ventral +aspect of female. +(×12)." /></a> +<span class="caption">69. Phthirius pubis. Ventral +aspect of female. +(×12).</span> +</div> + +<p><i>Phthirius pubis</i> (= <i>P. inguinalis</i>), the pubic louse, or so-called +"crab louse," differs greatly from the preceding in appearance. It is +characterized by its relatively short head which fits into a broad +depression in the thorax. The latter is broad and flat and merges +into the abdomen. The first pair of legs is slender and terminated +by a straight claw. The second and third pairs of legs are thicker<span class="pagenum"><a name="Page_86" id="Page_86">[Pg 86]</a></span> +and are provided with powerful claws fitted for clinging to hairs. +The females (<a href="#Fig_69">fig. 69</a>) measure 1.5 to 2 mm. in length by 1.5 mm. in +breadth. The male averages a +little over half as large. The eggs, +or nits, are fixed at the base of the +hairs. Only a few, ten to fifteen +are deposited by a single female, +and they hatch in about a week's +time. The young lice mature in +two weeks.</p> + +<p>The pubic louse usually infests +the hairs of the pubis and the +perineal region. It may pass to +the arm pits or even to the beard +or moustache. Rarely, it occurs +on the eyelids, and it has even +been found, in a very few instances, +occurring in all stages, on the scalp. +Infestation may be contracted +from beds or even from badly infested +persons in a crowd. We +have seen several cases which undoubtedly +were due to the use of +public water closets. It produces +papular eruption and an intense +pruritis. When abundant, there +occurs a grayish discoloration of +the skin which Duguet has shown +is due to a poisonous saliva injected by the louse, +as is the melanoderma caused by the body louse.</p> + +<p>The pubic louse may be exterminated by the +measures recommended for the head louse, or +by the use of officinal mercurial ointment.</p> + + +<h3><span class="smcap">Hemiptera</span></h3> + +<p>Several species of <b>Hemiptera-Heteroptera</b> are +habitual parasites of man, and others occur +as occasional or accidental parasites. Of all +these, the most important and widespread are the bed-bugs, belonging +to the genus <i>Cimex</i> (= <i>Acanthia</i>).<span class="pagenum"><a name="Page_87" id="Page_87">[Pg 87]</a></span></p> + + +<p><a name="The_Bed-bugs" id="The_Bed-bugs"></a><b>The Bed-bugs</b>—The bed-bugs are characterized by a much flattened +oval body, with the short, broad head unconstricted behind, +and fitting into the strongly excavated anterior margin of the thorax. +The compound eyes are prominent, simple eyes lacking. Antennæ +four-jointed, the first segment short, the second long and thick, and +the third and fourth slender. The tarsi are short and three segmented.</p> + +<p>It is often assumed in the literature of the subject that there is +but a single species of <i>Cimex</i> attacking man, but several such species +are to be recognized. These are distinguishable by the characters +given in Chapter XII. We shall consider +especially <i>Cimex lectularius</i>, the +most common and widespread species.</p> + +<p><i>Cimex lectularius</i> (= <i>Acanthia +lectularia</i>, <i>Clinocoris lectularius</i>), is +one of the most cosmopolitan of human +parasites but, like the lice, it has been +comparatively little studied until +recent years, when the possibility +that it may be concerned with the +transmission of various diseases has +awakened interest in the details of +its life-history and habits.</p> + +<div class="figleft" style="width: 350px;"><a name="Fig_70" id="Fig_70"></a> +<a href="images/f070-full.png"><img src="images/f070.png" width="350" height="382" alt="70. Cimex lectularius adult and eggs. +Photograph by M. V. S." title="70. Cimex lectularius adult and eggs. +Photograph by M. V. S." /></a> +<span class="caption">70. Cimex lectularius adult and eggs. +Photograph by M. V. S.</span> +</div> + +<p>The adult insect (<a href="#Fig_70">fig. 70</a>) is 4-5 +mm. long by 3 mm. broad, reddish +brown in color, with the beak and body appendages lighter in color. +The short, broad and somewhat rectangular head has no neck-like +constriction but fits into the broadly semilunar prothorax. The +four segmented labium or proboscis encloses the lancet-like maxillæ +and mandibles. The distal of the four antennal segments is slightly +club-shaped. The prothorax is characteristic of the species, being +deeply incised anteriorly and with its thin lateral margins somewhat +turned up. The mesothorax is triangular, with the apex posteriorly, +and bears the greatly atrophied first pair of wings. There is no trace +of the metathoracic pair. The greatly flattened abdomen has eight +visible segments, though in reality the first is greatly reduced and +has been disregarded by most writers. The body is densely covered +with short bristles and hairs, the former being peculiarly saber-shaped +structures sharply toothed at the apex and along the convex +side (<a href="#Fig_159">fig. 159<i>b</i></a>).<span class="pagenum"><a name="Page_88" id="Page_88">[Pg 88]</a></span></p> + +<p>The peculiar disagreeable odor of the adult bed-bug is due to the +secretion of the stink glands which lie on the inner surface of the +mesosternum and open by a pair of orifices in front of the metacoxæ, +near the middle line. In the nymphs, the thoracic glands are not +developed but in the abdomen there are to be found three unpaired +dorsal stink glands, which persist until the fifth molt, when they +become atrophied and replaced by the thoracic glands. The nymphal +glands occupy the median dorsal portion of the abdomen, opening +by paired pores at the anterior margin of the fourth, fifth and sixth +segments. The secretion is a clear, oily, volatile fluid, strongly acid +in reaction. Similar glands are to be found in most of the Hemiptera-Heteroptera +and their secretion is doubtless protective, through +being disagreeable to the birds. In the bed-bug, as Marlatt points +out, "it is probably an illustration of a very common phenomenon +among animals, i.e., the persistence of a characteristic which is no +longer of any special value to the possessor." In fact, its possession +is a distinct disadvantage to the bed-bug, as the odor frequently +reveals the presence of the bugs, before they are seen.</p> + +<p>The eggs of the bed-bug (<a href="#Fig_70">fig. 70</a>) are pearly white, oval in outline, +about a millimeter long, and possess a small operculum or cap +at one end, which is pushed off when the young hatches. They are +laid intermittently, for a long period, in cracks and crevices of beds +and furniture, under seams of mattresses, under loose wall paper, +and similar places of concealment of the adult bugs. Girault (1905) +observed a well-fed female deposit one hundred and eleven eggs +during the sixty-one days that she was kept in captivity. She had +apparently deposited some of her eggs before being captured.</p> + +<p>The eggs hatch in six to ten days, the newly emerged nymphs +being about 1.5 mm. in length and of a pale yellowish white color. +They grow slowly, molting five times. At the last molt the mesathoracic +wing pads appear, characteristic of the adult. The total +length of the nymphal stage varies greatly, depending upon conditions +of food supply, temperature and possibly other factors. Marlatt +(1907) found under most favorable conditions a period averaging +eight days between molting which, added to an equal egg period, +gave a total of about seven weeks from egg to adult insect. Girault +(1912) found the postembryonic period as low as twenty-nine days +and as high as seventy days under apparently similar and normal +conditions of food supply. Under optimum and normal conditions +of food supply, beginning August 27, the average nymphal life was<span class="pagenum"><a name="Page_89" id="Page_89">[Pg 89]</a></span> +69.9 days; average number of meals 8.75 and the molts 5. Under +conditions allowing about half the normal food supply the average +nymphal life was from 116.9 to 139 days. Nymphs starved from +birth lived up to 42 days. We have kept unfed nymphs, of the first +stage, alive in a bottle for 75 days. The interesting fact was brought +out that under these conditions of minimum food supply there were +sometimes six molts instead of the normal number.</p> + +<p>The adults are remarkable for their longevity, a factor which is +of importance in considering the spread of the insect and methods of +control. Dufour (1833) (not De Geer, as often stated) kept specimens +for a year, in a closed vial, without food. This ability, coupled +with their willingness to feed upon mice, bats, and other small mammals, +and even upon birds, accounts for the long periods that deserted +houses and camps may remain infested. There is no evidence that +under such conditions they are able to subsist on the starch of the +wall paper, juices of moistened wood, or the moisture in the accumulations +of dust, as is often stated.</p> + +<p>There are three or four generations a year, as Girault's breeding +experiments have conclusively shown. He found that the bed-bug +does not hibernate where the conditions are such as to allow it to +breed and that breeding is continuous unless interrupted by the lack +of food or, during the winter, by low temperature.</p> + +<p>Bed-bugs ordinarily crawl from their hiding places and attack +the face and neck or uncovered parts of the legs and arms of their +victims. If undisturbed, they will feed to repletion. We have +found that the young nymph would glut itself in about six minutes, +though some individuals fed continuously for nine minutes, while +the adult required ten to fifteen minutes for a full meal. When +gorged, it quickly retreats to a crack or crevice to digest its meal, +a process which requires two or three days. The effect of the bite +depends very greatly on the susceptibility of the individual attacked. +Some persons are so little affected that they may be wholly ignorant +of the presence of a large number of bugs. Usually the bite produces +a small hard swelling, or wheal, whitish in color. It may even be +accompanied by an edema and a disagreeable inflammation, and in +such susceptible individuals the restlessness and loss of sleep due to +the presence of the insects may be a matter of considerable importance. +Stiles (1907) records the case of a young man who underwent +treatment for neurasthenia, the diagnosis being agreed upon by +several prominent physicians; all symptoms promptly disappeared,<span class="pagenum"><a name="Page_90" id="Page_90">[Pg 90]</a></span> +however, immediately following a thorough fumigation of his rooms, +where nearly a pint of bed-bugs were collected.</p> + +<p>It is natural to suppose that an insect which throughout its whole +life is in such intimate relationship with man should play an important +rôle in the transmission of disease. Yet comparatively little is +definitely known regarding the importance of the bed-bug in this +respect. It has been shown that it is capable of transmitting the +bubonic plague, and South American trypanosomiasis. Nuttall +succeeded in transmitting European relapsing fever from mouse to +mouse by its bite. It has been claimed that Oriental sore, tuberculosis, +and even syphilis may be so carried. These phases of the +subject will be considered later.</p> + +<p>The sources of infestation are many, and the invasion of a house +is not necessarily due to neglect, though the continued presence of +the pests is quite another matter. In apartments and closely placed +houses they are known to invade new quarters by migration. They +are frequently to be met with in boat and sleeper berths, and even +the plush seats of day coaches, whence a nucleus may be carried in +baggage to residences. They may be brought in the laundry or +in clothes of servants.</p> + +<p>Usually they are a great scourge in frontier settlements and it is +generally believed that they live in nature under the bark of trees, +in lumber, and under similar conditions. This belief is founded upon +the common occurrence of bugs resembling the bed-bug, in such +places. As a matter of fact, they are no relation to bed-bugs but +belong to plant-feeding forms alone (<a href="#Fig_19">fig. 19</a> <i>c</i>, <i>d</i>).</p> + +<p>It is also often stated that bed-bugs live in poultry houses, in +swallows nests, and on bats, and that it is from these sources that they +gain access to dwellings. These bugs are specifically distinct from +the true bed-bug, but any of them may, rarely, invade houses. +Moreover, chicken houses are sometimes thoroughly infested with +the true <i>Cimex lectularius</i>.</p> + +<p>Control measures consist in the use of iron bedsteads and the +reduction of hiding places for the bugs. If the infestation is slight +they may be exterminated by a vigilant and systematic hunt, and +by squirting gasoline or alcohol into cracks and crevices of the beds, +and furniture. Fumigation must be resorted to in more general +infestations.</p> + +<p>The simplest and safest method of fumigation is by the use of +flowers of sulphur at the rate of two pounds to each one thousand<span class="pagenum"><a name="Page_91" id="Page_91">[Pg 91]</a></span> +cubic feet of room space. The sulphur should be placed in a pan, +a well made in the top of the pile and a little alcohol poured in, to +facilitate burning. The whole should be placed in a larger pan +and surrounded by water so as to avoid all danger of fire. Windows +should be tightly closed, beds, closets and drawers opened, and +bedding spread out over chairs in order to expose them fully to the +fumes. As metal is tarnished by the sulphur fumes, ornaments, +clocks, instruments, and the like should be removed. When all is +ready the sulphur should be fired, the room tightly closed and left +for twelve to twenty-four hours. Still more efficient in large houses, +or where many hiding places favor the bugs, is fumigation with +hydrocyanic acid gas. This is a deadly poison and must be used +under rigid precautions. Through the courtesy of Professor Herrick, +who has had much experience with this method, we give in the Appendix, +the clear and detailed directions taken from his bulletin on +"Household Insects."</p> + +<p>Fumigation with formaldehyde gas, either from the liquid or +"solid" formalin, so efficient in the case of contagious diseases, is +useless against bed-bugs and most other insects.</p> + + +<p><a name="Other_Bed-bugs" id="Other_Bed-bugs"></a><b>Other Bed-bugs</b>—<i>Cimex hemipterus</i> (= <i>C. rotundatus</i>) is a tropical +and subtropical species, occurring in both the old and new world. +Patton and Cragg state that it is distributed throughout India, +Burma, Assam, the Malay Peninsula, Aden, the Island of Mauritius, +Reunion, St. Vincent and Porto Rico. "It is widely distributed +in Africa, and is probably the common species associated there +with man." Brumpt also records it for Cuba, the Antilles, Brazil, +and Venezuela.</p> + +<p>This species, which is sometimes called the Indian bed-bug, +differs from <i>C. lectularius</i> in being darker and in having a more +elongate abdomen. The head also is shorter and narrower, and the +prothorax has rounded borders.</p> + +<p>It has the same habits and practically the same life cycle as +<i>Cimex lectularius</i>. Mackie, in India, has found that it is capable +of transmitting the Asiatic type of recurrent fever. Roger suggested +that it was also capable of transmitting Kala-azar and Patton has +described in detail the developmental stages of <i>Leishmania</i>, the +causative organism of Kala-azar, in the stomach of this bug, but +Brumpt declares that the forms described are those of a common, +non-pathogenic flagellate to be found in the bug, and have nothing<span class="pagenum"><a name="Page_92" id="Page_92">[Pg 92]</a></span> +to do with the human disease. Brumpt has shown experimentally +that <i>Cimex hemipterus</i> may transmit <i>Trypanosoma cruzi</i> in its excrement.</p> + +<p><i>Cimex boueti</i>, occurring in French Guinea, is another species +attacking man. Its habits and general life history are the same as +for the above species. It is 3 to 4.5 mm. in length, +has vestigial elytra, and much elongated antennæ and +legs. The extended hind legs are about as long as the +body.</p> + +<p><i>Cimex columbarius</i>, a widely distributed species normally +living in poultry houses and dove cotes, <i>C. inodorus</i>, +infesting poultry in Mexico, <i>C. hirundinis</i>, occurring in +the nests of swallows in Europe and <i>Oeciacus vicarius</i> +(<a href="#Fig_19">fig. 19i</a>) occurring in swallows' nests in this country, +are species which occasionally infest houses and attack +man.</p> + +<div class="figleft" style="width: 150px;"><a name="Fig_71" id="Fig_71"></a> +<a href="images/f071-full.png"><img src="images/f071.png" width="150" height="271" alt="71. Conorhinus +sanguisugus." title="71. Conorhinus +sanguisugus." /></a> +<span class="caption">71. Conorhinus +sanguisugus.</span> +</div> + +<p><i>Conorhinus sanguisugus</i>, the cone-nosed bed-bug. We have seen +in our consideration of poisonous insects, that various species of +Reduviid bugs readily attack man. Certain of these are nocturnal +and are so commonly found in houses that they have gained the +name, of "big bed-bugs." The most noted of these, in the United +States, is <i>Conorhinus <span title="for sangiusugus read sanguisugus"><a name="AC_5a" id="AC_5a"></a>sangiusugus</span></i> (<a href="#Fig_71">fig. 71</a>), which is widely distributed +in our Southern States.</p> + +<p>Like its near relatives, <i>Conorhinus +<span title="for sangiusugus read sanguisugus"><a name="AC_5b" id="AC_5b"></a>sangiusugus</span></i> is carnivorous in habit and +feeds upon insects as well as upon +mammalian and human blood. It is +reported as often occurring in poultry +houses and as attacking horses in +barns. The life history has been +worked out in considerable detail by +Marlatt, (1902), from whose account we +extract the following.</p> + +<div class="figright" style="width: 350px;"><a name="Fig_72" id="Fig_72"></a> +<a href="images/f072-full.png"><img src="images/f072.png" width="350" height="419" alt="72. Beak of Conorhinus sanguisugus. +After Marlatt." title="72. Beak of Conorhinus sanguisugus. +After Marlatt." /></a> +<span class="caption">72. Beak of Conorhinus sanguisugus. +After Marlatt.</span> +</div> + +<p>The eggs are white, changing to +yellow and pink before hatching. The +young hatch within twenty days +and there are four nymphal stages. +In all these stages the insect is active and predaceous, the mouth-parts +(<a href="#Fig_72">fig. 72</a>) being powerfully developed. The eggs are normally +deposited, and the early stages are undoubtedly passed, out of doors,<span class="pagenum"><a name="Page_93" id="Page_93">[Pg 93]</a></span> +the food of the immature forms being other insects. Immature +specimens are rarely found indoors. It winters both in the partly +grown and adult stage, often under the bark of trees or in any +similar protection, and only in its nocturnal spring and early +summer flights does it attack men. Marlatt states that this insect +seems to be decidedly on the increase in the region which it particularly +infests,—the plains region from Texas northward and westward. +In California a closely related species of similar habits is +known locally as the "monitor bug."</p> + +<p>The effect of the bite of the giant bed-bug on man is often very +severe, a poisonous saliva apparently being injected into the wound. +We have discussed this phase of the subject more fully under the +head of poisonous insects.</p> + +<p><i>Conorhinus megistus</i> is a Brazilian species very commonly attacking +man, and of special interest since Chagas has shown that it is +the carrier of a trypanosomiasis of man. Its habits and life history +have been studied in detail by Neiva, (1910).</p> + +<p>This species is now pre-eminently a household insect, depositing +its eggs in cracks and crevices in houses, though this is a relatively +recent adaptation. The nymphs emerge in from twenty to forty +days, depending upon the temperature. There are five nymphal +stages, and as in the case of true bed-bugs, the duration of these is +very greatly influenced by the availability of food and by temperature. +Neiva reckons the entire life cycle, from egg to egg, as requiring +a minimum of three hundred and twenty-four days.</p> + +<p>The nymphs begin to suck blood in three to five days after hatching. +They usually feed at night and in the dark, attacking especially +the face of sleeping individuals. The bite occasions but little pain. +The immature insects live in cracks and crevices in houses and +invade the beds which are in contact with walls, but the adults are +active flyers and attack people sleeping in hammocks. The males +as well as the females are blood suckers.</p> + +<p>Like many blood-sucking forms, <i>Conorhinus megistus</i> can endure +for long periods without food. Neiva received a female specimen +which had been for fifty-seven days alive in a tightly closed box. +They rarely feed on two consecutive days, even on small quantities +of blood, and were never seen to feed on three consecutive days.</p> + +<p>Methods of control consist in screening against the adult bugs, +and the elimination of crevices and such hiding places of the nymphs. +Where the infestation is considerable, fumigation with sulphur is +advisable.<span class="pagenum"><a name="Page_94" id="Page_94">[Pg 94]</a></span></p> + + +<h3><a name="Parasitic_Diptera_or_Flies" id="Parasitic_Diptera_or_Flies"></a><span class="smcap">Parasitic Diptera or Flies</span></h3> + +<p>Of the <b>Diptera</b> or two-winged flies, many species occasionally +attack man. Of these, a few are outstanding pests, many of them +may also serve to disseminate disease, a phase of our subject which +will be considered later. We shall now consider the most important +of the group from the viewpoint of their direct attacks on man.</p> + + +<h3><a name="Psychodidae_or_Moth-Flies" id="Psychodidae_or_Moth-Flies"></a>Psychodidæ or Moth-Flies</h3> + +<p>The <b>Psychodidæ</b> or Moth-flies, include a few species which attack +man, and at least one species, <i>Phlebotomus papatasii</i>, is known +to transmit the so-called "three-day fever" of man. Another species +is supposed to be the vector of Peruvian verruga.</p> + +<p>The family is made up of small, sometimes very small, nematocerous +Diptera, which are densely covered with hairs, giving them a +moth-like appearance. The wings are relatively large, oval or +lanceolate in shape, and when at rest are held in a sloping manner +over the abdomen, or are held horizontally in such a way as to give +the insect a triangular outline. Not only is the moth-like appearance +characteristic, but the venation of the wings (<a href="#Fig_163">fig. 163, d</a>) is very peculiar +and, according to Comstock, presents an extremely generalized form. +All of the longitudinal veins separate near the base of the wing +except veins R<sub>2</sub> and R<sub>3</sub> and veins M<sub>1 </sub>and M<sub>2</sub>. Cross veins are +wanting in most cases.</p> + +<p>Comparatively little is known regarding the life-history and +habits of the Psychodidæ, but one genus, <i>Phlebotomus</i>, contains +minute, blood-sucking species, commonly known as sand-flies. The +family is divided into two subfamilies, the <b>Psychodinæ</b> and the +<b>Phlebotominæ</b>. The second of these, the <b>Phlebotominæ</b>, is of +interest to us.</p> + + +<p><a name="The_Phlebotominae" id="The_Phlebotominae"></a><b>The Phlebotominæ</b>—The Phlebotominæ differ from the Psychodinæ +in that the radical sector branches well out into the wing rather +than at the base of the wing. They are usually less hairy than the +Psychodinæ. The ovipositor is hidden and less strongly chitinized. +The species attacking man belong to the genus <i>Phlebotomus</i>, small +forms with relatively large, hairy wings which are held upright, +and with elongate proboscis. The mandibles and maxillæ are serrated +and fitted for biting.</p> + +<p>According to Miss Summers (1913) there are twenty-nine known +species of the genus <i>Phlebotomus</i>, five European, eleven Asiatic,<span class="pagenum"><a name="Page_95" id="Page_95">[Pg 95]</a></span> +seven African and six American. One species only, <i>Phlebotomus +vexator</i>, has been reported for the United States. This was described +by Coquillett, (1907), from species taken on Plummer's Island, Maryland. +It measures only 1.5 mm. in length. As it is very probable +that this species is much more widely distributed, and that other +species of these minute flies will be found to occur in our fauna, we +quote Coquillett's description.</p> + +<p><i>Phlebotomus vexator</i>, Coq.: Yellow, the mesonotum brown, +hairs chiefly brown; legs in certain lights appear brown, but are +covered with a white tomentum; wings hyaline, unmarked; the first +vein (R<sub>1</sub>) terminates opposite one-fifth of the length of the first +submarginal cell (cell R<sub>2</sub>); this cell is slightly over twice as long +as its petiole; terminal, horny portion of male claspers slender, +bearing many long hairs; the apex terminated by two curved spines +which are more than one-half as long as the preceding part, and just +in front of these are two similar spines, while near the middle of the +length of this portion is a fifth spine similar to the others. Length +1.5 mm.</p> + +<p>The life-history of the Phlebotomus flies has been best worked out +for the European <i>Phlebotomus papatasii</i> and we shall briefly summarize +the account of Dœrr and Russ (1913) based primarily on work +on this species. The European Phlebotomus flies appear at the +beginning of the warm season, a few weeks after the cessation of the +heavy rains and storms of springtime. They gradually become more +abundant until they reach their first maximum, which in Italy is near +the end of July (Grassi). They then become scarcer but reach a +second maximum in September. At the beginning of winter they +vanish completely, hibernating individuals not being found.</p> + +<p>After fertilization there is a period of eight to ten days before oviposition. +The eggs are then deposited, the majority in a single mass +covered by a slimy secretion from the sebaceous glands. The larvæ +emerge in fourteen to twenty days. There is uncertainty as to the +length of larval life, specimens kept in captivity remaining fifty or +more days without transforming. Growth may be much more rapid +in nature. The larvæ do not live in fluid media but in moist detritus +in dark places. Marett believes that they live chiefly on the excrement +of pill-bugs (Oniscidæ) and lizards. Pupation always occurs +during the night. The remnants of the larval skin remain attached +to the last two segments of the quiescent pupa and serve to attach +it to the stone on which it lives. The pupal stage lasts eleven to +sixteen days, the adult escaping at night.<span class="pagenum"><a name="Page_96" id="Page_96">[Pg 96]</a></span></p> + +<p>Only the females suck blood. They attack not only man but all +warm-blooded animals and, according to recent workers, also cold-blooded +forms, such as frogs, lizards, and larvæ. Indeed, Townsend +(1914) believes that there is an intimate relation between <i>Phlebotomus</i> +and lizards, or other reptiles the world over. The Phlebotomus +passes the daylight hours within the darkened recesses of the loose +stone walls and piles of rock in order to escape wind and strong light. +Lizards inhabit the same places, and the flies, always ready to suck +blood in the absence of light and wind, have been found more prone +to suck reptilian than mammalian blood.</p> + +<p>On hot summer nights, when the wind is not stirring, the Phlebotomus +flies, or sand-flies, as they are popularly called, invade houses and +sleeping rooms in swarms and attack the inmates. As soon as light +begins to break the flies either escape to the breeding places, or cool, +dark places protected from the wind, or a part of them remain in the +rooms, hiding behind pictures, under garments, and in similar places. +Wherever the Phlebotomus flies occur they are an intolerable nuisance. +On account of their small size they can easily pass through +the meshes of ordinary screens and mosquito curtains. They attack +silently and inflict a very painful, stinging bite, followed by itching. +The ankles, dorsum of the feet, wrists, inner elbow, knee joint and +similar places are favorite places of attack, possibly on account of +their more delicate skin.</p> + +<p>Special interest has been attracted to these little pests in recent +years, since it has been shown that they transmit the European +"pappatici fever" or "three day fever." More recently yet, it +appears that they are the carriers of the virus of the Peruvian "verruga." +This phase of the subject will be discussed later.</p> + +<p>Control measures have not been worked out. As Newstead says, +"In consideration of the facts which have so far been brought to light +regarding the economy of Phlebotomus, it is clearly evident that the +task of suppressing these insects is an almost insurmountable one. +Had we to deal with insects as large and as accessible as mosquitoes, +the adoption of prophylactic measures would be comparatively easy, +but owing to the extremely minute size and almost flea-like habits of +the adult insects, and the enormous area over which the breeding-places +may occur, we are faced with a problem which is most difficult +of solution." For these reasons, Newstead considers that the only +really prophylactic measures which can at present be taken, are those +which are considered as precautionary against the bites of the insects.<span class="pagenum"><a name="Page_97" id="Page_97">[Pg 97]</a></span></p> + +<p>Of repellents, he cites as one of the best a salve composed of the +following:</p> + +<div class="center"> +<table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td align="left">Ol. Anisi</td><td align="left">3 grs.</td></tr> +<tr><td align="left">Ol. Eucalypti</td><td align="left">3 grs.</td></tr> +<tr><td align="left">Ol. Terebenth</td><td align="left">3 grs.</td></tr> +<tr><td align="left">Unq. Acid Borac.</td></tr> +</table></div> + +<p>Of sprays he recommends as the least objectionable and at the +same time one of the most effective, formalin. "The dark portions +and angles of sleeping apartments should be sprayed with a one per +cent. solution of this substance every day during the season in which +the flies are prevalent. A fine spraying apparatus is necessary for +its application and an excessive amount must not be applied. It is +considered an excellent plan also to spray the mosquito curtains +regularly every day towards sunset; nets thus treated are claimed to +repel the attacks of these insects." This effectiveness of formalin is +very surprising for, as we have seen, it is almost wholly ineffective +against bed-bugs, mosquitoes, house flies and other insects, where it +has been tried.</p> + +<p>A measure which promises to be very effective, where it can be +adopted, is the use of electric fans so placed as to produce a current +of air in the direction of the windows of sleeping apartments. On +account of the inability of the Phlebotomus flies to withstand even +slight breezes, it seems very probable that they would be unable to +enter a room so protected.</p> + + +<h3><a name="Culicidae_or_Mosquitoes" id="Culicidae_or_Mosquitoes"></a>Culicidæ or Mosquitoes</h3> + +<p>From the medical viewpoint, probably the most interesting and +important of the blood-sucking insects are the mosquitoes. Certainly +this is true of temperate zones, such as those of the United States. +The result is that no other group of insects has aroused such widespread +interest, or has been subjected to more detailed study than +have the mosquitoes, since their rôle as carriers of disease was made +known. There is an enormous literature dealing with the group, but +fortunately for the general student, this has been well summarized +by a number of workers. The most important and helpful of the +general works are those of Howard (1901), Smith (1904), Blanchard +(1905), Mitchell (1907), and especially of Howard, Dyar, and Knab, +whose magnificent monograph is still in course of publication.<span class="pagenum"><a name="Page_98" id="Page_98">[Pg 98]</a></span></p> + +<p>Aside from their importance as carriers of disease, mosquitoes are +notorious as pests of man, and the earlier literature on the group is +largely devoted to references to their enormous numbers and their +blood-thirstiness in certain regions. They are to be found in all +parts of the world, from the equator to the Arctic and Antarctic +regions. Linnæus, in the "Flora Lapponica," according to Howard, +Dyar and Knab, "dwells at some length upon the great abundance of +mosquitoes in Lapland and the torments they inflicted upon man and +beast. He states that he believes that nowhere else on earth are they +found in such abundance and he compares their numbers to the dust +of the earth. Even in the open, you cannot draw your breath without +having your mouth and nostrils filled with them; and ointments of +tar and cream or of fish grease are scarcely sufficient to protect even +the case-hardened cuticle of the Laplander from their bite. Even in +their cabins, the natives cannot take a mouthful of food or lie down +to sleep unless they are fumigated almost to suffocation." In some +parts of the Northwestern and Southwestern United States it is +necessary to protect horses working in the fields by the use of sheets or +burlaps, against the ferocious attacks of these insects. It is a surprising +fact that even in the dry deserts of the western United States +they sometimes occur in enormous numbers.</p> + +<p>Until comparatively recent years, but few species of mosquitoes +were known and most of the statements regarding their life-history +were based upon the classic work of Reaumur (1738) on the biology +of the rain barrel mosquito, <i>Culex pipiens</i>. In 1896, Dr. Howard +refers to twenty-one species in the United States, now over fifty are +known; Giles, in 1900, gives a total of two hundred and forty-two +for the world fauna, now over seven hundred species are known. +We have found eighteen species at Ithaca, N. Y.</p> + +<p>All of the known species of mosquitoes are aquatic in the larval +stage, but in their life-histories and habits such great differences occur +that we now know that it is not possible to select any one species as +typical of the group. For our present purpose we shall first discuss +the general characteristics and structure of mosquitoes, and shall +then give the life-history of a common species, following this by a +brief consideration of some of the more striking departures from what +have been supposed to be the typical condition.</p> + +<p>The <b>Culicidæ</b> are slender, nematocerous Diptera with narrow wings, +antennæ plumose in the males, and usually with the proboscis much +longer than the head, slender, firm and adapted for piercing in the<span class="pagenum"><a name="Page_99" id="Page_99">[Pg 99]</a></span> +female. The most characteristic feature is that the margins of the +wings and, in most cases, the wing veins possess a fringe of scale-like +hairs. These may also cover in part, or entirely, the head, thorax, +abdomen and legs. The females, only, suck blood.</p> + +<p>On account of the importance of the group in this country and the +desirability of the student being able to determine material in various +stages, we show in the accompanying figures the characters most +used in classification.</p> + +<p>The larvæ (<a href="#Fig_73">fig. 73</a>) are elongate, +with the head and thorax sharply +distinct. The larval antennæ are +prominent, consisting of a single +cylindrical and sometimes curved +segment. The outer third is often +narrower and bears at its base a +fan-shaped tuft of hairs, the arrangement +and abundance of which +is of systematic importance. About +the mouth are the so-called rotary +mouth brushes, dense masses of +long hairs borne by the labrum +and having the function of sweeping +food into the mouth. The +form and arrangement of thoracic, +abdominal, and anal tufts of hair +vary in different species and present +characteristics of value. On either +side of the eighth abdominal segment +is a patch of scales varying +greatly in arrangement and number and of much value in separating +species. Respiration is by means of tracheæ which open at the apex +of the so-called anal siphon, when it is present. In addition, there +are also one or two pairs of tracheal gills which vary much in appearance +in different species. On the ventral side of the anal siphon is a +double row of flattened, toothed spines whose number and shape are +likewise of some value in separating species. They constitute the +comb or pecten.</p> + +<div class="figright" style="width: 300px;"><a name="Fig_73" id="Fig_73"></a> +<a href="images/f073-full.png"><img src="images/f073.png" width="300" height="490" alt="73. Culex larva showing details of external +structure." title="73. Culex larva showing details of external +structure." /></a> +<span class="caption">73. Culex larva showing details of external +structure.</span> +</div> + +<p>The pupa (<a href="#Fig_139">fig. 139, b</a>) unlike that of most insects, is active, though it +takes no food. The head and thorax are not distinctly separated, but +the slender flexible abdomen in sharply marked off. The antennæ,<span class="pagenum"><a name="Page_100" id="Page_100">[Pg 100]</a></span> +mouth-parts, legs, and wings of the future adult are now external, but +enclosed in chitinous cases. On the upper surface, near the base of +the wings are two trumpets, or breathing tubes, for the pupal spiracles +are towards the anterior end instead of at the caudal end, as in the +larva. At the tip of the abdomen is a pair of large chitinous swimming +paddles.</p> + +<p>As illustrative of the life cycle of a mosquito we shall discuss the +development of a common house mosquito, <i>Culex pipiens</i>, often +referred to in the Northern United States as the rain barrel mosquito. +Its life cycle is often given as typical for the entire group, but, as we +have already emphasized, no one species can serve this purpose.</p> + +<p>The adults of <i>Culex pipiens</i> hibernate throughout the winter in +cellars, buildings, hollow trees, or similar dark shelters. Early in +the spring they emerge and deposit their eggs in a raft-like mass. +The number of eggs in a single mass is in the neighborhood of two +hundred, recorded counts varying considerably. A single female +may deposit several masses during her life time. The duration of +the egg stage is dependent upon temperature. In the warm summer +time the larvæ may emerge within a day. The larvæ undergo four +molts and under optimum conditions may transform into pupæ in +about a week's time. Under the same favorable conditions, the pupal +stage may be completed in a day's time. The total life cycle of <i>Culex +pipiens</i>, under optimum conditions, may thus be completed in a week +to ten days. This period may be considerably extended under less +favorable conditions of temperature and food supply.</p> + +<p><i>Culex pipiens</i> breeds continuously throughout the summer, +developing in rain barrels, horse troughs, tin cans, or indeed in any +standing water about houses, which lasts for a week or more. The +catch basins of sewers furnish an abundant supply of the pests under +some conditions. Such places, the tin gutters on residences, and all +possible breeding places must be considered in attempts to exterminate +this species.</p> + +<p>Other species of mosquitoes may exhibit radical departures from +<i>Culex pipiens</i> in life-history and habits. To control them it is essential +that the biological details be thoroughly worked out for, as +Howard, Dyar, and Knab have emphasized, "much useless labor and +expense can be avoided by an accurate knowledge of the habits of the +species." For a critical discussion of the known facts the reader is +referred to their monograph. We shall confine ourselves to a few +illustrations.<span class="pagenum"><a name="Page_101" id="Page_101">[Pg 101]</a></span></p> + +<p>The majority of mosquitoes in temperate climates hibernate in +the egg stage, hatching in the spring or even mild winter days in water +from melting snow. It is such single-brooded species which appear in +astounding numbers in the far North. Similarly, in dry regions the +eggs may stand thorough dessication, and yet hatch out with great +promptness when submerged by the rains. "Another provision to +insure the species against destruction in such a case, exists in the fact +* * * that not all the eggs hatch, a part of them lying over until +again submerged by subsequent rains." In temperate North +America, a few species pass the winter in the larval state. An interesting +illustration of this is afforded by <i>Wyeomia smithii</i>, whose +larvæ live in pitcher plants and are to be found on the coldest winter +days imbedded in the solid ice. Late in the spring, the adults emerge +and produce several broods during the summer.</p> + +<p>In the United States, one of the most important facts which has +been brought out by the intensive studies of recent years is that certain +species are migratory and that they can travel long distances and +become an intolerable pest many miles from their breeding places. +This was forcibly emphasized in Dr. Smith's work in New Jersey, +when he found that migratory mosquitoes, developing in the salt +marshes along the coast, are the dominant species largely responsible +for the fame of the New Jersey mosquito. The species concerned are +<i>Aedes sollicitans</i>, <i>A. cantator</i> and <i>A. tæniorhynchus</i>. Dr. Smith +decided that the first of these might migrate at least forty miles +inland. It is obvious that where such species are the dominant pest, +local control measures are a useless waste of time and money. Such +migratory habits are rare, however, and it is probable that the +majority of mosquitoes do not fly any great distance from their +breeding places.</p> + +<p>While mosquitoes are thought of primarily as a pest of man, there +are many species which have never been known to feed upon human +or mammalian blood, no matter how favorable the opportunity. +According to Howard, Dyar, and Knab, this is true of <i>Culex territans</i>, +one of the common mosquitoes in the summer months in the Northern +United States. There are some species, probably many, in which +the females, like the males, are plant feeders. In experimental work, +both sexes are often kept alive for long periods by feeding them upon +ripe banana, dried fig, raisins, and the like, and in spite of sweeping +assertions that mosquitoes must have a meal of blood in order to +stimulate the ovaries to development, some of the common blood-sucking<span class="pagenum"><a name="Page_102" id="Page_102">[Pg 102]</a></span> +species, notably <i>Culex pipiens</i>, have been bred repeatedly +without opportunity to feed upon blood.</p> + +<p>The effect of the bite varies greatly with different species and +depends upon the susceptibility of the individual bitten. Some +persons are driven almost frantic by the attacks of the pests when +their companions seem almost unconscious of any inconvenience. +Usually, irritation and some degree of inflammation appear shortly +following the bite. Not infrequently a hardened wheal or even a +nodule forms, and sometimes scratching leads to secondary infection +and serious results.</p> + +<p>The source of the poison is usually supposed to be the salivary +glands of the insect. As we have already pointed out, (<a href="#Page_34">p. 34</a>), +Macloskie believed that one lobe of the gland, on each side, was +specialized for forming the poison, while a radically different view is +that of Schaudinn, who believed that the irritation is due to the +expelled contents of the œsophageal diverticula, which contain a +gas and a peculiar type of fungi or bacteria. In numerous attempts, +Schaudinn was unable to produce any irritation by applying the +triturated salivary glands to a wound, but obtained the typical result +when he used the isolated diverticula.</p> + +<p>The irritation of the bite may be relieved to some extent by using +ammonia water, a one per cent. alcoholic solution of menthol, or +preparations of cresol, or carbolic acid. Dr. Howard recommends +rubbing the bite gently with a piece of moist toilet soap. Castellani +and Chalmers recommend cleansing inflamed bites with one in forty +carbolic lotion, followed by dressing with boracic ointment. Of +course, scratching should be avoided as much as possible.</p> + +<p>Repellents of various kinds are used, for warding off the attacks +of the insects. We have often used a mixture of equal parts of oil +of pennyroyal and kerosene, applied to the hands and face. Oil of +citronella is much used and is less objectionable to some persons. A +recommended formula is, oil of citronella one ounce, spirits of camphor +one ounce, oil of cedar one-half ounce. A last resort would seem to +be the following mixture recommended by Howard, Dyar, and Knab +for use by hunters and fishermen in badly infested regions, against +mosquitoes and blackflies.</p> + +<p>Take 2¼ lbs. of mutton tallow and strain it. While still hot add +½ lb. black tar (Canadian tar). Stir thoroughly and pour into the +receptacle in which it is to be contained. When nearly cool stir in +three ounces of oil of citronella and 1¼ oz. of pennyroyal.<span class="pagenum"><a name="Page_103" id="Page_103">[Pg 103]</a></span></p> + +<p>At night the surest protection is a good bed net. There are many +types of these in use, but in order to be serviceable and at the same +time comfortable it should be roomy and hung in such a way as to +be stretched tightly in every direction. We prefer one suspended +from a broad, square frame, supported by a right-angled standard +which is fastened to the head of the bed. It must be absolutely free +from rents or holes and tucked in securely under the mattress or it +will serve merely as a convenient cage to retain mosquitoes which gain +an entrance. While such nets are a convenience in any mosquito +ridden community, they are essential in regions where disease-carrying +species abound. Screening of doors, windows and porches, against +the pests is so commonly practiced in this country that its importance +and convenience need hardly be urged.</p> + +<p>Destruction of mosquitoes and prevention of breeding are of +fundamental importance. Such measures demand first, as we have +seen, the correct determination of the species which is to be dealt +with, and a knowledge of its life-history and habits. If it prove to be +one of the migratory forms, it is beyond mere local effort and becomes +a problem demanding careful organization and state control. An +excellent illustration of the importance and effectiveness of work +along these lines is afforded by that in New Jersey, begun by the late +Dr. John B. Smith and being pushed with vigor by his successor, +Dr. Headlee.</p> + +<p>In any case, there is necessity for community action. Even near +the coast, where the migratory species are dominant, there are the +local species which demand attention and which cannot be reached +by any measures directed against the species of the salt marshes. The +most important of local measures consist in the destruction of breeding +places by filling or draining ponds and pools, clearing up of more +temporary breeding places, such as cans, pails, water barrels and the +like. Under conditions where complete drainage of swamps is impracticable +or undesirable, judicious dredging may result in a pool or +series of steep-sided pools deep enough to maintain a supply of fish, +which will keep down the mosquito larvæ. Where water receptacles +are needed for storage of rain water, they should be protected by +careful screening or a film of kerosene over the top of the water, +renewed every two weeks or so, so as to prevent mosquitoes from +depositing their eggs. When kerosene is used, Water drawn from the +bottom of the receptacle will not be contaminated by it to any injurious +extent. Where ponds cannot be drained much good will be<span class="pagenum"><a name="Page_104" id="Page_104">[Pg 104]</a></span> +accomplished by spraying kerosene oil on the surface of the water, or +by the introduction of fish which will feed on the larvæ.</p> + +<div class="figleft" style="width: 350px;"><a name="Fig_74" id="Fig_74"></a> +<a href="images/f074-full.jpg"><img src="images/f074.jpg" width="350" height="244" alt="74. Mouth parts of Simulium. After +Grünberg." title="74. Mouth parts of Simulium. After +Grünberg." /></a> +<span class="caption">74. Mouth parts of Simulium. After +Grünberg.</span> +</div> + +<p>Detailed consideration of the +most efficient measures for controlling +mosquitoes is to be found +in Dr. Howard's Bulletin No. 88 +of the Bureau of Entomology, +"Preventive and remedial work +against mosquitoes" or, in more +summarized form, in Farmers' +Bulletin No. 444. One of these +should be obtained by any person +interested in the problems of mosquito +control and public health.</p> + + +<h3><a name="The_Simuliidae_or_Black_Flies" id="The_Simuliidae_or_Black_Flies"></a>The Simuliidæ, or Black Flies</h3> + +<div class="figright" style="width: 150px;"><a name="Fig_75" id="Fig_75"></a> +<a href="images/f075-full.png"><img src="images/f075.png" width="150" height="502" alt="75. Larva of Simulium, +(×8). +After Garman." title="75. Larva of Simulium, +(×8). +After Garman." /></a> +<span class="caption">75. Larva of Simulium, +(×8). +After Garman.</span> +</div> + +<p>The <b>Simuliidæ</b>, or black flies, are small, dark, or black flies, with +a stout body and a hump-back appearance. The +antennæ are short but eleven-segmented, the wings +broad, without scales or hairs, and with the anterior +veins stout but the others very weak. The mouth-parts +(<a href="#Fig_74">fig. 74</a>) are fitted for biting.</p> + +<p>The larvæ of the Simuliidæ (<a href="#Fig_75">fig. 75</a>) are aquatic +and, unlike those of mosquitoes, require a well ærated, +or swiftly running water. Here they attach to stones, +logs, or vegetation and feed upon various micro-organisms. +They pupate in silken cocoons open at +the top. Detailed life-histories have not been worked +out for most of the species. We shall consider as +typical that of <i>Simulium pictipes</i>, an inoffensive +species widely distributed in the Eastern United +States, which has been studied especially at Ithaca, +N.Y. (Johannsen, 1903).</p> + +<p>The eggs are deposited in a compact yellowish layer +on the surface of rock, on the brinks of falls and +rapids where the water is flowing swiftly. They are +elongate ellipsoidal in shape, about .4 by .18 mm. +As myriads of females deposit in the same place the +egg patches may be conspicuous coatings of a foot or much more +in diameter. When first laid they are enveloped in a yellowish<span class="pagenum"><a name="Page_105" id="Page_105">[Pg 105]</a></span> +white slime, which becomes darker, until finally it becomes black just +before the emerging of the larvæ. The egg stage lasts a week.</p> + +<p>The larvæ (<a href="#Fig_75">fig. 75</a>) are black, soft skinned, somewhat cylindrical +in shape, enlarged at both ends and attenuated in the middle. The +posterior half is much stouter than the anterior part and almost club-shaped. +The head bears two large fan-shaped organs which aid in +procuring food. Respiration is accomplished by means of three so-called +blood gills which are pushed out from the dorsal part of the +rectum. The larvæ occur in enormous numbers, in moss-like patches. +If removed from their natural habitat and placed in quiet water they +die within three or four hours. Fastened to the rock by means of a +disk-like sucker at the caudal end of +the body, they ordinarily assume an +erect position. They move about on +the surface of the rocks, to a limited +extent, with a looping gait similar to +that of a measuring worm, and a web +is secreted which prevents their being +washed away by the swiftly flowing +water. They feed chiefly upon algæ +and diatoms.</p> + +<p>The complete larval stage during +the summer months occupies about +four weeks, varying somewhat with the +temperature and velocity of the water. +At the end of this period they spin +from cephalic glands, boot-shaped +silken cocoons within which they pupate. The cocoon when spun +is firmly attached to the rock and also to adjacent cocoons. +Clustered continuously over a large area and sometimes one above +another, they form a compact, carpet-like covering on the rocks, +the reddish-brown color of which is easily distinguishable from the +jet-black appearance of the larvæ. The pupal stage lasts about +three weeks. The adult fly, surrounded by a bubble of air, quickly +rises to the surface of the water and escapes. The adults (<a href="#Fig_76">fig. 76</a>) +are apparently short lived and thus the entire life cycle, from egg +to egg is completed in approximately eight weeks.</p> + +<div class="figright" style="width: 350px;"><a name="Fig_76" id="Fig_76"></a> +<a href="images/f076-full.png"><img src="images/f076.png" width="350" height="450" alt="76. Simulium venustum, (×8). +After Garman." title="76. Simulium venustum, (×8). +After Garman." /></a> +<span class="caption">76. Simulium venustum, (×8). +After Garman.</span> +</div> + +<p>In the case of <i>Simulium pictipes</i> at Ithaca, N. Y., the first brood +of adults emerges early in May and successive generations are produced +throughout the summer and early autumn. This species winters in<span class="pagenum"><a name="Page_106" id="Page_106">[Pg 106]</a></span> +the larval condition. Most of the other species of <i>Simulium</i> which +have been studied seem to be single brooded.</p> + +<p>While <i>Simulium pictipes</i> does not attack man, there are a number +of the species which are blood-sucking and in some regions they are a +veritable scourge. In recent years the greatest interest in the group +has been aroused by Sambon's hypothesis that they transmit pellagra +from man to man. This has not been established, and, indeed, seems +very doubtful, but the importance of these insects as pests and the +possibility that they may carry disease make it urgent that detailed +life-histories of the hominoxious species be worked out.</p> + +<p>As pests a vivid account of their attacks is in Agassiz's "Lake +Superior" (p. 61), quoted by Forbes (1912).</p> + +<p>"Neither the love of the picturesque, however, nor the interests of +science, could tempt us into the woods, so terrible were the black flies. +This pest of flies which all the way hither had confined our ramblings +on shore pretty closely to the rocks and the beach, and had been +growing constantly worse, here reached its climax. Although detained +nearly two days, * * * we could only sit with folded +hands, or employ ourselves in arranging specimens, and such other +operations as could be pursued in camp, and under the protection of +a 'smudge.' One, whom scientific ardor tempted a little way up the +river in a canoe, after water plants, came back a frightful spectacle, +with blood-red rings round his eyes, his face bloody, and covered with +punctures. The next morning his head and neck were swollen as if +from an attack of erysipelas."</p> + +<p>There are even well authenticated accounts on record of death of +humans from the attacks of large swarms of these gnats. In some +regions, and especially in the Mississippi Valley in this country, certain +species of black flies have been the cause of enormous losses to +farmers and stockmen, through their attacks on poultry and domestic +animals. C. V. Riley states that in 1874 the loss occasioned in one +county in Tennessee was estimated at $500,000.</p> + +<p>The measures of prevention and protection against these insects +have been well summarized by Forbes (1912). They are of two kinds: +"the use of repellents intended to drive away the winged flies, and +measures for the local destruction of the aquatic larvæ. The repellents +used are either smudges, or surface applications made to keep +the flies from biting. The black-fly will not endure a dense smoke, +and the well-known mosquito smudge seems to be ordinarily sufficient +for the protection of man. In the South, leather, cloth, and other<span class="pagenum"><a name="Page_107" id="Page_107">[Pg 107]</a></span> +materials which will make the densest and most stifling smoke, are +often preserved for this use in the spring. Smudges are built in +pastures for the protection of stock, and are kept burning before the +doors of barns and stables. As the black-flies do not readily enter a +dark room, light is excluded from stables as much as possible during +the gnat season. If teams must be used in the open field while gnats +are abroad, they may be protected against the attacks of the gnats by +applying cotton-seed oil or axle grease to the surface, especially to the +less hairy parts of the animals, at least twice a day. A mixture of oil +and tar and, indeed, several other preventives, are of practical use in +badly infested regions; but no definite test or exact comparison has +been made with any them in a way to give a record of the precise +results."</p> + +<p>"It is easy to drive the flies from houses or tents by burning +pyrethrum powder inside; this either kills the flies or stupifies them +so that they do not bite for some time thereafter." * * * "Oil of +tar is commonly applied to the exposed parts of the body for the purpose +of repelling the insects, and this preparation is supplied by the +Hudson Bay Company to its employees. Minnesota fishermen +frequently grease their faces and hands with a mixture of kerosene +and mutton tallow for the same purpose." We have found a mixture +of equal parts of kerosene and oil of pennyroyal efficient.</p> + +<p>Under most circumstances very little can be done to destroy this +insect in its early stage, but occasionally conditions are such that a +larvicide can be used effectively. Weed (1904), and Sanderson (1910) +both report excellent results from the use of phinotas oil, a proprietary +compound. The first-mentioned also found that in some places the +larvæ could be removed by sweeping them loose in masses with stiff +stable brooms and then catching them downstream on wire netting +stretched in the water.</p> + + +<h3><a name="Chironomidae_or_Midges" id="Chironomidae_or_Midges"></a>Chironomidæ or Midges</h3> + +<p>The flies of this family, commonly known as midges, resemble +mosquitoes in form and size but are usually more delicate, and the +wing-veins, though sometimes hairy, are not fringed with scales. +The venation is simpler than in the mosquitoes and the veins are +usually less distinct.</p> + +<p>These midges, especially in spring or autumn, are often seen in +immense swarms arising like smoke over swamps and producing a +humming noise which can be heard for a considerable distance. At<span class="pagenum"><a name="Page_108" id="Page_108">[Pg 108]</a></span> +these seasons they are frequently to be found upon the windows of +dwellings, where they are often mistaken for mosquitoes.</p> + +<p>The larvæ are worm-like, but vary somewhat in form in the different +genera. Most of them are aquatic, but a few live in the earth, in +manure, decaying wood, under bark, or in the sap of trees, especially +in the sap which collects in wounds.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_77" id="Fig_77"></a> +<a href="images/f077-full.png"><img src="images/f077.png" width="500" height="348" alt="77. Culicoides guttipennis; (a) adult, (×15); (b) head of same; (c) larva; +(d) head; (e) pupa. After Pratt." title="77. Culicoides guttipennis; (a) adult, (×15); (b) head of same; (c) larva; +(d) head; (e) pupa. After Pratt." /></a> +<span class="caption">77. Culicoides guttipennis; (<i>a</i>) adult, (×15); (<i>b</i>) head of same; (<i>c</i>) larva; +(<i>d</i>) head; (<i>e</i>) pupa. After Pratt.</span> +</div> + +<p>Of the many species of <b>Chironomidæ</b>, (over eight hundred known), +the vast majority are inoffensive. The sub-family Ceratopogoninæ, +however, forms an exception, for some of the members of this group, +known as sandflies, or punkies, suck blood and are particularly troublesome +in the mountains, along streams, and at the seashore. Most of +these have been classed under the genus <i>Ceratopogon</i>, but the group +has been broken up into a number of genera and <i>Ceratopogon</i>, in the +strict sense, is not known to contain any species which sucks the blood +of vertebrates.</p> + +<p><b>The Ceratopogoninæ</b>—The Ceratopogoninæ are among the smallest +of the Diptera, many of them being hardly a millimeter long and some +not even so large. They are Chironomidæ in which the thorax is not +prolonged over the head. The antennæ are filiform with fourteen +(rarely thirteen) segments in both sexes, those of the male being brush-like. +The basal segment is enlarged, the last segment never longer<span class="pagenum"><a name="Page_109" id="Page_109">[Pg 109]</a></span> +than the two preceding combined, while the last five are sub-equal to, +or longer than the preceding segment. The legs are relatively stouter +than in the other Chironomidæ. The following three genera of this +subfamily are best known as blood suckers in this country.</p> + +<div class="figright" style="width: 300px;"><a name="Fig_78" id="Fig_78"></a> +<a href="images/f078-full.png"><img src="images/f078.png" width="300" height="352" alt="78. Culicoides guttipennis; mouth +parts of adult. After Pratt." title="78. Culicoides guttipennis; mouth +parts of adult. After Pratt." /></a> +<span class="caption">78. Culicoides guttipennis; mouth +parts of adult. After Pratt.</span> +</div> + +<p>Of the genus <i>Culicoides</i> there are many species occurring in various +parts of the world. A number are known to bite man and animals and +it is probable that all are capable of inflicting injury. In some +localities they are called punkies, in others, sand-flies, a name sometimes +also applied to the species of <i>Simulium</i> and <i>Phlebotomus</i>. +Owing to their very small size they are known by some tribes of +Indians as No-see-ums. The larvæ are found in ponds, pools, water +standing in hollow tree stumps, and the like. Though probably living +chiefly in fresh water, we have found a species occurring in salt water. +The larvæ are small, slender, legless, +worm-like creatures (<a href="#Fig_77">fig. 77<i>c</i></a>) with +small brown head and twelve body +segments. The pupæ (<a href="#Fig_77">fig. 77<i>e</i></a>) are +slender, more swollen at the anterior +end and terminating in a forked process. +They float nearly motionless in +a vertical position, the respiratory +tubes in contact with the surface film. +The adults are all small, rarely exceeding +2¼ mm. in length. The wings +are more or less covered with erect +setulæ or hairs and in many species +variously spotted and marked with +iridescent blotches. The antennæ have fourteen segments, the palpi +usually five. The wing venation and mouth-parts are shown in +figures <a href="#Fig_77">77</a> and <a href="#Fig_78">78</a>. Of the twenty or more species of this genus +occurring in the United States the following are known to bite: <i>C. cinctus</i>, +<i>C. guttipennis</i>, <i>C. sanguisuga</i>, <i>C. stellifer</i>, <i>C. variipennis</i>, +<i>C. unicolor</i>.</p> + +<p>One of the most widely distributed and commonest species in the +Eastern States is <i>C. guttipennis</i> (<a href="#Fig_77">fig. 77a</a>). It is black with brown +legs, a whitish ring before the apex of each femur and both ends of +each tibia; tarsi yellow, knobs of halteres yellow. Mesonotum +opaque, brown, two vittæ in the middle, enlarging into a large spot +on the posterior half, also a curved row of three spots in front of each +wing, and the narrow lateral margins, light gray pruinose. Wings<span class="pagenum"><a name="Page_110" id="Page_110">[Pg 110]</a></span> +nearly wholly covered with brown hairs, gray, with markings as +shown in the figure. Length one mm.</p> + +<p><i>Johannseniella</i> Will. is a +wide-spread genus related +to the foregoing. Its +mouth-parts are well +adapted for piercing and +it is said to be a persistent +blood sucker, particularly +in Greenland. This genus +is distinguished from <i>Culicoides</i> +by its bare wings, +the venation (<a href="#Fig_163">fig. 163</a>, c), +and the longer tarsal claws. +There are over twenty +North American species.</p> + +<div class="figcenter" style="width: 320px;"> +<a href="images/f079-full.png"><img src="images/f079.png" width="320" height="254" alt="79. Chrysops univittatus, (×4). After Osborn." title="79. Chrysops univittatus, (×4). After Osborn." /></a> +<span class="caption">79. Chrysops univittatus, (×4). After Osborn.</span> +</div> + +<p>In the Southwestern United States, <i>Tersesthes torrens</i> Towns. +occurs, a little gnat which annoys horses, and perhaps man also, by +its bite. It is related to <i>Culicoides</i> but differs in the number of +antennal segments and in its wing venation (<a href="#Fig_163">fig. 163</a>, e). The fly +measures but two mm. in length and is blackish in color. The +antennæ of the female have thirteen segments, the palpi but three, of +which the second is enlarged and swollen.</p> + + +<h3><a name="Tabanidae_or_Horse-Flies" id="Tabanidae_or_Horse-Flies"></a>Tabanidæ or Horse-Flies</h3> + +<div class="figleft" style="width: 250px;"><a name="Fig_80a" id="Fig_80a"></a> +<a href="images/f080a-full.jpg"><img src="images/f080a.png" width="250" height="806" alt="80. (a) Eggs of Tabanus. Photograph +by J. T. Lloyd." title="80. (a) Eggs of Tabanus. Photograph +by J. T. Lloyd." /></a> +<span class="caption">80. (<i>a</i>) Eggs of Tabanus. Photograph +by J. T. Lloyd.</span> +</div> + +<div class="figright" style="width: 250px;"><a name="Fig_80b" id="Fig_80b"></a> +<a href="images/f080b-full.jpg"><img src="images/f080b.png" width="250" height="601" alt="80. (b) Larva of Tabanus. +Photograph by M. V. S." title="80. (b) Larva of Tabanus. +Photograph by M. V. S." /></a> +<span class="caption">80. (<i>b</i>) Larva of Tabanus. +Photograph by M. V. S.</span> +</div> + +<p>The <b>Tabanidæ</b>,—horse-flies, ear-flies, and deer-flies,—are well-known +pests of cattle and horses and are often extremely annoying +to man. The characteristics of the family and of the principal North +American genera are given in the keys of Chapter XII. There are +over 2500 recorded species. As in the mosquitoes, the females +alone are blood suckers. The males are flower feeders or live on +plant juices. This is apparently true also of the females of some of +the genera.</p> + +<p>The eggs are deposited in masses on water plants or grasses and +sedges growing in marshy or wet ground. Those of a common +species of <i>Tabanus</i> are illustrated in <a href="#Fig_80a">figure 80, <i>a</i></a>. They are placed +in masses of several hundred, light colored when first deposited but +turning black. In a week or so the cylindrical larvæ, tapering at +both ends (<a href="#Fig_80b">fig. 80, <i>b</i></a>), escape to the water, or damp earth, and lead<span class="pagenum"><a name="Page_111" id="Page_111">[Pg 111]</a></span> +an active, carnivorous life, feeding mainly on insect larvæ, and worms. +In the forms which have been best studied the larval life is a long +one, lasting for months or even for more +than a year. Until recently, little was +known concerning the life-histories of this +group, but the studies of Hart (1895), +and Hine (1903 +) +have added +greatly to the +knowledge concerning +North +American +forms.</p> + +<p>Many of the +species attack +man with avidity +and are able +to inflict painful +bites, which +may smart for +hours. In some +instances the +wound is so +considerable +that blood will +continue to flow +after the fly has +left. We have +seen several +cases of secondary infection following +such bites.</p> + +<p>The horse-flies have been definitely +convicted of transferring the trypanosome +of surra from diseased to healthy animals +and there is good evidence that they transfer anthrax. The possibility +of their being important agents in the conveyal of human +diseases should not be overlooked. Indeed, Leiper has recently +determined that a species of <i>Chrysops</i> transfers the blood parasite +<i>Filaria diurna</i>.<span class="pagenum"><a name="Page_112" id="Page_112">[Pg 112]</a></span></p> + + +<h3><a name="Leptidae_or_Snipe-Flies" id="Leptidae_or_Snipe-Flies"></a>Leptidæ or Snipe-Flies</h3> + +<p>The family <b>Leptidæ</b> is made up of moderate or large sized flies, +predaceous in habit. They are sufficiently characterized in the keys +of Chapter XII. Four blood-sucking +species belonging to three +genera have been reported. Of +these <i>Symphoromyia pachyceras</i> is +a western species. Dr. J. C. +Bradley, from personal experience, +reports it as a vicious biter.</p> + +<div class="figcenter" style="width: 400px;"> +<a href="images/f080c-full.png"><img src="images/f080c.png" width="400" height="346" alt="80. (c) Mouth parts of Tabanus. After Grünberg." title="80. (c) Mouth parts of Tabanus. After Grünberg." /></a> +<span class="caption">80. (<i>c</i>) Mouth parts of Tabanus. After Grünberg.</span> +</div> + + +<h3><a name="Oestridae_or_Bot-flies" id="Oestridae_or_Bot-flies"></a>Oestridæ or Bot-flies</h3> + +<p>To the family <b>Oestridæ</b> belong +the bot and warble-flies so frequently +injurious to animals. +The adults are large, or of +medium size, heavy bodied, rather +hairy, and usually resemble bees in appearance.</p> + +<p>The larvæ live parasitically in various parts of the body of mammals, +such as the stomach (horse bot-fly), the subcutaneous connective +tissue (warble-fly of cattle), or the nasal passage (sheep bot-fly +or head maggot).</p> + +<p>There are on record many cases of the occurrence of the larvæ +of Oestridæ as occasional parasites of man. A number of these have +been collected and reviewed in a thesis by Mme. Pètrovskaia (1910). +The majority of them relate to the following species.</p> + +<p><i>Gastrophilus hæmorrhoidalis</i>, the red tailed bot-fly, is one of the +species whose larvæ are most commonly found in the stomach of the +horse. Schoch (1877) cites the case of a woman who suffered from +a severe case of chronic catarrh of the stomach, and who vomited, +and also passed from the anus, larvæ which apparently belonged +to this species. Such cases are exceedingly rare but instances of +subcutaneous infestation are fairly numerous. In the latter type +these larvæ are sometimes the cause of the peculiar "creeping myasis." +This is characterized at its beginning by a very painful swelling +which gradually migrates, producing a narrow raised line four to +twenty-five millimeters broad. When the larva is mature, sometimes +after several months, it becomes stationary and a tumor is formed +which opens and discharges the larva along with pus and serum.<span class="pagenum"><a name="Page_113" id="Page_113">[Pg 113]</a></span></p> + +<p><i>Gastrophilus equi</i> is the most widespread and common of the horse +bot-flies. Portschinsky reports it as commonly causing subcutaneous +myasis of man in Russia.</p> + +<p><i>Hypoderma bovis</i> (= <i>Oestrus bovis</i>), and <i>Hypoderma lineata</i> are +the so-called warble-flies of cattle. The latter species is the more +common in North America but Dr. C. G. Hewitt has recently shown +that <i>H. bovis</i> also occurs. Though warbles are very common in +cattle in this country, the adult flies are very rarely seen. They +are about half an inch in length, very hairy, dark, and closely resemble +common honey-bees.</p> + +<p>They deposit their eggs on the hairs of cattle and the animals in +licking themselves take in the young larvæ. These pass out through +the walls of the œsophagus and migrate through the tissues of the +animal, to finally settle down in the subcutaneous tissue of the back. +The possibility of their entering directly through the skin, especially +in case of infestation of man, is not absolutely precluded, although +it is doubtful.</p> + +<p>For both species of <i>Hypoderma</i> there are numerous cases on +record of their occurrence in man. Hamilton (1893) saw a boy, +six years of age, who had been suffering for some months from the +glands on one side of his neck being swollen and from a fetid ulceration +around the back teeth of the lower jaw of the same side. Three +months' treatment was of no avail and the end seemed near; one day +a white object, which was seen to move, was observed in the ulcer +at the root of the tongue, and on being extracted was recognized as a +full grown larva of <i>Hypoderma</i>. It was of usual tawny color, about +half an inch long when contracted, about one third that thickness, +and quite lively. The case resulted fatally. The boy had been on a +dairy farm the previous fall, where probably the egg (or larva) was +in some way taken into his mouth, and the larva found between the +base of the tongue and the jaw suitable tissue in which to develop.</p> + +<p>Topsent (1901) reports a case of "creeping myasis" caused by +<i>H. lineata</i> in the skin of the neck and shoulders of a girl eight years +of age. The larva travelled a distance of nearly six and a half inches. +The little patient suffered excruciating pain in the place occupied by +the larva.</p> + +<p><i>Hypoderma diana</i> infests deer, and has been known to occur in +man.</p> + +<p><i>Oestris ovis</i>, the sheep bot-fly, or head maggot, is widely distributed +in all parts of the world. In mid-summer the flies deposit<span class="pagenum"><a name="Page_114" id="Page_114">[Pg 114]</a></span> +living maggots in the nostrils of sheep. These larvæ promptly pass +up the nasal passages into the frontal and maxillary sinuses, where +they feed on the mucous to be found there. In their migrations +they cause great irritation to their host, and when present in numbers +may cause vertigo, paroxysms, and even death. Portschinsky in an +important monograph on this species, has discussed in detail its +relation to man. He shows that it is not uncommon for the fly to +attack man and that the minute living larvæ are deposited in the +eyes, nostrils, lips, or mouth. A typical case in which the larvæ +were deposited in the eye was described by a German oculist Kayser, +in 1905. A woman brought her six year old daughter to him and +said that the day before, about noontime, a flying insect struck the +eye of the child and that since then she had felt a pain which increased +towards evening. In the morning the pain ceased but the +eye was very red. She was examined at about noon, at which time +she was quiet and felt no pain. She was not sensitive to light, and +the only thing noticed was a slight congestion and accumulation of +secretion in the corner of the right eye. A careful examination of +the eye disclosed small, active, white larvæ that crawled out from +the folds of the conjunctiva and then back and disappeared. Five +of these larvæ were removed and although an uncomfortable feeling +persisted for a while, the eye became normal in about three weeks.</p> + +<p>Some of the other recorded cases have not resulted so favorably, +for the eyesight has been seriously affected or even lost.</p> + +<p>According to Edmund and Etienne Sergent (1907), myasis caused +by the larvæ of <i>Oestris ovis</i> is very common among the shepherds in +Algeria. The natives say that the fly deposits its larvæ quickly, +while on the wing, without pause. The greatest pain is caused when +these larvæ establish themselves in the nasal cavities. They then +produce severe frontal headaches, making sleep impossible. This +is accompanied by continuous secretion from the nasal cavities +and itching pains in the sinuses. If the larvæ happen to get into +the mouth, the throat becomes inflamed, swallowing is painful, +and sometimes vomiting results. The diseased condition may last +for from three to ten days or in the case of nasal infection, longer, +but recovery always follows. The natives remove the larvæ from +the eye mechanically by means of a small rag. When the nose is +infested, tobacco fumigations are applied, and in case of throat +infestation gargles of pepper, onion, or garlic extracts are used.<span class="pagenum"><a name="Page_115" id="Page_115">[Pg 115]</a></span></p> + +<p><i>Rhinœstrus nasalis</i>, the Russian gad-fly, parasitizes the nasopharyngeal +region of the horse. According to Portschinsky, it not +infrequently attacks man +and then, in all the known +cases deposits its larvæ +in the eye, only. This +is generally done while +the person is quiet, but +not during sleep. The +fly strikes without stopping +and deposits its larva +instantaneously. Immediately +after, the victim +experiences lancinating +pains which without intermission +increase in +violence. There is an intense +conjunctivitis and +if the larvæ are not removed promptly the envelopes +of the eye are gradually destroyed and the organ +lost.</p> + +<div class="figcenter" style="width: 360px;"><a name="Fig_81" id="Fig_81"></a> +<a href="images/f081-full.png"><img src="images/f081.png" width="360" height="323" alt="81. Larvæ of Dermatobia cyaniventris. After Blanchard." title="81. Larvæ of Dermatobia cyaniventris. After Blanchard." /></a> +<span class="caption">81. Larvæ of Dermatobia cyaniventris. After Blanchard.</span> +</div> + +<div class="figcenter" style="width: 125px;"><a name="Fig_82" id="Fig_82"></a> +<a href="images/f082-full.png"><img src="images/f082.png" width="125" height="519" alt="82. Young larva of +Dermatobia cyaniventris. +After Surcouf." title="82. Young larva of +Dermatobia cyaniventris. +After Surcouf." /></a> +<span class="caption">82. Young larva of +Dermatobia cyaniventris. +After Surcouf.</span> +</div> + +<p><i>Dermatobia cyaniventris</i>—This fly (<a href="#Fig_83">fig. 83</a>) is widely +distributed throughout tropical America, and in its +larval stage is well known as a parasite of man. The +larvæ (figs. <a href="#Fig_81">81</a> and <a href="#Fig_82">82</a>) which are known as the "ver +macaque," "torcel," "ver moyocuil" or by several other +local names, enter the skin and give rise to a boil-like +swelling, open at the top, and comparable with the swelling +produced by the warble fly larvæ, in cattle. They +cause itching and occasional excruciating pain. When +mature, nearly an inch in length, they voluntarily +leave their host, drop to the ground and complete their +development. The adult female is about 12 mm. in +length. The face is yellow, the frons black with a +grayish bloom; antennæ yellow, the third segment +four times as long as the second, the arista pectinate. +The thorax is bluish black with grayish bloom; the +abdomen depressed, brilliant metallescent blue with +violet tinge. The legs are yellowish, the squamæ and +wings brownish.<span class="pagenum"><a name="Page_116" id="Page_116">[Pg 116]</a></span></p> + +<p>The different types of larvæ represented in <a href="#Fig_81">figure 81</a> were formerly +supposed to belong to different species but Blanchard regards them +as merely various stages +of the same species. It +is only very recently +that the early stage and +the method by which +man becomes infested +were made known.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_83" id="Fig_83"></a> +<a href="images/f083-full.png"><img src="images/f083.png" width="500" height="353" alt="83. Dermatobia cyaniventris (×1¾). After Graham-Smith." title="83. Dermatobia cyaniventris (×1¾). After Graham-Smith." /></a> +<span class="caption">83. Dermatobia cyaniventris (×1¾). After <span title="for Graham-Smith read Manson">Graham-Smith</span>.</span> +</div> + +<div class="figcenter" style="width: 280px;"><a name="Fig_84" id="Fig_84"></a> +<a href="images/f084-full.png"><img src="images/f084.png" width="280" height="323" alt="84. Mosquito carrying eggs of Dermatobia +cyaniventris. After Surcouf." title="84. Mosquito carrying eggs of Dermatobia +cyaniventris. After Surcouf." /></a> +<span class="caption">84. Mosquito carrying eggs of Dermatobia +cyaniventris. After Surcouf.</span> +</div> + +<p>About 1900, Blanchard +observed the presence +of packets of large-sized +eggs under the +abdomen of certain mosquitoes +from Central +America; and in 1910, +Dr. Moralès, of Costa Rica, declared that the Dermatobia deposited +its eggs directly under the abdomen of the mosquito and that they +were thus carried to vertebrates. +Dr. Nunez Tovar observed the +mosquito carriers of the eggs and +placing larvæ from this source on +animals, produced typical tumors +and reared the adult flies. It +remained for Surcouf (1913) to +work out the full details. He +found that the Dermatobia deposits +its eggs in packets covered +by a very viscid substance, on +leaves. These become attached +to mosquitoes of the species +<i>Janthinosoma lutzi</i> (<a href="#Fig_84">fig. 84</a>) which +walk over the leaves. The eggs +which adhere to the abdomen, +remain attached and are thus +transported. The embryo develops, +but the young larva (<a href="#Fig_82">fig. 82</a>) remains in the egg until it has +opportunity to drop upon a vertebrate fed upon by the mosquito.<span class="pagenum"><a name="Page_117" id="Page_117">[Pg 117]</a></span></p> + + +<h3><a name="Muscidae" id="Muscidae"></a>Muscidæ</h3> + +<p>The following <b>Muscidæ</b>, characterized elsewhere, deserve special +mention under our present grouping of parasitic species. Other +important species will be considered as facultative parasites.</p> + +<p><i>Stomoxys calcitrans</i>, the stable-fly, or the biting house-fly, +is often confused with <i>Musca domestica</i> and therefore +is discussed especially in our consideration of the latter +species as an accidental carrier of disease. Its possible +relation to the spread of infantile paralysis is also considered +later.</p> + +<div class="figcenter" style="width: 100px;"><a name="Fig_85" id="Fig_85"></a> +<a href="images/f085-full.png"><img src="images/f085.png" width="100" height="320" alt="85. Larva of +Auchmeromyia +luteola. After +Graham-Smith." title="85. Larva of +Auchmeromyia +luteola. After +Graham-Smith." /></a> +<span class="caption">85. Larva of +Auchmeromyia +luteola. After +Graham-Smith.</span> +</div> + +<p>The <i>tsetse flies</i>, belonging to the genus <i>Glossina</i>, are +African species of blood-sucking Muscidæ which have +attracted much attention because of their rôle in transmitting +various trypanosome diseases of man and animals. +They are characterized in Chapter XII and are also +discussed in connection with the diseases which they +convey.</p> + +<p><a name="Chrysomyia_macellaria" id="Chrysomyia_macellaria"></a><i>Chrysomyia macellaria</i>, (= <i>Compsomyia</i>), the "screw worm"-fly +is one of the most important species of flies directly affecting man, +in North America. It is not normally parasitic, however, and hence +will be considered with other facultative parasites in Chapter IV.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_86" id="Fig_86"></a> +<a href="images/f086-full.png"><img src="images/f086.png" width="500" height="416" alt="86. Auchmeromyia luteola (×4). After Graham-Smith." title="86. Auchmeromyia luteola (×4). After Graham-Smith." /></a> +<span class="caption">86. Auchmeromyia luteola (×4). After Graham-Smith.</span> +</div> + +<p><i>Auchmeromyia luteola</i>, +the Congo floor +maggot. This is a +muscid of grewsome +habits, which has a wide +distribution throughout +Africa. The fly (<a href="#Fig_86">fig. 86</a>) +deposits its eggs on the +ground of the huts of the +natives. The whitish +larvæ (<a href="#Fig_85">fig. 85</a>) on hatching +are slightly flattened +ventrally, and +each segment bears +posteriorly three foot-pads +transversely arranged. At night the larvæ find their way into the +low beds or couches of the natives and suck their blood. The adult +flies do not bite man and, as far as known, the larvæ do not play any +rôle in the transmission of sleeping sickness or other diseases.<span class="pagenum"><a name="Page_118" id="Page_118">[Pg 118]</a></span></p> + +<div class="figleft" style="width: 350px;"><a name="Fig_87" id="Fig_87"></a> +<a href="images/f087-full.png"><img src="images/f087.png" width="350" height="327" alt="87. Cordylobia anthropophaga (×3). +After Fülleborn." title="87. Cordylobia anthropophaga (×3). +After Fülleborn." /></a> +<span class="caption">87. Cordylobia anthropophaga (×3). +After Fülleborn.</span> +</div> + +<div class="figright" style="width: 300px;"><a name="Fig_88" id="Fig_88"></a> +<a href="images/f088-full.png"><img src="images/f088.png" width="300" height="409" alt="88. Larva of Cordylobia anthropophaga. +After Blanchard." title="88. Larva of Cordylobia anthropophaga. +After Blanchard." /></a> +<span class="caption">88. Larva of Cordylobia anthropophaga. +After Blanchard.</span> +</div> + +<p>This habit of blood-sucking by muscid larvæ is usually referred +to as peculiar to <i>Auchmeromyia luteola</i> but it should be noted that the +larvæ of <i>Protocalliphora</i> frequent the +nests of birds and feed upon the +young. Mr. A. F. Coutant has studied +especially the life-history and habits +of <i>P. azurea</i>, whose larvæ he found +attacking young crows at Ithaca, N.Y. +He was unable to induce the larvæ to +feed on man.</p> + +<p><i>Cordylobia anthropophaga</i>, (<i>Ochromyia +anthropophaga</i>), or Tumbu-fly +(<a href="#Fig_87">fig. 87</a>) is an African species whose +larvæ affect man much as do those of +<i>Dermatobia cyniventris</i>, of Central and +South America. The larva (<a href="#Fig_88">fig. 88</a>), which is known as "ver du +Cayor" because it was first observed in Cayor, in Senegambia, +develops in the skin of man and of various animals, such as dogs, +cats, and monkeys. It is about 12 mm. in length, and of the form +of the larvæ of other muscids. Upon the intermediate segments are +minute, brownish recurved spines which give to the larva its characteristic +appearance. The life-history is not satisfactorily worked +out, but Fuller (1914), after reviewing +the evidence believes that, as a rule, it +deposits its young in the sleeping places +of man and animals, whether such be a +bed, a board, the floor, or the bare ground. +In the case of babies, the maggots may +be deposited on the scalp. The minute +maggots bore their way painlessly into +the skin. As many as forty parasites +have been found in one individual and +one author has reported finding more +than three hundred in a spaniel puppy. +Though their attacks are at times extremely +painful, it is seldom that any +serious results follow.<span class="pagenum"><a name="Page_119" id="Page_119">[Pg 119]</a></span></p> + + +<h3><a name="The_Siphonaptera_or_Fleas" id="The_Siphonaptera_or_Fleas"></a><span class="smcap">The Siphonaptera or Fleas</span></h3> + +<p>The <b>Siphonaptera</b>, or fleas (<a href="#Fig_89">fig. 89</a>) are wingless insects, with +highly chitinized and laterally compressed bodies. The mouth-parts +are formed for piercing and sucking. Compound eyes are lacking +but some species possess ocelli. The metamorphosis is complete.</p> + +<p>This group of parasites, concerning which little was known until +recently, has assumed a very great importance since it was learned +that fleas are the carriers of bubonic plague. Now over four hundred +species are known. Of these, several species commonly attack man. +The most common hominoxious species are <i>Pulex irritans</i>, <i>Xenopsylla +cheopis</i>, <i>Ctenocephalus canis</i>, <i>Ctenocephalus felis</i>, <i>Ceratophyllus +fasciatus</i> and <i>Dermatophilus penetrans</i>, but many others will feed +readily on human blood if occasion arises.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_89" id="Fig_89"></a> +<a href="images/f089-full.png"><img src="images/f089.png" width="500" height="467" alt="89. Xenopsylla cheopis, male (×25). After Jordan and Rothschild." title="89. Xenopsylla cheopis, male (×25). After Jordan and Rothschild." /></a> +<span class="caption">89. Xenopsylla cheopis, male (×25). After Jordan and Rothschild.</span> +</div> + +<p>We shall treat in this place of the general biology and habits of +the hominoxious forms and reserve for the systematic section the +discussion of the characteristics of the different genera.<span class="pagenum"><a name="Page_120" id="Page_120">[Pg 120]</a></span></p> + +<p>The most common fleas infesting houses in the Eastern United +States are the cosmopolitan dog and cat fleas, <i>Ctenocephalus canis</i> +(<a href="#Fig_90">fig. 90</a>) and <i>C. felis</i>. Their life +cycles will serve as typical. +These two species have until +recently been considered as one, +under the name <i>Pulex serraticeps</i>. +See <a href="#Fig_92">figure 92</a>.</p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_90" id="Fig_90"></a> +<a href="images/f090-full.png"><img src="images/f090.png" width="400" height="295" alt="90. Dog flea (×15). After Howard." title="90. Dog flea (×15). After Howard." /></a> +<span class="caption">90. Dog flea (×15). After Howard.</span> +</div> + +<p>The eggs are oval, slightly +translucent or pearly white, and +measure about .5 mm. in their +long diameter. They are deposited +loosely in the hairs of +the host and readily drop off as the animal moves around. Howard +found that these eggs hatch in one to two days. The larvæ are +elongate, legless, white, worm-like creatures. They are exceedingly +active, and avoid the light in every way possible. They +cast their first skin in from three to seven days and their second +in from three to four days. They commenced spinning in from +seven to fourteen days after hatching and the imago appeared +five days later. Thus in summer, at Washington, the entire life +cycle may be completed in about two weeks. (cf. fig. <a href="#Fig_91">91</a>, <a href="#Fig_92">92</a>).</p> + +<p>Strickland's (1914) studies on the biology of the rat flea, <i>Ceratophyllus +fasciatus</i>, have so important a general bearing that we shall +cite them in considerable detail.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_91" id="Fig_91"></a> +<a href="images/f091-full.png"><img src="images/f091.png" width="500" height="130" alt="91. Larva of Xenopsylla cheopis. After Bacot and Ridewood." title="91. Larva of Xenopsylla cheopis. After Bacot and Ridewood." /></a> +<span class="caption">91. Larva of Xenopsylla cheopis. After Bacot and Ridewood.</span> +</div> + +<p>He found, to begin with, that there is a marked inherent range +in the rate of development. Thus, of a batch of seventy-three eggs, +all laid in the same day and kept together under the same conditions, +one hatched in ten days; four in eleven days; twenty-five in +twelve days; thirty-one in thirteen days; ten in fourteen days; one +in fifteen days; and one in sixteen days. Within these limits the +duration of the egg period seems to depend mainly on the degree +of humidity. The incubation period is never abnormally prolonged<span class="pagenum"><a name="Page_121" id="Page_121">[Pg 121]</a></span> +as in the case of lice, (Warburton) and varying conditions of temperature +and humidity have practically no effect on the percentage of +eggs which ultimately hatch.</p> + +<p>The same investigator found that the most favorable condition +for the larva is a low temperature, combined with a high degree of +humidity; and that the presence of rubbish in which the larva may +bury itself is essential to its successful development. When larvæ +are placed in a bottle containing either wood-wool soiled by excrement, +or with feathers or filter paper covered with dried blood they +will thrive readily and pupate. They seem to have no choice between +dried blood and powdered rat feces for food, and also feed +readily on flea excrement. They possess the curious habit of always +devouring their molted skins.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_92" id="Fig_92"></a> +<a href="images/f092-full.png"><img src="images/f092.png" width="500" height="312" alt="92. Head and pronotum of (a) dog flea; (b) of cat flea; (c) of hen flea. After Rothschild. +(d) Nycteridiphilus (Ischnopsyllus) hexactenus. After Oudemans." title="92. Head and pronotum of (a) dog flea; (b) of cat flea; (c) of hen flea. After Rothschild. +(d) Nycteridiphilus (Ischnopsyllus) hexactenus. After Oudemans." /></a> +<span class="caption">92. Head and pronotum of (<i>a</i>) dog flea; (<i>b</i>) of cat flea; (<i>c</i>) of hen flea. After Rothschild. +(<i>d</i>) Nycteridiphilus (Ischnopsyllus) hexactenus. After Oudemans.</span> +</div> + +<p>An important part of Strickland's experiments dealt with the +question of duration of the pupal stage under the influence of temperature +and with the longevity and habits of the adult. In October, +he placed a batch of freshly formed cocoons in a small dish that was +kept near a white rat in a deep glass jar in the laboratory. Two +months later one small and feeble flea had emerged, but no more +until February, four months after the beginning of the experiment. +Eight cocoons were then dissected and seven more found to contain +the imago fully formed but in a resting state. The remainder of<span class="pagenum"><a name="Page_122" id="Page_122">[Pg 122]</a></span> +the batch was then placed at 70° F. for one night, near a white +rat. The next day all the cocoons were empty and the fleas were +found on the white rat.</p> + +<p>Thus, temperature greatly influences the duration of the pupal +period, which in <i>Ceratophyllus fasciatus</i> averages seventeen days. +Moreover, when metamorphosis is complete a low temperature will +cause the imago to remain within the cocoon.</p> + +<p>Sexually mature and ovipositing fleas, he fed at intervals and kept +alive for two months, when the experiment was discontinued. In +the presence of rubbish in which they could bury themselves, unfed +rat fleas were kept alive for many months, whereas in the absence of +any such substratum they rarely lived a month. In the former case, +it was found that the length of life is influenced to some degree by the +temperature and humidity. In an experiment carried out at 70° F. +and 45 per cent humidity, the fleas did not live for more than four +months, while in an experiment at 60° F. and 70 per cent humidity +they lived for at least seventeen months. There was no indication +that fleas kept under these conditions sucked moisture from surrounding +objects, and those kept in bell jars, with an extract of flea-rubbish +on filter paper, did not live any longer than those which were not so +supplied.</p> + +<p>Curiously enough, although the rat is the normal host of <i>Ceratophyllus +fasciatus</i>, it was found that when given the choice these fleas +would feed upon man in preference to rats. However, none of the +fleas laid eggs unless they fed on rat blood.</p> + +<p>The experiments of Strickland on copulation and oviposition in +the rat flea showed that fleas do not copulate until they are sexually +mature and that, at least in the case of <i>Ceratophyllus fasciatus</i>, the +reproductive organs are imperfectly developed for some time (more +than a week) after emerging from the pupa. When mature, copulation +takes place soon after the fleas have fed on their true host—the +rat—but not if they have fed on a facultative host only, such as man. +Copulation is always followed by oviposition within a very short +time.</p> + +<p>The effect of the rat's blood on the female with regard to egg-laying, +Strickland concludes, is stimulating rather than nutritive, +as fleas that were without food for many months were observed to +lay eggs immediately after one feed. Similarly, the male requires +the stimulus of a meal of rat's blood before it displays any copulatory +activity.<span class="pagenum"><a name="Page_123" id="Page_123">[Pg 123]</a></span></p> + +<p>Mitzmain (1910) has described in detail the act of biting on man, +as observed in the squirrel flea, <i>Ceratophyllus acutus</i>. "The flea +when permitted to walk freely on the arm selects a suitable hairy +space where it ceases abruptly in its locomotion, takes a firm hold +with the tarsi, projects its proboscis, and prepares to puncture the +skin. A puncture is drilled by the pricking epipharynx, the saw-tooth +mandibles supplementing the movement by lacerating the +cavity formed. The two organs of the rostrum work alternately, +the middle piece boring, while the two lateral elements execute a +sawing movement. The mandibles, owing to their basal attachments, +are, as is expressed by the advisory committee on plague +investigations in India (<i>Journal of Hygiene</i>, vol. 6, No. 4, p. 499), +'capable of independent action, sliding up and down but maintaining +their relative positions and preserving the lumen of the aspiratory +channel.' The labium doubles back, the V-shaped groove of this +organ guiding the mandibles on either side."</p> + +<p>"The action of the proboscis is executed with a forward movement +of the head and a lateral and downward thrust of the entire body. +As the mouth-parts are sharply inserted, the abdomen rises simultaneously. +The hind and middle legs are elevated, resembling oars. +The forelegs are doubled under the thorax, the tibia and tarsi resting +firmly on the epidermis serve as a support for the body during the +feeding. The maxillary palpi are retracted beneath the head and +thorax. The labium continues to bend, at first acting as a sheath +for the sawing mandibles, and as these are more deeply inserted, it +bends beneath the head with the elasticity of a bow, forcing the +mandibles into the wound until the maxillæ are embedded in the skin +of the victim. When the proboscis is fully inserted, the abdomen +ceases for a time its lateral swinging."</p> + +<p>"The acute pain of biting is first felt when the mandibles have +not quite penetrated and subsequently during each distinct movement +of the abdomen. The swinging of the abdomen gradually +ceases as it becomes filled with blood. The sting of the biting +gradually becomes duller and less sensitive as feeding progresses. +The movements of the elevated abdomen grow noticeably feebler +as the downward thrusts of the springy bow-like labium becomes less +frequent."</p> + +<p>"As the feeding process advances one can discern through the +translucent walls of the abdomen a constant flow of blood, caudally +from the pharynx, accompanied by a peristaltic movement. The<span class="pagenum"><a name="Page_124" id="Page_124">[Pg 124]</a></span> +end of the meal is signified in an abrupt manner. The flea shakes +its entire body, and gradually withdraws its proboscis by lowering +the abdomen and legs and violently twisting the head."</p> + +<p>"When starved for several days the feeding of the rat fleas is +conducted in a rather vigorous manner. As soon as the proboscis +is buried to the full length the abdomen is raised and there ensues a +gradual lateral swaying motion, increasing the altitude of the raised +end of the abdomen until it assumes the perpendicular. The flea is +observed at this point to gain a better foothold by advancing the +fore tarsi, and then, gradually doubling back the abdomen, it turns +with extreme agility, nearly touching with its dorsal side the skin +of the hand upon which it is feeding. Meanwhile, the hungry parasite +feeds ravenously."</p> + +<p>"It is interesting to note the peculiar nervous action which the +rodent fleas exhibit immediately when the feeding process is completed +or when disturbed during the biting. Even while the rostrum +is inserted to the fullest the parasite shakes its head spasmodically; +in a twinkling the mouth is withdrawn and then the flea hops away."</p> + +<p>A habit of fleas which we shall see is of significance in considering +their agency in the spread of bubonic plague, is that of ejecting blood +from the anus as they feed.</p> + +<p>Fleas are famous for their jumping powers, and in control measures +it is of importance to determine their ability along this line. It is +often stated that they can jump about four inches, or, according to +the Indian Plague Commission <i>Xenopsylla cheopis</i> cannot hop farther +than five inches. Mitzmain (1910) conducted some careful experiments +in which he found that the human flea, <i>Pulex irritans</i>, was +able to jump as far as thirteen inches on a horizontal plane. The +mean average of five specimens permitted to jump at will was seven +and three-tenths inches. The same species was observed to jump +perpendicularly to a height of at least seven and three-fourths inches. +Other species were not able to equal this record.</p> + +<p>The effect of the bite of fleas on man varies considerably according +to the individual susceptibility. According to Patton and Cragg, +this was borne out in a curious manner by the experiments of Chick +and Martin. "In these, eight human hosts were tried; in seven, +little or no irritation was produced, while in one quite severe inflammation +was set up around each bite." Of two individuals, equally +accustomed to the insects, going into an infested room, one may be +literally tormented by them while the other will not notice them.<span class="pagenum"><a name="Page_125" id="Page_125">[Pg 125]</a></span> +Indeed it is not altogether a question of susceptibility, for fleas seem +to have a special predilection for certain individuals. The typical +itching wheals produced by the bites are sometimes followed, especially +after scratching, by inflammatory papules.</p> + +<p>The itching can be relieved by the use of lotions of carbolic acid +(2-3 per cent), camphor, menthol lotion, or carbolated vaseline. +If forced to sleep in an infested room, protection from attacks can +be in a large measure gained by sprinkling pyrethrum, bubach, or +California insect powder between the sheets. The use of camphor, +menthol, or oil of eucalyptus, or oil of pennyroyal is also said to afford +protection to a certain extent.</p> + +<p>In the Eastern United States the occurrence of fleas as household +pests is usually due to infested cats and dogs which have the run of +the house. We have seen that the eggs are not attached to the host +but drop to the floor when they are laid. Verrill, cited by Osborn, +states that on one occasion he was able to collect fully a teaspoonful +of eggs from the dress of a lady in whose lap a half-grown kitten had +been held for a short time. Patton and Cragg record seeing the +inside of a hat in which a kitten had spent the night, so covered with +flea eggs that it looked "as if it had been sprinkled with sugar from +a sifter." It is no wonder that houses in which pets live become +overrun with the fleas.</p> + +<p>One of the first control measures, then, consists in keeping such +animals out of the house or in rigorously keeping them free from fleas. +The latter can best be accomplished by the use of strong tar soap +or Armour's "Flesope," which may be obtained from most druggists. +The use of a three per cent solution of creolin, approximately four +teaspoonfuls to a quart of warm water, has also been recommended. +While this is satisfactory in the case of dogs, it is liable to sicken cats, +who will lick their fur in an effort to dry themselves. Howard +recommends thoroughly rubbing into the fur a quantity of pyrethrum +powder. This partially stupifies the fleas which should be promptly +swept up and burned.</p> + +<p>He also recommends providing a rug for the dog or cat to sleep +on and giving this rug a frequent shaking and brushing, afterwards +sweeping up and burning the dust thus removed.</p> + +<p>Since the larvæ of fleas are very susceptible to exposure, the use +of bare floors, with few rugs, instead of carpets or matting, is to be +recommended. Thorough sweeping, so as to allow no accumulation +of dust in cracks and crevices will prove efficient. If a house is once<span class="pagenum"><a name="Page_126" id="Page_126">[Pg 126]</a></span> +infested it may be necessary to thoroughly scrub the floors with hot +soapsuds, or to spray them with gasoline. If the latter method is +adopted, care must be taken to avoid the possibility of fire.</p> + +<p>To clear a house of fleas Skinner recommends the use of flake +naphthalene. In a badly infested house he took one room at a time, +scattering on the floor five pounds of flake naphthalene, and closed +it for twenty-four hours. It proved to be a perfect and effectual +remedy and very inexpensive, as the naphthalene could be swept up +and transferred to other rooms. Dr. Skinner adds, "so far as I am +concerned, the flea question is solved and if I have further trouble +I know the remedy. I intend to keep the dog and cat."</p> + +<p>The late Professor Slingerland very effectively used hydrocyanic +acid gas fumigation in exterminating fleas in houses. In one case, +where failure was reported, he found on investigation that the house +had become thoroughly reinfested from pet cats, which had been left +untreated. Fumigation with sulphur is likewise efficient.</p> + +<p>The fact that adult fleas are usually to be found on the floor, +when not on their hosts, was ingeniously taken advantage of by +Professor S. H. Gage in ridding an animal room at Cornell University +of the pests. He swathed the legs of a janitor with sticky fly-paper +and had him walk back and forth in the room. Large numbers of +the fleas were collected in this manner.</p> + +<p>In some parts of the southern United States hogs are commonly +infested and in turn infest sheds, barns and even houses. Mr. H. E. +Vick informs us that it is a common practice to turn sheep into barn-lots +and sheds in the spring of the year to collect in their wool, the +fleas which abound in these places after the hogs have been turned +out.</p> + +<p>It is a common belief that adult fleas are attracted to fresh meat +and that advantage of this can be taken in trapping them. Various +workers, notably Mitzman (1910), have shown that there is no basis +for such a belief.</p> + + +<p><a name="The_true_chiggers" id="The_true_chiggers"></a><b>The true chiggers</b>—The chigoes, or true chiggers, are the most +completely parasitic of any of the fleas. Of the dozen or more known +species, one commonly attacks man. This is <i>Dermatophilus penetrans</i>, +more commonly known as <i>Sarcopsylla penetrans</i> or <i>Pulex penetrans</i>.</p> + +<p>This species occurs in Mexico, the West Indies, Central and South +America. There are no authentic records of its occurrence in the +United States although, as Baker has pointed out, there is no reason<span class="pagenum"><a name="Page_127" id="Page_127">[Pg 127]</a></span> +why it should not become established in Florida and Texas. It is +usually believed that Brazil was its original home. Sometime about +the middle of the nineteenth century it was introduced into West +Africa and has spread across that continent.</p> + +<p>The males and the immature females of <i>Dermatophilus penetrans</i> +(<a href="#Fig_93">fig. 93</a>) closely resemble those of other fleas. They are very active +little brown insects about 1-1.2 mm. in size, which live in the dust of +native huts and stables, and in dry, sandy soil. In such places they +often occur in enormous numbers and become a veritable plague.</p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_93" id="Fig_93"></a> +<a href="images/f093-full.png"><img src="images/f093.png" width="400" height="283" alt="93. Dermatophilus penetrans. Much enlarged. After Karsten." title="93. Dermatophilus penetrans. Much enlarged. After Karsten." /></a> +<span class="caption">93. Dermatophilus penetrans. Much enlarged. After Karsten.</span> +</div> + +<p>They attack not only man but various animals. According to +Castellani and Chalmers, "Perhaps the most noted feature is the way +in which it attacks pigs. On the Gold Coast it appeared to be largely +kept in existence by these animals. It is very easily captured in +the free state by taking a little pig with a pale abdomen, and placing +it on its back on the ground on which infected pigs are living. After +watching a few moments, a black speck will appear on the pig's +abdomen, and quickly another and another. These black specks are +jiggers which can easily be transferred to a test tube. On examination +they will be found to be males and females in about equal +numbers."</p> + +<p>Both the males and females suck blood. That which characterizes +this species as distinguished from other fleas attacking man is that +when the impregnated female attacks she burrows into the skin +and there swells until in a few days she has the size and appearance of +a small pea (<a href="#Fig_94">fig. 94</a>). Where they are abundant, hundreds of the<span class="pagenum"><a name="Page_128" id="Page_128">[Pg 128]</a></span> +pests may attack a single individual (<a href="#Fig_95">fig. 95</a>). Here they lie with the +apex of the abdomen blocking the opening. According to Fülleborn +(1908) they do not +penetrate beneath the +epidermis. The eggs are +not laid in the flesh of +the victim, as is sometimes +stated, but are +expelled through this +opening. The female +then dies, withers and +falls away or is expelled +by ulceration. According +to Brumpt, she first +quits the skin and then, +falling to the ground, +deposits her eggs. The +subsequent development +in so far as known, +is like that of other fleas.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_94" id="Fig_94"></a> +<a href="images/f094-full.png"><img src="images/f094.png" width="350" height="266" alt="94. Dermatophilus penetrans, gravid female. After Moniez." title="94. Dermatophilus penetrans, gravid female. After Moniez." /></a> +<span class="caption">94. Dermatophilus penetrans, gravid female. After Moniez.</span> +</div> + +<div class="figcenter" style="width: 250px;"><a name="Fig_95" id="Fig_95"></a> +<a href="images/f095-full.png"><img src="images/f095.png" width="250" height="334" alt="95. Chiggers in the sole of foot of man. Manson's +Tropical Diseases. Permission of Cassell and Co." title="95. Chiggers in the sole of foot of man. Manson's +Tropical Diseases. Permission of Cassell and Co." /></a> +<span class="caption">95. Chiggers in the sole of foot of man. Manson's +Tropical Diseases. Permission of Cassell and Co.</span> +</div> + +<div class="figleft" style="width: 250px;"><a name="Fig_96" id="Fig_96"></a> +<a href="images/f096-full.png"><img src="images/f096.png" width="250" height="327" alt="96. Echidnophaga gallinacea." title="96. Echidnophaga gallinacea." /></a> +<span class="caption">96. Echidnophaga gallinacea.</span> +</div> + +<p>The chigoe usually +enters between the toes, +the skin about the roots +of the nails, or the soles<span class="pagenum"><a name="Page_129" id="Page_129">[Pg 129]</a></span> +of the feet, although it may attack other parts of the body. Mense +records the occurrence in folds of the epidermis, as in the neighborhood +of the anus. They give rise to irritation +and unless promptly and aseptically removed +there often occurs pus formation and the +development of a more or less serious abscess. +Gangrene and even tetanus may ensue.</p> + +<p>Treatment consists in the careful removal +of the insect, an operation more easily accomplished +a day or two after its entrance, than +at first, when it is unswollen. The ulcerated +point should then be treated with weak carbolic +acid, or tincture of iodine, or dusted +thoroughly with an antiseptic powder.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_97" id="Fig_97"></a> +<a href="images/f097-full.png"><img src="images/f097.png" width="350" height="272" alt="97. Echidnophaga gallinacea infesting head of chicken. After Enderlein." title="97. Echidnophaga gallinacea infesting head of chicken. After Enderlein." /></a> +<span class="caption">97. Echidnophaga gallinacea infesting head of chicken. After Enderlein.</span> +</div> + +<p>Castellani and Chalmers recommend as +prophylactic measures, keeping the house clean and keeping pigs, +poultry, and cattle away therefrom. "High boots should be used, +and especial care should be taken not to go to a ground floor bathroom +with bare feet. The feet, especially the toes, and under the +nails, should be carefully examined every morning to see if any black +dots can be discovered, when the jigger should be at once removed, +and in this way suppuration will be prevented. It is advisable,<span class="pagenum"><a name="Page_130" id="Page_130">[Pg 130]</a></span> +also, to sprinkle the floors with carbolic lotion, Jeyes' fluid, or with +pyrethrum powder, or with a strong infusion of native tobacco, as +recommended by Law and Castellani."</p> + +<p><i>Echidnophaga gallinacea</i> (<a href="#Fig_96">fig. 96</a>) is a widely distributed Hectopsyllid +attacking poultry (<a href="#Fig_97">fig. 97</a>). It occurs in the Southern and Southwestern +United States and has been occasionally reported as attacking +man, especially children. It is less highly specialized than +<i>Dermatophilus penetrans</i>, and does not ordinarily cause serious +trouble in man.<span class="pagenum"><a name="Page_131" id="Page_131">[Pg 131]</a></span></p> + + + + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_IV" id="CHAPTER_IV"></a>CHAPTER IV</h2> + +<h2>ACCIDENTAL OR FACULTATIVE PARASITES</h2> + + +<p>In addition to the many species of Arthropods which are normally +parasitic on man and animals, there is a considerable number of those +which may be classed as <i>accidental</i> or <i>facultative</i> parasites.</p> + +<p>Accidental or facultative parasites are species which are normally +free-living, but which are able to exist as parasites when accidentally +introduced into the body of man or other animal. A wide range of +forms is included under this grouping.</p> + + +<h3><a name="Acarina" id="Acarina"></a><span class="smcap">Acarina</span></h3> + +<p>A considerable number of mites have been reported as accidental +or even normal, endoparasites of man, but the authentic cases are +comparatively few.</p> + +<p>In considering such reports it is well to keep in mind von Siebold's +warning that in view of the universal distribution of mites one should +be on his guard. In vessels in which animal and other organic +fluids and moist substances gradually dry out, mites are very abundantly +found. If such vessels are used without very careful preliminary +cleaning, for the reception of evacuations of the sick, or for the +reception of parts removed from the body, such things may be readily +contaminated by mites, which have no other relation whatever to +them.</p> + +<p>Nevertheless, there is no doubt but that certain mites, normally +free-living, have occurred as accidental parasites of man. Of these +the most commonly met with is <i>Tyroglyphus siro</i>, the cheese-mite.</p> + +<p><i>Tyroglyphus siro</i> is a small mite of a whitish color. The male +measures about 500µ long by 250µ wide, the female slightly larger. +They live in cheese of almost any kind, especially such as is a little +decayed. "The individuals gather together in winter in groups or +heaps in the hollows and chinks of the cheese and there remain +motionless. As soon as the temperature rises a little, they gnaw +away at the cheese and reduce it to a powder. The powder is composed +of excrement having the appearance of little grayish microscopic +balls; eggs, old and new, cracked and empty; larvæ, nymphs, and +perfect mites, cast skins and fragments of cheese, to which must be +added numerous spores of microscopic fungi."—Murray.<span class="pagenum"><a name="Page_132" id="Page_132">[Pg 132]</a></span></p> + +<p><i>Tyroglyphus siro</i>, and related species, have been found many +times in human feces, under conditions which preclude the explanation +that the contamination occurred outside of the body. They +have been supposed to be the cause of dysentery, or diarrhœa, and +it is probable that the <i>Acarus dysenteriæ</i> of Linnæus, and Latreille, +was this species. However, there is little evidence that the mites +cause any noteworthy symptoms, even when taken into the body +in large numbers.</p> + +<p><i>Histiogaster spermaticus</i> (<a href="#Fig_152">fig. 152</a>) is a Tyroglyphid mite which +was reported by Trouessart (1902) as having been found in a cyst +in the groin, adherent to the testis. When the cyst was punctured, +it yielded about two ounces of opalescent fluid containing spermatozoa +and numerous mites in all stages of development. The evidence +indicated that a fecundated female mite had been introduced into +the urethra by means of an unclean catheter. Though Trouessart +reported the case as that of a Sarcoptid, Banks places the genus +<i>Histiogaster</i> with the Tyroglyphidæ. He states that our species +feeds on the oyster-shell bark louse, possibly only after the latter is +dead, and that in England a species feeds within decaying reeds.</p> + +<p><i>Nephrophages sanguinarius</i> is a peculiarly-shaped, angular mite +which was found by Miyake and Scriba (1893) for eight successive +days in the urine of a Japanese suffering from fibrinuria. Males, +.117 mm. long by .079 mm. wide, females .36 mm. by. 12 mm., +and eggs were found both in the spontaneously emitted urine and in +that drawn by means of a catheter. All the mites found were dead. +The describers regarded this mite as a true endoparasite, but it is +more probable that it should be classed as an accidental parasite.</p> + + +<h3><a name="Myriapoda" id="Myriapoda"></a><span class="smcap">Myriapoda</span></h3> + +<p>There are on record a number of cases of myriapods occurring as +accidental parasites of man. The subject has been treated in detail +by Blanchard (1898 and 1902), who discussed forty cases. Since +then at least eight additions have been made to the list.</p> + +<p>Neveau-Lamaire (1908) lists thirteen species implicated, representing +eight different genera. Of the <i>Chilognatha</i> there are three, +<i>Julus terrestris</i>, <i>J. londinensis</i> and <i>Polydesmus complanatus</i>. The +remainder are <i>Chilopoda</i>, namely, <i>Lithobius forficatus</i>, <i>L. malenops</i>, +<i>Geophilus carpophagus</i>, <i>G. electricus</i>, <i>G. similis</i>, <i>G. cephalicus</i>, <i>Scutigera +coleoptrata</i>, <i>Himantarium gervaisi</i>, <i>Chætechelyne vesuviana</i> and +<i>Stigmatogaster subterraneus</i>.<span class="pagenum"><a name="Page_133" id="Page_133">[Pg 133]</a></span></p> + +<p>The majority of the cases relate to infestation of the nasal fossæ, +or the frontal sinus, but intestinal infestation also occurs and there +is one recorded case of the presence of a species in <i>Julus</i> (<a href="#Fig_13">fig. 13</a>) in +the auditory canal of a child.</p> + +<p>In the nose, the myriapods have been known to live for months +and according to some records, even for years. The symptoms +caused by their presence are inflammation, with or without increased +flow of mucus, itching, more or less intense headache, and at times +general symptoms such as vertigo, delirium, convulsions, and the +like. These symptoms disappear suddenly when the parasites are +expelled.</p> + +<p>In the intestine of man, myriapods give rise to obscure symptoms +suggestive of infestation by parasitic worms. In a case reported by +Verdun and Bruyant (1912), a child twenty months of age had been +affected for fifteen days by digestive disturbances characterized by +loss of appetite, nausea and vomiting. The latter had been particularly +pronounced for three days, when there was discovered in the +midst of the material expelled a living myriapod of the species +<i>Chætechelyne vesuviana</i>. Anthelminthics had been administered +without result. In some of the other cases, the administration of +such drugs had resulted in the expulsion of the parasite through the +anus.</p> + +<p>One of the extreme cases on record is that reported by Shipley +(1914). Specimens of <i>Geophilus gorizensis</i> (= <i>G. subterraneus</i>) +"were vomited and passed by a woman of 68 years of age. Some of +the centipedes emerged through the patient's nose, and it must be +mentioned that she was also suffering from a round worm. One of +her doctors was of the opinion that the centipedes were certainly +breeding inside the lady's intestines, and as many as seven or eight, +sometimes more, were daily leaving the alimentary canal."</p> + +<p>"According to her attendant's statements those centipedes had +left the body in some hundreds during a period of twelve or eighteen +months. Their presence produced vomiting and some hæmatemesis, +and treatment with thymol, male-fern and turpentine had no effect +in removing the creatures."</p> + +<p>The clinical details, as supplied by Dr. Theodore Thompson were +as follows:</p> + +<p>"Examined by me July, 1912, her tongue was dry and glazed. +There was bleeding taking place from the nose and I saw a living +centipede she had just extracted from her nostril. Her heart, lungs<span class="pagenum"><a name="Page_134" id="Page_134">[Pg 134]</a></span> +and abdomen appeared normal. She was not very wasted, and did +not think she had lost much flesh, nor was there any marked degree +of anemia."</p> + +<p>Shipley gives the following reasons for believing it impossible +that these centipedes could have multiplied in the patient's intestine. +"The breeding habits of the genus <i>Geophilus</i> are peculiar, and ill +adapted for reproducing in such a habitat. The male builds a small +web or nest, in which he places his sperm, and the female fertilizes +herself from this nest or web, and when the eggs are fertilized they +are again laid in a nest or web in which they incubate and in two or +three weeks hatch out. The young <i>Geophilus</i> differ but very little +from the adult, except in size. It is just possible, but improbable, +that a clutch of eggs had been swallowed by the host when eating +some vegetables or fruit, but against this is the fact that the <i>Geophilus</i> +does not lay its eggs upon vegetables or fruit, but upon dry wood or +earth. The egg-shell is very tough and if the eggs had been swallowed +the egg-shells could certainly have been detected if the dejecta were +examined. The specimens of the centipede showed very little signs +of being digested, and it is almost impossible to reconcile the story +of the patient with what one knows of the habits of the centipedes."</p> + +<p>In none of the observed cases have there been any clear indications +as to the manner of infestation. It is possible that the myriapods +have been taken up in uncooked fruit or vegetables.</p> + + +<h3><a name="Lepidopterous_Larvae" id="Lepidopterous_Larvae"></a><span class="smcap">Lepidopterous Larvæ</span></h3> + + +<p><b>Scholeciasis</b>—Hope (1837) brought together six records of infestation +of man by lepidopterous larvæ and proposed to apply the name +scholeciasis to this type of parasitism. The clearest case was that +of a young boy who had repeatedly eaten raw cabbage and who +vomited larvæ of the cabbage butterfly, <i>Pieris brassicæ</i>. Such cases +are extremely rare, and there are few reliable data relative to the +subject. In this connection it may be noted that Spuler (1906) has +described a moth whose larvæ live as ectoparasites of the sloth.</p> + + +<h3><a name="Coleoptera" id="Coleoptera"></a><span class="smcap">Coleoptera</span></h3> + + +<p><b>Canthariasis</b>—By this term Hope designated instances of accidental +parasitism by the larvæ or adults of beetles. Reports of +such cases are usually scouted by parasitologists but there seems no +good basis for wholly rejecting them. Cobbold refers to a half +dozen cases of accidental parasitism by the larvæ of <i>Blaps mortisaga</i>.<span class="pagenum"><a name="Page_135" id="Page_135">[Pg 135]</a></span> +In one of these cases upwards of 1200 larvæ and several perfect +insects were said to have been passed <i>per annum</i>. French (1905) +reports the case of a man +who for a considerable period +voided adult living beetles +of the species <i>Nitidula +bipustulata</i>. Most of the +other cases on record relate +to the larvæ of <i>Dermestidæ</i> +(larder beetles <i>et al.</i>) or +<i>Tenebrionidæ</i> (meal infesting species). Infestation probably occurs +through eating raw or imperfectly cooked foods containing eggs or +minute larvæ of these insects.</p> + +<div class="figcenter" style="width: 450px;"><a name="Fig_98" id="Fig_98"></a> +<a href="images/f098-full.png"><img src="images/f098.png" width="450" height="181" alt="98. Larva of Piophila casei. Caudal aspect of larva. +Posterior stigmata." title="98. Larva of Piophila casei. Caudal aspect of larva. +Posterior stigmata." /></a> +<span class="caption">98. Larva of Piophila casei. Caudal aspect of larva. +Posterior stigmata.</span> +</div> + +<p>Brumpt cites a curious case of accidental parasitism by a coleopterous +larva belonging to the genus <i>Necrobia</i>. This larva was extracted +from a small tumor, several millimeters long, on the surface of the +conjunctiva of the eye. The larvæ of this genus ordinarily live in +decomposing flesh and cadavers.</p> + + +<h3><a name="Dipterous_Larvae" id="Dipterous_Larvae"></a><span class="smcap">Dipterous Larvæ</span></h3> + +<div class="figcenter" style="width: 500px;"><a name="Fig_99" id="Fig_99"></a> +<a href="images/f099-full.png"><img src="images/f099.png" width="500" height="380" alt="99. Piophila casei. After Graham-Smith." title="99. Piophila casei. After Graham-Smith." /></a> +<span class="caption">99. Piophila casei. After Graham-Smith.</span> +</div> + + +<p><b>Myasis</b>—By this term (spelled also myiasis, and myiosis), is +meant parasitism by dipterous larvæ. Such parasitism may be +normal, as in the cases already described under the heading parasitic +Diptera, or it may be facultative, due to free-living larvæ being +accidentally introduced +into wounds or the body-cavities +of man. Of this +latter type, there is a +multitude of cases on +record, relating to comparatively +few species. +The literature of the subject, +like that relating +to facultative parasitism +in general, is unsatisfactory, +for most of the +determinations of species +have been very loose. +Indeed, so little has been known regarding the characteristics of +the larvæ concerned that in many instances they could not be exactly<span class="pagenum"><a name="Page_136" id="Page_136">[Pg 136]</a></span> +determined. Fortunately, several workers have undertaken comparative +studies along this line. The most comprehensive publication +is that of Banks (1912), entitled "The structure of certain dipterous +larvæ, with particular reference to those in human food."</p> + +<p>Without attempting an exhaustive list, we shall discuss here the +more important species of Diptera whose larvæ are known to cause +myasis, either external or internal. The following key will serve +to determine those most likely to be encountered. The writers +would be glad to examine specimens not readily identifiable, if +accompanied by exact data relative to occurrence.</p> + +<div class="blockquot"><div class="hanging"> +<p><i>a.</i> Body more or less flattened, depressed; broadest in the middle, each segment +with dorsal, lateral, and ventral fleshy processes, of which the laterals, +at least, are more or less spiniferous (<a href="#Fig_101">fig. 101</a>). <i>Fannia</i> (= <i>Homalomyia</i>).</p> + +<div class="hanging"> +<p>In <i>F. canicularis</i> the dorsal processes are nearly as long as the laterals; +in <i>F. scalaris</i> the dorsal processes are short spinose tubercles.</p> +</div> + +<p><i>aa.</i> Body cylindrical, or slender conical tapering toward the head; without +fleshy lateral processes (<a href="#Fig_105">fig. 105</a>).</p> + +<p><i>b.</i> With the posterior stigmata at the end of shorter or longer tubercles, or if not +placed upon tubercles, then not in pit; usually without a "marginal button" +and without a chitinous ring surrounding the three slits; the slits narrowly +or broadly oval, not bent (<a href="#Fig_171">fig. 171 i</a>). <i>Acalyptrate muscidæ</i> and some species +of <i>Anthomyiidæ</i>. To this group belong the cheese skipper (<i>Piophila casei</i>, +figs. <a href="#Fig_98">98</a>, <a href="#Fig_99">99</a>), the pomace-fly (<i>Drosophila ampelophila</i>), the apple maggot +(<i>Rhagoletis pomonella</i>), the cherry fruit fly (<i>Rhagoletis cingulata</i>), the small +dung fly (<i>Sepsis violacea</i>, <a href="#Fig_170">fig. 170</a>), the beet leaf-miner (<i>Pegomyia vicina</i>, +<a href="#Fig_171">fig. 171 i</a>), the cabbage, bean and onion maggots (<i>Phorbia</i> spp.) et. al.</p> + +<p><i>bb.</i> Posterior stigmata of various forms, if the slits are narrowly oval (<a href="#Fig_171">fig. 171</a>) +then they are surrounded by a chitin ring which may be open ventro-mesally.</p> + +<p><i>c.</i> Integument leathery and usually strongly spinulose; larvæ hypodermatic or +endoparasitic. Bot flies (<a href="#Fig_171">fig. 171, f, g, k</a>).—<i>Oestridæ</i></p> + +<p><i>cc.</i> Integument not leathery and (except in <i>Protocalliphora</i>) spinulæ restricted +to transverse patches near the incisures of the segments.</p> + +<p><i>d.</i> The stigmal plates in a pit; the lip-like margin of the pit with a number of +fleshy tubercles; <span title="insert "ring" after "chitin""><a name="AC_7" id="AC_7"></a>chitin</span> of the stigma not complete; open ventro-mesally, +button absent (<a href="#Fig_171">fig. 171 e</a>). Flesh flies.—<i>Sarcophaga</i></p> + +<p><i>dd.</i> Stigmata not in a pit.</p> + +<p><i>e.</i> The chitin ring open ventra-mesally; button absent (<a href="#Fig_171">fig. 171 c</a>). Screw-worm +fly. <i>Chrysomyia</i></p> + +<p><i>ee.</i> The chitin ring closed.</p> + +<p><i>f.</i> Slits of the posterior stigmata straight; marginal "button" present (<a href="#Fig_171">fig. 171 b</a>); +two distinct mouth hooks, fleshy tubercles around the anal area. <i>Phormia</i> +(<a href="#Fig_171">fig. 171 f</a>), <i>Lucilia</i> and <i>Calliphora</i> (<a href="#Fig_172">fig. 172, a, b</a>), <i>Protocalliphora</i> (<a href="#Fig_171">fig. 171, j</a>), +<i>Cynomyia</i> (<a href="#Fig_171">fig. 171, a</a>). Blow flies, bluebottle flies. <i>Calliphorinæ</i><span class="pagenum"><a name="Page_137" id="Page_137">[Pg 137]</a></span></p> + +<p><i>ff.</i> Slits of the posterior stigmata sinuous or bent. Subfamily Muscinæ.</p> + +<p><i>g.</i> Slits of the posterior stigmata bent; usually two mouth hooks. <i>Muscina +stabulans</i> (<a href="#Fig_171">fig. 171, l</a>), <i>Muscina similis</i>, <i>Myiospila <span title="for meditatunda read meditabunda"><a name="AC_8" id="AC_8"></a>meditatunda</span></i> (<a href="#Fig_172">fig. 172, i</a>), +and some of the higher <i>Anthomyiidæ</i>.</p> + +<p><i>gg.</i> Slits of the posterior stigmata sinuous; mouth hooks usually consolidated +into one. The house-fly (<i>Musca domestica</i> <a href="#Fig_171">fig. 171, d</a>), the stable fly +(<i>Stomoxys calcitrans</i>), the horn fly (<i>Lyperosia irritans</i>), <i>Pyrellia</i>, <i>Pseudopyrellia</i>, +<i>Morellia</i>, <i>Mesembrina</i>. <i>Polietes</i>, et. al. (<a href="#Fig_172">fig. 172</a> in part).</p> +</div></div> + +<p><i>Eristalis</i>—The larvæ of <i>Eristalis</i> are the so-called rat-tailed maggots, +which develop in foul water. In a few instances these larvæ +have been known to pass through the human alimentary canal +uninjured. Hall and Muir (1913) report the case of a boy five years +of age, who had been ailing for ten weeks and who was under treatment +for indigestion and chronic constipation. For some time he +had vomited everything he ate. On administration of a vermifuge +he voided one of the rat-tailed maggots of <i>Eristalis</i>. He admitted +having drunk water from a ditch full of all manner of rotting matter. +It was doubtless through this that he became infested. It is worth +noting that the above described symptoms may have been due to +other organisms or substances in the filthy water.</p> + +<p><a name="Piophila_casei" id="Piophila_casei"></a><i>Piophila casei</i>, the cheese-fly (<a href="#Fig_99">fig. 99</a>), deposits its eggs not only +in old cheeses, but on ham, bacon, and other fats. The larvæ (<a href="#Fig_98">fig. 98</a>) +are the well-known cheese skippers, which sometimes occur in great +abundance on certain kinds of cheese. Indeed, some people have +a comfortable theory that such infested cheese is especially good. +Such being the case, it is small wonder that this species has been +repeatedly reported as causing intestinal myasis. Thebault (1901) +describes the case of a girl who, shortly after consuming a large piece +of badly infested cheese, became ill and experienced severe pains +in the region of the navel. Later these extended through the entire +alimentary canal, the excrement was mixed with blood and she +suffered from vertigo and severe headaches. During the four following +days the girl felt no change, although the excretion of the blood +gradually diminished and stopped. On the fourth day she voided +two half-digested larvæ and, later, seven or eight, of which two were +alive and moving.</p> + +<p>That these symptoms may be directly attributed to the larvæ, +or "skippers," has been abundantly shown by experimental evidence. +Portschinsky cites the case of a dog fed on cheese containing the +larvæ. The animal suffered much pain and its excrement contained +blood. On <i>post mortem</i> it was found that the small intestine throughout<span class="pagenum"><a name="Page_138" id="Page_138">[Pg 138]</a></span> +almost its entire length was marked by bloody bruises. The +papillæ on these places were destroyed, although the walls were +not entirely perforated. In the appendix were found two or three +dead larvæ. Alessandri (1910) has likewise shown that the larvæ +cause intestinal lesions.</p> + +<p>According to Graham-Smith, Austen (1912) has recorded a case +of myasis of the nose, attended with a profuse watery discharge of +several weeks duration and pain, due to the larvæ of <i>Piophila casei</i>.</p> + + +<p><a name="Anthyomyiidae" id="Anthyomyiidae"></a><b>Anthyomyiidæ</b>—The characteristic larvæ of two species of <i>Fannia</i> +(= <i>Homalomyia</i> or <i>Anthomyia</i>, in part) (<a href="#Fig_101">fig. 101</a>) are the most commonly +reported of dipterous +larvæ causing intestinal +myasis. Hewitt +(1912) has presented a +valuable study of the bionomics +and of the larvæ +of these flies, a type of +what is needed for all the +species concerned in myasis. +We have seen two +cases of their having been +passed in stools, without having caused any special symptoms. +In other instances their presence in the alimentary canal has given +rise to symptoms vaguely described as those of tapeworm infestation, +or helminthiasis. More specifically, they have been described as +causing vertigo, severe headache, nausea and vomiting, severe +abdominal pains, and in some instances, bloody diarrhœa.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_100" id="Fig_100"></a> +<a href="images/f100-full.png"><img src="images/f100.png" width="500" height="306" alt="100. Fannia canicularis (×4). After Graham-Smith." title="100. Fannia canicularis (×4). After Graham-Smith." /></a> +<span class="caption">100. Fannia canicularis (×4). After Graham-Smith.</span> +</div> + +<p>One of the most striking cases is that reported by Blankmeyer +(1914), of a woman whose illness began fourteen years previously +with nausea and vomiting. After several months of illness she began +passing larvæ and was compelled to resort to enemas. Three years +previous to the report, she noticed frequent shooting pains in the +rectal region and at times abdominal tenderness was marked. There +was much mucus in the stools and she "experienced the sensation +of larvæ crawling in the intestine." Occipital headaches were +marked, with remissions, and constipation became chronic. The +appetite was variable, there was a bad taste in the mouth, tongue +furred and ridged, and red at the edges. Her complexion was sallow, +and general nervousness was marked. As treatment, there +were given doses of magnesium sulphate before breakfast and at<span class="pagenum"><a name="Page_139" id="Page_139">[Pg 139]</a></span> +4 <span class="smcap">P. M.</span>, with five grain doses of salol four times a day. The customary +parasiticides yielded no marked benefit. At the time of the report +the patient passed from four to fifty larvæ per day, and was showing +some signs of improvement. The nausea had disappeared, her +nervousness was less evident, and there was a slight gain in weight.</p> + +<div class="figleft" style="width: 200px;"><a name="Fig_101" id="Fig_101"></a> +<a href="images/f101-full.png"><img src="images/f101.png" width="200" height="442" alt="101. Larva of Fannia +scalaris." title="101. Larva of Fannia +scalaris." /></a> +<span class="caption">101. Larva of Fannia +scalaris.</span> +</div> + +<p>The case was complicated by various other disorders, but the +symptoms given above seem to be in large part attributable to the +myasis. There is nothing in the case to justify the assumption +that larvæ were continuously present, for years. It seems more +reasonable to suppose that something in the habits of the patient +favored repeated infestation. Nevertheless, a study of the various +cases of intestinal myasis caused by these and +other species of dipterous larvæ seems to indicate +that the normal life cycle may be considerably +prolonged under the unusual conditions.</p> + +<p>The best authenticated cases of myasis of the +urinary passage have been due to larvæ of +<i>Fannia</i>. Chevril (1909) collected and described +twenty cases, of which seven seemed beyond +doubt. One of these was that of a woman of +fifty-five who suffered from albuminuria, and +urinated with much difficulty, and finally passed +thirty to forty larvæ of <i>Fannia canicularis</i>.</p> + +<p>It is probable that infestation usually occurs +through eating partially decayed fruit or vegetables +on which the flies have deposited their +eggs. Wellman points out that the flies may +deposit their eggs in or about the anus of +persons using outside privies and Hewitt +believes that this latter method of infection is probably the common +one in the case of infants belonging to careless mothers. "Such +infants are sometimes left about in an exposed and not very clean +condition, in consequence of which flies are readily attracted to them +and deposit their eggs."</p> + + +<p><a name="Muscinae" id="Muscinae"></a><b>Muscinæ</b>—The larvæ of the common house-fly, <i>Musca domestica</i>, +are occasionally recorded as having been passed with the feces or +vomit of man. While such cases may occur, it is probable that in +most instances similar appearing larvæ of other insects have been +mistakenly identified.<span class="pagenum"><a name="Page_140" id="Page_140">[Pg 140]</a></span></p> + +<p><i>Muscina stabulans</i> is regarded +by Portschinsky +(1913) as responsible for +many of the cases of intestinal +myasis attributed to other +species. He records the case +of a peasant who suffered from +pains in the lower part of the +breast and intestines, and +whose stools were mixed with +blood. From November until +March he had felt particularly +ill, being troubled with +nausea and vomiting in addition +to the pain in his intestines. In March, his physician prescribed +injections of a concentrated solution of tannin, which resulted in the +expulsion of fifty living larvæ of <i>Muscina stabulans</i>. Thereafter +the patient felt much better, although he suffered from intestinal +catarrh in a less severe form.</p> + +<div class="figcenter" style="width: 450px;"><a name="Fig_102" id="Fig_102"></a> +<a href="images/f102-full.png"><img src="images/f102.png" width="450" height="386" alt="102. Muscina stabulans (×4). After Graham-Smith." title="102. Muscina stabulans (×4). After Graham-Smith." /></a> +<span class="caption">102. Muscina stabulans (×4). After Graham-Smith.</span> +</div> + + +<p><a name="Calliphorinae" id="Calliphorinae"></a><b>Calliphorinæ</b>—Closely related to the Sarcophagidæ are the +<i>Calliphorinæ</i>, to which group belong many of the so-called "blue +bottle" flies. Their larvæ feed upon dead animals, and upon fresh +and cooked meat. Those of <i>Protocalliphora</i>, +already mentioned, +are ectoparasitic on living nestling +birds. Larva of <i>Lucilia</i>, we have +taken from tumors on living turtles. +To this sub-family belongs also +<i>Auchmeromyia luteola</i>, the Congo +floor maggot. Some of these, +and at least the last mentioned, +are confirmed, rather than faculative +parasites. Various species of +Calliphorinæ are occasionally met +with as facultative parasites of +man.</p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_103" id="Fig_103"></a> +<a href="images/f103-full.png"><img src="images/f103.png" width="400" height="407" alt="103. Lucilia cæsar, (×3). After Howard." title="103. Lucilia cæsar, (×3). After Howard." /></a> +<span class="caption">103. Lucilia cæsar, (×3). After Howard.</span> +</div> + +<p><i>Chrysomyia macellaria</i>, the screw worm fly (<a href="#Fig_107">fig. 107</a>), is the fly +which is responsible for the most serious cases of human myasis in +the United States. It is widely distributed in the United States<span class="pagenum"><a name="Page_141" id="Page_141">[Pg 141]</a></span> +but is especially abundant in the south. While the larvæ breed in +decaying matter in general, they so commonly breed in the living +flesh of animals that they merit rank as true parasites. The females +are attracted to open wounds of all kinds on cattle and other animals +and quickly deposit large numbers of eggs. Animals which have +been recently castrated, dehorned, or branded, injured by barbed +wire, or even by the attacks of ticks are promptly attacked in the +regions where the fly abounds. Even the navel of young calves or +discharges from the vulva of cows may attract the insect.</p> + +<div class="figcenter" style="width: 500px;"> +<a href="images/f104-full.png"><img src="images/f104.png" width="500" height="300" alt="104. Calliphora erythrocephala, (×6). After Graham-Smith." title="104. Calliphora erythrocephala, (×6). After Graham-Smith." /></a> +<span class="caption">104. Calliphora erythrocephala, (×6). After Graham-Smith.</span> +</div> + +<p>Not infrequently the fly attacks man, being attracted by an offensive +breath, a chronic catarrh, or a purulent discharge from the +ears. Most common are the cases where the eggs are deposited in +the nostrils. The larvæ, which are hatched in a day or two, are +provided with strong spines and proceed to bore into the tissues +of the nose, even down into or through the bone, into the frontal +sinus, the pharynx, larynx, and neighboring parts.</p> + +<p>Osborn (1896) quotes a number of detailed accounts of the attacks +of the <i>Chrysomyia</i> on man. A vivid picture of the symptomology +of rhinal myasis caused by the larvæ of this fly is given by Castellani +and Chalmers: "Some couple of days after a person suffering from +a chronic catarrh, foul breath, or ozæna, has slept in the open or has +been attacked by a fly when riding or driving,—<i>i.e.</i>, when the hands +are engaged—signs of severe catarrh appear, accompanied with +inordinate sneezing and severe pain over the root of the nose or the +frontal bone. Quickly the nose becomes swollen, and later the face +also may swell, while examination of the nose may show the presence<span class="pagenum"><a name="Page_142" id="Page_142">[Pg 142]</a></span> +of the larvæ. Left untreated, the patient rapidly becomes worse, +and pus and blood are discharged from the nose, from which an +offensive odor issues. Cough appears as well as fever, and often +some delirium. If the patient lives long enough, the septum of the +nose may fall in, the soft and hard palates may be pierced, the wall +of the pharynx may be destroyed. By this time, however, the course +of the disease will have become quite evident by the larvæ dropping +out of the nose, and if the patient continues to live all the larvæ +may come away naturally."</p> + +<p>For treatment of rhinal myasis these writers recommend douching +the nose with chloroform water or a solution of chloroform in +sweet milk (10-20 per cent), followed by douches of mild antiseptics. +Surgical treatment may be necessary.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_105" id="Fig_105"></a> +<a href="images/f105-full.png"><img src="images/f105.png" width="500" height="296" alt="105. Larva of a flesh fly (Sarcophaga). Caudal aspect. Anterior stigmata. Pharyngeal skeleton." title="105. Larva of a flesh fly (Sarcophaga). Caudal aspect. Anterior stigmata. Pharyngeal skeleton." /></a> +<span class="caption">105. Larva of a flesh fly (Sarcophaga). Caudal aspect. Anterior stigmata. Pharyngeal skeleton.</span> +</div> + + +<p><a name="Sarcophagidae" id="Sarcophagidae"></a><b>Sarcophagidæ</b>—The larvæ (<a href="#Fig_105">fig. 105</a>) of flies of this family usually +feed upon meats, but have been found in cheese, oleomargerine, +pickled herring, dead and living insects, cow dung and human feces. +Certain species are parasitic in insects. Higgins (1890) reported +an instance of "hundreds" of larvæ of <i>Sarcophaga</i> being vomited by a +child eighteen months of age. There was no doubt as to their origin +for they were voided while the physician was in the room. There +are many other reports of their occurrence in the alimentary canal. +We have recorded elsewhere (Riley, 1906) a case in which some ten +or twelve larvæ of <i>Sarcophaga</i> were found feeding on the diseased +tissues of a malignant tumor. The tumor, a melanotic sarcoma, +was about the size of a small walnut, and located in the small of the +back of an elderly lady. Although they had irritated and caused a +slight hæmorrhage, neither the patient nor others of the family knew<span class="pagenum"><a name="Page_143" id="Page_143">[Pg 143]</a></span> +of their presence. Any discomfort which they had caused had been +attributed to the sarcomatous growth. The infestation occurred +in mid-summer. It is probable that the adult was attracted by the +odor of the discharges and deposited the living maggots upon the +diseased tissues.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_106" id="Fig_106"></a> +<a href="images/f106-full.png"><img src="images/f106.png" width="500" height="347" alt="106. A flesh fly (Sarcophaga), (×4). After Graham-Smith." title="106. A flesh fly (Sarcophaga), (×4). After Graham-Smith." /></a> +<span class="caption">106. A flesh fly (Sarcophaga), (×4). After Graham-Smith.</span> +</div> + +<p>According to Küchenmeister, <i>Sarcophaga carnaria</i> (<a href="#Fig_106">fig. 106</a>), +attracted by the odor, deposits its eggs and larvæ in the vagina of +girls and women when they lie naked in hot summer days upon dirty +clothes, or when they have a discharge from the vagina. In malignant +inflammations of the eyes the larvæ +even nestle under the eyelids and in +Egypt, for example, produce a very +serious addition to the effects of small-pox +upon the cornea, as according to +Pruner, in such cases perforation of the +cornea usually takes place.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_107" id="Fig_107"></a> +<a href="images/f107-full.png"><img src="images/f107.png" width="350" height="475" alt="107. Chrysomyia macellaria, (×3)." title="107. Chrysomyia macellaria, (×3)." /></a> +<span class="caption">107. Chrysomyia macellaria, (×3).</span> +</div> + +<p><i>Wohlfartia magnifica</i> is another +Sarcophagid which commonly infests +man in the regions where it is abundant. +It is found in all Europe but is +especially common in Russia, where +Portschinsky has devoted much attention +to its ravages. It deposits living +larvæ in wounds, the nasal fossæ, the +ears and the eyes, causing injuries +even more revolting than those described for <i>Chrysomyia</i>.<span class="pagenum"><a name="Page_144" id="Page_144">[Pg 144]</a></span></p> + + + + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_V" id="CHAPTER_V"></a>CHAPTER V</h2> + +<h2>ARTHROPODS AS SIMPLE CARRIERS OF DISEASE</h2> + + +<p>The fact that certain arthropods are poisonous, or may affect the +health of man as direct parasites has always received attention in +the medical literature. We come now to the more modern aspect +of our subject,—the consideration of insects and other arthropods +as transmitters and disseminators of disease.</p> + +<p>The simplest way in which arthropods may function in this +capacity is as <i>simple carriers</i> of pathogenic organisms. It is conceivable +that any insect which has access to, and comes in contact +with such organisms and then passes to the food, or drink, or to the +body of man, may in a wholly accidental and incidental manner +convey infection. That this occurs is abundantly proved by the +work of recent years. We shall consider as typical the case against +the house-fly, which has attracted so much attention, both popular +and scientific. The excellent general treatises of Hewitt (1910), +Howard (1911), and Graham-Smith (1913), and the flood of bulletins +and popular literature render it unnecessary to consider the topic +in any great detail.</p> + + +<h3><a name="The_House-fly_As_a_Carrier_of_Disease" id="The_House-fly_As_a_Carrier_of_Disease"></a><span class="smcap">The House-fly As a Carrier of Disease</span></h3> + +<p>Up to the past decade the house-fly has usually been regarded as a +mere pest. Repeatedly, however, it had been suggested that it +might disseminate disease. We have seen that as far back as the +sixteenth century, Mercurialis suggested that it was the agent in the +spread of bubonic plague, and in 1658, Kircher reiterated this view. +In 1871, Leidy expressed the opinion that flies were probably a means +of communicating contagious diseases to a greater degree than was +generally suspected. From what he had observed regarding gangrene +in hospitals, he thought flies should be carefully excluded from +wounds. In the same year, the editor of the <i>London Lancet</i>, referring +to the belief that they play a useful rôle in purifying the air said, +"Far from looking upon them as dipterous angels dancing attendance +on Hygeia, regard them rather in the light of winged sponges spreading +hither and thither to carry out the foul behests of Contagion."</p> + +<p>These suggestions attracted little attention from medical men, for +it is only within very recent years that the charges have been supported +by direct evidence. Before considering this evidence, it is<span class="pagenum"><a name="Page_145" id="Page_145">[Pg 145]</a></span> +necessary that we define what is meant by "house-fly" and that we +then consider the life-history of the insect.</p> + +<p>There are many flies which are occasionally to be found in houses, +but according to various counts, from 95 per cent to 99 per cent of +these in warm weather in the Eastern United States belong to the +one species <i>Musca domestica</i> (<a href="#Fig_108">fig. 108</a>). This is the dominant house-fly +the world over and is the one which merits the name. It has been +well characterized by Schiner (1864), whose description has been +freely translated by Hewitt, as follows:</p> + +<p>"Frons of male occupying a fourth part of the breadth of the head. +Frontal stripe of female narrow in front, so broad behind that it +entirely fills up the width of the frons. The dorsal region of the +thorax dusty grey in color with four equally broad longitudinal +stripes. Scutellum gray with black sides. The light regions of +the abdomen yellowish, transparent, the darkest parts at least at +the base of the ventral side yellow. The last segment and a dorsal +line blackish brown. Seen from behind and against the light, the +whole abdomen shimmering yellow, and only on each side of the +dorsal line on each segment a dull transverse band. The lower part +of the face silky yellow, shot with blackish brown. Median stripe +velvety black. Antennæ brown. Palpi black. Legs blackish +brown. Wings tinged with pale gray with yellowish base. The +female has a broad velvety back, often reddishly shimmering frontal +stripe, which is not broader at the anterior end than at the bases of +the antennæ, but become so very much broader above that the light +dustiness of the sides is entirely obliterated. The abdomen gradually +becoming darker. The shimmering areas on the separate segments +generally brownish. All the other parts are the same as in +the male."</p> + +<p>The other species of flies found in houses in the Eastern United +States which are frequently mistaken for the house or typhoid fly +may readily be distinguished by the characters of the following key:</p> + +<div class="blockquot"><div class="hanging"> +<p><i>a.</i> Apical cell (<span title="for Rs read R_5"><a name="AC_9" id="AC_9"></a>R<sub>s</sub></span>) of the wide wing open, i.e., the bounding veins +parallel or divergent (<a href="#Fig_100">fig. 100</a>). Their larvæ are flattened, the +intermediate body segments each fringed with fleshy, more or +less spinose, processes. <span class="rightalign"><i>Fannia</i></span></p> + +<div class="hanging"> +<p>b. Male with the sides of the second and third abdominal segments +translucent yellowish. The larva with three pairs +of nearly equal spiniferous appendages on each segment,<span class="pagenum"><a name="Page_146" id="Page_146">[Pg 146]</a></span> +arranged in a longitudinal series and in addition two pairs +of series of smaller processes (<a href="#Fig_100">fig. 100</a>) <span class="rightalign"><i>F. canicularis</i></span></p> + +<p>bb. Male with blackish abdomen, middle tibia with a tubercle +beyond the middle. The larva with spiniferous appendages +of which the dorsal and ventral series are short, the +lateral series long and feathered (<a href="#Fig_101">fig. 101</a>) <span class="rightalign"><i>F. scalaris</i></span></p> +</div> + +<p>aa. Apical cell (R) of the wing more or less narrowed in the +margin; i. e., the bounding veins more or less converging +(<a href="#Fig_108">fig. 108</a>).</p> + +<div class="hanging"> +<p>b. The mouth-parts produced and pointed, fitted for piercing.</p> + +<div class="hanging"> +<p>c. Palpi much shorter than the proboscis; a brownish gray +fly, its thorax with three rather broad whitish stripes; +on each border of the middle stripe and on the mesal +borders of the lateral stripes is a blackish brown line. +Abdomen yellowish brown; on the second, third and +fourth segments are three brown spots which may be +faint or even absent. The larvæ live in dung. The +stable-fly (<a href="#Fig_110">fig. 110</a>) <span class="rightalign"><i>Stomoxys calcitrans</i></span></p> + +<p>cc. Palpi nearly as long as the proboscis. Smaller species +than the house-fly. The horn-fly (<a href="#Fig_167">fig. 167</a>) +<span class="rightalign"><i>Hæmatobia irritans</i></span></p> +</div> + +<p>bb. Mouth-parts blunt, fitted for lapping.</p> + +<div class="hanging"> +<p>c. Thorax, particularly on the sides and near the base of the +wings with soft golden yellow hairs among the bristles. +This fly is often found in the house in very early spring +or even in the winter. The cluster-fly, <span class="rightalign"><i>Pollenia rudis</i></span></p> + +<p>cc. Thorax without golden yellow hairs among the bristles.</p> + +<div class="hanging"> +<p>d. The last segment of the vein M with an abrupt +angle. (<a href="#Fig_108">fig. 108</a>). The larvæ live in manure, +etc. <span class="rightalign">House-fly, <i>Musca domestica</i></span></p> + +<p>dd. The last segment of vein M with a broad, gentle +curve (<a href="#Fig_102">fig. 102</a>).</p> + +<div class="hanging"> +<p>e. Eyes microscopically hairy; each abdominal +segment with two spots. Larvæ in dung. +<span class="rightalign"><i>Myiospila meditabunda</i></span></p> + +<p>ee. Eyes bare; abdomen gray and brown marbled. +<span class="rightalign"><i>Muscina</i></span></p> + +<div class="hanging"> +<p>f. With black legs and palpi. <span class="rightalign"><i>M. assimilis</i></span><span class="pagenum"><a name="Page_147" id="Page_147">[Pg 147]</a></span></p> + +<p>ff. With legs more or less yellowish; palpi +yellow. Larvæ in decaying vegetable +substances, dung, etc. <span class="rightalign"><i>M. stabulans</i></span></p> +</div></div></div></div></div></div></div> + +<p>It is almost universally believed that the adults of <i>Musca domestica</i> +hibernate, remaining dormant throughout the winter in attics, +around chimneys, and in sheltered but cold situations. This belief +has been challenged by Skinner (1913), who maintains that all the +adult flies die off during the fall and early winter and that the species +is carried over in the pupal stage, and in no other way. The cluster-fly, +<i>Pollenia rudis</i>, undoubtedly does hibernate in attics and similar +situations and is often mistaken for the house-fly. In so far as +concerns <i>Musca domestica</i>, the important question as to hibernation +in the adult stage is an open one. Many observations by one of the +writers (Johannsen) tend to confirm Dr. Skinner's conclusion, in so +far as it applies to conditions in the latitude of New York State. +Opposed, is the fact that various experimenters, notably Hewitt +(1910) and Jepson (1909) wholly failed to carry pupæ through the +winter.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_108" id="Fig_108"></a> +<a href="images/f108-full.png"><img src="images/f108.png" width="500" height="259" alt="108. The house or typhoid fly (Musca domestica (×4)). After Howard." title="108. The house or typhoid fly (Musca domestica (4×)). After Howard." /></a> +<span class="caption">108. The house or typhoid fly (Musca domestica (4×)). After Howard.</span> +</div> + +<p>The house-fly breeds by preference in horse manure. Indeed, +Dr. Howard, whose extensive studies of the species especially qualify +him for expressing an opinion on the subject, has estimated that under +ordinary city and town conditions, more than ninety per cent of the +flies present in houses have come from horse stables or their vicinity. +They are not limited to such localities, by any means, for it has been +found that they would develop in almost any fermenting organic +substance. Thus, they have been bred from pig, chicken, and cow<span class="pagenum"><a name="Page_148" id="Page_148">[Pg 148]</a></span> +manure, dirty waste paper, decaying vegetation, decaying meat, +slaughter-house refuse, sawdust-sweepings, and many other sources. +A fact which makes them especially dangerous as disease-carriers +is that they breed readily in human excrement.</p> + +<p>The eggs are pure white, elongate ovoid, somewhat broader at +the anterior end. They measure about one millimeter (1-25 inch) +in length. They are deposited in small, irregular clusters, one +hundred and twenty to one hundred and fifty from a single fly. A +female may deposit as many as four batches in her life time. The +eggs hatch in from eight to twenty-four hours.</p> + +<p>The newly hatched larva, or maggot (<a href="#Fig_108">fig. 108</a>), measures about two +millimeters (1-12 inch) in length. It is pointed at the head end and +blunt at the opposite end, where the spiracular openings are borne. +It grows rapidly, molts three times and reaches maturity in from six +to seven days, under favorable conditions.</p> + +<p>The pupal stage, like that of related flies, is passed in the old +larval skin which, instead of being molted, becomes contracted and +heavily chitinized, forming the so-called <i>puparium</i> (<a href="#Fig_108">fig. 108</a>). The +pupal stage may be completed in from three to six days.</p> + +<p>Thus during the warm summer months a generation of flies may +be produced in ten to twelve days. Hewitt at Manchester, England, +found the minimum to be eight days but states that larvæ bred in +the open air in horse manure which had an average daily temperature +of 22.5° C., occupied fourteen to twenty days in their development, +according to the air temperature.</p> + +<p>After emergence, a period of time must elapse before the fly is +capable of depositing eggs. This period has been tuned the <i>preoviposition</i> +period. Unfortunately we have few exact data regarding +this period. Hewitt found that the flies became sexually mature in +ten to fourteen days after their emergence from the pupal state and +four days after copulation they began to deposit their eggs; in other +words the preoviposition stage was fourteen days or longer. Griffith +(1908) found this period to be ten days. Dr. Howard believes that +the time "must surely be shorter, and perhaps much shorter, under +midsummer conditions, and in the freedom of the open air." He +emphasizes that the point is of great practical importance, since it is +during this period that the trapping and other methods of destroying +the adult flies, will prove most useful.</p> + +<p>Howard estimates that there may be nine generations of flies a +year under outdoor conditions in places comparable in climate to<span class="pagenum"><a name="Page_149" id="Page_149">[Pg 149]</a></span> +Washington. The number may be considerably increased in warmer +climates.</p> + +<p>The rate at which flies may increase under favorable conditions is +astounding. Various writers have given estimates of the numbers of +flies which may develop as the progeny of a single individual, providing +all the eggs and all the individual flies survived. Thus, Howard +estimates that from a single female, depositing one hundred and +twenty eggs on April 15th, there may be by September 10th, +5,598,720,000,000 adults. Fortunately, living forms do not produce +in any such mathematical manner and the chief value of the figures +is to illustrate the enormous struggle for existence which is constantly +taking place in nature.</p> + +<p>Flies may travel for a considerable distance to reach food and +shelter, though normally they pass to dwellings and other sources +of food supply in the immediate neighborhood of their breeding +places. Copeman, Howlett and Merriman (1911) marked flies by +shaking them in a bag containing colored chalk. Such flies were +repeatedly recovered at distances of eight to one thousand yards +and even at a distance of seventeen hundred yards, nearly a mile.</p> + +<p>Hindle and Merriman (1914) continued these experiments on a +large scale at Cambridge, England. They "do not think it likely +that, as a rule, flies travel more than a quarter of a mile in thickly-housed +areas." In one case a single fly was recovered at a distance +of 770 yards but a part of this distance was across open fen-land. +The surprising fact was brought out that flies tend to travel either +<i>against</i> or across the wind. The actual direction followed may be +determined either directly by the action of the wind (positive anemotropism), +or indirectly owing to the flies being attracted by any odor +that it may convey from a source of food. They conclude that it is +likely that the chief conditions favoring the disposal of flies are fine +weather and a warm temperature. The nature of the locality is +another considerable factor. Hodge (1913) has shown that when +aided by the wind they may fly to much greater distances over the +water. He reports that at Cleveland, Ohio, the cribs of the water +works, situated a mile and a quarter, five miles, and six miles out in +Lake Erie are invaded by a regular plague of flies when the wind +blows from the city. Investigation showed that there was absolutely +nothing of any kind in which flies could breed on the crib.</p> + +<p>The omnivorous habits of the house-fly are matters of everyday +observation. From our view point, it is sufficient to emphasize<span class="pagenum"><a name="Page_150" id="Page_150">[Pg 150]</a></span> +that from feeding on excrement, on sputum, on open sores, or on +putrifying matter, the flies may pass to the food or milk upon the table +or to healthy mucous membranes, or uncontaminated wounds. +There is nothing in its appearance to tell whether the fly that comes +blithely to sup with you is merely unclean, or whether it has just +finished feeding upon dejecta teeming with typhoid bacilli.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_109" id="Fig_109"></a> +<a href="images/f109-full.png"><img src="images/f109.png" width="350" height="350" alt="109. Pulvillus of foot of house-fly, showing glandular hairs." title="109. Pulvillus of foot of house-fly, showing glandular hairs." /></a> +<span class="caption">109. Pulvillus of foot of house-fly, showing glandular hairs.</span> +</div> + +<p>The method of feeding of the house-fly has an important bearing +on the question of its ability to transmit pathogenic organisms. +Graham-Smith (1910) has shown that when feeding, flies frequently +moisten soluble substances with "vomit" which is regurgitated from +the crop. This is, of course, loaded with bacteria from previous +food. When not sucked up again these drops of liquid dry, and produce +round marks with an opaque center and rim and an intervening +less opaque area. Fly-specks, then, consist of both vomit spots +and feces. Graham-Smith shows a photograph of a cupboard window +where, on an area six inches square, there were counted eleven hundred +and two vomit marks and nine fecal deposits.<span class="pagenum"><a name="Page_151" id="Page_151">[Pg 151]</a></span></p> + +<p>From a bacteriologist's viewpoint a discussion of the possibility +of a fly's carrying bacteria would seem superfluous. Any exposed +object, animate or inanimate, is contaminated by bacteria and will +transfer them if brought into contact with suitable culture media, +whether such substance be food, or drink, open wounds, or the sterile +culture media of the laboratory. A needle point may convey enough +germs to produce disease. Much more readily may the house-fly +with its covering of hairs and its sponge-like pulvilli (<a href="#Fig_109">fig. 109</a>) pick +up and transfer bits of filth and other contaminated material.</p> + +<p>For popular instruction this inevitable transfer of germs by the +house-fly is strikingly demonstrated by the oft copied illustration +of the tracks of a fly on a sterile culture plate. Two plates of gelatine +or, better, agar medium are prepared. Over one of these a fly +(with wings clipped) is allowed to walk, the other is kept as a check. +Both are put aside at room temperature, to be examined after twenty-four +to forty-eight hours. At the end of that time, the check plate +is as clear as ever, the one which the fly has walked is dotted with +colonies of bacteria and fungi. The value in the experiment consists +in emphasizing that by this method we merely render visible what is +constantly occurring in nature.</p> + +<p>A comparable experiment which we use in our elementary laboratory +work is to take three samples of <i>clean</i> (preferably, sterile) fresh +milk in sterile bottles. One of them is plugged with a pledget of +cotton, into the second is dropped a fly from the laboratory and into +the third is dropped a fly which has been caught feeding upon garbage +or other filth. After a minute or two the flies are removed and +the vials plugged as was number one. The three are then set aside +at room temperature. When examined after twenty-four hours +the milk in the first vial is either still sweet or has a "clean" sour odor; +that of the remaining two is very different, for it has a putrid odor, +which is usually more pronounced in the case of sample number +three.</p> + +<p>Several workers have carried out experiments to determine the +number of bacteria carried by flies under natural conditions. One +of the most extended and best known of these is the series by Esten +and Mason (1908). These workers caught flies from various sources +in a sterilized net, placed them in a sterile bottle and poured over +them a known quantity of sterilized water, in which they were shaken +so as to wash the bacteria from their bodies. They found the number +of bacteria on a single fly to range from 550 to 6,600,000. Early in<span class="pagenum"><a name="Page_152" id="Page_152">[Pg 152]</a></span> +the fly season the numbers of bacteria on flies are comparatively +small, while later the numbers are comparatively very large. The +place where flies live also determines largely the numbers that they +carry. The lowest number, 550, was from a fly caught in the +bacteriological laboratory, the highest number, 6,600,000 was the +average from eighteen swill-barrel flies. Torrey (1912) made examination +of "wild" flies from a tenement house district of New York +City. He found "that the surface contamination of these 'wild' +flies may vary from 570 to 4,400,000 bacteria per insect, and the +intestinal bacterial content from 16,000 to 28,000,000."</p> + +<p>Less well known in this country is the work of Cox, Lewis, and +Glynn (1912). They examined over four hundred and fifty naturally +infected house-flies in Liverpool during September and early October. +Instead of washing the flies they were allowed to swim on the surface +of sterile water for five, fifteen, or thirty minutes, thus giving natural +conditions, where infection occurs from vomit and dejecta of the +flies, as well as from their bodies. They found, as might be expected, +that flies from either insanitary or congested areas of the city contain +far more bacteria than those from the more sanitary, less congested, +or suburban areas. The number of aerobic bacteria from the former +varied from 800,000 to 500,000,000 per fly and from the latter from +21,000 to 100,000. The number of intestinal forms conveyed by +flies from insanitary or congested areas was from 10,000 to 333,000,000 +as compared with from 100 to 10,000 carried by flies from the more +sanitary areas.</p> + +<p>Pathogenic bacteria and those allied to the food poisoning group +were only obtained from the congested or moderately congested +areas and not from the suburban areas, where the chances of infestation +were less.</p> + +<p>The interesting fact was brought out that flies caught in milk +shops apparently carry and obtain more bacteria than those from +other shops with exposed food in a similar neighborhood. The +writers explained this as probably due to the fact that milk when +accessible, especially during the summer months, is suitable culture +medium for bacteria, and the flies first inoculate the milk and later +reinoculate themselves, and then more of the milk, so establishing a +vicious circle.</p> + +<p>They conclude that in cities where food is plentiful flies rarely +migrate from the locality in which they are bred, and consequently +the number of bacteria which they carry depends upon the general<span class="pagenum"><a name="Page_153" id="Page_153">[Pg 153]</a></span> +standard of cleanliness in that locality. Flies caught in a street of +modern, fairly high class, workmen's dwellings forming a sanitary +oasis in the midst of a slum area, carried far less bacteria than those +caught in the adjacent neighborhood.</p> + +<p>Thus, as the amount of dirt carried by flies in any particular +locality, measured in the terms of bacteria, bears a definite relation +to the habits of the people and to the state of the streets, it demonstrates +the necessity of efficient municipal and domestic cleanliness, +if the food of the inhabitants is to escape pollution, not only with +harmless but also with occasional pathogenic bacteria.</p> + +<p>The above cited work is of a general nature, but, especially in +recent years, many attempts have been made to determine more +specifically the ability of flies to transmit pathogenic organisms. +The critical reviews of Nuttall and Jepson (1909), Howard (1911), +and Graham-Smith (1913) should be consulted by the student of +the subject. We can only cite here a few of the more striking experiments.</p> + +<p>Celli (1888) fed flies on pure cultures of <i>Bacillus typhosus</i> and declared +that he was able to recover these organisms from the intestinal +contents and excrement.</p> + +<p>Firth and Horrocks (1902), cited by Nuttall and Jepson, "kept +<i>Musca domestica</i> (also bluebottles) in a large box measuring 4 × 3 × 3 +feet, with one side made of glass. They were fed on material +contaminated with cultures of <i>B. typhosus</i>. Agar plates, litmus, +glucose broth and a sheet of clean paper were at the same time +exposed in the box. After a few days the plates and broth were +removed and incubated with a positive result." Graham-Smith +(1910) "carried out experiments with large numbers of flies kept +in gauze cages and fed for eight hours on emulsions of <i>B. typhosus</i> +in syrup. After that time the infested syrup was removed and the +flies were fed on plain syrup. <i>B. typhosus</i> was isolated up to 48 +hours (but not later) from emulsions of their feces and from plates +over which they walked."</p> + +<p>Several other workers, notably Hamilton (1903), Ficker (1903), +Bertarelli (1910) Faichnie (1909), and Cochrane (1912), have isolated +<i>B. typhosus</i> from "wild" flies, naturally infected. The papers +of Faichnie and of Cochrane we have not seen, but they are quoted +in <i>extenso</i> by Graham-Smith (1913).</p> + +<p>On the whole, the evidence is conclusive that typhoid germs not +only may be accidentally carried on the bodies of house-flies but<span class="pagenum"><a name="Page_154" id="Page_154">[Pg 154]</a></span> +may pass through their bodies and be scattered in a viable condition +in the feces of the fly for at least two days after feeding. Similar, +results have been reached in experiments with cholera, tuberculosis +and yaws, the last-mentioned being a spirochæte disease. Darling +(1913) has shown that murrina, a trypanosome disease of horses +and mules in the Canal zone is transmitted by house-flies which feed +upon excoriated patches of diseased animals and then pass to cuts +and galls of healthy animals.</p> + +<p>Since it is clear that flies are abundantly able to disseminate +viable pathogenic bacteria, it is important to consider whether they +have access to such organisms in nature. A consideration of the +method of spread of typhoid will serve to illustrate the way in which +flies may play an important rôle.</p> + +<p>Typhoid fever is a specific disease caused by <i>Bacillus typhosus</i>, +and by it alone. The causative organism is to be found in the excrement +and urine of patients suffering from the disease. More than +that, it is often present in the dejecta for days, weeks, or even months +and years, after the individual has recovered from the disease. +Individuals so infested are known as "typhoid carriers" and they, +together with those suffering from mild cases, or "walking typhoid," +are a constant menace to the health of the community in which they +are found.</p> + +<p>Human excrement is greedily visited by flies, both for feeding and +for ovipositing. The discharges of typhoid patients, or of chronic +"carriers," when passed in the open, in box privies, or camp latrines, +or the like, serve to contaminate myriads of the insects which may +then spread the germ to human food and drink. Other intestinal +diseases may be similarly spread. There is abundant epidæmiological +evidence that infantile diarrhœa, dysentery, and cholera may be +so spread.</p> + +<p>Stiles and Keister (1913) have shown that spores of <i>Lamblia +intestinalis</i>, a flagellate protozoan living in the human intestine, +may be carried by house-flies. Though this species is not normally +pathogenic, one or more species of <i>Entamœba</i> are the cause of a type +of a highly fatal tropical dysentery. Concerning it, and another +protozoan parasite of man, they say, "If flies can carry <i>Lamblia</i> +spores measuring 10 to 7µ, and bacteria that are much smaller, and +particles of lime that are much larger, there is no ground to assume +that flies may not carry <i>Entamœba</i> and <i>Trichomonas</i> spores."<span class="pagenum"><a name="Page_155" id="Page_155">[Pg 155]</a></span></p> + +<p>Tuberculosis is one of the diseases which it is quite conceivable +may be carried occasionally. The sputum of tubercular patients +is very attractive to flies, and various workers, notably Graham-Smith, +have found that <i>Musca domestica</i> may distribute the bacillus +for several days after feeding on infected material.</p> + +<p>A type of purulent opthalmia which is very prevalent in Egypt +is often said to be carried by flies. Nuttall and Jepson (1909) +consider that the evidence regarding the spread of this disease by +flies is conclusive and that the possibility of gonorrhœal secretions +being likewise conveyed cannot be denied.</p> + +<p>Many studies have been published, showing a marked agreement +between the occurrence of typhoid and other intestinal diseases +and the prevalence of house-flies. The most clear-cut of these are +the studies of the Army Commission appointed to investigate the +cause of epidemics of enteric fever in the volunteer camps in the +Southern United States during the Spanish-American War. Though +their findings as presented by Vaughan (1909), have been quoted +very many times, they are so germane to our discussion that they +will bear repetition:</p> + +<p>"Flies swarmed over infected fecal matter in the pits and fed +upon the food prepared for the soldiers in the mess tents. In some +instances where lime had recently been sprinkled over the contents +of the pits, flies with their feet whitened with lime were seen walking +over the food." Under such conditions it is no wonder that "These +pests had inflicted greater loss upon American soldiers than the arms +of Spain."</p> + +<p>Similar conditions prevailed in South Africa during the Boer War. +Seamon believes that very much of the success of the Japanese in +their fight against Russia was due to the rigid precautions taken to +prevent the spread of disease by these insects and other means.</p> + +<p>Veeder has pointed out that the characteristics of a typical fly-borne +epidemic of typhoid are that it occurs in little neighborhood +epidemics, extending by short leaps from house to house, without +regard to water supply or anything else in common. It tends to +follow the direction of prevailing winds (cf. the conclusions of Hindle +and Merriman). It occurs during warm weather. Of course, when +the epidemic is once well under way, other factors enter into its spread.</p> + +<p>In general, flies may be said to be the chief agency in the spread of +typhoid in villages and camps. In cities with modern sewer systems +they are less important, though even under the best of such conditions,<span class="pagenum"><a name="Page_156" id="Page_156">[Pg 156]</a></span> +they are important factors. Howard has emphasized that in +such cities there are still many uncared-for box privies and that, in +addition, the deposition of feces overnight in uncared-for waste lots +and alleys is common.</p> + +<p>Not only unicellular organisms, such as bacteria and protozoa, +but also the eggs, embryos and larvæ of parasitic worms have been +found to be transported by house-flies. Ransom (1911) has found +that <i>Habronema muscæ</i>, a nematode worm often found in adult flies, +is the immature stage of a parasite occurring in the stomach of the +horse. The eggs or embryos passing out with the feces of the horse, +are taken up by fly larvæ and carried over to the imago stage.</p> + +<p>Grassi (1883), Stiles (1889), Calandruccio (1906), and especially +Nicoll (1911), have been the chief investigators of the ability of +house-flies to carry the ova and embryos of human intestinal parasites. +Graham-Smith (1913) summarizes the work along this line as follows:</p> + +<p>"It is evident from the investigations that have been quoted that +house-flies and other species are greatly attracted to the ova of +parasitic worms contained in feces and other materials, and make +great efforts to ingest them. Unless the ova are too large they often +succeed, and the eggs are deposited uninjured in their feces, in some +cases up to the third day at least. The eggs may also be carried on +their legs or bodies. Under suitable conditions, food and fluids +may be contaminated with the eggs of various parasitic worms by flies, +and in one case infection of the human subject has been observed. +Feces containing tape-worm segments may continue to be a source of +infection for as long as a fortnight. Up to the present, however, +there is no evidence to show what part flies play in the dissemination +of parasitic worms under natural conditions."</p> + +<p>Enough has been said to show that the house-fly must be dealt +with as a direct menace to public health. Control measures are +not merely matters of convenience but are of vital importance.</p> + +<p>Under present conditions the speedy elimination of the house-fly +is impossible and the first thing to be considered is methods of protecting +food and drink from contamination. The first of these +methods is the thorough screening of doors and windows to prevent +the entrance of flies. In the case of kitchen doors, the flies, attracted +by odors, are likely to swarm onto the screen and improve the first +opportunity for gaining an entrance. This difficulty can be largely +avoided by screening-in the back porch and placing the screen door +at one end rather than directly before the door.<span class="pagenum"><a name="Page_157" id="Page_157">[Pg 157]</a></span></p> + +<p>The use of sticky fly paper to catch the pests that gain entrance +to the house is preferable to the various poisons often used. Of the +latter, formalin (40 per cent formaldehyde) in the proportion of two +tablespoonfuls to a pint of water is very efficient, if all other liquids +are removed or covered, so that the flies must depend on the formalin +for drink. The mixture is said to be made more attractive by the +addition of sugar or milk, though we have found the plain solution +wholly satisfactory, under proper conditions. It should be emphasized +that this formalin mixture is not perfectly harmless, as so +often stated. There are on record cases of severe and even fatal +poisoning from the accidental drinking of solutions.</p> + +<p>When flies are very abundant in a room they can be most readily +gotten rid of by fumigation with sulphur, or by the use of pure +pyrethrum powder either burned or puffed into the air. Herrick +(1913) recommends the following method: "At night all the doors +and windows of the kitchen should be closed; fresh powder should +be sprinkled over the stove, on the window ledges, tables, and in the +air. In the morning flies will be found lying around dead or stupified. +They may then be swept up and burned." This method has proved +very efficaceous in some of the large dining halls in Ithaca.</p> + +<p>The writers have had little success in fumigating with the vapors +of carbolic acid, or carbolic acid and gum camphor, although these +methods will aid in driving flies from a darkened room.</p> + +<p>All of these methods are but makeshifts. As Howard has so well +put it, "the truest and simplest way of attacking the fly problem +is to prevent them from breeding, by the treatment or abolition of +all places in which they can breed. To permit them to breed undisturbed +and in countless numbers, and to devote all our energy to +the problem of keeping them out of our dwellings, or to destroy them +after they have once entered in spite of all obstacles, seems the +wrong way to go about it."</p> + +<p>We have already seen that <i>Musca domestica</i> breeds in almost any +fermenting organic material. While it prefers horse manure, it +breeds also in human feces, cow dung and that of other animals, +and in refuse of many kinds. To efficiently combat the insect, +these breeding places must be removed or must be treated in some +such way as to render them unsuitable for the development of the +larvæ. Under some conditions individual work may prove effective, +but to be truly efficient there must be extensive and thorough coöperative +efforts.<span class="pagenum"><a name="Page_158" id="Page_158">[Pg 158]</a></span></p> + +<p>Manure, garbage, and the like should be stored in tight receptacles +and carted away at least once a week. The manure may be carted +to the fields and spread. Even in spread manure the larvæ may continue +their development. Howard points out that "it often happens +that after a lawn has been heavily manured in early summer the +occupants of the house will be pestered with flies for a time, but +finding no available breeding place these disappear sooner or later. +Another generation will not breed in the spread manure."</p> + +<p>Hutchinson (1914) has emphasized that the larvæ of houseflies +have deeply engrained the habit of migrating in the prepupal +stage and has shown that this offers an important point of attack +in attempts to control the pest. He has suggested that maggot +traps might be developed into an efficient weapon in the warfare +against the house-fly. Certain it is that the habit greatly simplifies the +problem of treating the manure for the purpose of killing the larvæ.</p> + +<p>There have been many attempts to find some cheap chemical +which would destroy fly larvæ in horse manure without injuring the +bacteria or reducing the fertilizing values of the manure. The literature +abounds in recommendations of kerosene, lime, chloride of lime, +iron sulphate, and other substances, but none of them <span title="for have read has"><a name="AC_10" id="AC_10"></a>have</span> met the +situation. The whole question has been gone into thoroughly by +Cook, Hutchinson and Scales (1914), who tested practically all of the +substances which have been recommended. They find that by far +the most effective, economical, and practical of the substances is +borax in the commercial form in which it is available throughout the +country.</p> + +<p>"Borax increases the water-soluble nitrogen, ammonia and alkalinity +of manure and apparently does not permanently injure the +bacterial flora. The application of manure treated with borax at the +rate of 0.62 pound per eight bushels (10 cubic feet) to soil does not +injure the plants thus far tested, although its cumulative effect, if +any, has not been determined."</p> + +<p>As their results clearly show that the substances so often recommended +are inferior to borax, we shall quote in detail their directions +for treating manure so as to kill fly eggs and maggots.</p> + +<p>"Apply 0.62 pound borax or 0.75 pound calcined colemanite to +every 10 cubic feet (8 bushels) of manure immediately on its removal +from the barn. Apply the borax particularly around the outer +edges of the pile with a flour sifter or any fine sieve, and sprinkle two +or three gallons of water over the borax-treated manure.<span class="pagenum"><a name="Page_159" id="Page_159">[Pg 159]</a></span></p> + +<p>"The reason for applying the borax to the fresh manure immediately +after its removal from the stable is that the flies lay their eggs +on the fresh manure, and borax, when it comes in contact with the +eggs, prevents their hatching. As the maggots congregate at the +outer edge of the pile, most of the borax should be applied there. +The treatment should be repeated with each addition of fresh manure, +but when the manure is kept in closed boxes, less frequent applications +will be sufficient. When the calcined colemanite is available, +it may be used at the rate of 0.75 pound per 10 cubic feet of manure, +and is a cheaper means of killing the maggots. In addition to the +application of borax to horse manure to kill fly larvæ, it may be +applied in the same proportion to other manures, as well as to refuse +and garbage. Borax may also be applied to the floors and crevices in +barns, stables, markets, etc., as well as to street sweepings, and water +should be added as in the treatment of horse manure. After estimating +the amount of material to be treated and weighing the necessary +amount of borax, a measure may be used which will hold the proper +amount, thus avoiding the subsequent weighings.</p> + +<p>"While it can be safely stated that no injurious action will follow +the application of manure treated with borax at the rate of 0.62 +pound for eight bushels, or even larger amounts in the case of some +plants, nevertheless the borax-treated manure has not been studied +in connection with the growth of all crops, nor has its cumulative +effect been determined. It is therefore recommended that not more +than 15 tons per acre of the borax-treated manure should be applied +to the field. As truckmen use considerably more than this amount, +it is suggested that all cars containing borax-treated manure be so +marked, and that public-health officials stipulate in their directions +for this treatment that not over 0.62 pound for eight bushels of manure +be used, as it has been shown that larger amounts of borax will +injure most plants. It is also recommended that all public-health +officials and others, in recommending the borax treatment for killing +fly eggs and maggots in manure, warn the public against the +injurious effects of large amounts of borax on the growth of plants."</p> + +<p>"The amount of manure from a horse varies with the straw or +other bedding used, but 12 or 15 bushels per week represent the +approximate amount obtained. As borax costs from five to six +cents per pound in 100-pound lots in Washington, it will make the +cost of the borax practically one cent per horse, per day. And if +calcined colemanite is purchased in large shipments the cost should +be considerably less."<span class="pagenum"><a name="Page_160" id="Page_160">[Pg 160]</a></span></p> + +<p>Hodge (1910) has approached the problem of fly extermination +from another viewpoint. He believes that it is practical to trap +flies out of doors during the preoviposition period, when they are +sexually immature, and to destroy such numbers of them that the +comparatively few which survive will not be able to lay eggs in sufficient +numbers to make the next generation a nuisance. To the end +of capturing them in enormous numbers he has devised traps to be +fitted over garbage cans, into stable windows, and connected with the +kitchen window screens. Under some conditions this method of +attack has proved very satisfactory.</p> + +<p>One of the most important measures for preventing the spread +of disease by flies is the abolition of the common box privy. In +villages and rural districts this is today almost the only type to be +found. It is the chief factor in the spread of typhoid and other +intestinal diseases, as well as intestinal parasites. Open and exposed +to myriads of flies which not only breed there but which feed +upon the excrement, they furnish ideal conditions for spreading contamination. +Even where efforts are made to cover the contents +with dust, or ashes, or lime, flies may continue to breed unchecked. +Stiles and Gardner have shown that house-flies buried in a screened +stand-pipe forty-eight inches under sterile sand came to the surface. +Other flies of undetermined species struggled up through seventy-two +inches of sand.</p> + +<p>So great is the menace of the ordinary box privy that a number of +inexpensive and simple sanitary privies have been designed for use +where there are not modern sewer systems. Stiles and Lumsden +(1911) have given minute directions for the construction of one of the +best types, and their bulletin should be obtained by those interested.</p> + +<p>Another precaution which is of fundamental importance in +preventing the spread of typhoid, is that of disinfecting all discharges +from patients suffering with the disease. For this purpose, quick-lime +is the cheapest and is wholly satisfactory. In chamber vessels +it should be used in a quantity equal to that of the discharge to be +treated. It should be allowed to act for two hours. Air-slaked +lime is of no value whatever. Chloride of lime, carbolic acid, or +formalin may be used, but are more expensive. Other intestinal +diseases demand similar precautions.</p> + + +<p><a name="Stomoxys_calcitrans_the_stable-fly" id="Stomoxys_calcitrans_the_stable-fly"></a><b>Stomoxys calcitrans, the stable-fly</b>—It is a popular belief that +house-flies bite more viciously just before a rain. As a matter of<span class="pagenum"><a name="Page_161" id="Page_161">[Pg 161]</a></span> +fact, the true house-flies never bite, for their mouth-parts are not +fitted for piercing. The basis of the misconception is the fact that a +true biting fly, <i>Stomoxys calcitrans</i> (<a href="#Fig_110">fig. 110</a>), closely resembling the +house-fly, is frequently found in houses and may be driven in in +greater numbers by muggy weather. From its usual habitat this +fly is known as the "stable-fly" or, sometimes as the "biting house-fly."</p> + +<p><i>Stomoxys calcitrans</i> may be separated from the house-fly by the use +of the key on p. 145. It may be more fully characterized as follows:</p> + +<p>The eyes of the male are separated by a distance equal to one-fourth +of the diameter of the head, in the female by one-third. The +frontal stripe is black, the cheeks and margins of the orbits silvery-white. +The antennæ are black, the arista feathered on the upper +side only. The proboscis is black, slender, fitted for piercing and +projects forward in front of the head. The thorax is grayish, marked +by four conspicuous, more or less complete black longitudinal stripes; +the scutellum is paler; the macrochætæ are black. The abdomen is +gray, dorsally with three brown spots on the second and third segments +and a median spot on the fourth. These spots are more +pronounced in the female. The legs are black, the pulvilli distinct. +The wings are hyaline, the vein M<sub>1+2</sub> less sharply curved than in +the house-fly, the apical cell being thus more widely open (cf. <a href="#Fig_110">fig. 110</a>). +Length 7 mm.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_110" id="Fig_110"></a> +<a href="images/f110-full.png"><img src="images/f110.png" width="500" height="271" alt="110. Stomoxys calcitrans; adult, larva, puparium and details, (×5). After Howard." title="110. Stomoxys calcitrans; adult, larva, puparium and details, (×5). After Howard." /></a> +<span class="caption">110. Stomoxys calcitrans; adult, larva, puparium and details, (×5). After Howard.</span> +</div> + +<p>This fly is widely distributed, being found the world over. It was +probably introduced into the United States, but has spread to all<span class="pagenum"><a name="Page_162" id="Page_162">[Pg 162]</a></span> +parts of the country. Bishopp (1913) regards it as of much more +importance as a pest of domestic animals in the grain belt than elsewhere +in the United States. The life-history and habits of this +species have assumed a new significance since it has been suggested +that it may transmit the human diseases, infantile paralysis and +pellagra. In this country, the most detailed study of the fly is that +of Bishopp (1913) whose data regarding the life cycle are as follows:</p> + +<p>The eggs like those of the house-fly, are about one mm. +in length. Under a magnifying glass they show a distinct furrow +along one side. When placed on any moist substance they hatch +in from one to three days after being deposited.</p> + +<p>The larva or maggots (<a href="#Fig_110">fig. 110</a>) have the typical shape and actions +of most maggots of the Muscid group. They can be distinguished +from those of the house-fly as the stigma-plates are smaller, much +further apart, with the slits less sinuous. Development takes place +fairly rapidly when the proper food conditions are available and +the growth is completed within eleven to thirty or more days.</p> + +<p>The pupa (<a href="#Fig_110">fig. 110</a>), like that of related flies, undergoes its development +within the contracted and hardened last larval skin, or puparium. +This is elongate oval, slightly thicker towards the head end, +and one-sixth to one-fourth of an inch in length. The pupal stage +requires six to twenty days, or in cool weather considerably longer.</p> + +<p>The life-cycle of the stable-fly is therefore considerably longer +than that of <i>Musca domestica</i>. Bishopp found that complete +development might be undergone in nineteen days, but that the +average period was somewhat longer, ranging from twenty-one to +twenty-five days, where conditions are very favorable. The longest +period which he observed was forty-three days, though his finding +of full grown larvæ and pupæ in straw during the latter part of +March, in Northern Texas, showed that development may require +about three months, as he considered that these stages almost certainly +developed from eggs deposited the previous December.</p> + +<p>The favorite breeding place, where available, seems to be straw or +manure mixed with straw. It also breeds in great numbers in horse-manure, +in company with <i>Musca domestica</i>.</p> + +<p>Newstead considers that in England the stable-fly hibernates in +the pupal stage. Bishopp finds that in the southern part of the +United States there is no true hibernation, as the adults have been +found to emerge at various times during the winter. He believes +that in the northern United States the winter is normally passed<span class="pagenum"><a name="Page_163" id="Page_163">[Pg 163]</a></span> +in the larval and pupal stages, and that the adults which have been +observed in heated stables in the dead of winter were bred out in +refuse within the warm barns and were not hibernating adults.</p> + +<p>Graham-Smith (1913) states that although the stable-fly frequents +stable manure, it is probably not an important agent in +distributing the organisms of intestinal diseases. Bishopp makes the +important observation that "it has never been found breeding in +human excrement and does not frequent malodorous places, which +are so attractive to the house-fly. Hence it is much less likely to +carry typhoid and other germs which may be found in such places."</p> + +<p>Questions of the possible agency of <i>Stomoxys calcitrans</i> in the transmission +of infantile paralysis and of pellagra, we shall consider later.</p> + + +<p><a name="Other_arthropods_which_may_serve_as_simple_carriers_of_pathogenic_organisms" id="Other_arthropods_which_may_serve_as_simple_carriers_of_pathogenic_organisms"></a><b>Other arthropods which may serve as simple carriers of pathogenic +organisms</b>—It should be again emphasized that any insect which +has access to, and comes in contact with, pathogenic organisms +and then passes to the food, or drink, or the body of man, may serve +as a simple carrier of disease. In addition to the more obvious +illustrations, an interesting one is the previously cited case of the +transfer of <i>Dermatobia cyaniventris</i> by a mosquito (<a href="#Fig_81">fig. 81-84</a>). +Darling (1913) has shown that in the tropics, the omnipresent ants +may be important factors in the spread of disease.<span class="pagenum"><a name="Page_164" id="Page_164">[Pg 164]</a></span></p> + + + + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_VI" id="CHAPTER_VI"></a>CHAPTER VI</h2> + +<h2>ARTHROPODS AS DIRECT INOCULATORS OF DISEASE GERMS</h2> + + +<p>We have seen that any insect which, like the house-fly, has access +to disease germs and then comes into contact with the food or drink +of man, may serve to disseminate disease. Moreover, it has been +clearly established that a contaminated insect, alighting upon +wounded or abraded surfaces, may infect them. These are instances +of mere accidental, mechanical transfer of pathogenic organisms.</p> + +<p>Closely related are the instances of direct inoculation of disease +germs by insects and other arthropods. In this type, a blood-sucking +species not only takes up the germs but, passing to a healthy +individual, it inserts its contaminated mouth-parts and thus directly +inoculates its victim. In other words, the disease is transferred +just as blood poisoning may be induced by the prick of a contaminated +needle, or as the laboratory worker may inoculate an experimental +animal.</p> + +<p>Formerly, it was supposed that this method of the transfer of +disease by arthropods was a very common one and many instances +are cited in the earlier literature of the subject. It is, however, +difficult to draw a sharp line between such cases and those in which, +on the one hand, the arthropod serves as a mere passive carrier or, +on the other hand, serves as an essential host of the pathogenic +organism. More critical study of the subject has led to the belief +that the importance of the rôle of arthropods as direct inoculators +has been much overestimated.</p> + +<p>The principal reason for regarding this phase of the subject as +relatively unimportant, is derived from a study of the habits of the +blood-sucking species. It is found that, in general, they are intermittent +feeders, visiting their hosts at intervals and then abstaining +from feeding for a more or less extended period, while digesting their +meal. In the meantime, most species of bacteria or of protozoan +parasites with which they might have contaminated their mouth-parts, +would have perished, through inability to withstand drying.</p> + +<p>In spite of this, it must be recognized that this method of transfer +does occur and must be reckoned with in any consideration of the +relations of insects to disease. We shall first cite some general +illustrations and shall then discuss the rôle of fleas in the spreading +of bubonic plague, an illustration which cannot be regarded as typical, +since it involves more than mere passive carriage.<span class="pagenum"><a name="Page_165" id="Page_165">[Pg 165]</a></span></p> + + +<h3><a name="Some_Illustrations_of_Direct_Inoculation_of_Disease_Germs_by_Arthropods" id="Some_Illustrations_of_Direct_Inoculation_of_Disease_Germs_by_Arthropods"></a><span class="smcap">Some Illustrations of Direct Inoculation of Disease Germs +by Arthropods</span></h3> + +<p>In discussing poisonous arthropods, we have already emphasized +that species which are of themselves innocuous to man, may occasionally +introduce bacteria by their bite or sting and thus cause more or +less severe secondary symptoms. That such cases should occur, is +no more than is to be expected. The mouth-parts or the sting of +the insect are not sterile and the chances of their carrying pyogenic +organisms are always present.</p> + +<p>More strictly falling in the category of transmission of disease +germs by direct inoculation are the instances where the insect, or +related form, feeds upon a diseased animal and passes promptly to a +healthy individual which it infects. Of such a nature are the following:</p> + +<p>Various species of biting flies are factors in the dissemination of +anthrax, an infectious and usually fatal disease of animals and, +occasionally, of man. That the bacteria with which the blood of +diseased animals teem shortly before death might be transmitted +by such insects has long been contended, but the evidence in support +of the view has been unsatisfactory. Recently, Mitzmain (1914) +has reported a series of experiments which show conclusively that the +disease may be so conveyed by a horse-fly, <i>Tabanus striatus</i>, and by +the stable-fly, <i>Stomoxys calcitrans</i>.</p> + +<p>Mitzmain's experiments were tried with an artificially infected +guinea pig, which died of the disease upon the third day. The flies +were applied two and one-half hours, to a few minutes, before the +death of the animal. With both species the infection was successfully +transferred to healthy guinea pigs by the direct method, in +which the flies were interrupted while feeding on the sick animal. +The evidence at hand does not warrant the conclusion that insect +transmission is the rule in the case of this disease.</p> + +<p>The nagana, or tsetse-fly disease of cattle is the most virulent +disease of domestic animals in certain parts of Africa. It is caused +by a protozoan blood parasite, <i>Trypanosoma brucei</i>, which is conveyed +to healthy animals by the bite of <i>Glossina morsitans</i> and possibly +other species of tsetse-flies. The flies remain infective for +forty-eight hours after feeding on a diseased animal. The insect +also serves as an essential host of the parasite.</p> + +<p>Surra, a similar trypanosomiasis affecting especially horses and +mules, occurs in southern Asia, Malaysia, and the Philippines where<span class="pagenum"><a name="Page_166" id="Page_166">[Pg 166]</a></span> +the tsetse-flies are not to be found. It is thought to be spread by +various species of blood-sucking flies belonging to the genera <i>Stomoxys</i>, +<i>Hæmatobia</i>, and <i>Tabanus</i>. Mitzmain (1913) demonstrated that in +the Philippines it is conveyed mechanically by <i>Tabanus striatus</i>.</p> + +<p>The sleeping sickness of man, in Africa, has also been supposed +to be directly inoculated by one, or several, species of tsetse-flies. +It is now known that the fly may convey the disease for a short +time after feeding, but that there is then a latent period of from +fourteen to twenty-one days, after which it again becomes infectious. +This indicates that in the meantime the parasite has been undergoing +some phase of its life-cycle and that the fly serves as an intermediate +host. We shall therefore consider it more fully under that +grouping.</p> + +<p>These are a few of the cases of direct inoculation which may be +cited as of the simpler type. We shall next consider the rôle of the +flea in the dissemination of the bubonic plague, an illustration +complicated by the fact that the bacillus multiples within the insect +and may be indirectly inoculated.</p> + + +<h3><a name="The_Role_of_Fleas_in_the_Transmission_of_the_Plague" id="The_Role_of_Fleas_in_the_Transmission_of_the_Plague"></a><span class="smcap">The Rôle of Fleas in the Transmission of the Plague</span></h3> + +<p>The plague is a specific infectious disease caused by <i>Bacillus pestis</i>. +It occurs in several forms, of which the bubonic and the pneumonic +are the most common. According to Wyman, 80 per cent of the +human cases are of the bubonic type. It is a disease which, under +the name of oriental plague, the pest, or the black death, has ravaged +almost from time immemorial the countries of Africa, Asia, and +Europe. The record of its ravages are almost beyond belief. In 542 +A. D. it caused in one day ten thousand deaths in Constantinople. +In the 14th century it was introduced from the East and prevailed +throughout Armenia, Asia Minor, Egypt and Northern Africa and +Europe. Hecker estimates that one-fourth of the population of +Europe, or twenty-five million persons, died in the epidemic of that +century. From then until the 17th century it was almost constantly +present in Europe, the great plague of London, in 1665 killing 68,596 +out of a population of 460,000. Such an epidemic would mean for +New York City a proportionate loss of over 600,000 in a single year. +It is little wonder that in the face of such an appalling disaster suspicion +and credulity were rife and the wildest demoralization ensued.</p> + +<p>During the 14th century the Jews were regarded as responsible +for the disease, through poisoning wells, and were subjected to the<span class="pagenum"><a name="Page_167" id="Page_167">[Pg 167]</a></span> +most incredible persecution and torture. In Milan the visitation +of 1630 was credited to the so-called anointers,—men who were +supposed to spread the plague by anointing the walls with magic +ointment—and the most horrible tortures that human ingenuity +could devise were imposed on scores of victims, regardless of rank +or of public service (<a href="#Fig_112a">fig. 112, a</a>). Manzoni's great historical novel, +"The Betrothed" has well pictured conditions in Italy during this +period.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_111" id="Fig_111"></a> +<a href="images/f111-full.png"><img src="images/f111.png" width="350" height="217" alt="111. A contemporaneous engraving of the pest hospital in Vienna in 1679. +After Peters." title="111. A contemporaneous engraving of the pest hospital in Vienna in 1679. +After Peters." /></a> +<span class="caption">111. A contemporaneous engraving of the pest hospital in Vienna in 1679. +After Peters.</span> +</div> + +<p>In modern times the plague is confined primarily to warm climates, +a condition which has been brought about largely through general +improvement in sanitary conditions.</p> + +<p>At present, the hotbed of the disease is India, where there were +1,040,429 deaths in 1904 and where in a period of fifteen years, +ending with January 1912, there were over 15,000,000 deaths. The +reported deaths in that country for 1913 totaled 198,875.</p> + +<p>During the winter of 1910-11 there occurred in Manchuria and +North China a virulent epidemic of the pneumonic plague which +caused the death of nearly 50,000 people. The question as to its +origin and means of spread will be especially referred to later.</p> + +<div class="figcenter" style="width: 350px;"><a name="Fig_112a" id="Fig_112a"></a> +<a href="images/f112a-full.png"><img src="images/f112a.png" width="350" height="257" alt="112 a. A medieval method of combating the plague. The persecution of the anointers in Milan in 1630. From a +copy of "Il processi originale degli untori" in the library of Cornell University." title="112 a. A medieval method of combating the plague. The persecution of the anointers in Milan in 1630. From a +copy of "Il processi originale degli untori" in the library of Cornell University." /></a> +<span class="caption">112 a. A medieval method of combating the plague. The persecution of the anointers in Milan in 1630. From a +copy of "Il processi originale degli untori" in the library of Cornell University.</span> +</div><span class="pagenum"><a name="Page_168" id="Page_168">[Pg 168]</a></span> + +<p>Until recent years, the plague had not been known to occur in +the New World but there were outbreaks in Brazil and Hawaii in +1899, and in 1900 there occurred the first cases in San Francisco.<span class="pagenum"><a name="Page_169" id="Page_169">[Pg 169]</a></span> +In California there were 125 cases in the period 1900-04; three cases +in the next three years and then from May 1907 to March 1908, +during the height of the outbreak, 170 cases. Since that time there +have been only sporadic cases, the last case reported being in May +1914. Still more recent were the outbreaks in the Philippine Islands, +Porto Rico, and Cuba.</p> + +<p>On June 24, 1914, there was recognized a case of human plague +in New Orleans. The Federal Health Service immediately took +charge, and measures for the eradication of the disease were vigorously +enforced. Up to October 10, 1914 there had been reported +30 cases of the disease in man, and 181 cases of plague in rats.</p> + +<div class="figcenter" style="width: 450px;"> +<a href="images/f112b-full.png"><img src="images/f112b.png" width="450" height="225" alt="112 b. The modern method of combating the plague. A day's catch of rats in the fight +against plague in San Francisco. Courtesy of Review of Reviews." title="112 b. The modern method of combating the plague. A day's catch of rats in the fight +against plague in San Francisco. Courtesy of Review of Reviews." /></a> +<span class="caption">112 b. The modern method of combating the plague. A day's catch of rats in the fight +against plague in San Francisco. Courtesy of Review of Reviews.</span> +</div> + +<p>The present-day methods of combating bubonic plague are well +illustrated by the fight in San Francisco. Had it not been for the +strenuous and radical anti-plague campaign directed by the United +States Marine Hospital Service we might have had in our own +country an illustration of what the disease can accomplish. On what +newly acquired knowledge was this fight based?</p> + +<p>The basis was laid in 1894, when the plague bacillus was first +discovered. All through the centuries, before and during the Christian +era, down to 1894, the subject was enveloped in darkness and there +had been a helpless, almost hopeless struggle in ignorance on the part +of physicians, sanitarians, and public health officials against the +ravages of this dread disease. Now its cause, method of propagation +and means to prevent its spread are matters of scientific certainty.<span class="pagenum"><a name="Page_170" id="Page_170">[Pg 170]</a></span></p> + +<p>After the discovery of the causative organism, one of the first +advances was the establishment of the identity of human plague +and that of rodents. It had often been noted that epidemics of the +human disease were preceded by great epizootics among rats and +mice. So well established was this fact that with the Chinese, +unusual mortality among these rodents was regarded as foretelling +a visitation of the human disease. That there was more than an +accidental connection between the two was obvious when Yersin, +the discoverer of <i>Bacillus pestis</i>, announced that during an epidemic +the rats found dead in the houses and in the streets almost always +contain the bacillus in great abundance in their organs, and that many +of them exhibit veritable buboes.</p> + +<p>Once it was established that the diseases were identical, the attention +of the investigators was directed to a study of the relations +between that of rats and of humans, and evidence accumulated to +show that the bubonic plague was primarily a disease of rodents +and that in some manner it was conveyed from them to man.</p> + +<p>There yet remained unexplained the method of transfer from rat +to man. As long ago as the 16th century, Mercuralis suggested +that house-flies were guilty of disseminating the plague but modern +investigation, while blaming the fly for much in the way of spreading +disease, show that it is an insignificant factor in this case.</p> + +<p>Search for blood-sucking insects which would feed on both rodents +and man, and which might therefore be implicated, indicated that +the fleas most nearly met the conditions. At first it was urged that +rat fleas would not feed upon man and that the fleas ordinarily attacking +man would not feed upon rats. More critical study of the habits +of fleas soon showed that these objections were not well-founded. +Especially important was the evidence that soon after the death of +their host, rat fleas deserted its body and might then become a pest +in houses where they had not been noticed before.</p> + +<p>Attention was directed to the fact that while feeding, fleas are in +the habit of squirting blood from the anus and that in the case of those +which had fed upon rats and mice dying of the plague, virulent plague +bacilli were to be found in such blood. Liston (1905) even found, +and subsequent investigations confirmed, that the plague bacilli +multiply in the stomach of the insect and that thus the blood ejected +was richer in the organisms than was that of the diseased animal. +It was found that a film of this infected blood spread out under the +body of the flea and that thus the bacilli might be inoculated by the +bite of the insect and by scratching.<span class="pagenum"><a name="Page_171" id="Page_171">[Pg 171]</a></span></p> + +<p>Very recently, Bacot and Martin (1914) have paid especial +attention to the question of the mechanism of the transmission of +the plague bacilli by fleas. They believe that plague infested fleas +regurgitate blood through the mouth, and that under conditions +precluding the possibility of infection by dejecta, the disease may be +thus transmitted. The evidence does not seem sufficient to establish +that this is the chief method of transmission.</p> + +<p>Conclusive experimental proof that fleas transmit the disease is +further available from a number of sources. The most extensive +series of experiments is that of the English Plague Commission in +India, which reported in 1906 that:</p> + +<p>On thirty occasions a healthy rat contracted plague in sequence +of living in the neighborhood of a plague infected rat under circumstances +which prevented the healthy rat coming in contact with +either the body or excreta of the diseased animal.</p> + +<p>In twenty-one experiments out of thirty-eight, healthy rats living +in flea-proof cages contracted plague when exposed to rat fleas +(<i>Xenopsylla cheopis</i>), collected from rats dead or dying of septicæmic +plague.</p> + +<p>Close contact of plague-infected with healthy animals, if fleas +are excluded, does not give rise to an epizootic among the latter. +As the huts were never cleaned out, close contact included contact +with feces and urine of infected animals, and contact with, and eating +of food contaminated with feces and urine of infected animals, +as well as pus from open plague ulcers. Close contact of young, +even when suckled by plague-infected mothers, did not give the +disease to the former.</p> + +<p>If fleas are present, then the epizootic, once started, spreads from +animal to animal, the rate of progress being in direct proportion to +the number of fleas.</p> + +<p>Aerial infection was excluded. Thus guinea-pigs suspended in a +cage two feet above the ground did not contract the disease, while +in the same hut those animals allowed to run about and those placed +two inches above the floor became infected. It had previously +been found that a rat flea could not hop farther than about five +inches.</p> + +<p>Guinea pigs and monkeys were placed in plague houses in pairs, +both protected from soil contact infection and both equally exposed +to aerial infection, but one surrounded with a layer of tangle-foot +paper and the other surrounded with a layer of sand. The following +observations were made:<span class="pagenum"><a name="Page_172" id="Page_172">[Pg 172]</a></span></p> + +<p>(<i>a</i>) Many fleas were caught in the tangle-foot, a certain proportion +of which were found on dissection to contain in their stomachs +abundant bacilli microscopically identical with plague bacilli. Out +of eighty-five human fleas dissected only one contained these bacilli, +while out of seventy-seven rat fleas twenty-three were found thus +infected.</p> + +<p>(<i>b</i>) The animals surrounded with tangle-foot in no instance +developed plague, while several (24 per cent) of the non-protected +animals died of the disease.</p> + +<p>Thus, the experimental evidence that fleas transmit the plague +from rat to rat, from rats to guinea pigs, and from rats to monkeys +is indisputable. There is lacking direct experimental proof of its +transfer from rodents to man but the whole chain of indirect evidence +is so complete that there can be no doubt that such a transfer +does occur so commonly that in the case of bubonic plague it must +be regarded as the normal method.</p> + +<p>Rats are not the only animals naturally attacked by the plague +but as already suggested, it occurs in various other rodents. In +California the disease has spread from rats to ground squirrels +(<i>Otospermophilus beecheyi</i>), a condition readily arising from the +frequency of association of rats with the squirrels in the neighborhood +of towns, and from the fact that the two species of fleas found +on them are also found on rats. While the danger of the disease +being conveyed from squirrels to man is comparatively slight, the +menace in the situation is that the squirrels may become a more or +less permanent reservoir of the disease and infect rats, which may +come into more frequent contact with man.</p> + +<p>The tarbagan (<i>Arctomys bobac</i>), is a rodent found in North Manchuria, +which is much prized for its fur. It is claimed that this animal +is extremely susceptible to the plague and there is evidence to +indicate that it was the primary source of the great outbreak of +pneumonic plague which occurred in Manchuria and North China +during the winter of 1910-11.</p> + +<p>Of fleas, any species which attacks both rodents and man may be +an agent in the transmission of the plague. We have seen that in +India the species most commonly implicated is the rat flea, <i>Xenopsylla +cheopis</i>, (= <i>Lœmopsylla</i> or <i>Pulex cheopis</i>) (<a href="#Fig_89">fig. 89</a>). This species has +also been found commonly on rats in San Francisco. The cat flea, +<i>Ctenocephalus felis</i>, the dog flea, <i>Ctenocephalus canis</i>, the human flea, +<i>Pulex irritans</i>, the rat fleas, <i>Ceratophyllus fasciatus</i> and <i>Ctenopsyllus +musculi</i> have all been shown to meet the conditions.<span class="pagenum"><a name="Page_173" id="Page_173">[Pg 173]</a></span></p> + +<p>But, however clear the evidence that fleas are the most important +agent in the transfer of plague, it is a mistake fraught with danger +to assume that they are the only factor in the spread of the disease. +The causative organism is a bacillus and is not dependent upon any +insect for the completion of its development.</p> + +<p>Therefore, any blood-sucking insect which feeds upon a plague +infected man or animal and then passes to a healthy individual, +conceivably might transfer the bacilli. Verjbitski (1908) has shown +experimentally that bed-bugs may thus convey the disease. Hertzog +found the bacilli in a head-louse, <i>Pediculus humanus</i>, taken from a +child which had died from the plague, and McCoy found them in a +louse taken from a plague-infected squirrel. On account of their +stationary habits, the latter insects could be of little significance in +spreading the disease.</p> + +<p>Contaminated food may also be a source of danger. While this +source, formerly supposed to be the principal one, is now regarded as +unimportant, there is abundant experimental evidence to show that +it cannot be disregarded. It is believed that infection in this way +can occur only when there is some lesion in the alimentary canal.</p> + +<p>Still more important is the proof that in pneumonic plague the +patient is directly infective and that the disease is spread from man +to man without any intermediary. Especially conclusive is the +evidence obtained by Drs. Strong and Teague during the Manchurian +epidemic of 1910-11. They found that during coughing, in pneumonic +plague cases, even when sputum visible to the naked eye is +not expelled, plague bacilli in large numbers may become widely +disseminated into the surrounding air. By exposing sterile plates +before patients who coughed a single time, very numerous colonies +of the bacillus were obtained.</p> + +<p>But the great advance which has been made rests on the discovery +that bubonic plague is in the vast majority of cases transmitted +by the flea. The pneumonic type forms a very small percentage +of the human cases and even with it, the evidence indicates that the +original infection is derived from a rodent through the intermediary +of the insect.</p> + +<p>So modern prophylactic measures are directed primarily against +the rat and fleas. Ships coming from infected ports are no longer +disinfected for the purpose of killing the plague germs, but are fumigated +to destroy the rats and the fleas which they might harbor. +When anchored at infected ports, ships must observe strenuous<span class="pagenum"><a name="Page_174" id="Page_174">[Pg 174]</a></span> +precautions to prevent the ingress of rats. Cargo must be inspected +just before being brought on board, in order to insure its freedom from +rats. Even lines and hawsers must be protected by large metal discs +or funnels, for rats readily run along a rope to reach the ship. Once +infested, the ship must be thoroughly fumigated, not only to avoid +carrying the disease to other ports but to obviate an outbreak on +board.</p> + +<p>When an epidemic begins, rats must be destroyed by trapping +and poisoning. Various so-called biological poisons have not proved +practicable. Sources of food supply should be cut off by thorough +cleaning up, by use of rat-proof garbage cans and similar measures. +Hand in hand with these, must go the destruction of breeding places, +and the rat-proofing of dwellings, stables, markets, warehouses, docks +and sewers. All these measures are expensive, and a few years ago +would have been thought wholly impossible to put into practice +but now they are being enforced on a large scale in every fight against +the disease.</p> + +<p>Rats and other rodents are regularly caught in the danger zone +and examined for evidence of infection, for the sequence of the epizootic +and of the human disease is now understood. In London, rats +are regularly trapped and poisoned in the vicinity of the principal +docks, to guard against the introduction of infected animals in shipping. +During the past six years infected rats have been found +yearly, thirteen having been found in 1912. In Seattle, Washington, +seven infected rats were found along the water front in October, 1913, +and infected ground squirrels are still being found in connection with +the anti-plague measures in California.</p> + +<p>The procedure during an outbreak of the human plague was well +illustrated by the fight in San Francisco. The city was districted, +and captured rats, after being dipped in some fluid to destroy the fleas, +were carefully tagged to indicate their source, and were sent to the +laboratory for examination. If an infected rat was found, the officers +in charge of the work in the district involved were immediately +notified by telephone, and the infected building was subjected to a +thorough fumigation. In addition, special attention was given to +all the territory in the four contiguous blocks.</p> + +<p>By measures such as these, this dread scourge of the human race +is being brought under control. Incidentally, the enormous losses +due to the direct ravages of rats are being obviated and this alone +would justify the expenditure many times over of the money and +labor involved in the anti-rat measures.<span class="pagenum"><a name="Page_175" id="Page_175">[Pg 175]</a></span></p> + + + + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_VII" id="CHAPTER_VII"></a>CHAPTER VII</h2> + +<h2>ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC ORGANISMS</h2> + + +<p>We now have to consider the cases in which the arthropod acts +as the essential host of a pathogenic organism. In other words, +cases in which the organism, instead of being passively carried or +merely accidentally inoculated by the bite of its carrier, or vector, is +taken up and undergoes an essential part of its development within +the arthropod.</p> + +<p>In some cases, the sexual cycle of the parasite is undergone in the +arthropod, which then serves as the <i>definitive</i> or +<i>primary host</i>. In other cases, it is the asexual stage +of the parasite which is undergone, and the arthropod +then acts as the <i>intermediate host</i>. This distinction +is often overlooked and all the cases incorrectly +referred to as those in which the insect or other +arthropod acts as intermediate host.</p> + +<div class="figcenter" style="width: 150px;"><a name="Fig_113" id="Fig_113"></a> +<a href="images/f113-full.png"><img src="images/f113.png" width="150" height="472" alt="113. Dipylidium +caninum. The +double pored +tapeworm of the +dog." title="113. Dipylidium +caninum. The +double pored +tapeworm of the +dog." /></a> +<span class="caption">113. Dipylidium +caninum. The +double pored +tapeworm of the +dog.</span> +</div> + +<p>We have already emphasized that this is the most +important way in which insects may transmit disease, +for without them the particular organisms concerned +could never complete their development. Exterminate +the arthropod host and the life cycle of the +parasite is broken, the disease is exterminated.</p> + +<p>As the phenomenon of alternation of generations, +as exhibited by many of the parasitic protozoa, is a +complicated one and usually new to the student, we +shall first take up some of the grosser cases illustrated +by certain parasitic worms. There is the additional +reason that these were the first cases known of arthropod +transmission of pathogenic organisms.</p> + + +<h3><a name="Insects_as_Intermediate_Hosts_of_Tapeworms" id="Insects_as_Intermediate_Hosts_of_Tapeworms"></a><span class="smcap">Insects as Intermediate Hosts of Tapeworms</span></h3> + +<p>A number of tapeworms are known to undergo their sexual stage +in an insect or other arthropod. Of these at least two are occasional +parasites of man.</p> + +<p><i>Dipylidium caninum</i> (figs. <a href="#Fig_113">113</a> and <a href="#Fig_114">114</a>), more generally known as +<i>Taenia cucumerina</i> or <i>T. elliptica</i>, is the commonest intestinal parasite +of pet dogs and cats. It is occasionally found as a human parasite, +70 per cent of the cases reported being in young children.<span class="pagenum"><a name="Page_176" id="Page_176">[Pg 176]</a></span></p> + +<div class="figleft" style="width: 250px;"><a name="Fig_114" id="Fig_114"></a> +<a href="images/f114-full.png"><img src="images/f114.png" width="250" height="264" alt="114. Dipylidium caninum. +Rostrum evaginated and +invaginated. After +Blanchard." title="114. Dipylidium caninum. +Rostrum evaginated and +invaginated. After +Blanchard." /></a> +<span class="caption">114. Dipylidium caninum. +Rostrum evaginated and +invaginated. After +Blanchard.</span> +</div> + +<p>In 1869, Melnikoff found in a dog louse, <i>Trichodectes canis</i>, some +peculiar bodies which Leuckart identified as the larval form of this +tapeworm. The worm is, however, much more +common in dogs and cats than is the skin parasite, +and hence it appears that the <i>Trichodectes</i> +could not be the only intermediate host. In +1888, Grassi found that it could also develop +in the cat and dog fleas, <i>Ctenocephalus felis</i> +and <i>C. canis</i>, and in the human flea, <i>Pulex +irritans</i>.</p> + +<div class="figright" style="width: 200px;"><a name="Fig_115" id="Fig_115"></a> +<a href="images/f115-full.png"><img src="images/f115.png" width="200" height="161" alt="115. Dipylidium caninum. +Immature cysticercoid. +After Grassi and Rovelli." title="115. Dipylidium caninum. +Immature cysticercoid. +After Grassi and Rovelli." /></a> +<span class="caption">115. Dipylidium caninum. +Immature cysticercoid. +After Grassi and Rovelli.</span> +</div> + +<p>The eggs, scattered among the hairs of the +dog or cat, are ingested by the insect host and +in its body cavity they develop into pyriform +bodies, about 300µ in length, almost entirely destitute of a bladder, +but in the immature stage provided with a caudal appendage (<a href="#Fig_115">fig. 115</a>). +Within the pear-shaped body (<a href="#Fig_116">fig. 116</a>) are the invaginated head and +suckers of the future tapeworm. This larval +form is known as a cysticercoid, in contradistinction +to the bladder-like cysticercus of many +other cestodes. It is often referred to in literature +as <i>Cryptocystis trichodectis</i> Villot.</p> + +<p>As many as fifty of the cysticercoids have +been found in the body cavity of a single flea. +When the dog takes up an infested flea or louse, +by biting itself, or when the cat licks them up, the +larvæ quickly develop into tapeworms, reaching sexual maturity in +about twenty days in the intestine of their host. Puppies and +kittens are quickly infested when suckling a flea-infested mother, the +developing worms having been found in the intestines of puppies not +more than five or six days old.</p> + +<div class="figleft" style="width: 200px;"><a name="Fig_116" id="Fig_116"></a> +<a href="images/f116-full.png"><img src="images/f116.png" width="200" height="138" alt="116. Dipylidium caninum. +Cysticercoid. After +Villet." title="116. Dipylidium caninum. +Cysticercoid. After +Villet." /></a> +<span class="caption">116. Dipylidium caninum. +Cysticercoid. After +Villet.</span> +</div> + +<p>Infestation of human beings occurs only +through accidental ingestion of an infested flea. +It is natural that such cases should occur largely +in children, where they may come about in +some such way as illustrated in the accompanying +figures <a href="#Fig_117">117</a> and <a href="#Fig_118">118</a>.</p> + +<p><i>Hymenolepis diminuta</i>, very commonly living in the intestine +of mice and rats, is also known to occur in man. Its cysticercoid +develops in the body cavity of a surprising range of meal-infesting +insects. Grassi and Rovelli (abstract in Ransom, 1904) found it in the<span class="pagenum"><a name="Page_177" id="Page_177">[Pg 177]</a></span> +larvæ and adult of a moth, <i>Asopia farinalis</i>, in the earwig, <i>Anisolabis +annulipes</i>, the Tenebrionid beetles <i>Akis spinosa</i> and <i>Scaurus striatus</i>. +Grassi considers that the lepidopter is the normal intermediate +host. The insect takes +up the eggs scattered by rats +and mice. It has been experimentally +demonstrated that +man may develop the tapeworm +by swallowing infested +insects. Natural infection +probably occurs by ingesting +such insects with cereals, or +imperfectly cooked foods.</p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_117" id="Fig_117"></a> +<a href="images/f117-full.png"><img src="images/f117.png" width="400" height="290" alt="117. One way in which Dipylidium infection in +children may occur. After Blanchard." title="117. One way in which Dipylidium infection in +children may occur. After Blanchard." /></a> +<span class="caption">117. One way in which Dipylidium infection in +children may occur. After Blanchard.</span> +</div> + +<p><i>Hymenolepis lanceolata</i>, a parasite of geese and ducks, has been +reported once for man. The supposed cysticercoid occurs in various +small crustaceans of the family Cyclopidæ.</p> + +<div class="figcenter" style="width: 450px;"><a name="Fig_118" id="Fig_118"></a> +<a href="images/f118-full.png"><img src="images/f118.png" width="450" height="353" alt="118. The probable method by which Dipylidium infection usually occurs." title="118. The probable method by which Dipylidium infection usually occurs." /></a> +<span class="caption">118. The probable method by which Dipylidium infection usually occurs.</span> +</div><p><span class="pagenum"><a name="Page_178" id="Page_178">[Pg 178]</a></span></p> + +<p>Several other cestode parasites of domestic animals are believed +to develop their intermediate stage in certain arthropods. Among +these may be mentioned:</p> + +<p><i>Choanotænia infundibulformis</i>, of chickens, developing in the house-fly +(Grassi and Rovelli);</p> + +<p><i>Davainea cesticillus</i>, of chickens, in some lepidopter or coleopter +(Grassi and Rovelli);</p> + +<p><i>Hymenolepis anatina</i>, <i>H. gracilis</i>, <i>H. sinuosa</i>, <i>H. coronula</i> and +<i>Fimbriaria fasciolaris</i>, all occurring in ducks, have been reported as +developing in small aquatic crustaceans. In these cases, cysticercoids +have been found which, on account of superficial characters, +have been regarded as belonging to the several species, but direct +experimental evidence is scant.</p> + + +<h3><a name="Arthropods_as_Intermediate_Hosts_of_Nematode_Worms" id="Arthropods_as_Intermediate_Hosts_of_Nematode_Worms"></a><span class="smcap">Arthropods as Intermediate Hosts of Nematode Worms</span></h3> + + +<p><a name="Filariasis_and_Mosquitoes" id="Filariasis_and_Mosquitoes"></a><b>Filariasis and Mosquitoes</b>—A number of species of Nematode +worms belonging to the genus <i>Filaria</i>, infest man and other vertebrates +and in the larval condition are to be found in the blood. +Such infestation is known as <i>filariasis</i>. The sexually mature worms +are to be found in the blood, the lymphatics, the mesentery and subcutaneous +connective tissue. In the cases best studied it has been +found that the larval forms are taken up by mosquitoes and undergo +a transformation before they can attain maturity in man.</p> + +<p>The larvæ circulating in the blood are conveniently designated +as microfilariæ. In this stage they are harmless and only one species, +<i>Filaria bancrofti</i>, appears to be of any great pathological significance +at any stage.</p> + +<p><i>Filaria bancrofti</i> in its adult state, lives in the lymphatics of man. +Though often causing no injury it has been clearly established that +they and their eggs may cause various disorders due to stoppage +of the lymphatic trunks (<a href="#Fig_119">fig. 119</a>). Manson lists among other effects, +abscess, varicose groin glands, lymph scrotum, chyluria, and elephantiasis.</p> + +<p>The geographical distribution of this parasite is usually given as +coextensive with that of elephantiasis, but it is by no means certain +that it is the only cause of this disease and so actual findings of the +parasites are necessary. Manson reports that it is "an indigenous +parasite in almost every country throughout the tropical and subtropical +world, as far north as Spain in Europe and Charlestown in<span class="pagenum"><a name="Page_179" id="Page_179">[Pg 179]</a></span> +the United States, and as far south as Brisbane in Australia." In +some sections, fully 50 per cent of the natives are infested. Labredo +(1910) found 17.82 per cent infestation in Havana.</p> + +<div class="figcenter" style="width: 317px;"><a name="Fig_119" id="Fig_119"></a> +<a href="images/f119-full.png"><img src="images/f119.png" width="317" height="390" alt="119. Elephantiasis in Man. From "New +Sydenham Society's Atlas."" title="119. Elephantiasis in Man. From "New +Sydenham Society's Atlas."" /></a> +<span class="caption">119. Elephantiasis in Man. From "New +Sydenham Society's Atlas."</span> +</div> + +<p>The larval forms of <i>Filaria bancrofti</i> were first discovered in 1863, +by Demarquay, in a case of chylous dropsy. They were subsequently +noted under similar conditions, by several workers, and by +Wücherer in the urine of twenty-eight cases of tropical chyluria, +but in 1872 Lewis found that the blood of man was the normal +habitat, and gave them the name <i>Filaria sanguinis hominis</i>. The +adult worm was found in 1876 +by Bancroft, and in 1877, +Cobbold gave it the name <i>Filaria +bancrofti</i>. It has since been +found repeatedly in various parts +of the lymphatic system, and its +life-history has been the subject +of detailed studies by Manson +(1884), Bancroft (1899), Low +(1900), Grassi and Noé (1900), +Noé (1901) and Fülleborn (1910).</p> + +<p>The larvæ as they exist in +the circulating blood, exhibit a +very active wriggling movement, +without material progression. +They may exist in enormous +numbers, as many as five or +six hundred swarming in a +single drop of blood. This is the more surprising when we consider +that they measure about 300µ × 8µ, that is, their width is +equal to the diameter of the red blood corpuscle of their host and +their length over thirty-seven times as great.</p> + +<p>Their organs are very immature and the structure obscure. When +they have quieted down somewhat in a preparation it may be seen +that at the head end there is a six-lipped and very delicate prepuce, +enclosing a short "fang" which may be suddenly exserted and +retracted. Completely enclosing the larva is a delicate sheath, +which is considerably longer than the worm itself. To enter into +further details of anatomy is beyond the scope of this discussion +and readers interested are referred to the work of Manson and of +Fülleborn.<span class="pagenum"><a name="Page_180" id="Page_180">[Pg 180]</a></span></p> + +<p>One of the most surprising features of the habits of these larvæ +is the periodicity which they exhibit in their occurrence in the peripheral +blood. If a preparation be made during the day time there +may be no evidence whatever of filarial infestation, whereas a preparation +from the same patient taken late in the evening or during +the night may be literally swarming with the parasites. Manson +quotes Mackenzie as having brought out the further interesting +fact that should a "filarial subject be made to sleep during the day +and remain awake at night, the periodicity is reversed; that is to say, +the parasites come into the blood during the day and disappear from +it during the night." There have been numerous attempts to explain +this peculiar phenomenon of periodicity but in spite of objections +which have been raised, the most plausible remains that of Manson, +who believes that it is an adaptation correlated with the life-habits +of the liberating agent of the parasite, the mosquito.</p> + +<p>The next stages in the development of <i>Filaria nocturna</i> occur in +mosquitoes, a fact suggested almost simultaneously by Bancroft +and Manson in 1877, and first demonstrated by the latter very soon +thereafter. The experiments were first carried out with <i>Culex +quinquefasciatus</i> (= <i>fatigans</i>) as a host, but it is now known that a +number of species of mosquitoes, both anopheline and culicine, may +serve equally well.</p> + +<p>When the blood of an infested individual is sucked up and reaches +the stomach of such a mosquito, the larvæ, by very active movements, +escape from their sheaths and within a very few hours actively migrate +to the body cavity of their new host and settle down primarily +in the thoracic muscles. There in the course of sixteen to twenty +days they undergo a metamorphosis of which the more conspicuous +features are the formation of a mouth, an alimentary canal and a +trilobed tail. At the same time there is an enormous increase in +size, the larvæ which measured .3 mm. in the blood becoming 1.5 mm. +in length. This developmental period may be somewhat shortened +in some cases and on the other hand may be considerably extended. +The controlling factor seems to be the one of temperature.</p> + +<p>The transformed larvæ then reenter the body cavity and finally +the majority of them reach the interior of the labium (<a href="#Fig_120">fig. 120</a>). A +few enter the legs and antennæ, and the abdomen, but these are +wanderers which, it is possible, may likewise ultimately reach the +labium, where they await the opportunity to enter their human host.<span class="pagenum"><a name="Page_181" id="Page_181">[Pg 181]</a></span></p> + +<p>It was formerly supposed that when the infested mosquito punctured +the skin of man, the mature larvæ were injected into the circulation. +The manner in which this occurred was not obvious, for +when the insect feeds it inserts only the stylets, the labium itself +remaining on the surface of the skin. Fülleborn has cleared up the +question by showing that at this time the filariæ escape and, like +the hookworm, actively bore into the skin of their new host.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_120" id="Fig_120"></a> +<a href="images/f120-full.png"><img src="images/f120.png" width="500" height="343" alt="120. Filaria in the muscles and labium of Culex. After Blanchard." title="120. Filaria in the muscles and labium of Culex. After Blanchard." /></a> +<span class="caption">120. Filaria in the muscles and labium of Culex. After Blanchard.</span> +</div> + +<p>Once entered, they migrate to the lymphatics and there quickly +become sexually mature. The full grown females measure 85-90 mm. +in length by .24-.28 mm. in diameter, while the males are less than +half this size, being about 40 mm. by .1 mm. Fecundation occurs +and the females will be found filled with eggs in various stages of +development, for they are normally viviparous.</p> + +<p><i>Filaria philippinensis</i> is reported by Ashburn and Craig (1907) as +a common blood filaria in the Philippine Islands. As they describe +it, it differs from <i>Filaria bancrofti</i> primarily in that it does not exhibit +periodicity. Its development has been found to occur in <i>Culex +quinquefasciatus</i>, where it undergoes metamorphosis in about fourteen +or fifteen days. There is doubt as to the species being distinct from +<i>bancrofti</i>.</p> + +<p>Several other species occur in man and are thought to be transferred +by various insects, among which have been mentioned Tabanidæ +and tsetse-flies, but there is no experimental proof in support +of such conjectures.<span class="pagenum"><a name="Page_182" id="Page_182">[Pg 182]</a></span></p> + +<div class="figleft" style="width: 200px;"><a name="Fig_121" id="Fig_121"></a> +<a href="images/f121-full.png"><img src="images/f121.png" width="200" height="343" alt="121. Dracunculus +medinensis; female; +mouth; embryo. +After Bastian and +Leuckart." title="121. Dracunculus +medinensis; female; +mouth; embryo. +After Bastian and +Leuckart." /></a> +<span class="caption">121. Dracunculus +medinensis; female; +mouth; embryo. +After Bastian and +Leuckart.</span> +</div> + +<p><i>Filaria immitis</i> is a dangerous parasite of the dog, the adult worm +living in the heart and veins of this animal. It is one of the species +which has been clearly shown to undergo its development in the +mosquito, particularly in <i>Anopheles maculipennis</i> and <i>Aedes calopus</i> +(= Stegomyia). The larval form occurs in the peripheral blood, +especially at night. When taken up by mosquitoes they differ from +<i>Filaria bancrofti</i> in that they undergo their development in the +Malpighian tubules rather than in the thoracic muscles. In +about twelve days they have completed their growth in the tubules, +pierce the distal end, and pass to the labium. This species occurs +primarily in China and Japan, but is also found in Europe and in the +United States. It is an especially favorable species for studying +the transformations in the mosquito.</p> + +<p><i>Filariæ</i> are also commonly found in birds, and in this country +this is the most available source of laboratory material. We have +found them locally (Ithaca, N. Y.) in the blood of +over sixty per cent of all the crows examined, at +any season of the year, and have also found them +in English sparrows.</p> + +<p>In the crows, they often occur in enormous +numbers, as many as two thousand having been +found in a single drop of the blood of the most +heavily infested specimen examined. For study, a +small drop of blood should be mounted on a clean +slide and the coverglass rung with vaseline or oil +to prevent evaporation. In this way they can +be kept for hours.</p> + +<p>Permanent preparations may be made by +spreading out the blood in a film on a perfectly +clean slide and staining. This is easiest done by touching the fresh +drop of blood with the end of a second slide which is then held at +an angle of about 45° to the first slide and drawn over it without +pressure. Allow the smear to dry in the air and stain in the usual +way with hæmatoxylin.</p> + + +<h3><a name="Other_Nematode_Parasites_of_Man_and_Animals_Developing_in_Arthropods" id="Other_Nematode_Parasites_of_Man_and_Animals_Developing_in_Arthropods"></a><span class="smcap">Other Nematode Parasites of Man and Animals Developing +in Arthropods</span></h3> + +<p><i>Dracunculus medinensis</i> (<a href="#Fig_121">fig. 121</a>), the so-called guinea-worm, is +a nematode parasite of man which is widely distributed in tropical +Africa, Asia, certain parts of Brazil and is occasionally imported +into North America.<span class="pagenum"><a name="Page_183" id="Page_183">[Pg 183]</a></span></p> + +<div class="figright" style="width: 175px;"><a name="Fig_122" id="Fig_122"></a> +<a href="images/f122-full.png"><img src="images/f122.png" width="175" height="303" alt="122. Cyclops, the intermediate host of +Dracunculus." title="122. Cyclops, the intermediate host of +Dracunculus." /></a> +<span class="caption">122. Cyclops, the intermediate host of +Dracunculus.</span> +</div> + +<p>The female worm is excessively long and slender, measuring nearly +three feet in length and not more than one-fifteenth of an inch in +diameter. It is found in the subcutaneous connective tissue and when +mature usually migrates to some part of the leg. +Here it pierces the skin and there is formed a small +superficial ulcer through which the larvæ reach the +exterior after bursting the body of the mother.</p> + +<p>Fedtschenko (1879) found that when these larvæ +reach the water they penetrate the carapace of the +little crustacean, <i>Cyclops</i> (<a href="#Fig_122">fig. 122</a>). Here they molt +several times and undergo a metamorphosis. Fedtschenko, +in Turkestan, found that these stages required +about five weeks, while Manson who confirmed these +general results, found that eight or nine weeks were +required in the cooler climate of England.</p> + +<p>Infection of the vertebrate host probably occurs through swallowing +infested cyclops in drinking water. Fedtschenko was unable to +demonstrate this experimentally and objection has been raised against +the theory, but Leiper (1907), and Strassen (1907) succeeded in infesting +monkeys by feeding them on cyclops containing the larvæ.</p> + +<p><i>Habronema muscæ</i> is a worm which has long been known in its +larval stage, as a parasite of the house-fly. Carter found them in +33 per cent of the house-flies examined in Bombay during July, 1860, +and since that time they have been shown to be very widely distributed. +Italian workers reported them in 12 per cent to 30 per cent +of the flies examined. Hewitt reported finding it rarely in England. +In this country it was first reported by Leidy who found it in about +20 per cent of the flies examined at Philadelphia, Pa. Since then it +has been reported by several American workers. We have found it +at Ithaca, N. Y., but have not made sufficient examinations to justify +stating percentage. Ransom (1913) reports it in thirty-nine out of +one hundred and thirty-seven flies, or 28 per cent.</p> + +<div class="figcenter" style="width: 500px;"> +<a href="images/f123-full.png"><img src="images/f123.png" width="500" height="491" alt="123. An Echinorhynchid, showing the spinose retractile proboscis." title="123. An Echinorhynchid, showing the spinose retractile proboscis." /></a> +<span class="caption">123. An Echinorhynchid, showing the spinose retractile proboscis.</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_124" id="Fig_124"></a> +<a href="images/f124-full.png"><img src="images/f124.png" width="500" height="309" alt="124. June beetle (Lachnosterna). Larva." title="124. June beetle (Lachnosterna). Larva." /></a> +<span class="caption">124. June beetle (Lachnosterna). Larva.</span> +</div> + +<p>Until very recently the life-history of this parasite was unknown +but the thorough work of Ransom (1911, 1913) has shown clearly +that the adult stage occurs in the stomach of horses. The embryos, +produced by the parent worms in the stomach of the horse, pass +out with the feces and enter the bodies of fly larvæ which are developing +in the manure. In these they reach their final stage of larval +development at about the time the adult flies emerge from the pupal +stage. In the adult fly they are commonly found in the head.<span class="pagenum"><a name="Page_184" id="Page_184">[Pg 184]</a></span> +frequently in the proboscis, but they occur also in the thorax and +abdomen. Infested flies are accidentally swallowed by horses and +the parasite completes its development to maturity in the stomach of +its definitive host.<span class="pagenum"><a name="Page_185" id="Page_185">[Pg 185]</a></span></p> + +<p><i>Gigantorhynchus hirudinaceus</i> (= <i>Echinorhynchus gigas</i>) is a common +parasite of the pig and has been reported as occurring in man. +The adult female is 20-35 cm. long and 4-9 mm. in diameter. +It lacks an alimentary canal and is provided with a strongly spined +protractile rostrum, by means of which it attaches to the intestinal +mucosa of its host.</p> + +<p>The eggs are scattered with the feces of the host and are taken +up by certain beetle larvæ. In Europe the usual intermediate hosts +are the larvæ of the cockchafer, <i>Melolontha vulgaris</i>, or of the flower +beetle, <i>Cetonia aurata</i>. Stiles has shown that in the United States +the intermediate host is the larva of the June bug, <i>Lachnosterna</i> +(<a href="#Fig_124">fig. 124</a>). It is probable that several of the native species serve in +this capacity.</p> + +<p>A number of other nematode parasites of birds and mammals +have been reported as developing in arthropods but here, as in the +case of the cestodes, experimental proof is scant. The cases above +cited are the better established and will serve as illustrations.<span class="pagenum"><a name="Page_186" id="Page_186">[Pg 186]</a></span></p> + + + + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_VIII" id="CHAPTER_VIII"></a>CHAPTER VIII</h2> + +<h2>ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC +PROTOZOA</h2> + + +<h3><a name="Mosquitoes_and_Malaria" id="Mosquitoes_and_Malaria"></a><span class="smcap">Mosquitoes and Malaria</span></h3> + +<p>Under the name of malaria is included a group of morbid symptoms +formerly supposed to be due to a miasm or bad air, but now +known to be caused by protozoan parasites of the genus <i>Plasmodium</i>, +which attack the red blood corpuscles. It occurs in paroxysms, +each marked by a chill, followed by high fever and sweating. The +fever is either intermittent or remittent.</p> + +<p>There are three principal types of the disease, due to different +species of the parasite. They are:</p> + +<p>1. The benign-tertian, caused by <i>Plasmodium vivax</i>, which undergoes +its schizogony or asexual cycle in the blood in forty-eight hours +or even less. This type of the disease,—characterized by fever +every two days, is the most wide-spread and common.</p> + +<p>2. The quartan fever is due to the presence of <i>Plasmodium +malariæ</i>, which has an asexual cycle of seventy-two hours, and therefore +the fever recurs every three days. This type is more prevalent +in temperate and sub-tropical regions, but appears to be rare everywhere.</p> + +<p>3. The sub-tertian "æstivo-autumnal," or "pernicious" fever +is caused by <i>Plasmodium falciparum</i>. Schizogony usually occurs +in the internal organs, particularly in the spleen, instead of in the +peripheral circulation, as is the case of the tertian and quartan forms. +The fever produced is of an irregular type and the period of schizogony +has not been definitely determined. It is claimed by some that the +variations are due to different species of malignant parasites.</p> + +<p>It is one of the most wide-spread of human diseases, occurring +in almost all parts of the world, except in the polar regions and in +waterless deserts. It is most prevalent in marshy regions.</p> + +<p>So commonplace is malaria that it causes little of the dread +inspired by most of the epidemic diseases, and yet, as Ross says, +it is perhaps the most important of human diseases. Figures regarding +its ravages are astounding. Celli estimated that in Italy it +caused an average annual mortality of fifteen thousand, representing +about two million cases. In India alone, according to Ross (1910)<span class="pagenum"><a name="Page_187" id="Page_187">[Pg 187]</a></span> +"it has been officially estimated to cause a mean annual death-rate +of five per thousand; that is, to kill every year, on the average, one +million one hundred and thirty thousand." In the United States +it is widespread and though being restricted as the country develops, +it still causes enormous losses. During the year 1911, "in Alabama +alone there were seventy thousand cases and seven hundred and +seventy deaths." The weakening effects of the disease, the invasion +of other diseases due to the attacks of malaria, are among the very +serious results, but they cannot be estimated.</p> + +<p>Not only is there direct effect on man, but the disease has been one +of the greatest factors in retarding the development of certain regions. +Everywhere pioneers have had to face it, and the most fertile regions +have, in many instances been those most fully dominated by it. +Herrick (1903) has presented an interesting study of its effects on +the development of the southern United States and has shown that +some parts, which are among the most fertile in the world, are +rendered practically uninhabitable by the ravages of malaria. Howard +(1909) estimates that the annual money loss from the disease +in the United States is not less than $100,000,000.</p> + +<p>It was formerly supposed that the disease was due to a miasm, +to a noxious effluvia, or infectious matter rising in the air from +swamps. In other words its cause was, as the name indicated +"mal aria," and the deep seated fear of night air is based largely on +the belief that this miasm was given off at night. Its production +was thought to be favored by stirring of the soil, dredging operations +and the like.</p> + +<p>The idea of some intimate connection between malaria and +mosquitoes is not a new one. According to Manson, Lancisi noted +that in some parts of Italy the peasants for centuries have believed +that malaria is produced by the bite of mosquitoes. Celli states +that one not rarely hears from such peasants the statement that +"In such a place, there is much fever, because it is full of mosquitoes." +Koch points out that in German East Africa the natives call malaria +and the mosquito by the same name, <i>Mbù</i>. The opinion was not +lacking support from medical men. Celli quotes passages from the +writings of the Italian physician, Lancisi, which indicate that he +favored the view in 1717.</p> + +<p>Dr. Josiah Nott is almost universally credited with having supported +the theory, in 1848, but as we have already pointed out +his work has been misinterpreted. The statements of Beauperthuy, +(1853) were more explicit.<span class="pagenum"><a name="Page_188" id="Page_188">[Pg 188]</a></span></p> + +<p>The clearest early presentation of the circumstantial evidence in +favor of the theory of mosquito transmission was that of A. F. A. +King, an American physician, in 1883. He presented a series of +epidemiological data and showed "how they may be explicable by +the supposition that the mosquito is the real source of the disease, +rather than the inhalation or cutaneous absorption of a marsh vapor." +We may well give the space to summarizing his argument here for +it has been so remarkably substantiated by subsequent work:</p> + +<p>1. Malaria, like mosquitoes, affects by preference low and moist +localities, such as swamps, fens, jungles, marshes, etc.</p> + +<p>2. Malaria is hardly ever developed at a lower temperature +than 60° Fahr., and such a temperature is necessary for the development +of the mosquito.</p> + +<p>3. Mosquitoes, like malaria, may both accumulate in and be +obstructed by forests lying in the course of winds blowing from +malarious localities.</p> + +<p>4. By atmospheric currents malaria and mosquitoes are alike +capable of being transported for considerable distances.</p> + +<p>5. Malaria may be developed in previously healthy places by +turning up the soil, as in making excavations for the foundation of +houses, tracks for railroads, and beds for canals, because these operations +afford breeding places for mosquitoes.</p> + +<p>6. In proportion as countries, previously malarious, are cleared +up and thickly settled, periodical fevers disappear, because swamps +and pools are drained so that the mosquito cannot readily find a place +suitable to deposit her eggs.</p> + +<p>7. Malaria is most dangerous when the sun is down and the +danger of exposure after sunset is greatly increased by the person +exposed sleeping in the night air. Both facts are readily explicable +by the mosquito malaria theory.</p> + +<p>8. In malarial districts the use of fire, both indoors and to those +who sleep out, affords a comparative security against malaria, because +of the destruction of mosquitoes.</p> + +<p>9. It is claimed that the air of cities in some way renders the +poison innocuous, for, though a malarial disease may be raging outside, +it does not penetrate far into the interior. We may easily +conceive that mosquitoes, while invading cities during their nocturnal +pilgrimages will be so far arrested by walls and houses, as well as +attracted by lights in the suburbs, that many of them will in this +way be prevented from penetrating "far into the interior."<span class="pagenum"><a name="Page_189" id="Page_189">[Pg 189]</a></span></p> + +<p>10. Malarial diseases and likewise mosquitoes are most prevalent +toward the latter part of summer and in the autumn.</p> + +<p>11. Various writers have maintained that malaria is arrested by +canvas curtains, gauze veils and mosquito nets and have recommended +the rise of mosquito curtains, "through which malaria can +seldom or never pass." It can hardly be conceived that these +intercept marsh-air but they certainly do protect from mosquitoes.</p> + +<p>12. Malaria spares no age, but it affects infants much less +frequently than adults, because young infants are usually carefully +housed and protected from mosquito inoculation.</p> + +<p>Correlated with the miasmatic theory was the belief that some +animal or vegetable organism which lived in marshes, produced +malaria, and frequent searches were made for it. Salisbury (1862) +thought this causative organism to be an alga, of the genus <i>Palmella</i>; +others attributed it to certain fungi or bacteria.</p> + +<p>In 1880, the French physician, Laveran, working in Algeria, +discovered an amœboid organism in the blood of malarial patients +and definitely established the parasitic nature of this disease. Pigmented +granules had been noted by Meckel as long ago as 1847, in +the spleen and blood of a patient who had died of malaria, and his +observations had been repeatedly verified, but the granules had been +regarded as degeneration products, and the fact that they occurred +in the body of a foreign organism had been overlooked.</p> + +<p>Soon after the discovery of the parasites in the blood, Gerhardt +(1884) succeeded in transferring the disease to healthy individuals +by inoculation of malarious blood, and thus proved that it is a true +infection. This was verified by numerous experimenters and it +was found that inoculation with a very minute quantity of the diseased +blood would not only produce malaria but the particular type +of disease.</p> + +<p>Laveran traced out the life cycle of the malarial parasite as it +occurs in man. The details as we now know them and as they are +illustrated by the accompanying <a href="#Fig_125">figure 125</a>, are as follows:</p> + +<p>The infecting organism or <i>sporozoite</i>, is introduced into the circulation, +penetrates a red blood corpuscle, and forms the amœboid +<i>schizont</i>. This lives at the expense of the corpuscle and as it develops +there are deposited in its body scattered black or reddish black +particles. These are generally called melanin granules, but are +much better referred to as hæmozoin, as they are not related to<span class="pagenum"><a name="Page_190" id="Page_190">[Pg 190]</a></span> +melanin. The hæmozoin is the most conspicuous part of the parasite, +a feature of advantage in diagnosing from unstained preparations.</p> + +<div class="figcenter" style="width: 325px;"><a name="Fig_125" id="Fig_125"></a> +<a href="images/f125-full.png"><img src="images/f125.png" width="325" height="335" alt="125. Life cycle of the malaria parasite. Adapted from Leuckart's chart, +by Miss Anna Stryke." title="125. Life cycle of the malaria parasite. Adapted from Leuckart's chart, +by Miss Anna Stryke." /></a> +<span class="caption">125. Life cycle of the malaria parasite. Adapted from Leuckart's chart, +by Miss Anna Stryke.</span> +</div> + +<p>As the schizont matures, its nucleus breaks up into a number of +daughter nuclei, each with a rounded mass of protoplasm about it, +and finally the corpuscles are broken down and these rounded bodies +are liberated in the plasma as <i>merozoites</i>. These merozoites infect +new corpuscles and thus the asexual cycle is continued. The malarial +paroxysm is coincident with sporulation.</p> + +<p>As early as Laveran's time it was known that under conditions +not yet determined there are to be found in the blood of malarious +patients another phase of the parasite, differing in form according +to the type of the disease. In the pernicious type these appear as +large, crescent-shaped organisms which have commonly been called +"crescents." We now know that these are sexual forms.<span class="pagenum"><a name="Page_191" id="Page_191">[Pg 191]</a></span></p> + +<p>When the parasite became known there immediately arose speculations +as to the way in which it was transferred from man to man. +It was thought by some that in nature it occurred as a free-living +amœba, and that it gained access to man through being taken up +with impure water. However, numerous attempts to infect healthy +persons by having them drink or inhale marsh water, or by injecting +it into their circulation resulted in failure, and influenced by Leuckart's +and Melnikoff's work on <i>Dipylidium</i>, that of Fedtschenko on <i>Dracunculus</i>, +and more especially by that of Manson on <i>Filaria</i>, search was +made for some insect which might transfer the parasite.</p> + +<p>Laveran had early suggested that the rôle of carrier might be +played by the mosquito, but Manson first clearly formulated the +hypothesis, and it was largely due to his suggestions that Ross in +India, undertook to solve the problem. With no knowledge of the +form or of the appearance in this stage, or of the species of mosquito +concerned, Ross spent almost two and a half years of the most arduous +work in the search and finally in August, 1897, seventeen years +after the discovery of the parasite in man, he obtained his first +definite clue. In dissecting a "dappled-winged mosquito," "every +cell was searched and to my intense disappointment nothing whatever +was found, until I came to the insect's stomach. Here, however, +just as I was about to abandon the examination, I saw a very delicate +circular cell, apparently lying amongst the ordinary cells of the organ +and scarcely distinguishable from them. On looking further, +another and another similar object presented itself. I now focused +the lens carefully on one of these, and found that it contained a few +minute granules of some black substance, exactly like the pigment of +the parasite of malaria. I counted altogether twelve of these cells +in the insect."</p> + +<p>Further search showed that "the contents of the mature pigment +cells did not consist of clear fluid but of a multitude of delicate, +thread-like bodies which on the rupture of the parent cell, were poured +into the body cavity of the insect. They were evidently spores."</p> + +<p>With these facts established, confirmation and extension of +Ross's results quickly followed, from many different sources. We +cannot trace this work in detail but will only point out that much +of the credit is due to the Italian workers, Grassi, Bignami, and +Bastianelli, and to Koch and Daniels.</p> + +<p>It had already been found that when fresh blood was mounted and +properly protected against evaporation, a peculiar change occurred<span class="pagenum"><a name="Page_192" id="Page_192">[Pg 192]</a></span> +in these crescents after about half an hour's time. From certain +of them there were pushed out long whip-like processes which moved +with a very active, lashing movement. The parasite at this stage +is known as the "flagellated body." Others, differing somewhat in +details of structure, become rounded but do not give off "flagella."</p> + +<p>The American worker, MacCallum (1897), in studying bird +malaria as found in crows, first recognized the true nature of these +bodies. He regarded them as sexual forms and believed that the +so-called flagella played the part of spermatozoa. Thus, the "flagellated +body" is in reality a <i>microgametoblast</i>, producing <i>microgametes</i>, +or the male sexual element, while the others constitute the <i>macrogametes</i>, +or female elements.</p> + +<p>It was found that when blood containing these sexual forms was +sucked up by an Anopheline mosquito and taken into its stomach, a +microgamete penetrated and fertilized a macrogamete in a way +analogous to what takes place in the fertilization of the egg in higher +forms. The resultant, mobile organism is known as the <i>migratory +ookinete</i>. In this stage the parasite bores through the epithelial +lining of the "stomach" (mid-intestine) of the mosquito and becomes +encysted under the muscle layers. Here the <i>oocyst</i>, as it is now +known, matures and breaks up into the body cavity and finally +its products come to lie in the salivary glands of the mosquito. Ten +to twelve days are required for these changes, after which the mosquito +is infective, capable of introducing the parasite with its saliva, +when feeding upon a healthy person.</p> + +<p>Thus the malarial parasite is known to have a double cycle, an +alternation of generations, of which the asexual stage is undergone in +man, the sexual in certain species of mosquitoes. The mosquito is +therefore the definitive host rather than the <i>intermediate</i>, as usually +stated.</p> + +<p>The complicated cycle may be made clearer by the diagram of +Miss Stryke (1912) which, by means of a double-headed mosquito +(<a href="#Fig_126">fig. 126</a>) endeavors to show how infection takes place through the +biting of the human victim, (at A), in whom asexual multiplication +then takes place, and how the sexual stages, taken up at B in the +diagram, are passed in the body of the mosquito.</p> + +<div class="figcenter" style="width: 450px;"><a name="Fig_126" id="Fig_126"></a> +<a href="images/f126-full.png"><img src="images/f126.png" width="450" height="567" alt="126. Life cycle of the malarial parasite. After Miss Anna Stryke." title="126. Life cycle of the malarial parasite. After Miss Anna Stryke." /></a> +<span class="caption">126. Life cycle of the malarial parasite. After Miss Anna Stryke.</span> +</div> + +<p>The experimental proof that mosquitoes of the Anopheline group +are necessary agents in the transmission of malaria was afforded in +1900 when two English physicians, Drs. Sambon and Low lived for +the three most malarial months in the midst of the Roman Campagna,<span class="pagenum"><a name="Page_193" id="Page_193">[Pg 193]</a></span> +a region famous for centuries as a hot-bed of malaria. The two +experimenters moved about freely throughout the day, exposed +themselves to rains and all kinds of weather, drank marsh water, +slept exposed to the marsh air, and, in short, did everything which +was supposed to cause malaria, except that they protected themselves +thoroughly from mosquito bites, retiring at sunset to a mosquito-proof<span class="pagenum"><a name="Page_194" id="Page_194">[Pg 194]</a></span> +hut. Though they took no quinine and all of their neighbors +suffered from malaria, they were absolutely free from the disease.</p> + +<p>To complete the proof, mosquitoes which had fed in Rome on +malarious patients were sent to England and allowed to bite two +volunteers, one of them Dr. Manson's own son, who had not been +otherwise exposed to the disease. Both of these gentlemen contracted +typical cases of malaria and the parasites were to be found in +abundance in their blood.</p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_127" id="Fig_127"></a> +<a href="images/f127-full.png"><img src="images/f127.png" width="400" height="489" alt="127. Eggs of Anopheles. After Howard." title="127. Eggs of Anopheles. After Howard." /></a> +<span class="caption">127. Eggs of Anopheles. After Howard.</span> +</div> + +<p>Since that time there have been many practical demonstrations +of the fact that malaria is transmitted exclusively by the bite of +mosquitoes and that the destruction +of the mosquitoes means the +elimination of the disease.</p> + +<p>We have said that the malarial +parasite is able to undergo its +development only in certain +species of mosquitoes belonging +to the Anopheline group. It is +by no means certain that all of +this group even, are capable of +acting as the definitive host of +the parasites, and much careful +experiment work is still needed +along this line. In the United +States, several species have been +found to be implicated, <i>Anopheles +quadrimaculatus</i> and <i>Anopheles +crucians</i> being the most common. The characteristics of these species +and the distinctions between them and other mosquitoes will be +discussed in <a href="#CHAPTER_XII">Chapter XII</a>.</p> + +<p>In antimalarial work it is desirable to distinguish the anopheline +mosquitoes from the culicine species in all stages. The following +tabulation presents the more striking distinctions between the groups +as represented in the United States.</p> + + +<div class="center"> +<table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td align="center"><i>Anopheles</i></td><td align="center"><i>Culex, Aedes, etc.</i></td></tr> +<tr> +<td align="left"> +<p><i>Eggs</i>: Laid singly in small +numbers upon the surface of the +water. Eggs lie upon their sides +and float by means of lateral +<span class="pagenum"><a name="Page_195" id="Page_195">[Pg 195]</a></span>expansions (<a href="#Fig_127">fig. 127</a>).</p> +</td> +<td align="left"> +<p>Deposited in clumps in the +form of a raft (Culex group) or +deposited singly in the water or +on the ground in places which +may later be submerged.</p> +</td> +</tr> +<tr> +<td align="left"> +<p><i>Larva</i>: When at rest floats in +a horizontal position beneath the +surface film. No respiratory +tube but instead a flattened +area on the eighth abdominal +segment into which the two +spiracles open (<a href="#Fig_128a">fig. 128</a>).</p> +</td> +<td align="left"> +<p>When at rest (with few exceptions) +floats suspended in an +oblique or vertical position, or +more rarely nearly horizontal, +with the respiratory tube in +contact with the surface film +(<a href="#Fig_128a">fig. 128</a>).</p> +</td> +</tr> +<tr> +<td align="left"> +<p><i>Adults</i>: Palpi in both sexes +nearly or quite as long as the +proboscis. Proboscis projecting +forward nearly on line with the +axis of the body. When at rest +on a vertical wall the body is +usually held at an angle with the +vertical (<a href="#Fig_128b">fig. 128</a>). Wings frequently +spotted (<a href="#Fig_130">fig. 130</a>).</p> +</td> +<td align="left"> +<p>Palpi short in the female, in +the male usually elongate. Proboscis +projects forward at an +angle with the axis of the body. +When at rest on a vertical wall +the body is usually held parallel +or the tip of the abdomen inclined +towards the wall (<a href="#Fig_128b">fig. 128</a>). +Wings usually not spotted.</p> +</td> +</tr> +</table></div> + +<div class="figcenter" style="width: 425px;"><a name="Fig_128a" id="Fig_128a"></a> +<a href="images/f128a-full.png"><img src="images/f128a.png" width="425" height="142" alt="128. (a) Normal position of the larvæ +of Culex and Anopheles in +the water. Culex, left; Anopheles, +middle; Culex pupa, +right hand figure." title="128. (a) Normal position of the larvæ +of Culex and Anopheles in +the water. Culex, left; Anopheles, +middle; Culex pupa, +right hand figure." /></a> +<span class="caption">128. (<i>a</i>) Normal position of the larvæ +of Culex and Anopheles in +the water. Culex, left; Anopheles, +middle; Culex pupa, +right hand figure.</span> +</div> + +<p>These malarial-bearing species are essentially domesticated +mosquitoes. They develop in any accumulation of water which +stands for a week or more. +Ponds, puddles, rain barrels, +horse troughs, cess-pools, cans, +even the foot-prints of animals +in marshy ground may +afford them breeding places.</p> + +<div class="figcenter" style="width: 150px;"><a name="Fig_128b" id="Fig_128b"></a> +<a href="images/f128b-full.png"><img src="images/f128b.png" width="150" height="402" alt="128. (b) Normal +position of +Culex and +Anopheles on +the wall." title="128. (b) Normal +position of +Culex and +Anopheles on +the wall." /></a> +<span class="caption">128. (<i>b</i>) Normal +position of +Culex and +Anopheles on +the wall.</span> +</div> + +<p>It is clear from what has been said regarding the life cycle of the +malarial parasite that the mosquito is harmless if not itself diseased. +Hence malarial-bearing species may abound in the +neighborhood where there is no malaria, the disease +being absent simply because the mosquitoes are uninfected. +Such a locality is potentially malarious and +needs only the introduction of a malarial patient who is +exposed to the mosquitoes. It is found that such patients +may harbor the parasites in their blood long after they +are apparently well and thus may serve as a menace, +just as do the so-called typhoid carriers. In some +malarious regions as high as 80-90 per cent of the natives +are such malaria-carriers and must be reckoned with in +antimalaria measures.</p> + +<p>Based upon our present day knowledge of the life cycle +of the malarial parasite the fight against the disease<span class="pagenum"><a name="Page_196" id="Page_196">[Pg 196]</a></span> +becomes primarily a problem in economic entomology,—it is a question +of insect control, in its broadest interpretation.</p> + +<div class="figcenter" style="width: 350px;"> +<a href="images/f129-full.png"><img src="images/f129.png" width="350" height="682" alt="129. Larva of Anopheles. After Howard." title="129. Larva of Anopheles. After Howard." /></a> +<span class="caption">129. Larva of Anopheles. After Howard.</span> +</div> + +<p>The lines of defence and offence +against the disease as outlined by +Boyce (1909) are:</p> + +<div class="blockquot"><div class="hanging"> +<p>1. Measures to avoid the reservoir +(man):</p> + +<div class="hanging"> +<p>Segregation.</p> + +<p>Screening of patients.</p> +</div> + +<p>2. Measures to avoid Anopheles:</p> + +<div class="hanging"> +<p>Choice of suitable locality, +when possible.</p> + +<p>Screening of houses and +porches.</p> + +<p>Sleeping under mosquito nets.</p> +</div> + +<p>3. Measures to exterminate the +Anopheles:</p> + +<div class="hanging"> +<p>Use of natural enemies.</p> + +<p>Use of culicides, oiling ponds, +etc.</p> + +<p>Drainage and scavenging to +destroy breeding places.</p> + +<p>Enforcement of penalties for +harboring larvæ or keeping +stagnant water.</p> + +<p>Educational methods.</p> +</div> + +<p>4. Systematic treatment with quinine to exterminate the parasites.</p> +</div></div> + + +<h3><a name="Mosquitoes_and_Yellow_Fever" id="Mosquitoes_and_Yellow_Fever"></a><span class="smcap">Mosquitoes and Yellow Fever</span></h3> + +<p>Yellow fever was until recently one of the most dreaded of epidemic +diseases. It is an acute, specific and infectious disease, non-contagious +in character but occurring in epidemics, or endemics, +within a peculiarly limited geographical area. It is highly fatal, +but those who recover are generally immune from subsequent attacks.</p> + +<p>It is generally regarded as an American disease, having been +found by Cortez, in Mexico, and being confined principally to the +American continents and islands. It also occurs in Africa and attempts +have been made to show that it was originally an African +disease but there is not sufficient evidence to establish this view.<span class="pagenum"><a name="Page_197" id="Page_197">[Pg 197]</a></span></p> + +<p>There have been many noted outbreaks in the United States. +Boston suffered from it in 1691 and again in 1693; New York in +1668 and as late as 1856; Baltimore in 1819. In 1793 occurred the +great epidemic in Philadelphia, with a death rate of one in ten of the +population. In the past century it was present almost every year in +some locality of our Southern States, New Orleans being the greatest +sufferer. In the latter city there were 7848 deaths from the disease +in 1853, 4854 in 1858, and 4046 in 1878. The last notable outbreak +was in 1905. Reed and Carroll (1901) estimated that during the +period from 1793 to 1900 there had not been less than 500,000 cases +in the United States.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_130" id="Fig_130"></a> +<a href="images/f130-full.png"><img src="images/f130.png" width="500" height="377" alt="130. Anopheles quadrimaculatus, male and female, (×3½). After Howard." title="130. Anopheles quadrimaculatus, male and female, (×3½). After Howard." /></a> +<span class="caption">130. Anopheles quadrimaculatus, male and female, (×3½). After Howard.</span> +</div> + +<p>As in the case of the plague, the most stringent methods of control +proved ineffective and helplessness, almost hopelessness marked +the great epidemics. A vivid picture of conditions is that given by +Mathew Cary, 1793 (quoted by Kelly, 1906) in "A Short Account of +the Malignant Fever Lately Prevalent in Philadelphia."</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_131" id="Fig_131"></a> +<a href="images/f131-full.png"><img src="images/f131.png" width="500" height="534" alt="131. Anopheles punctipennis. Female, (×4). After Howard." title="131. Anopheles punctipennis. Female, (×4). After Howard." /></a> +<span class="caption">131. Anopheles punctipennis. Female, (×4). After Howard.</span> +</div> + +<p>"The consternation of the people of Philadelphia at this period +was carried beyond all bounds. Dismay and affright were visible<span class="pagenum"><a name="Page_198" id="Page_198">[Pg 198]</a></span> +in the countenance of almost every person. Of those who remained, +many shut themselves in their houses and were afraid to walk the +streets. * * * The corpses of the most respectable citizens, +even those who did not die of the epidemic, were carried to the grave +on the shafts of a chair (chaise), the horse driven by a negro, unattended +by friends or relative, and without any sort of ceremony. +People hastily shifted their course at the sight of a hearse coming +toward them. Many never walked on the footpath, but went into +the middle of the streets to avoid being infected by passing by houses +wherein people had died. Acquaintances and friends avoided each +other in the streets and only signified their regard by a cold nod. +The old custom of shaking hands fell into such disuse that many +shrunk back with affright at even the offer of the hand. A person<span class="pagenum"><a name="Page_199" id="Page_199">[Pg 199]</a></span> +with a crape, or any appearance of mourning was shunned like a +viper. And many valued themselves highly on the skill and address +with which they got to the windward of every person they met. +Indeed, it is not probable that London, at the last stage of the plague, +exhibited stronger marks of terror than were to be seen in Philadelphia +from the 24th or 25th of August until pretty late in September."</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_132" id="Fig_132"></a> +<a href="images/f132-full.png"><img src="images/f132.png" width="500" height="511" alt="132. Anopheles crucians. Female (×4). After Howard." title="132. Anopheles crucians. Female (×4). After Howard." /></a> +<span class="caption">132. Anopheles crucians. Female (×4). After Howard.</span> +</div> + +<p>Such was the condition in Philadelphia in 1793 and, as far as +methods of control of the disease were concerned, there was practically +no advance during the last century. The dominant theory +was that yellow fever was spread by <i>fomites</i>, that is, exposed bedding, +clothing, baggage, and the like. As late as 1898 a bulletin of the +United States Marine Hospital Service stated:<span class="pagenum"><a name="Page_200" id="Page_200">[Pg 200]</a></span></p> + +<p>"While yellow fever is a communicable disease, it is not contagious +in the ordinary acceptance of the term, but is spread by the +infection of places and articles of bedding, clothing, and furniture."</p> + +<p>Based upon this theory, houses, baggage, freight, even mail, +were disinfected, and the most rigid quarantine regulations were +enforced. The hardships to which people of the stricken regions +were subjected and the financial losses are incalculable. And withal, +the only efficient check upon the disease seemed to be the heavy frosts. +It was found that for some reason, the epidemic abated with cold +weather,—a measure beyond human control.</p> + +<div class="figcenter" style="width: 500px;"> +<a href="images/f133-full.png"><img src="images/f133.png" width="500" height="518" alt="133. Culex sollicitans. Female (×4). After Howard." title="133. Culex sollicitans. Female (×4). After Howard." /></a> +<span class="caption">133. Culex sollicitans. Female (×4). After Howard.</span> +</div> + +<p>It is not strange that among the multitude of theories advanced to +explain the cause and method of dissemination of the disease there +should be suggestions that yellow fever was transmitted by the +mosquito. We have seen that Beauperthuy (1855) clearly urged +this theory.</p> + +<p>More detailed, and of the greatest influence in the final solution +of the problem were the arguments of Dr. Cárlos Finlay, of Havana. +In 1881, in a paper presented before the "Real Academia de Ciencias +Médicas, Físicas y Naturales de la Habana," he said:<span class="pagenum"><a name="Page_201" id="Page_201">[Pg 201]</a></span></p> + +<p>"I feel convinced that any theory which attributes the origin and +the propagation of yellow fever to atmospheric influences, to miasmatic +or meteorological conditions, to filth, or to the neglect of general +hygienic precautions, must be considered as utterly indefensible."</p> + +<p>He postulated the existence of a material transportable substance +causing yellow fever,—"something tangible which requires to be +conveyed from the sick to the healthy before the disease can be +propagated" and after discussing the peculiarities of the spread of +the disease and the influence of meteorological conditions, he decides +that the carriers of the disease must be sought among insects. He +continues:</p> + +<p>"On the other hand, the fact of yellow fever being characterized +both clinically and (according to recent findings) histologically, by +lesions of the blood vessels and by alterations of the physical and +chemical conditions of the blood, suggested that the insect which +should convey the infectious particles from the patient to the healthy +should be looked for among those which drive their sting into blood +vessels in order to suck human blood. Finally, by reason of other +considerations which need not be stated here, I came to think that +the mosquito might be the transmitter of yellow fever."</p> + +<p>"Assimilating the disease to small-pox and to vaccination, it +occurred to me that in order to inoculate yellow fever it would be +necessary to pick out the inoculable material from within the blood +vessels of a yellow fever patient and to carry it likewise into the +interior of a blood vessel of a person who was to be inoculated. All +of which conditions the mosquito satisfies most admirably through +its bite."</p> + +<p>In the course of his study of the problem, Finlay made detailed +studies of the life history and habits of the common mosquitoes at +Havana, and arrived at the conclusion that the carrier of the yellow +fever was the <i>Culex mosquito</i> or <i>Aëdes calopus</i>, as it is now known. +With this species he undertook direct experimental tests, and believed +that he succeeded in transmitting the disease by the bite of +infected mosquitoes in three cases. Unfortunately, possibility +of other exposure was not absolutely excluded, and the experiments +attracted little attention.</p> + +<p>Throughout the next twenty years Finlay continued his work on +yellow fever, modifying his original theory somewhat as time went on. +Among his later suggestions was that in the light of Smith's work +on Texas fever, his theory must be "somewhat modified so as to<span class="pagenum"><a name="Page_202" id="Page_202">[Pg 202]</a></span> +include the important circumstance that the faculty of transmitting +the yellow fever germ need not be limited to the parent insect, +directly contaminated by stinging a yellow fever patient (or perhaps +by contact with or feeding from his discharges), but may be +likewise inherited by the next generation of mosquitoes issued from +the contaminated parent." He believed that the bite of a single +mosquito produced a light attack of the disease and was thus effective +in immunizing the patient. Throughout the period, many +apparently successful attempts to transmit the disease by mosquitoes +were made. In the light of present day knowledge we must +regard these as defective not only because possibility of other infection +was not absolutely excluded but because no account was taken +of the incubation period within the body of the mosquito.</p> + +<p>In 1900, while the American army was stationed in Cuba there +occurred an epidemic of yellow fever and an army medical board was +appointed for "the purpose of pursuing scientific investigations with +reference to the acute infectious diseases prevalent on the island." +This was headed by Walter Reed and associated with him were James +Carroll, Jesse W. Lazear and Aristides Agramonte, the latter a Cuban +immune. For a detailed summary of this work the lay reader cannot +do better than read Dr. Kelly's fascinating biography "Walter +Reed and Yellow Fever."</p> + +<p>Arriving at the army barracks near Havana the Commission first +took up the study of <i>Bacillus icteroides</i>, the organism which Sanarelli, +an Italian physician, had declared the causative agent in yellow fever. +They were unable to isolate this bacillus either from the blood during +life or from the blood and organs of cadavers and therefore turned +their attention to Finlay's theory of the propagation of yellow fever +by means of the mosquito. In this work they had the unselfish +and enthusiastic support of Dr. Finlay himself, who not only consulted +with them and placed his publications at their disposal, but furnished +eggs from which their experimental mosquitoes were obtained. +Inoculations of eleven non-immunes through the bite of infected +mosquitoes were made, and of these, two gave positive results. The +first of the two was Dr. Carroll who allowed himself to be bitten +by a mosquito which had been caused to feed upon four cases of +yellow fever, two of them severe and two mild. The first patient +had been bitten twelve days before.</p> + +<p>Three days after being bitten, Dr. Carroll came down with a +typical case of yellow fever. So severe was the attack that for three<span class="pagenum"><a name="Page_203" id="Page_203">[Pg 203]</a></span> +days his life hung in the balance. During his convalescence an +incident occurred which showed how the theory of mosquito transmission +of the disease was generally regarded. We quote from Dr. +Kelly: "One of his nurses who came from Tennessee had had considerable +experience with yellow fever, having indeed, lost her husband +and several children from it. One day early in his illness Dr. +Carroll mentioned to her that he had contracted the disease through +the bite of a mosquito, and noticed that she looked surprised. Some +time later, when well enough to look over the daily records of his +condition, he found this entry: 'Says he got his illness through the +bite of a mosquito,—delirious'."</p> + +<p>The second case was that of an American who was bitten by four +mosquitoes, two of which had bitten severe (fatal) cases of yellow +fever twelve days previously, one of which had bitten a severe case +(second day) sixteen days before and one which had bitten a severe +case eight days before. Five days later, the subject developed a well +pronounced but mild case of the disease.</p> + +<p>In the meantime, another member of the Commission, Dr. Lazear, +was accidentally bitten by a mosquito while collecting blood from +yellow fever patients. Five days later he contracted a typical case +which resulted fatally.</p> + +<p>So clear was the evidence from these preliminary experiments +that the commission felt warranted in announcing, October 27, 1900, +that, "The mosquito serves as the intermediate host for the parasite +of yellow fever, and it is highly probable that the disease is only +propagated through the bite of this insect."</p> + +<p>In order to extend the experimental evidence under conditions +which could leave no possibility of infection from other sources, a +special experimental sanitary station, named in honor of the deceased +member of the Commission, was established in an open field near +the town of Quemados, Cuba. Here there were constructed two small +buildings known respectively as the "infected clothing building" +and the "infected mosquito building."</p> + +<p>The infected clothing building, 14 × 20 feet in size, was purposely +so constructed as to exclude anything like efficient ventilation, but +was thoroughly screened to prevent the entrance of mosquitoes. +Into this building were brought sheets, pillow-slips, blankets, etc., +contaminated by contact with cases of yellow fever and their discharges,—many +of them purposely soiled with a liberal quantity of +black vomit, urine, and fecal matter from patients sick with yellow<span class="pagenum"><a name="Page_204" id="Page_204">[Pg 204]</a></span> +fever. Nothing could better serve as the fomites which were supposed +to convey the dread disease.</p> + +<p>Three non-immunes unpacked these articles, giving each a +thorough handling and shaking in order to disseminate through the +air of the room the specific agent of the disease. They were then +used in making up the beds which the volunteers occupied each night +for a period of twenty days. The experiment was repeated three +times, volunteers even sleeping in the soiled garments of yellow fever +victims but in not a single case was there the slightest symptom of +disease. The theory of the spread of yellow fever by fomites was +completely demolished.</p> + +<p>The infected mosquito building, equal in size to its companion, +was the antithesis as far as other features were concerned. It was +so constructed as to give the best possible ventilation, and bedding +which was brought into it was thoroughly sterilized. Like the +infected clothing building it was carefully screened, but in this case +it was in order to keep mosquitoes in it as well as to prevent entrance +of others. Through the middle of the room ran a mosquito-proof +screen.</p> + +<p>On December 5, 1900, a non-immune volunteer who had been in +the quarantine camp for fifteen days and had had no other possible +exposure, allowed himself to be bitten by five mosquitoes which had +fed on yellow fever patients fifteen or more days previously. The +results were fully confirmatory of the earlier experiments of the +Commission—at the end of three days, nine and a half hours, the +subject came down with a well marked case of yellow fever.</p> + +<p>In all, ten cases of experimental yellow fever, caused by the bite +of infected mosquitoes were developed in Camp Lazear. Throughout +the period of the disease, other non-immunes slept in the little +building, separated from the patient only by the mosquito-proof +screen, but in no circumstances did they suffer any ill effects.</p> + +<p>It was found that a yellow fever patient was capable of infecting +mosquitoes only during the first three or four days after coming +down with the disease. Moreover, after the mosquito has bitten +such a patient, a period of at least twelve days must elapse before +the insect is capable of transmitting the disease.</p> + +<p>Once the organism has undergone its twelve day development, +the mosquito may remain infective for weeks. In experiments of +the Commission, two of the mosquitoes transmitted the disease to a +volunteer fifty-seven days after their contamination. No other<span class="pagenum"><a name="Page_205" id="Page_205">[Pg 205]</a></span> +volunteers presenting themselves, one of these mosquitoes died the +sixty-ninth and one the seventy-first day after their original contamination, +without it being determined whether they were still +capable of transmitting the disease.</p> + +<p>So carefully carried out was this work and so conclusive were the +results that Dr. Reed was justified in writing:</p> + +<p>"Six months ago, when we landed on this island, absolutely nothing +was known concerning the propagation and spread of yellow +fever—it was all an unfathomable mystery—but today the curtain +has been drawn—its mode of propagation is established and we know +that a case minus mosquitoes is no more dangerous than one of +chills and fever."</p> + +<p>The conclusions of the Commission were fully substantiated by +numerous workers, notably Dr. Guiteras of the Havana Board of +Health, who had taken a lively interest in the work and whose +results were made known in 1901, and by the Brazilian and French +Commission at Sao Paulo, Brazil, in 1903.</p> + +<p>Throughout the work of the Army Commission and down to the +present time many fruitless efforts have been made to discover the +specific organism of yellow fever. It was clearly established that +the claims of Sanarelli for <i>Bacillus icteroides</i> were without foundation. +It was found, too, that whatever the infective agent might +be it was capable of passing through a Berkefeld filter and thus belongs +to the puzzling group of "filterable viruses." It was further +found that the virus was destroyed by heating up to 55° C for ten +minutes. It is generally believed that the organism is a Protozoan.</p> + +<p>The question of the hereditary transmission of the yellow fever +organism within the mosquito was left unsettled by the Army Commission, +though, as we have seen, it was raised by Finlay. Marchoux +and Simond, of the French Commission devoted much attention to +this phase of the problem and basing their conclusions on one apparently +positive case, they decided that the disease could be transmitted +through the egg of an infected <i>Aëdes calopus</i> to the second +generation and thence to man. The conclusion, which is of very +great importance in the control of yellow fever, has not been verified +by other workers.</p> + +<p>Once clearly established that yellow fever was transmitted solely +by mosquitoes, the question of the characteristics, habits, and geographical +distribution of the insect carrier became of vital importance.<span class="pagenum"><a name="Page_206" id="Page_206">[Pg 206]</a></span></p> + +<p><i>Aëdes calopus</i>, more commonly known as <i>Stegomyia fasciata</i> or +<i>Stegomyia calopus</i> (<a href="#Fig_134">fig. 134</a>) is a moderate sized, rather strikingly +marked mosquito. The general color is dark-brown or reddish-brown, +but the thorax has a conspicuous broad, silvery-white curved +line on each side, with two parallel median silvery lines. Between +the latter there is a +slender, broken line. +The whole gives a lyre-shaped +pattern to the +thorax. The abdomen +is dark with silvery-white +basal bands and +silvery white spots on +each side of the abdominal +segments. +Legs black with rings +of pure white at the +base of the segments.</p> + +<p>Size of the female +3.3 to 5 mm.; male 3 +to 4.5 mm.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_134" id="Fig_134"></a> +<a href="images/f134-full.png"><img src="images/f134.png" width="500" height="702" alt="134. The yellow fever mosquito (Aëdes calopus). (×7). +After Howard." title="134. The yellow fever mosquito (Aëdes calopus). (×7). +After Howard." /></a> +<span class="caption">134. The yellow fever mosquito (Aëdes calopus). (×7). +After Howard.</span> +</div> + +<p>It is preeminently +a domesticated species, +being found almost +exclusively about the +habitation of man. +"Its long association +with man is shown by +many of its habits. It +approaches stealthily +from behind. It retreats +upon the slightest +alarm. The ankles and, when one is sitting at a table or desk, +the underside of the hands and wrists are favorable points of attack. +It attacks silently, whereas other mosquitoes have a piping or humming +note. The warning sound has doubtless been suppressed in +the evolutionary process of its adaptation to man. It is extremely +wary. It hides whenever it can, concealing itself in garments, +working into the pockets, and under the lapels of coats, and crawling +up under the clothes to bite the legs. In houses, it will hide<span class="pagenum"><a name="Page_207" id="Page_207">[Pg 207]</a></span> +in dark corners, under picture moldings and behind the heads of +old-fashioned bedsteads. It will enter closets and hide in the folds +of garments."—Howard.</p> + +<p>It was claimed by the French Commission, and subsequently +often stated in discussions of the relation of the mosquito to yellow +fever that the mature <i>Aëdes calopus</i> will bite only at night. If this +were true it would be of the greatest importance in measures to +avoid the disease. Unfortunately, the claim was illy founded and +numerous workers have clearly established that the exact converse +is more nearly true, this mosquito being pre-eminently a day species, +feeding most actively in early morning, +about sunrise, and late in the afternoon. +On cloudy days it attacks at any time +during the day. Thus there is peril in +the doctrine that infected regions may +be visited with perfect safety during +the daytime and that measures to +avoid the mosquito attack need be +taken only at night.</p> + +<div class="figright" style="width: 350px;"> +<a href="images/f135a-full.png"><img src="images/f135a.png" width="350" height="430" alt="135a. Aëdes calopus. Pupa. +After Howard." title="135a. Aëdes calopus. Pupa. +After Howard." /></a> +<span class="caption">135<i>a</i>. Aëdes calopus. Pupa. +After Howard.</span> +</div> + +<p>Dr. Finlay maintained that the +adult, even when starved, would not +bite when the temperature was below +23° C, but subsequent studies have +shown that this statement needs modification. +The French Commission, +working at Rio de Janeiro, found that <i>Aëdes calopus</i> would bite +regularly at temperatures between 22° and 25° and that the optimum +temperature was between 27° and 30° C, but their experiments led +them to believe that it would bite in nature at a temperature as +low as 17° C.</p> + +<p>The yellow fever mosquito breeds in cisterns, water barrels, +pitchers and in the various water receptacles about the house. In +our own Southern States it very commonly breeds in the above-ground +cisterns which are in general use. Often the larvæ (<a href="#Fig_135b">fig. 135b</a>) +are found in flower vases, or even in the little cups of water which +are placed under the legs of tables to prevent their being overrun by +ants. They have been repeatedly found breeding in the holy water +font in churches. In short, they breed in any collection of water in +close proximity to the dwellings or gathering places of man.<span class="pagenum"><a name="Page_208" id="Page_208">[Pg 208]</a></span></p> + +<p>The life cycle under favorable conditions is completed in from +twelve to fifteen days. These figures are of course very dependent +upon the temperature. The Army Commission in Cuba found that +the cycle might be completed in as brief a period as nine and a half +days. Under less favorable conditions it may be greatly lengthened.</p> + +<p>The adults are long lived. We have +seen that during the experimental work +in Cuba specimens were kept in captivity +for sixty-nine and seventy-one +days, respectively, and that they were +proved to retain their infectivity for at +least fifty-seven days. Dr. Guiteras +subsequently kept an infected adult for +one hundred and fifty-four days.</p> + +<p>Low temperatures have a very great +effect not only on development, but on +the activity and even life of the adults. +Long before the method of transmission +of yellow fever was discovered it was well +known that the epidemics were brought +to a close by heavy frosts, and it is now +known that this is due to the killing of +the mosquitoes which alone could spread +the disease.</p> + +<div class="figleft" style="width: 300px;"><a name="Fig_135b" id="Fig_135b"></a> +<a href="images/f135b-full.png"><img src="images/f135b.png" width="300" height="695" alt="135b. Aëdes calopus; larva. (×7). +After Howard." title="135b. Aëdes calopus; larva. (×7). +After Howard." /></a> +<span class="caption">135<i>b</i>. Aëdes calopus; larva. (×7). +After Howard.</span> +</div> + +<p><i>Aëdes calopus</i> has a very wide distribution +since, as Howard says, being a +domestic mosquito, having a fairly long +life in the adult stage, and having the +custom of hiding itself in the most ingenious +ways, it is particularly subject to carriage +for long distances on board vessels, +in railway trains, even packed in baggage. In general, its permanent +distribution is from 40 degrees north latitude to 40 degrees south +latitude (Brumpt), in a belt extending around the world. In the +United States it breeds in most of our Southern States.</p> + +<p>Thus, as in the case of malaria, there are many places where the +insect carrier is abundant but where yellow fever does not occur. +Such, for instance, are Hawaii, Australia and Asia. An outbreak may +occur at any time that a patient suffering from the disease is allowed +to enter and become a source of infection for the mosquitoes. In<span class="pagenum"><a name="Page_209" id="Page_209">[Pg 209]</a></span> +this connection various writers have called attention to the menace +from the Panama Canal. When it is completed, it will allow of +direct passage from regions where yellow fever is endemic and this +will greatly increase the possibility of its introduction into these places +where it is now unknown. The result, with a wholly non-immune +population, would be appalling.</p> + +<p>On the other hand, there are places wholly outside of the normal +range of <i>Aëdes calopus</i> where the disease has raged. Such are New +York, Boston, and even Philadelphia, which have suffered notable +epidemics. These outbreaks have been due to the introduction of +infected mosquitoes during the heat of summer, when they have not +only conveyed the disease but have found conditions favorable +for their multiplication. Or, uninfected mosquitoes have been thus +accidentally brought in and developed in large numbers, needing +then only the accidental introduction of cases of the disease to start +an epidemic.</p> + +<p>Methods of control of various diseases have been revolutionized +by the discovery that they were insect-borne, but in no other case +has the change been as radical or the results as spectacular as in the +case of yellow fever. The "shot-gun quarantine," the sufferings and +horrors, the hopelessness of fighting absolutely blindly have given +way to an efficient, clear-cut method of control, based upon the knowledge +that the disease is carried from man to man solely by the mosquito, +<i>Aëdes calopus</i>. The lines of defense and offense are essentially +as follows:</p> + +<p>In the first place, when a case of yellow fever occurs, stringent +precautions must be adopted to prevent the infection of mosquitoes +and the escape of any already infected. This means that the patient +must be removed to a mosquito-proof room, or ward beyond reach of +the insects, and that the infected room must be thoroughly fumigated +at once, to kill the mosquitoes hiding within it. All cracks +and openings should be closed with strips of paper and fumigation +with burning sulphur or pyrethrum carefully carried out.</p> + +<p>It should be remembered that if the first case noted is that of a +resident rather than imported, it means that the mosquito carriers +became infected more than two weeks before the case was diagnosed, +for as we have seen, the germ must undergo a twelve-day period of +development within its insect host. Therefore a careful search must +be made for mild cases which, though unrecognized, may serve as +foci for the spread of the disease.<span class="pagenum"><a name="Page_210" id="Page_210">[Pg 210]</a></span></p> + +<p>In face of a threatened epidemic one of the most essential measures +is to educate the citizens and to gain their complete coöperation in +the fight along modern lines. This may be done through the schools, +the pulpit, places of amusement, newspapers and even bulletin +boards.</p> + +<p>Emphasis should be placed on the necessity of both non-immunes +and immunes using mosquito curtains, and in all possible ways +avoiding exposure to the mosquitoes.</p> + +<p>Then the backbone of the fight must be the anti-mosquito measures. +In general, these involve screening and fumigating against +adults, and control of water supply, oiling, and drainage against the +larvæ. The region involved must be districted and a thorough survey +undertaken to locate breeding places, which must, if possible, +be eradicated. If they are necessary for water supplies, such as +casks, or cisterns, they should be carefully screened to prevent +access of egg-laying adults.</p> + +<p>The practical results of anti-mosquito measures in the fight +against yellow fever are well illustrated by the classic examples of +the work in Havana, immediately following the discoveries of the +Army Commission and by the stamping out of the New Orleans +epidemic in 1905.</p> + +<p>The opportunities for an immediate practical application of the +theories of the Army Commission in Havana were ideal. The city +had always been a hotbed of yellow fever and was the principal +source from which the disease was introduced year after year into +our Southern States. It was under martial law and with a military +governor who was himself a physician and thoroughly in sympathy +with the views of the Commission, the rigid enforcement of the +necessary regulations was possible. The story of the first campaign +has been often told, but nowhere more clearly than in Dr. Reed's +own account, published in the <i>Journal of Hygiene</i> for 1902.</p> + +<p>Closer home was the demonstration of the efficacy of these +measures in controlling the yellow fever outbreak in New Orleans +in 1905. During the spring and early summer of the year the disease +had, unperceived, gained a firm foothold in that city and when, in +early July the local Board of Health took cognizance of its existence, +it was estimated that there had been in the neighborhood of one +hundred cases.</p> + +<p>Conditions were not as favorable as they had been under martial +law in Havana for carrying on a rigid fight along anti-mosquito lines.<span class="pagenum"><a name="Page_211" id="Page_211">[Pg 211]</a></span> +The densely populated city was unprepared, the public had to be +educated, and an efficient organization built up. The local authorities +actively began a general fight against the mosquito but in spite +of their best efforts the disease continued to spread. It was recognized +that more rigid organization was needed and on August 12th +the United States Public Health and Marine Hospital Service was +put in absolute charge of the fight. Up to this time there had been +one hundred and forty-two deaths from a total of nine hundred and +thirteen cases and all of the conditions seemed to threaten an outbreak +to exceed the memorable one of 1878 when, as we have seen +there were four thousand and forty-six deaths.</p> + +<p>With the hearty coöperation of the citizens,—physicians and +laymen alike,—the fight was waged and long before frost or any near +approach thereto the disease was stamped out,—a thing unheard of +in previous epidemics. The total loss of life was four hundred and +sixty—about 11 per cent as great as that from the comparable epidemic +of 1878. If the disease had been promptly recognized and +combated with the energy which marked the fight later in the summer, +the outbreak would have made little headway and the great +proportion of these lives would have been saved.<span class="pagenum"><a name="Page_212" id="Page_212">[Pg 212]</a></span></p> + + + + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_IX" id="CHAPTER_IX"></a>CHAPTER IX</h2> + +<h2><span title="insert "continued"">ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA</span></h2> + + +<h3><span class="smcap"><a name="Insects_and_Trypanosomiases" id="Insects_and_Trypanosomiases"></a>Insects and Trypanosomiases</span></h3> + +<p>By trypanosomiasis is meant a condition of animal parasitism, +common to man and the lower animals, in which trypanosomes, +peculiar flagellate protozoa, infest the blood. Depending upon the +species, they may be harmless, producing no appreciable ill-effect, +or pathogenic, giving rise to conditions of disease. A number of +these are known to be transferred by insects.</p> + +<p>In order that we may +consider more fully the +developmental stage of +these parasites within +their insect host, it is +necessary that we describe +briefly the structure +of the blood-inhabiting +stage.</p> + +<div class="figcenter" style="width: 360px;"><a name="Fig_136" id="Fig_136"></a> +<a href="images/f136-full.png"><img src="images/f136.png" width="360" height="290" alt="136. Trypanosome brucei. After Bruce." title="136. Trypanosome brucei. After Bruce." /></a> +<span class="caption">136. Trypanosoma brucei. After Bruce.</span> +</div> + +<p>The trypanosomes are +elongated, usually pointed, +flagellated protozoa +(<a href="#Fig_136">fig. 136</a>) in which the +single flagellum, bent +under the body, forms the +outer limit of a delicate undulating membrane. It arises near +one end of the organism from a minute centrosome-like body +which is known as the blepheroplast, and at the opposite end extends +for a greater or less distance as a free flagellum. Enclosing, or +close beside the blepheroplast is the small kinetonucleus. The +principal nucleus, round or oval in form, is situated near the center +of the body. Asexual reproductions occurs in this stage, by longitudinal +fission, the nucleus and the blepheroplast dividing independently +of one another. From the blepheroplast of one of the daughter +cells a new flagellum is formed.</p> + +<p>Among the pathogenic species are to be found the causative +organisms of some of the most serious diseases of domestic animals +and even of man. It is probable that these pathogenic species secrete<span class="pagenum"><a name="Page_213" id="Page_213">[Pg 213]</a></span> +a specific poison. The majority of them are tropical in distribution.</p> + +<p>Though we are concerned especially with the species which infest +man, we shall first consider two of the trypanosomes of lower animals, +known long before any of those of man had been found.</p> + + +<p><a name="Fleas_and_Lice_as_Carriers_of_Trypanosoma_lewisi" id="Fleas_and_Lice_as_Carriers_of_Trypanosoma_lewisi"></a><b>Fleas and Lice as Carriers of Trypanosoma lewisi.</b>—<i>Trypanosoma +lewisi</i>, the first mammalian trypanosome known, is to be found in the +blood of wild rats. Like its host, it appears to be cosmopolitan in +distribution, having been reported from several localities in the +United States, Brazil, Argentine, England, Germany, France, Italy, +Russia, Asia and Africa.</p> + +<p>This species is usually regarded as non-pathogenic, but in experimental +work, especially with white rats, heavy infestations often +result fatally, and naturally infested specimens sometimes show +evidence of injury. Rats which have been infested exhibit at least +temporary immunity against new infection.</p> + +<p><i>Trypanosoma lewisi</i> is transmitted from rat to rat by fleas and +by lice. Rabinowitsch and Kempner (1899) first found that healthy +rats which were kept with infested rats, showed trypanosomes in +their blood after about two weeks. They found the trypanosomes +in the alimentary canal of fleas which had fed on the diseased rats. +On teasing such fleas in physiological salt solution and inoculating +them into fresh rats they were able to produce the infection. Finally, +they showed that the fleas which had fed upon infested rats were +able to carry the parasites to healthy rats. Corresponding experiments +with lice were not successful. Prowazek (1905) found in the +rat louse (<i>Hæmatopinus spinulosus</i>) organisms which he regarded +as developmental stages of the <i>Trypanosoma lewisi</i>. He believed +that the sexual cycle was undergone in this insect.</p> + +<p>Nuttall (1908) readily transmitted the trypanosomes through the +agency of fleas, (<i>Ceratophyllus fasciatus</i> and <i>Ctenopthalmus agyrtes</i>). +He believes that these insects are probably the chief transmitters +of the parasite. He was also able to transmit it from diseased to +healthy rats through the agency of the rat louse. He was unable +to trace any developmental stages in the louse and inclined to the +opinion that Prowazek was deceived by the presence of extraneous +flagellates such as are known to exist in a number of blood-sucking +arthropods.</p> + +<p>Nuttall concludes that since three distinct kinds of blood-sucking +insects are capable of transmitting <i>Trypanosoma lewisi</i> it appears<span class="pagenum"><a name="Page_214" id="Page_214">[Pg 214]</a></span> +doubtful that this flagellate is a parasite of the invertebrate "host" +in the sense claimed by Prowazek and other investigators.</p> + + +<p><a name="Tsetse-flies_and_Nagana" id="Tsetse-flies_and_Nagana"></a><b>Tsetse-flies and Nagana</b>—One of the greatest factors in retarding +the development of certain regions of Africa has been the presence +of a small fly, little larger than the common house-fly. This is the +tsetse-fly, <i>Glossina morsitans</i> (<a href="#Fig_165">fig. 165</a>) renowned on account of the +supposed virulence of its bite for cattle, horses and other domestic +mammals.</p> + +<p>The technical characteristics of the tsetse-flies, or Glossinas, and +their several species, will be found in a later chapter. We need +emphasize only that they are blood-sucking Muscidæ and that, +unlike the mosquitoes, the sexes resemble each other closely in structure +of the mouth-parts, and in feeding habits.</p> + +<p>In 1894, Colonel David Bruce discovered that the fly was not in +itself poisonous but that the deadly effect of its bite was due to the +fact that it transmitted a highly pathogenic blood parasite, <i>Trypanosoma +brucei</i>. This trypanosome Bruce had discovered in the blood +of South African cattle suffering from a highly fatal disease known as +"nagana". On inoculating the blood of infected cattle into horses +and dogs he produced the disease and found the blood teeming with +the causative organism. In the course of his work he established +beyond question that the "nagana" and the tsetse-fly disease were +identical.</p> + +<p>Tsetse-flies of the species <i>Glossina morsitans</i>, which fed upon +diseased animals, were found capable of giving rise to the disease +in healthy animals up to forty-eight hours after feeding. Wild +tsetse-flies taken from an infected region to a region where they did +not normally occur were able to transmit the disease to healthy +animals. It was found that many of the wild animals in the tsetse-fly +regions harbored <i>Trypanosoma brucei</i> in their blood, though they +showed no evidence of disease. As in the case of natives of malarial +districts, these animals acted as reservoirs of the parasite. Non-immune +animals subjected to the attacks of the insect carrier, quickly +succumbed to the disease.</p> + +<p>A question of prime importance is as to whether the insect serves +as an essential host of the pathogenic protozoan or whether it is a +mere mechanical carrier. Bruce inclined to the latter view. He was +unable to find living trypanosomes in the intestines or excrements +of the fly or to produce the disease on the many occasions when he<span class="pagenum"><a name="Page_215" id="Page_215">[Pg 215]</a></span> +injected the excrement into healthy animals. Moreover, he had +found that the experimental flies were infective only during the first +forty-eight hours and that if wild flies were taken from the infected +region, "kept without food for three days and then fed on a healthy +dog, they never gave rise to the disease."</p> + +<p>Koch had early described what he regarded as sexual forms from +the intestine of the fly but it remained for Kleine (1909) to experimentally +demonstrate that a part of the life cycle of the parasite +was undergone in the fly. Working with <i>Glossina palpalis</i>, he found +that for a period of ten days or longer after feeding on an animal +suffering from nagana it was non-infective, but that then it became +infective and was able to transmit the disease for weeks thereafter. +He discovered and described developmental stages of the parasite +within the intestine of the insect. In other words, the tsetse-fly +(in nature, <i>Glossina morsitans</i>), serves as an essential host, within +which an important part of the life cycle of the parasite is undergone. +These conclusions were quickly verified by Bruce and numerous +other workers and are no longer open to question. Klein and Taute +are even inclined to think that mechanical transmission plays practically +no rôle in nature, unless the fly is interrupted while feeding +and passes immediately to a new animal.</p> + + +<p><a name="Tsetse-flies_and_Sleeping_Sickness_of_Man" id="Tsetse-flies_and_Sleeping_Sickness_of_Man"></a><b>Tsetse-flies and Sleeping Sickness of Man</b>—About the beginning +of the present century a hitherto little known disease of man began +to attract great attention on account of its ravages in Uganda and +the region of Victoria Nyanza in South Africa. It was slow, insiduous +and absolutely fatal, characterized in its later stages by dullness, +apathy, and finally absolute lethargy all day long, symptoms which +gave it the name of "sleeping sickness."</p> + +<p>It was soon found that the disease was not a new one but that it +had been known for over a hundred years on the west coast of Africa. +Its introduction into Central and East Africa and its rapid spread +have been attributed primarily to the development of the country, +the formation of new trade routes and the free mingling of native +tribes formerly isolated. It is estimated that in the first ten years +of the present century there were approximately two hundred +thousand deaths from the disease in the Uganda protectorate. In +the British province Bugosa, on the Victoria Nyanza there were +thirty thousand deaths in the period from 1902-1905.<span class="pagenum"><a name="Page_216" id="Page_216">[Pg 216]</a></span></p> + +<p>While the disease is peculiarly African there are a number of +instances of its accidental introduction into temperate regions. +Slaves suffering from it were occasionally brought to America in +the early part of the last century and cases have sometimes been +imported into England. In none of the cases did the disease gain a +foothold or spread at all to other individuals.</p> + +<p>In 1902 Dutton described a trypanosome, <i>T. gambiense</i>, which he +and Forde had found the year before in the blood of a patient suffering +from a peculiar type of fever in Gambia. In 1902-1903 Castellani +found the same parasite in the cerebro-spinal fluid of sleeping-sickness +patients and definitely reported it as the causative organism +of the disease. His work soon found abundant confirmation, and +it was discovered that the sleeping sickness was but the ultimate +phase of the fever discovered by Dutton and Forde.</p> + +<p>When Castellani made known his discovery of the trypanosome +of sleeping sickness, Brumpt, in France, and Sambon, in England, +independently advanced the theory that the disease was transmitted +by the tsetse-fly, <i>Glossina palpalis</i>. This theory was based upon the +geographical distribution and epidemiology of the disease. Since +then it has been abundantly verified by experimental evidence.</p> + +<p>Fortunately for the elucidation of problems relating to the methods +of transfer of sleeping sickness, <i>Trypanosoma gambiense</i> is pathogenic +for many species of animals. In monkeys it produces symptoms +very similar to those caused in man. Bruce early showed that +<i>Glossina palpalis</i> "fed on healthy monkeys eight, twelve, twenty-four +and forty-eight hours after having fed on a native suffering from +trypanosomiasis, invariably transmitted the disease. After three +days the flies failed to transmit it." In his summary in Osler's +Modern Medicine, he continues "But this is not the only proof that +these flies can carry the infective agent. On the lake shore there +was a large native population among whom we had found about +one-third to be harboring trypanosomes in their blood. The tsetse-flies +caught on this lake shore, brought to the laboratory in cages, +and placed straightway on healthy monkeys, gave them the disease +in every instance, and furnished a startling proof of the danger of +loitering along the lake shore among those infected flies."</p> + +<p>As in the case of nagana, Bruce and most of the earlier investigators +supposed the transmission of the sleeping sickness trypanosome +by <i>Glossina palpalis</i> to be purely mechanical. The work of +Kleine (1909) clearly showed that for <i>Trypanosoma gambiense</i> as<span class="pagenum"><a name="Page_217" id="Page_217">[Pg 217]</a></span> +well as for <i>Trypanosoma brucei</i> the fly served as an essential host. +Indeed, Kleine and many subsequent investigators are inclined to +think that there is practically no mechanical transmission of trypanosomes +from animal to animal by <i>Glossina</i> in nature, and that the +many successful experiments of the earlier investigators were due +to the fact that they used wild flies which already harbored the +transformed parasite rather than directly inoculated it from the +blood of the diseased experimental animals. While the criticism +is applicable to some of the work, this extreme view is not fully +justified by the evidence at hand.</p> + +<p>Kleine states (1912) that <i>Glossina palpalis</i> can no longer be +regarded as the sole transmitter of sleeping sickness. Taute (1911) +had shown that under experimental conditions <i>Glossina morsitans</i> +was capable of transferring the disease and Kleine calls attention to +the fact that in German East Africa, in the district of the Rovuma +River, at least a dozen cases of the disease have occurred recently, +though only <i>Glossina morsitans</i> exists in the district. It appears, +however, that these cases are due to a different parasite, <i>Trypanosoma +rhodesiense</i>. This species, found especially in north-east +Rhodesia and in Nyassaland, is transferred by <i>Glossina morsitans</i>.</p> + +<p>Other workers maintain that the disease may be transmitted by +various blood-sucking flies, or even bugs and lice which attack man. +Fülleborn and Mayer (1907) have shown by conclusive experiments +that <i>Aedes (Stegomyia) calopus</i> may transmit it from one +animal to another if the two bites immediately succeed each other.</p> + +<p>It is not possible that insects other than the tsetse-flies (and only +certain species of these), play an important rôle in the transmission +of the disease, else it would be much more wide-spread. Sambon +(1908) pointed out that the hypothesis that is spread by <i>Aedes +calopus</i> is opposed by the fact that the disease never spread in the +Antilles, though frequently imported there by West African slaves. +The same observation would apply also to conditions in our own +Southern States in the early part of the past century.</p> + +<p>Since <i>Glossina palpalis</i> acts as an essential host of the parasite +and the chief, if not the only, transmitter, the fight against sleeping +sickness, like that against malaria and yellow fever, becomes primarily +a problem in economic entomology. The minutest detail +of the life-history, biology, and habits of the fly, and of its parasites +and other natural enemies becomes of importance in attempts to +eradicate the disease. Here we can consider only the general features +of the subject.<span class="pagenum"><a name="Page_218" id="Page_218">[Pg 218]</a></span></p> + +<p><i>Glossina palpalis</i> lives in limited areas, where the forest and undergrowth +is dense, along the lake shore or river banks. According to +Hodges, the natural range from shore is under thirty yards, though +the distance to which the flies may follow man greatly exceed this.</p> + +<p>It is a day feeder, a fact which may be taken advantage of in +avoiding exposure to its attacks. The young are brought forth alive +and full-grown, one every nine or ten days. Without feeding, they +enter the ground and under favorable conditions, complete their +development in a month or more.</p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_137" id="Fig_137"></a> +<a href="images/f137-full.png"><img src="images/f137.png" width="400" height="277" alt="137. Sleeping sickness concentration camp in German East Africa. Report of German +Commission." title="137. Sleeping sickness concentration camp in German East Africa. Report of German +Commission." /></a> +<span class="caption">137. Sleeping sickness concentration camp in German East Africa. Report of German +Commission.</span> +</div> + +<p>Methods of control of the disease must look to the prevention +of infection of the flies, and to their avoidance and destruction. +Along the first line, much was hoped from temporary segregation +of the sick in regions where the fly was not found. On the assumption +that the flies acted as carriers only during the first two or three +days, it was supposed that even the "fly belts" would become safe +within a few days after the sick were removed. The problem was +found to be a much more difficult one when it was learned that after +a given brief period the fly again became infective and remained so +for an indeterminate period. Nevertheless, isolation of the sick +is one of the most important measures in preventing the spread of<span class="pagenum"><a name="Page_219" id="Page_219">[Pg 219]</a></span> +the disease into new districts. Much, too, is being accomplished +by moving native villages from the fly belts. (c.f. <a href="#Fig_137">fig. 137</a>.)</p> + +<p>All measures to avoid the flies should be adopted. This means +locating and avoiding the fly belts as far as possible, careful screening +of houses, and protection of the body against bites.</p> + +<p>Clearing the jungle along the water courses for some yards beyond +the natural range of the fly has proved a very important measure. +Castellani recommends that the area be one hundred yards and +around a village three hundred yards at least.</p> + +<p>Detailed studies of the parasites and the natural enemies of the +tsetse-fly are being undertaken and may ultimately yield valuable +results.</p> + + +<p><a name="South_American_Trypanosomiasis" id="South_American_Trypanosomiasis"></a><b>South American Trypanosomiasis</b>—The tsetse-flies are distinctively +African in distribution and until recently there were no trypanosomes +known to infest man in America. In 1909 Dr. Chagas, of +Rio de Janeiro described a new species, <i>Trypanosoma cruzi</i>, pathogenic +to man.</p> + +<p><i>Trypanosoma cruzi</i> is the causative organism of a disease common +in some regions of Brazil, where it is known as "opilacao." It is +especially to be met with in children and is characterized by extreme +anemia, wasting, and stunted development associated with fever, +and enlargement of the thyroid glands. The disease is transmitted +by the bites of several species of assassin-bugs, or Reduviidæ, notably +by <i>Conorhinus megistus</i>. The evolution of the parasite within +the bug has been studied especially by Chagas and by Brumpt. +From the latter's text we take the following summary.</p> + +<p>The adult trypanosomes, ingested by a <i>Conorhinus megistus</i>, of +any stage, first change into Crithidia-like forms and then those +which remain in the stomach become ovoid and non-motile. Brumpt +found these forms in immense numbers, in a <span title="for Cornohinus read Conorhinus"><a name="AC_12" id="AC_12"></a><i>Cornohinus</i></span> which had +been infested fourteen months before. The forms which pass into +the intestine quickly assume the <i>Crithidia</i> form and continue to +develop rapidly under this form. Some weeks later they evolve +into the trypanosome forms, pathogenic for man. They then pass +out with the excrement of the bug and infect the vertebrate host +as soon as they come in contact with any mucous layer (buccal, +ocular or rectal). More rarely they enter through the epidermis.</p> + +<p>Brumpt showed that the development could take place in three +species; bed-bugs (<i>Cimex lectularius</i>, <i>C. hemipterus</i>) and in the tick<span class="pagenum"><a name="Page_220" id="Page_220">[Pg 220]</a></span> +<i>Ornithodoros moubata</i>. The evolution proceeds in the first two +species of bed-bugs as rapidly as in Conorhinus, or even more rapidly, +but they remain infective for a much shorter time and hence Brumpt +considers that they play a much less important rôle in the spread of +the disease.</p> + +<p><i>Conorhinus megistus</i>, like related forms in our Southern States, +very commonly frequents houses and attacks man with avidity. +Chagas states that the bites are painless and do not leave any traces. +They are usually inflicted on the lips, or the cheeks and thus the +buccal mucosa of a sleeper may be soiled by the dejections of the +insect and the bite serving as a port of entry of the virus, remain +unnoticed.</p> + +<p>The possibility of some of our North American Reduviidæ playing +a similar rôle in the transmission of disease should not be overlooked.</p> + + +<p><a name="Leishmanioses_and_Insects" id="Leishmanioses_and_Insects"></a><b>Leishmanioses and Insects</b>—Closely related to the trypanosomes +is a group of intracellular parasites which have recently been grouped +by Ross under the genus <i>Leishmania</i>. Five species are known to +affect man. Three of these produce local skin infestations, but two +of them, <i>Leishmania donovani</i> and <i>L. infantum</i>, produce serious and +often fatal systemic diseases.</p> + +<p>The first of these, that produced by <i>L. donovani</i>, is an exceedingly +virulent disease common in certain regions of India and China. It +is commonly known as "Kala-azar," or "dum-dum" fever, and more +technically as tropical leishmaniasis. Patton (1907) believes that +the parasite is transmitted by the bed-bug <i>Cimex hemipterus</i>, and has +described a developmental cycle similar to that which can be found +in artificial cultures. On the other hand, Donovan was unable to +confirm Patton's work and believes that the true intermediate host is +a Reduviid bug, <i>Conorhinus rubrofasciatus</i>.</p> + +<p><i>Leishmania infantum</i> is the cause of the so-called infantile splenic +leishmaniasis, occurring in northern Africa, Spain, Portugal, Italy, +and possibly other parts of Europe. The parasite occurs habitually +in the dog and is only accidentally transferred to children. Alvares +and da Silva, in Portugal (according to Brumpt, 1913) have found +that the excrement of a flea from a diseased dog contains flagellates, +and they suggest that the infection may be transmitted by the accidental +inoculation of this excrement by means of the proboscis of the +flea, as has been thought to occur in the case of the plague. To this<span class="pagenum"><a name="Page_221" id="Page_221">[Pg 221]</a></span> +Brumpt objects that they and other workers who thought to trace +the development of <i>Leishmania infantum</i> were apparently misled by +the presence of a harmless <i>Herpetomonas</i> which infests dog fleas in all +countries, even where the leishmaniasis is unknown.</p> + +<p>Basile (1910 and 1911) however, carried on numerous experiments +indicating that the disease was transferred from children to dogs +and from dog to dog by the dog flea, and was able to find in the +tissues of the insects forms perfectly identical with those found in +children and in dogs suffering from leishmaniasis. He also found +that <i>Pulex irritans</i> was capable of acting as the carrier.</p> + +<p>Of the cutaneous type of leishmaniasis, the best known is the so-called +"Oriental sore," an ulcerative disease of the skin which is +epidemic in many tropical and subtropical regions. The causative +organism is <i>Leishmania tropica</i>, which occurs in the diseased tissues +as bodies very similar to those found in the spleen in cases of +Kala-azar. The disease is readily inoculable and there is no doubt +that it may be transferred from the open sores to abraded surfaces of +a healthy individual by house-flies. It is also believed by a number +of investigators that it may be transferred and directly inoculated +by various blood-sucking insects.</p> + + +<h3><a name="Ticks_and_Diseases_of_Man_and_Animals" id="Ticks_and_Diseases_of_Man_and_Animals"></a><span class="smcap">Ticks and Diseases of Man and Animals</span></h3> + +<p>We have seen that the way to the discoveries of the relations of +arthropods to disease was pointed out by the work of Leuckart and +Melnikoff on the life cycle of <i>Dipylidium</i>, and of Fedtschenko and +Manson on that of <i>Filaria</i>. They dealt with grosser forms, belonging +to well-recognized parasitic groups.</p> + +<p>This was long before the rôle of any insect as a carrier of pathogenic +micro-organisms had been established, and before the Protozoa +were generally regarded as of importance in the causation of disease. +The next important step was taken in 1889 when Smith and Kilbourne +conclusively showed that the so-called Texas fever of cattle, +in the United States, is due to an intracorpuscular blood parasite +transmitted exclusively by a tick. This discovery, antedating by +eight years the work on the relation of the mosquito to malaria, had a +very great influence on subsequent studies along these lines.</p> + +<p>While much of the recent work has dealt with the true insects, +or hexapods, it is now known that several of the most serious diseases +of animals, and at least two important diseases of man are tick +borne. These belong to the types known collectively as <i>babesioses</i> +(or "<i>piroplasmoses</i>"), and <i>spirochætoses</i>.<span class="pagenum"><a name="Page_222" id="Page_222">[Pg 222]</a></span></p> + +<p>The term <i>babesiosis</i> is applied to a disease of man or animals +which is caused by minute protozoan parasites of the genus <i>Babesia</i>, +living in the red blood corpuscles. These parasites have usually been +given the generic name <i>Piroplasma</i> and hence the type of disease +which they cause is often referred to as "<i>piroplasmosis</i>." The best +known illustration is the disease known in this country as Texas +fever of cattle.</p> + + +<p><a name="Cattle_Ticks_and_Texas_Fever" id="Cattle_Ticks_and_Texas_Fever"></a><b>Cattle Ticks and Texas Fever</b>—The cattle disease, which in the +United States is known as Texas fever, is a widely distributed, exceedingly +acute disease. In Australia it is known as <i>redwater fever</i> and +in Europe as hæmoglobinuria, due to the fact that the urine of the +diseased animals is discolored by the breaking down of the red blood +corpuscles infested by the parasite.</p> + +<p>In their historical discussion, Smith and Kilbourne, point out that +as far back as 1796 it was noted that Southern cattle, in a state of +apparent health, might spread a fatal disease among Northern herds. +As observations accumulated, it was learned that this infection was +carried only during the warm season of the year and in the depth of +winter Southern cattle were harmless. Moreover, Southern cattle +after remaining for a short time in the North lost their power to +transmit the disease, and the same was true of cattle which had been +driven for a considerable distance.</p> + +<p>Very significant was the fact that the infection was not communicated +directly from the Southern to Northern cattle but that +the ground over which the former passed was infected by them, and +that the infection was transmitted thence to susceptible cattle <i>after +a period of not less than thirty days had elapsed</i>.</p> + +<p>Of course a disease as striking as this, and which caused such +enormous losses of cattle in the region invaded was fruitful in theories +concerning its causation. The most widespread was the belief that +pastures were infected by the saliva, urine, or manure of Southern +cattle. There were not wanting keen observers who suggested that +the disease was caused by ticks, but little weight was given to their +view.</p> + +<p>Various workers had described bacteria which they had isolated +from the organs of the diseased animals, but their findings could not +be verified. In 1889, Smith and Kilbourne discovered a minute, +pear-shaped organism (<a href="#Fig_138">fig. 138</a>) in the red blood corpuscles of a cow +which had succumbed to Texas fever. On account of their shape<span class="pagenum"><a name="Page_223" id="Page_223">[Pg 223]</a></span> +they were given the generic name <i>Pyrososma</i> and because they were +usually found two in a corpuscle, the specific name, <i>bigeminum</i>. It +is now generally accepted that +the parasite is the same which +Babes had observed the year +before in Roumanian cattle +suffering from hæmoglobinuria, +and should be known as <i>Babesia +bovis</i> (Babes).</p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_138" id="Fig_138"></a> +<a href="images/f138-full.png"><img src="images/f138.png" width="400" height="241" alt="138. Babesia bovis in blood corpuscles. +After Calli." title="138. Babesia bovis in blood corpuscles. +After Calli." /></a> +<span class="caption">138. Babesia bovis in blood corpuscles. +After Calli.</span> +</div> + +<div class="figcenter" style="width: 245px;"><a name="Fig_139" id="Fig_139"></a> +<a href="images/f139-full.png"><img src="images/f139.png" width="245" height="522" alt="139. The cattle tick (Boophilus annulatus). +(a) Female; (b) male. After +Comstock." title="139. The cattle tick (Boophilus annulatus). +(a) Female; (b) male. After +Comstock." /></a> +<span class="caption">139. The cattle tick (Boophilus annulatus). +(<i>a</i>) Female; (<i>b</i>) male. After +Comstock.</span> +</div> + +<p>By a series of perfectly conclusive +experiments carried on +near Washington, D.C., Smith +and Kilbourne showed that +this organism was carried from Southern cattle to non-immune animals +by the so-called Southern cattle +tick, <i>Boophilus annulatus</i> (= <i>Margaropus +annulatus</i>) (<a href="#Fig_139">fig. 139</a>).</p> + +<p>Of fourteen head of native cattle +placed in a field with tick-infested +Northern cattle all but two contracted +the disease. This experiment was +repeated with similar results. Four +head of native cattle kept in a plot +with three North Carolina cattle +which had been carefully freed from +ticks remained healthy. A second +experiment the same year gave similar +results.</p> + +<p>Still more conclusive was the experiment +showing that fields which +had not been entered by Southern +cattle but which had been infected by +mature ticks taken from such animals +would produce Texas fever in native +cattle. On September 13, 1889, several +thousand ticks collected from +cattle in North Carolina three and +four days before, were scattered in a +small field near Washington. Three +out of four native animals placed in<span class="pagenum"><a name="Page_224" id="Page_224">[Pg 224]</a></span> +this field contracted the disease. The fourth animal was not +examined as to its blood but it showed no external symptoms of +the disease.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_140" id="Fig_140"></a> +<a href="images/f140-full.png"><img src="images/f140.png" width="500" height="383" alt="140. Hyalomma ægypticum. After Nuttall and Warburton." title="140. Hyalomma ægypticum. After Nuttall and Warburton." /></a> +<span class="caption">140. Hyalomma ægypticum. After Nuttall and Warburton.</span> +</div> + +<p>In these earlier experiments it was believed that the cattle tick +acted as a carrier of the disease between the Southern cattle and the +<i>soil</i> of the Northern pastures. "It was believed that the tick obtained +the parasite from the blood of its host and in its dissolution +on the pasture a certain resistant spore form was set free which +produced the disease when taken in with the food." The feeding of +one animal for some time with grass from the most abundantly +infected field, without any appearance of the disease, made this +hypothesis untenable.</p> + +<p>In the experimental work in 1890 the astonishing fact was brought +out that the disease was conveyed neither by infected ticks disintegrating +nor by their directly transferring the parasite, but that +it was conveyed by the young hatched from eggs of infected ticks. +In other words, the disease was hereditarily transferred to ticks of +the second generation and they alone were capable of conveying it.</p> + +<p>Thus was explained the fact that Texas fever did not appear +immediately along the route of Southern cattle being driven to +Northern markets but that after a certain definite period it manifested +itself. It was conveyed by the progeny of ticks which had +dropped from the Southern cattle and deposited their eggs on the +ground.<span class="pagenum"><a name="Page_225" id="Page_225">[Pg 225]</a></span></p> + +<p>These results have been fully confirmed by workers in different +parts of the world,—notably by Koch, in Africa, and by Pound, in +Australia.</p> + +<p>The disease is apparently transmitted by <i>Boophilus annulatus</i> +alone, in the United States, but it, or an almost identical disease, +is conveyed by <i>Ixodes hexagonus</i> in Norway, <i>Ixodes ricinus</i> in Finland +and France and by the three species, <i>Boophilus decoloratus</i>, <i>Hyalomma +ægypticum</i> (fig. <a href="#Fig_140">140</a> and <a href="#Fig_141">141</a>), and <i>Hæmaphysalis punctata</i> in Africa.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_141" id="Fig_141"></a> +<a href="images/f141-full.png"><img src="images/f141.png" width="500" height="413" alt="141. Hyalomma aegypticum. Capitulum of female; +(a) dorsal, (b) ventral aspect." title="141. Hyalomma aegypticum. Capitulum of female; +(a) dorsal, (b) ventral aspect." /></a> +<span class="caption">141. Hyalomma aegypticum. Capitulum of female; +(<i>a</i>) dorsal, (<i>b</i>) ventral aspect.</span> +</div> + +<p>In spite of the detailed study which it has received, the life cycle +of <i>Babesia bovis</i> has not been satisfactorily worked out. The asexual +reproduction in the +blood of the vertebrate +host has been described +but the cycle in the tick +is practically unknown.</p> + +<p>More successful +attempts have been +made to work out the life +cycle of a related species, +<i>Babesia canis</i>, which +causes malignant jaundice +in dogs in Africa +and parts of Southern +Europe. In this instance, +also, the disease +is transmitted by heredity to the ticks of the second generation. +Yet the larval, or "seed ticks," from an infected female are not +capable of conveying the disease, but only the nymphs and adults. +Still more complicated is the condition in the case of <i>Babesia ovis</i> of +sheep, which Motas has shown can be conveyed solely by the adult, +sexually mature ticks of the second generation.</p> + +<p>In <i>Babesia canis</i>, Christopher (1907) observed developmental +stages in the tick. He found in the stomach of adult ticks, large +motile club-shaped bodies which he considered as oökinetes. These +bodies pass to the ovaries of the tick and enter the eggs where they +become globular in form and probably represent an oocyst. This +breaks up into a number of sporoblasts which enter the tissues of +the developing tick and give rise to numerous sporozoites, which +collect in the salivary glands and thence are transferred to the +vertebrate host. A number of other species of <i>Babesia</i> are known<span class="pagenum"><a name="Page_226" id="Page_226">[Pg 226]</a></span> +to infest vertebrates and in all the cases where the method has been +worked out it has been found that the conveyal was by ticks. We +shall not consider the cases more fully here, as we are concerned +especially with the method of transfer of human diseases.</p> + + +<p><a name="Ticks_and_Rocky_Mountain_Spotted_Fever_of_Man" id="Ticks_and_Rocky_Mountain_Spotted_Fever_of_Man"></a><b>Ticks and Rocky Mountain Spotted Fever of Man</b>—Ever since +1873 there has been known in Montana and Idaho a peculiar febrile +disease of man, which has gained the name of "Rocky Mountain +spotted fever." Its onset is marked by chills and fever which rapidly +become acute. In about four to seven days there appears a characteristic +eruption on the wrists, ankles or back, which quickly covers +the body.</p> + +<p>McClintic (1912) states that the disease has now been reported +from practically all of the Rocky Mountain States, including Arizona, +California, Colorado, Idaho, Montana, Nevada, Oregon, Utah, +Washington, and Wyoming. "Although the disease is far more +prevalent in Montana and Idaho than in any of the other States, +its spread has assumed such proportions in the last decade as to call +for the gravest consideration on the part of both the state and national +health authorities. In fact, the disease has so spread from state +to state that it has undoubtedly become a very serious interstate +problem demanding the institution of measures for its control and +suppression."</p> + +<p>A peculiar feature of the Rocky Mountain spotted fever is a +marked variation in its severity in different localities. In Montana, +and especially in the famous Bitter Root Valley, from 33 per cent to +75 per cent of the cases result fatally. On the other hand, the fatality +does not exceed four per cent in Idaho.</p> + +<p>In 1902, Wilson and Chowning reported the causative organism +of spotted fever to be a blood parasite akin to the <i>Babesia</i> of Texas +fever, and made the suggestion that the disease was tick-borne. +The careful studies of Stiles (1905) failed to confirm the supposed +discovery of the organism, and the disease is now generally classed +as due to an invisible virus. On the other hand, the accumulated +evidence has fully substantiated the hypothesis that it is tick-borne.</p> + +<p>According to Ricketts (1907) the experimental evidence in support +of this hypothesis was first afforded by Dr. L. P. McCalla and +Dr. H. A. Brereton, in 1905. These investigators transmitted the +disease from man to man in two experiments. "The tick was +obtained 'from the chest of a man very ill with spotted fever' and<span class="pagenum"><a name="Page_227" id="Page_227">[Pg 227]</a></span> +'applied to the arm of a man who had been in the hospital for two +months and a half, and had lost both feet from gangrene due to +freezing.' On the eighth day the patient became very ill and passed +through a mild course of spotted fever, leaving a characteristic +eruption. The experiment was repeated by placing the tick on a +woman's leg and she likewise was infected with spotted fever."</p> + +<p>The most detailed studies were those of the late Dr. H. T. Ricketts, +and it was he who clearly established the tick hypothesis. In the +summer of 1906 he found that guinea pigs and monkeys are very +susceptible to spotted fever and can readily be infected by inoculation +of blood from patients suffering from the disease. This opened +the way to experimental work on tick transmission. A female tick +was fed upon an infected guinea pig for two days, removed and +isolated for two days and then placed upon a healthy guinea pig. +After an incubation period of three and a half days the experimental +animal contracted a well-marked case of the disease.</p> + +<p>A similar result was obtained at the same time by King, and later +in the season Ricketts proved that the male tick was also capable +of transmitting the disease. He found that there was a very intimate +relation of the virus to the tick and that the transmission must +be regarded as biological throughout. Ticks remained infective as +long as they lived and would feed for a period of several months. If +they acquired the disease in the larval or nymphal stage they retained +it during molting and were infective in the subsequent stages. In a +few cases the larvæ from an infected female were infective.</p> + +<p>The evidence indicated that the tick suffers from a relatively +harmless, generalized infection and the virus proliferates in its +body. The disease probably is transferred through the salivary +secretion of the tick since inoculation experiments show that the +salivary glands of the infected adult contain the virus.</p> + +<p>It is probable that in nature the reservoir of the virus of spotted +fever is some one or more of the native small animals. Infected +ticks have been found in nature, and as various wild animals are +susceptible to the disease, it is obvious that it may exist among them +unnoticed. Wilson and Chowning suggested that the ground squirrel +plays the principal rôle.</p> + +<p>Unfortunately, much confusion exists regarding the correct +name of the tick which normally conveys the disease. In the medical +literature it is usually referred to as <i>Dermacentor occidentalis</i>, +but students of the group now agree that it is specifically distinct.<span class="pagenum"><a name="Page_228" id="Page_228">[Pg 228]</a></span> +Banks has designated it as <i>Dermacentor venustus</i> and this name is +used in the publications of the Bureau of Entomology. On the other +hand, Stiles maintains that the common tick of the Bitter Root +Valley, and the form which has been collected by the authors who +have worked on Rocky Mountain spotted fever in that region, is +separable from <i>D. venustus</i>, and he has described it under the name of +<i>Dermacentor andersoni</i>.</p> + +<p>Mayer (1911) has shown experimentally that spotted fever may +be transmitted by several different species of ticks, notably <i>Dermacentor +marginatus</i>, <i>Dermacentor variabilis</i> and <i>Amblyomma americanum</i>. +This being the case, the question of the exact systematic status of +the species experimented upon in the Bitter Root Valley becomes +less important, for since <i>Dermacentor occidentalis</i>, <i>Dermacentor +venustus</i> and <i>Dermacentor andersoni</i> all readily attack man, it is +probable that either species would readily disseminate the disease +if it should spread into their range.</p> + +<p>Hunter and Bishop (1911) have emphasized the fact that in the +eastern and southern United States there occur several species which +attack man, and any one of which might transmit the disease from +animal to animal and from animal to man. The following species, +they state, would probably be of principal importance in the Southern +and Eastern States: the lone star tick (<i>Amblyomma americanum</i>); +the American dog tick (<i>Dermacentor variabilis</i>); and the gulf-coast +tick (<i>Amblyomma maculatum</i>). In the extreme southern portions of +Texas, <i>Amblyomma cajennense</i>, is a common pest of man.</p> + +<p>Since the evidence all indicates that Rocky Mountain spotted +fever is transmitted solely by the tick, and that some of the wild +animals serve as reservoirs of the virus, it is obvious that personal +prophylaxis consists in avoiding the ticks as fully as possible, and in +quickly removing those which do attack. General measures along +the line of tick eradication must be carried out if the disease is to be +controlled. That such measures are feasible has been shown by the +work which has been done in controlling the tick-borne Texas fever +of cattle, and by such work as has already been done against the +spotted fever tick, which occurs on both wild and domestic animals. +Detailed consideration of these measures is to be found in the +publications of the Public Health and Marine Hospital Service, +and the Bureau of Entomology. Hunter and Bishopp give the +following summarized recommendations for control or eradication +measures in the Bitter Root Valley.<span class="pagenum"><a name="Page_229" id="Page_229">[Pg 229]</a></span></p> + +<p>(1) A campaign of education, whereby all the residents of the +valley will be made thoroughly familiar with the feasibility of the +plan of eradication, and with what it will mean in the development of +the valley.</p> + +<p>(2) The obtaining of legislation to make it possible to dip or oil +all live stock in the Bitter Root Valley.</p> + +<p>(3) The obtaining of an accurate census of the horses, cattle, +sheep, mules, and dogs in the valley.</p> + +<p>(4) The construction of ten or more dipping vats.</p> + +<p>(5) The providing of materials to be used in the dipping mixture.</p> + +<p>(6) The organization of a corps of workers to carry on the operations.</p> + +<p>(7) The systematic dipping of the horses, cattle, sheep, and dogs +of the valley on a definite weekly schedule from approximately March +10 to June 9.</p> + +<p>(8) The treatment by hand of the animals in localities remote +from vats, on the same schedule.</p> + +<p>They estimate that after three seasons' operations a very small +annual expenditure would provide against reinfestation of the valley +by the incoming of cattle from other places.</p> + +<p>Supplementary measures consist in the killing of wild mammals +which may harbor the tick; systematic burning of the brush and +debris on the mountain side; and in clearing, since the tick is seldom +found on land under cultivation.<span class="pagenum"><a name="Page_230" id="Page_230">[Pg 230]</a></span></p> + + + + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_X" id="CHAPTER_X"></a>CHAPTER X</h2> + +<h2>ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA +[<i>Continued</i>]</h2> + + +<h3><a name="Arthropods_and_Spirochaetoses_of_Man_and_Animals" id="Arthropods_and_Spirochaetoses_of_Man_and_Animals"></a><span class="smcap">Arthropods and Spirochætoses of Man and Animals</span></h3> + +<p>The term spirochætoses is applied to diseases of man or animals +which are due to protistan parasites belonging to the group of slender, +spiral organisms known as spirochætes.</p> + +<p>There has been much discussion concerning the relationship Of +the spirochætes. Formerly, they were regarded as bacteria closely +related to the forms grouped in the genus <i>Spirillum</i>. The results +of the detailed study which the spirochætes have received in +recent years, have led most of the workers to consider them as belonging +to the protozoa. The merits of the discussion we are not concerned +with here, but rather with the fact that a number of diseases +caused by spirochætes are arthropod-borne. The better known of +these we shall discuss.</p> + + +<p><a name="African_Relapsing_Fever_of_Man" id="African_Relapsing_Fever_of_Man"></a><b>African Relapsing Fever of Man</b>—It has long been known to the +natives of Africa and to travelers in that country, that the bite of a +certain tick, <i>Ornithodoros moubata</i>, may be followed by severe or +even fatal fever of the relapsing type. Until recent years, it was +supposed that the effect was due to some special virulence of the tick, +just as nagana of cattle was attributed to the direct effect of the bite +of the tsetse-fly. The disease is commonly known as "tick-fever" +or by the various native names of the tick.</p> + +<p>In 1904, Ross and Milne, in Uganda, and Dutton and Todd on the +Congo, discovered that the cause of the disease is a spirochæte which +is transmitted by the tick. This organism has been designated by +Novy and Knapp as <i>Spirochæta duttoni</i>.</p> + +<p><i>Ornithodoros moubata</i> (<a href="#Fig_142">fig. 142</a>), the carrier of African relapsing +fever, or "tick-fever," is widely distributed in tropical Africa, and +occurs in great numbers in the huts of natives, in the dust, cracks +and crevices of the dirt floors, or the walls. It feeds voraciously +on man as well as upon birds and mammals. Like others of the +<i>Argasidæ</i>, it resembles the bed-bug in its habit of feeding primarily +at night. Dutton and Todd observed that the larval stage is undergone +in the egg and that the first free stage is that of the octopod +nymph.<span class="pagenum"><a name="Page_231" id="Page_231">[Pg 231]</a></span></p> + +<div class="figcenter" style="width: 375px;"><a name="Fig_142" id="Fig_142"></a> +<a href="images/f142-full.png"><img src="images/f142.png" width="375" height="661" alt="142. Ornithodoros moubata. (a) Anterior part of venter; (b) second stage +nymph; (c) capitulum; (d) dorsal and (e) ventral aspect of female; +(f) ventral aspect of nymph; (g) capitulum of nymph. After Nuttall +and Warburton." title="142. Ornithodoros moubata. (a) Anterior part of venter; (b) second stage +nymph; (c) capitulum; (d) dorsal and (e) ventral aspect of female; +(f) ventral aspect of nymph; (g) capitulum of nymph. After Nuttall +and Warburton." /></a> +<span class="caption">142. Ornithodoros moubata. (<i>a</i>) Anterior part of venter; (<i>b</i>) second stage +nymph; (<i>c</i>) capitulum; (<i>d</i>) dorsal and (<i>e</i>) ventral aspect of female; +(<i>f</i>) ventral aspect of nymph; (<i>g</i>) capitulum of nymph. After Nuttall +and Warburton.</span> +</div><p><span class="pagenum"><a name="Page_232" id="Page_232">[Pg 232]</a></span></p> + +<p>The evidence that the fever is transmitted by this tick is conclusive. +Koch found that from five per cent to fifteen per cent, and +in some places, fifty per cent of the ticks captured, harbored the +spirochæte. The disease is readily transmitted to monkeys, rats, +mice and other animals and the earlier experiments along these lines +have been confirmed by many workers.</p> + +<p>Not only are the ticks which have fed on infected individuals +capable of conveying the disease to healthy animals but they transmit +the causative organism to their progeny. Thus Möllers (1907), +working in Berlin, repeatedly infected monkeys through the bites +of nymphs which had been bred in the laboratory from infected ticks. +Still more astonishing was his discovery that ticks of the third generation +were infective. In other words, if the progeny of infected ticks +were fed throughout life on healthy animals, and on maturity deposited +eggs, the nymphs which hatched from these eggs would still +be capable of carrying the infection.</p> + +<p>The developmental cycle of the spirochæte within the tick has not +been fully worked out, though the general conclusions of Leishman +(1910) have been supported by the recent works of Balfour (1911 +and 1912), and Hindle (1912), on the life cycle of spirochætes affecting +fowls.</p> + +<p><i>Spirochæta duttoni</i> ingested by <i>Ornithodoros moubata</i> apparently +disappear within a few days, but Leishman believed that in reality +they break up into minute granules which are to be found in the +alimentary canal, the salivary glands and the Malpighian tubes of +the tick. These granules, or "coccoid bodies," as Hindle calls them, +are supposed to be the form in which the spirochætes infect the new +host. We shall see later that Marchoux and Couvy (1913) dissent +wholly from this interpretation.</p> + +<p>According to Leishman, and Hindle, the coccoid bodies are not +injected into the vertebrate host with the saliva of the tick, as are +the sporozoites of malaria with that of the mosquito. Instead, they +pass out with the excrement and secondarily gain access to the +wound inflicted by the tick.</p> + +<p>Nuttall (1912) calls attention to the fact that the geographical +distribution of <i>Ornithodoros moubata</i> is far wider than our present +records show for the distribution of the relapsing fever in man and +that there is every reason to fear the extension of the disease. Huts +where the ticks occur should be avoided and it should be remembered +that in infected localities there is special danger in sleeping on the +ground.<span class="pagenum"><a name="Page_233" id="Page_233">[Pg 233]</a></span></p> + + +<p><a name="European_Relapsing_Fever" id="European_Relapsing_Fever"></a><b>European Relapsing Fever</b>—There is widely distributed in Europe +a type of relapsing fever which is caused by <i>Spirochæta recurrentis</i>. +It has long been supposed that this disease is spread by the bed-bug +and there is some experimental evidence to show that it may be +conveyed by these insects.</p> + +<p>In 1897, Tictin found that he could infect monkeys by inoculating +the contents of bed-bugs which had fed upon a patient within forty-eight +hours. Nuttall, in 1907, in one experiment succeeded in transmitting +<i>Spirochæta recurrentis</i> from mouse to mouse by bites of bed-bugs. +The bugs, thirty-five in number, were transferred at short +intervals from one mouse to another, not being allowed to take a +full meal on the first, or infected mouse.</p> + +<p>On the other hand, there is much clinical evidence to show that +the European relapsing fever like various other types of the disease +is transmitted from man to man by head and body lice (<i>Pediculus +humanus</i> and <i>Pediculus corporis</i>).</p> + +<p>Interesting supplementary evidence is that of Bayon's observations +(1912), in Moscow. "Having visited the big municipal night hospitals +at Moscow I soon noticed that they were kept with such scrupulous +cleanliness, disinfected so lavishly, the beds of iron, the floor cemented, +that it was not possible for bed-bugs to thrive to any extent on +the premises. The people sleeping there were allowed, however, +to sleep in their own clothes. The introduction of these model homes +had not had any effect on the incidence of relapsing fever, for the +places were still hot-beds of the fever during winter. On the other +hand, though I changed my rooms several times, I found bugs in +every successive lodging, and I was told in Moscow, this can hardly +be avoided. Yet no foreigner, or Russian of the better class, ever +catches relapsing fever. To this may be added the fact that when +I asked for clothes-lice and promised to pay a kopec for two, the +attendants from the night hostel brought me next morning a small +ounce bottle crammed with <i>Pediculus capitis</i> (= <i>P. humanus</i>), and +<i>Pediculus vestimentorum</i> (= <i>P. corporis</i>) collected off the sleepers. +If relapsing fever were transmitted by bed-bugs, it would be much +more disseminated than it is at present in Moscow."</p> + +<p>Direct experimental evidence of the agency of lice in transmitting +relapsing fever is especially clear in the case of a type of the disease +prevalent in parts of North Africa. We shall consider this evidence +later.<span class="pagenum"><a name="Page_234" id="Page_234">[Pg 234]</a></span></p> + + +<p><a name="Other_Types_of_Relapsing_Fever_of_Man" id="Other_Types_of_Relapsing_Fever_of_Man"></a><b>Other Types of Relapsing Fever of Man</b>—In addition to the three +types of human relapsing fever already referred to, several others +have been distinguished and have been attributed to distinct species +of spirochætes. The various spirochætoses of man are:</p> + +<p>African, caused by <i>S. duttoni</i>; European, caused by <i>S. recurrentis</i>; +North African, caused by <i>S. berbera</i>; East African, caused +by <i>S. rossi</i>; East Indian, caused by <i>S. carteri</i>; North American, +caused by <i>S. novyi</i>; South American, caused by <i>S. duttoni</i> (?).</p> + +<p>Nuttall (1912) in his valuable résumé of the subject, has emphasized +that "in view of the morphological similarity of the supposedly +different species of spirochætes and their individual variations +in virulence, we may well doubt if any of the 'species' are valid. +As I pointed out four years ago, the various specific names given to +the spirochætes causing relapsing fever in man may be used merely +for convenience <i>to distinguish strains or races</i> of different origin. +They cannot be regarded as valid names, in the sense of scientific +nomenclature, for virulence and immunity reactions are not adequate +tests of specificity."</p> + + +<p><a name="North_African_Relapsing_Fever_of_Man" id="North_African_Relapsing_Fever_of_Man"></a><b>North African Relapsing Fever of Man</b>—The type of human +relapsing fever to be met with in Algeria, Tunis, and Tripoli, is due to +a <i>Spirochæta</i> to which does not differ morphologically from <i>Spirochæta +duttoni</i>, but which has been separated on biological grounds as +<i>Spirochæta berberi</i>.</p> + +<p>Experimenting with this type of disease in Algeria, Sergent and +Foly (1910), twice succeeded in transmitting it from man to monkeys +by inoculation of crushed body lice and in two cases obtained infection +of human subjects who had received infected lice under their +clothing and who slept under coverings harboring many of the lice +which had fed upon a patient. Their results were negative with +<i>Argas persicus</i>, <i>Cimex lectularius</i>, <i>Musca domestica</i>, <i>Hæmatopinus +spinulosus</i> and <i>Ceratophyllus fasciatus</i>. They found body lice +associated with every case of relapsing fever which they found in +Algeria.</p> + +<p>Nicolle, Blaizot, and Conseil (1912) showed that the louse did +not transmit the parasite by its bite. Two or three hours after it +has fed on a patient, the spirochætes begin to break up and finally +they disappear, so that after a day, repeated examinations fail to +reveal them. They persist, nevertheless, in some unknown form, +for if the observations are continued they reappear in eight to twelve<span class="pagenum"><a name="Page_235" id="Page_235">[Pg 235]</a></span> +days. These new forms are virulent, for a monkey was infected +by inoculating a single crushed louse which had fed on infected blood +fifteen days before.</p> + +<p>Natural infection is indirect. Those attacked by the insect +scratch, and in this act they excoriate the skin, crush the lice and +contaminate their fingers. The least abrasion of the skin serves for +the entrance of the spirochætes. Even the contact of the soiled +fingers on the various mucosa, such as the conjunctive of the eye, +is sufficient.</p> + +<p>As in the case of <i>Spirochæta duttoni</i>, the organism is transmitted +hereditarily in the arthropod vector. The progeny of lice which +have fed on infected blood may themselves be infective.</p> + + +<p><a name="Spirochaetosis_of_Fowls" id="Spirochaetosis_of_Fowls"></a><b>Spirochætosis of Fowls</b>—One of the best known of the spirochætes +transmitted by arthropods is <i>Spirochæta gallinarum</i>, the cause of a +very fatal disease of domestic fowls in widely separated regions of +the world. According to Nuttall, it occurs in Southeastern Europe, +Asia, Africa, South America and Australia.</p> + +<p>In 1903, Marchoux and Salimbeni, working in Brazil, made the +first detailed study of the disease, and showed that the causative +organism is transmitted from fowl to fowl by the tick <i>Argas persicus</i>. +They found that the ticks remained infective for at least five months. +Specimens which had fed upon diseased birds in Brazil were sent to +Nuttall and he promptly confirmed the experiments. Since that +date many investigators, notably Balfour and Hindle, have contributed +to the elucidating of the life-cycle of the parasite. Since it +has been worked out more fully than has that of any of the human +spirochætes, we present Hindle's diagram (<a href="#Fig_143">fig. 143</a>) and quote the +brief summary from his preliminary paper (1911<i>b</i>).</p> + +<p>"Commencing with the ordinary parasite in the blood of the fowl, +the spirochæte grows until it reaches a certain length (16-19µ) and +then divides by transverse division. This process is repeated, and +is probably the only method of multiplication of the parasite within +the blood. When the spirochætes disappear from the circulation, +some of them break up into the coccoid bodies which, however, +do not usually develop in the fowl. When the spirochætes are +ingested by <i>Argas persicus</i>, some of them pass through the gut wall +into the cœlomic fluid. From this medium they bore their way into +the cells of the various organs of the tick and there break up into a +number of coccoid bodies. These intracellular forms multiply by<span class="pagenum"><a name="Page_236" id="Page_236">[Pg 236]</a></span> +ordinary fission in the cells of the Malpighian tubules and gonads. +Some of the coccoid bodies are formed in the lumen of the gut and +Malpighian tubules. The result is that some of the coccoid bodies +may be present in the Malpighian secretion and excrement of an +infected tick and when mixed with the coxal fluid may gain entry +into another fowl by the open wound caused by the tick's bite. They +then elongate and redevelop into ordinary spirochætes in the blood +of the fowl, and the cycle may be repeated."</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_143" id="Fig_143"></a> +<a href="images/f143-full.png"><img src="images/f143.png" width="500" height="483" alt="143. Spirochæta gallinarum. After Hindle." title="143. Spirochæta gallinarum. After Hindle." /></a> +<span class="caption">143. Spirochæta gallinarum. After Hindle.</span> +</div> + +<p>Hindle's account is clear cut and circumstantial, and is quite in +line with the work of Balfour, and of Leishman. Radically different +is the interpretation of Marchoux and Couvy (1913). These investigators +maintain that the granules localized in the Malpighian tubules +in the larvæ and, in the adult, also in the ovules and the genital ducts +of the male and female, are not derived from spirochætes but that they +exist normally in many acariens. They interpret the supposed<span class="pagenum"><a name="Page_237" id="Page_237">[Pg 237]</a></span> +disassociation of the spirochæte into granules as simply the first +phase, not of a process of multiplication, but of a degeneration +ending in the death of the parasite. The fragmented chromatin +has lost its affinity for stains, remaining always paler than that of +the normal spirochætes. On the other hand, the granules of Leishman +stain energetically with all the basic stains.</p> + +<p>Further, according to Marchoux and Couvy, infection takes +place without the emission of the coxal fluid and indeed, soiling of the +host by the coxal fluid diluting the excrement is exceptional. All +of the organs of the Argasid are invaded by the parasites, but they +pass from the cœlom into the acini of the salivary glands and collect +in its efferent canal. The saliva serves as the vehicle of infection.</p> + +<p>Thus, the question of the life cycle of <i>Spirochæta gallinarum</i>, and +of spirochætes in general, is an open one.</p> + +<p>It should be noted that <i>Argas persicus</i>, the carrier of <i>Spirochæta +gallinarum</i>, is a common pest of poultry in the southwestern United +States. Though the disease has not been reported from this country, +conditions are such that if accidentally introduced, it might do great +damage.</p> + + +<p><a name="Other_Spirochaete_Diseases_of_Animals" id="Other_Spirochaete_Diseases_of_Animals"></a><b>Other Spirochæte Diseases of Animals</b>—About a score of other +blood inhabiting spirochætes have been reported as occurring in +mammals, but little is known concerning their life-histories. One +of the most important is <i>Spirochæta theileri</i> which produces a spirochætosis +of cattle in the Transvaal. Theiler has determined that it +is transmitted by an Ixodid tick, <i>Margaropus decoloratus</i>.</p> + + +<h3><a name="Typhus_Fever_and_Pediculidae" id="Typhus_Fever_and_Pediculidae"></a><span class="smcap">Typhus Fever and Pediculidæ</span></h3> + +<p>Typhus is an acute, and continued fever, formerly epidemically +prevalent in camps, hospitals, jails, and similar places where persons +were crowded together under insanitary conditions. It is accompanied +by a characteristic rash, which gives the disease the common +name of "spotted" or "lenticular" fever. The causative organism +is unknown.</p> + +<p>Typhus fever has not generally been supposed to occur in the +United States, but there have been a few outbreaks and sporadic +cases recognized. According to Anderson and Goldberger (1912<i>a</i>), +it has been a subject of speculation among health authorities why, +in spite of the arrival of occasional cases in this country and of many +persons from endemic foci of the disease, typhus fever apparently +does not gain a foothold in the United States. These same workers<span class="pagenum"><a name="Page_238" id="Page_238">[Pg 238]</a></span> +showed that the so-called Brill's disease, studied especially in New +York City, is identical with the typhus fever of Mexico and of +Europe.</p> + +<p>The conditions under which the disease occurs and under which +it spreads most rapidly are such as to suggest that it is carried by +some parasitic insect. On epidemiological grounds the insects most +open to suspicion are the lice, bed-bugs and fleas.</p> + +<p>In 1909, Nicolle, Comte and Conseil, succeeded in transmitting +typhus fever from infected to healthy monkeys by means of the +body louse (<i>Pediculus corporis</i>). Independently of this work, +Anderson and Goldberger had undertaken work along this line in +Mexico, and in 1910 reported two attempts to transmit the disease +to monkeys by means of body lice. The first experiment resulted +negatively, but the second resulted in a slight rise in temperature, +and in view of later results it seems that this was due to infection +with typhus.</p> + +<p>Shortly after, Ricketts and Wilder (1910) succeeded in transmitting +the disease to the monkey by the bite of body lice in two experiments, +the lice in one instance deriving their infection from a man +and in another from the monkey. Another monkey was infected +by typhus through the introduction of the feces and abdominal +contents of infested lice into small incisions. Experiments with +fleas and bed-bugs resulted negatively.</p> + +<p>Subsequently, Goldberger and Anderson (1912<i>b</i>) indicated that +the head louse (<i>Pediculus humanus</i>) as well, may become infected +with typhus. In an attempt to transmit typhus fever (Mexican +virus) from man to monkey by subcutaneous injection of a saline +suspension of crushed head lice, the monkeys developed a typical +febrile reaction with subsequent resistance to an inoculation of +virulent typhus (Mexican) blood. In one of the three experiments +to transmit the disease from man to monkey by means of the bite +of the head louse, the animal bitten by the presumably infected head +lice proved resistant to two successive immunity tests with virulent +typhus blood.</p> + +<p>In 1910, Ricketts and Wilder reported an experiment undertaken +with a view to determining whether the young of infected lice were +themselves infected. Young lice were reared to maturity on the +bodies of typhus patients, so that if the eggs were susceptible to +infection at any stage of their development, they would have every +opportunity of being infected within the ovary. The eggs of these +infected lice were obtained, they were incubated, and the young lice<span class="pagenum"><a name="Page_239" id="Page_239">[Pg 239]</a></span> +of the second generation were placed on a normal rhesus monkey. +The experimenters were unable to keep the monkey under very +close observation during the following three or four weeks, but from +the fact that he proved resistant to a subsequent immunity test +they concluded that he probably owed this immunity to infection +by these lice of the second generation.</p> + +<p>Anderson and Goldberger (1912<i>b</i>) object that due consideration +was not given to the possibility of a variable susceptibility of the +monkey to typhus. Their similar experiment was "frankly negative."</p> + +<p>Prophylaxis against typhus fever is, therefore, primarily a question +of vermin extermination. A brief article by Dr. Goldberger +(1914) so clearly shows the practical application of his work and that +of the other investigators of the subject, that we abstract from it +the following account:</p> + +<p>"In general terms it may be stated that association with a case of +typhus fever in the absence of the transmitting insect is no more +dangerous than is association with a case of yellow fever in the +absence of the yellow fever mosquito. Danger threatens only when +the insect appears on the scene."</p> + +<p>"We may say, therefore, that to prevent infection of the individual +it is necessary for him only to avoid being bitten by the louse. +In theory this may readily be done, for we know that the body louse +infests and attaches itself almost entirely to the body linen, and that +boiling kills this insect and its eggs. Individual prophylaxis is +based essentially, therefore, on the avoidance of contact with individuals +likely to harbor lice. Practically, however, this is not +always as easy as it may seem, especially under the conditions of +such intimate association as is imposed by urban life. Particularly +is this the case in places such as some of the large Mexican cities, +where a large proportion of the population harbors this vermin. +Under such circumstances it is well to avoid crowds or crowded places, +such as public markets, crowded streets, or public assemblies at +which the 'peon' gathers."</p> + +<p>"Community prophylaxis efficiently and intelligently carried out +is, from a certain point of view, probably easier and more effective +in protecting the individual than is the individual's own effort to +guard himself. Typhus emphasizes, perhaps better than any other +disease, the fact that fundamentally, sanitation and health are +economic problems. In proportion as the economic condition of the +masses has improved—that is, in proportion as they could afford<span class="pagenum"><a name="Page_240" id="Page_240">[Pg 240]</a></span> +to keep clean—the notorious filth disease has decreased or disappeared. +In localities where it still prevails, its further reduction +or complete eradication waits on a further improvement in, or extension +of, the improved economic status of those afflicted. Economic +evolution is very slow process, and, while doing what we can to hasten +it, we must take such precautions as existing conditions permit, +looking to a reduction in or complete eradication of the disease."</p> + +<p>"When possible, public bath houses and public wash houses, +where the poor may bathe and do their washings at a minimum or +without cost, should be provided. Similar provision should be +made in military and construction camps. Troops in the field should +be given the opportunity as frequently as possible to wash and <i>scald</i> +or <i>boil</i> their body linen."</p> + +<p>"Lodging houses, cheap boarding houses, night shelters, hospitals, +jails and prisons, are important factors in the spread and frequently +constitute foci of the disease. They should receive rigid sanitary +supervision, including the enforcement of measures to free all inmates +of such institutions of lice on admission."</p> + +<p>"So far as individual foci of the disease are concerned these +should be dealt with by segregating and keeping under observation +all exposed individuals for 14 days—the period of incubation—from +the last exposure, by disinfecting (boiling or steaming) the suspected +bedding, body linen, and clothes, for the destruction of any possible +vermin that they may harbor, and by fumigating (with sulphur) +the quarters that they may have occupied."</p> + +<p>"It will be noted that nothing has been said as to the disposition +of the patient. So far as the patient is concerned, he should be +removed to 'clean' surroundings, making sure that he does not +take with him any vermin. This may be done by bathing, treating +the hair with an insecticide (coal oil, tincture of larkspur), and a +complete change of body linen. Aside from this, the patient may +be treated or cared for in a general hospital ward or in a private house, +provided the sanitary officer is satisfied that the new surroundings +to which the patient has been removed are 'clean,' that is, free +from vermin. Indeed, it is reasonably safe to permit a 'clean' +patient to remain in his own home if this is 'clean,' for, as has already +been emphasized, there can be no spread in the absence of lice. +This is a common experience in native families of the better class +and of Europeans in Mexico City."</p> + +<p>"Similarly the sulphur fumigation above prescribed may be +dispensed with as unnecessary in this class of cases."<span class="pagenum"><a name="Page_241" id="Page_241">[Pg 241]</a></span></p> + + + + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_XI" id="CHAPTER_XI"></a>CHAPTER XI</h2> + +<h2>SOME POSSIBLE, BUT IMPERFECTLY ESTABLISHED CASES OF +ARTHROPOD TRANSMISSION OF DISEASE</h2> + + +<h3><a name="Infantile_Paralysis_or_Acute_Anterior_Poliomyelitis" id="Infantile_Paralysis_or_Acute_Anterior_Poliomyelitis"></a><span class="smcap">Infantile Paralysis or Acute Anterior Poliomyelitis</span></h3> + +<p>The disease usually known in this country as infantile paralysis +or, more technically, as acute anterior poliomyelitis, is one which +has aroused much attention in recent years.</p> + +<p>The causative organism of infantile paralysis is unknown, but +it has been demonstrated that it belongs to the group of filterable +viruses. It gives rise to a general infection, producing characteristic +lesions in the central nervous system. The result of the injury to +the motor nerves is a more or less complete paralysis of the corresponding +muscle. This usually manifests itself in the legs and arms. +The fatal cases are usually the result of paralysis of the muscles +of respiration. Of the non-fatal cases about 60 per cent remain +permanently crippled in varying degrees.</p> + +<p>Though long known, it was not until about 1890 that it was +emphasized that the disease occurs in epidemic form. At this time +Medin reported his observations on an epidemic of forty-three cases +which occurred in and around Stockholm in 1887. Since then, +according to Frost (1911), epidemics have been observed with increasing +frequency in various parts of the world. The largest recorded +epidemics have been those in Vermont, 1894, 126 cases; Norway and +Sweden, 1905, about 1,500 cases; New York City, 1907, about +2,500 cases. Since 1907 many epidemics have been reported in the +United States, and especially in the Northern States east of the +Dakotas. In 1912 there were over 300 cases of the disease in Buffalo, +N. Y., with a mortality of somewhat over 11 per cent.</p> + +<p>In view of the sudden prominence and the alarming spread of +infantile paralysis, there have been many attempts to determine +the cause, and the manner in which the disease spreads and develops +in epidemic form. In the course of these studies, the question of +possible transmission by insects was naturally suggested.</p> + +<p>C. W. Howard and Clark (1912) presented the results of studies +in this phase of the subject. They dealt especially with the house-fly, +bedbug, head, and body lice, and mosquitoes. It was found +that the house-fly (<i>Musca domestica</i>) can carry the virus of poliomyelitis +in an active state for several days upon the surface of the body<span class="pagenum"><a name="Page_242" id="Page_242">[Pg 242]</a></span> +and for several hours within the gastro-intestinal tract. Mosquitoes +and lice were found not to take up or maintain the virus. On the +other hand, the bedbug (<i>Cimex lectularius</i>) was found to take the +virus from the infected monkeys and to maintain it in a living state +within the body for a period of seven days. This was demonstrated +by grinding up in salt solution, insects which had fed on poliomyelitic +animals and injecting the filtrate into a healthy monkey. The experimenters +doubted that the bedbug is a carrier of the virus in nature.</p> + +<p>Earlier in the same year, Brues and Sheppard published the results +of an intensive epidemiological study of the outbreak of 1911, in +Massachusetts. Special attention had been paid to the possibility +of insect transfer and the following conclusion was reached:</p> + +<p>"Field work during the past summer together with a consideration +of the epidemiology of the disease so far as known, points strongly +toward biting flies as possible carriers of the virus. It seems probable +that the common stable-fly (<i>Stomoxys calcitrans</i> L.) may be responsible +to a certain extent for the spread of acute epidemic poliomyelitis, +possibly aided by other biting flies, such as <i>Tabanus lineola</i>. No +facts which disprove such a hypothesis have as yet been adduced, +and experiments based upon it are now in progress."</p> + +<p>As stated by Brues (1913), especial suspicion fell upon the stable-fly +because:</p> + +<p>1. The blood-sucking habits of the adult fly suit it for the transfer +of virus present in the blood.</p> + +<p>2. The seasonal abundance of the fly is very closely correlated +with the incidence of the disease, rising rapidly during the summer +and reaching a maximum in July and August, then slowly declining +in September and October.</p> + +<p>3. The geographical distribution of the fly is, so far as can be +ascertained, wider, or at least co-extensive with that of poliomyelitis.</p> + +<p>4. <i>Stomoxys</i> is distinctly more abundant under rural conditions, +than in cities and thickly populated areas.</p> + +<p>5. While the disease spreads over districts quickly and in a +rather erratic way, it often appears to follow along lines of travel, +and it is known that <i>Stomoxys</i> flies will often follow horses for long +distances along highways.</p> + +<p>6. In a surprisingly large number of cases, it appeared probable +that the children affected had been in the habit of frequenting places +where <i>Stomoxys</i> is particularly abundant, i.e., about stables, barnyards, +etc.<span class="pagenum"><a name="Page_243" id="Page_243">[Pg 243]</a></span></p> + +<p>The experiments referred to were carried on during the summer of +1912 and in September Dr. Rosenau announced that the disease was +transferred by the bite of the stable-fly.</p> + +<p>A monkey infected by inoculation was exposed to the bites of +upwards of a thousand of the <i>Stomoxys</i> flies daily, by stretching it +at full length and rolling it in a piece of chicken wire, and then placing +it on the floor of the cage in which the flies were confined. The flies +fed freely from the first, as well as later, after paralysis had set in. +Alternating with the inoculated monkey, healthy monkeys were +similarly introduced into the cage at intervals. New monkeys were +inoculated to keep a supply of such infected animals and additional +healthy ones were exposed to the flies, which fed willingly and in +considerable numbers on each occasion. "Thus the flies were given +every opportunity to obtain infection from the monkeys, since the +animals were bitten during practically every stage of the disease +from the time of the inoculation of the virus till their death following +the appearance of paralysis. By the same arrangement the +healthy monkeys were likely to be bitten by flies that had previously +fed during the various stages of the disease on the infected monkeys. +The flies had meanwhile enjoyed the opportunity of incubating the +virus for periods varying from the day or two which usually elapses +between consecutive feedings, to the two or three-week period for +which at least some (although a very small percentage) of the flies +lived in the cage."</p> + +<p>"In all, twelve apparently healthy monkeys of a small Japan +species were exposed to the flies in the manner described for the infected +monkeys. Some were placed in the cage only once or twice +and others a number of times after varying intervals. These exposures +usually lasted for about half an hour, but were sometimes +more protracted. No results were apparent until two or three +weeks after the experiment was well under way, and then in rather +rapid succession six of the animals developed symptoms of poliomyelitis. +In three, the disease appeared in a virulent form, resulting +in death, while the other three experienced transient tremblings, +diarrhœa, partial paralysis and recovery."—Brues, 1913.</p> + +<p>Very soon after the announcement of the results of experiments +by Rosenau and Brues, they were apparently conclusively confirmed +by Anderson and Frost (1912), who repeated the experiments, at +Washington. They announced that through the bites of the <i>Stomoxys</i> +flies that had previously fed on infected monkeys, they had succeeded +in experimentally infecting three healthy monkeys.<span class="pagenum"><a name="Page_244" id="Page_244">[Pg 244]</a></span></p> + +<p>The results of these experiments gained much publicity and in +spite of the conservative manner in which they had been announced, +it was widely proclaimed that infantile paralysis was conveyed in +nature by the stable-fly and by it alone.</p> + +<p>Serious doubt was cast on this theory by the results of further +experiments by Anderson and Frost, reported in May of 1913. +Contrary to the expectations justified by their first experience, the +results of all the later, and more extended, experiments were wholly +negative. Not once were these investigators again able to transmit +the infection of poliomyelitis through <i>Stomoxys</i>. They concluded that +it was extremely doubtful that the insect was an important factor +in the natural transmission of the disease, not only because of their +series of negative results, "but also because recent experiments have +afforded additional evidence of the direct transmissibility or contagiousness +of poliomyelitis, and because epidemiological studies +appear to us to indicate that the disease is more likely transmitted +largely through passive human virus carriers."</p> + +<p>Soon after this, Kling and Levaditi (1913) published their detailed +studies on acute anterior poliomyelitis. They considered that the +experiments of Flexner and Clark (and Howard and Clark), who fed +house-flies on emulsion of infected spinal cord, were under conditions +so different from what could occur in nature that one could not +draw precise conclusions from them regarding the epidemiology of +the disease. They cited the experiments of Josefson (1912), as +being under more reasonable conditions. He sought to determine +whether the inoculation of monkeys with flies caught in the wards of +the Hospital for Contagious Diseases at Stockholm, where they had +been in contact with cases of poliomyelitis, would produce the +disease. The results were completely negative.</p> + +<p>Kling and Lavaditi made four attempts of this kind. The flies +were collected in places where poliomyelitics had dwelt, three, four +and twenty-four after the beginning of the disease in the family and +one, three, and fifteen days after the patient had left the house. +These insects were for the greater part living and had certainly been +in contact with the infected person. In addition, flies were used +which had been caught in the wards of the Hospital for Contagious +Diseases at Söderkoping, when numbers of poliomyelitics were confined +there. Finally, to make the conditions as favorable as possible, +the emulsions prepared from these flies were injected without previous +filtering, since filtration often causes a weakening of the virus. In<span class="pagenum"><a name="Page_245" id="Page_245">[Pg 245]</a></span> +spite of these precautions, all their results were negative, none of the +inoculated animals having contracted poliomyelitis. They also +experimented with bedbugs which had fed upon infected patients at +various stages of the disease, but the results in these cases also were +wholly negative.</p> + +<p>Kling and Levaditi considered at length the possibility of transmission +of the disease by <i>Stomoxys</i>. As a result of their epidemiological +studies, they found that infantile paralysis continued to spread +in epidemic form in the dead of winter, when these flies were very +rare and torpid, or were even completely absent. Numerous cases +developed in the northern part of Sweden late in October and +November, long after snow had fallen. On account of the rarity +of the Stomoxys flies during the period of their investigations they +were unable to conduct satisfactory experiments. In one instance, +during a severe epidemic, they found a number of the flies in a stable +near a house inhabited by an infected family, though none was +found in the house itself. These flies were used in preparing an +emulsion which, after filtering, was injected into the peritoneal +cavity of a monkey. The result was wholly negative.</p> + +<p>As for the earlier experiments, Kling and Levaditi believe if the +flies were responsible for the transmission of the disease in the cases +reported by Rosenau and Brues, and the first experiments of Anderson +and Frost, it was because the virus of infantile paralysis is eliminated +with the nasal secretions of paralyzed monkeys and the flies, +becoming contaminated, had merely acted as accidental carriers.</p> + +<p>Still further evidence against the hypothesis of the transmission +of acute anterior poliomyelitis by <i>Stomoxys calcitrans</i> was brought +forward by Sawyer and Herms (1913). Special precautions were +used to prevent the transference of saliva or other possibly infectious +material from the surface of one monkey to that of another, and to +avoid the possibility of complicating the experiments by introducing +other pathogenic organisms from wild flies, only laboratory-bred +flies were used. In a series of seven carefully performed experiments, +in which the conditions were varied, Sawyer and Herms were +unable to transmit poliomyelitis from monkey to monkey through +the agency of <i>Stomoxys</i>, or to obtain any indication that the fly is the +usual agent for spreading the disease in nature.</p> + +<p>The evidence at hand to date indicates that acute anterior poliomyelitis, +or infantile paralysis, is transmitted by contact with infected +persons. Under certain conditions insects may be agents in +spreading the disease, but their rôle is a subordinate one.<span class="pagenum"><a name="Page_246" id="Page_246">[Pg 246]</a></span></p> + + +<h3><a name="Pellagra" id="Pellagra"></a>Pellagra</h3> + +<p><b>Pellagra</b> is an endemic and epidemic disease characterized by a +peculiar eruption or erythema of the skin (figs <a href="#Fig_144">144</a> and <a href="#Fig_145">145</a>), digestive +disturbances and nervous trouble. +Insanity is a common result, rather +than a precursor of the disease. +The manifestations of pellagra are +periodic and its duration indeterminate.</p> + +<div class="figleft" style="width: 350px;"><a name="Fig_144" id="Fig_144"></a> +<a href="images/f144-full.png"><img src="images/f144.png" width="350" height="540" alt="144. Pellagrous eruption on the face. +After Watson." title="144. Pellagrous eruption on the face. +After Watson." /></a> +<span class="caption">144. Pellagrous eruption on the face. +After Watson.</span> +</div> + +<p>The disease is one the very name +of which was almost unknown in the +United States until within the past +decade. It has usually been regarded +as tropical, though it occurs commonly +in Italy and in various parts of Europe. +Now it is known that it not only +occurs quite generally in the United +States but that it is spreading. Lavinder +(1911) says that "There are +certainly many thousand cases of the +disease in this country, and the present +situation must be looked upon +with grave concern."</p> + +<p>It is not within the scope of this book to undertake a general +discussion of pellagra. The subject is of such importance to every +medical man that we cannot do better than refer to Lavinder's +valuable précis. We can only touch briefly upon the entomological +phases of the problems presented.</p> + +<p>The most commonly accepted theories regarding the etiology +of the disease have attributed it to the use of Indian corn as an article +of diet. This supposed relationship was explained either on the +basis of, (a) insufficiency of nutriment and inappropriateness of +corn as a prime article of food; (b) toxicity of corn or, (c) parasitism +of certain organisms—fungi or bacteria—ingested with either sound +or deteriorated corn.</p> + +<p>In 1905, Sambon proposed the theory of the protozoal origin of +pellagra and in 1910 he marshalled an imposing array of objections +to the theory that there existed any relationship between corn and +the disease. He presented clear evidence that pellagra existed in +Europe before the introduction of Indian corn from America, as an<span class="pagenum"><a name="Page_247" id="Page_247">[Pg 247]</a></span> +article of diet, and that its spread was not <i>pari passu</i> with that of the +use of corn. Cases were found in which the patients had apparently +never used corn, though that is obviously difficult to establish. He +showed that preventive measures based on the theory had been a +failure. Finally, he believed that the recurrence of symptoms of +the disease for successive springs, in patients who abstained absolutely +from the use of corn, militated against the theory.</p> + +<div class="figcenter" style="width: 500px;"><a name="Fig_145" id="Fig_145"></a> +<a href="images/f145-full.png"><img src="images/f145.png" width="500" height="340" alt="145. Pellagrous eruption on the hand. After Watson." title="145. Pellagrous eruption on the hand. After Watson." /></a> +<span class="caption">145. Pellagrous eruption on the hand. After Watson.</span> +</div> + +<p>On the other hand, Sambon believed that the periodicity of the +symptoms, peculiarities of distribution and seasonal incidence, and +analogies of the symptoms to those of other parasitic diseases indicated +that pellagra was of protozoal origin, and that it was insect-borne.</p> + +<p>The insect carriers, he believed to be one or more species of +Simuliidæ, or black-flies. In support of this he stated that <i>Simulium</i> +appears to effect the same topographical conditions as pellagra, +that in its imago stage it seems to present the same seasonal incidence, +that it has a wide geographical distribution which seems to cover +that of pellagra, and that species of the genus are known to cause +severe epizootics. Concluding from his studies in Italy, that pellagra +was limited almost wholly to agricultural laborers, he pointed +out that the Simulium flies are found only in rural districts, and as a +rule do not enter towns, villages, or houses.</p> + +<div class="figcenter" style="width: 250px;"> +<a href="images/f146-full.png"><img src="images/f146.png" width="250" height="367" alt="146. A favorite breeding place of Simulium. Ithaca, N. Y." title="146. A favorite breeding place of Simulium. Ithaca, N. Y." /></a> +<span class="caption">146. A favorite breeding place of Simulium. Ithaca, N. Y.</span> +<span class="pagenum"><a name="Page_248" id="Page_248">[Pg 248]</a></span></div> + +<p>When Sambon's detailed report was published in 1910, his theory +was seized upon everywhere by workers who were anxious to test it<span class="pagenum"><a name="Page_249" id="Page_249">[Pg 249]</a></span> +and who, in most cases, were favorably disposed towards it because +of the wonderful progress which had been made in the understanding +of other insect-borne diseases. In this country, the entomological +aspects of the subject have been dealt with especially by Forbes +(1912), and by King and Jennings, under the direction of W. D. +Hunter, of the Bureau of Entomology, and in coöperation with +the Thompson-McFadden Pellagra Commission of the Department +of Tropical Medicine of the New York Post-Graduate Medical +School. An important series of experiments with monkeys has +been undertaken by S. J. Hunter, of Kansas, but unfortunately we have +as yet no satisfactory evidence that these animals are susceptible +to the disease—a fact which renders the whole problem difficult.</p> + +<p>The accumulated evidence is increasingly opposed to Sambon's +hypothesis of the transmission of pellagra by <i>Simulium</i>. This has +been so clearly manifested in the work of the Thompson-McFadden +Commission that we quote here from the report by Jennings (1914):</p> + +<p>"Our studies in 1912 convinced us that there was little evidence +to support the incrimination of any species of <i>Simulium</i> in South +Carolina in the transmission of pellagra. Reviewing the group as a +whole, we find that its species are essentially "wild" and lack those +habits of intimate association with man which would be expected +in the vector of such a disease as pellagra. Although these flies are +excessively abundant in some parts of their range and are moderately +so in Spartanburg County, man is merely an incidental host, and no +disposition whatever to seek him out or to invade his domicile seems +to be manifested. Critically considered, it is nearer the fact that +usually man is attacked only when he invades their habitat."</p> + +<p>"As our knowledge of pellagra accumulates, it is more and more +evident that its origin is in some way closely associated with the +domicile. The possibility that an insect whose association with man +and his immediate environment is, at the best, casual and desultory, +can be active in the causation of the disease becomes increasingly +remote."</p> + +<p>"Our knowledge of the biting habits of <i>Simulium</i> is not complete, +but it is evident, as regards American species at least, that these are +sometimes not constant for the same species in different localities. +Certain species will bite man freely when opportunity offers, while +others have never been known to attack him. To assume that the +proximity of a <i>Simulium</i>-breeding stream necessarily implies that +persons in its vicinity must be attacked and bitten is highly fallacious.<span class="pagenum"><a name="Page_250" id="Page_250">[Pg 250]</a></span> +In Spartanburg County attacks by <i>Simulium</i> seems to be +confined to the immediate vicinity of the breeding-places. Our +records and observations, exceedingly few in number, refer almost +exclusively to such locations. Statements regarding such attacks, +secured with much care and discrimination from a large number of +persons, including many pellagrins, indicate conclusively that these +insects are seldom a pest of man in this county. A certain number +of the persons questioned were familiar with the gnats in other +localities, but the majority were seemingly ignorant of the existence +of such flies with biting habits. This is especially striking, in view +of the fact that the average distance of streams from the homes of +the pellagra cases studied was about 200 yards, many being at a +distance of less than 200 yards, and that 78 per cent of these streams +were found to be infested by larval <i>Simulium</i>. Such ignorance in a +large number of persons cannot be overlooked and indicates strongly +that our belief in the negligible character of local attacks by <i>Simulium</i> +is well founded."</p> + +<p>"In localities infested by 'sand-flies,' mosquitoes, etc., these +pests are always well known and the ignorance described above is +very significant."</p> + +<p>"Such positive reports as we received nearly always referred to +bites received in the open, along streams, etc., and observations made +of their attack were of those on field laborers in similar situations. +Males engaged in agricultural pursuits are almost exempt from +pellagra in Spartanburg County. During the season of 1913, in +some two or three instances, observations were made of the biting +of <i>Simulium</i> and some additional and entirely creditable reports +were received. These observations and reports were under conditions +identical with those referred to in the reports of 1912 and confirm +the conclusions based on the observations of that year. I +would repeat with emphasis that it is inconceivable that a fly of the +appearance and habits of the prevalent species of <i>Simulium</i> could be +present in such a region, especially about the haunts of man and +attack him with sufficient frequency and regularity to satisfactorily +account for so active and prevalent a disease as pellagra without +being a well-known and recognized pest."</p> + +<p>"In connection with the conditions in the Piedmont region of +South Carolina, it may be well to cite the results of a study of those +in the arid region of western Texas."<span class="pagenum"><a name="Page_251" id="Page_251">[Pg 251]</a></span></p> + +<p>"In May, 1913, in company with Capt. J. F. Siler of the Thompson-McFadden +Pellagra Commission, I visited the region of which +Midland in Midland County is the center. This region is very dry +and totally devoid of running water for a long distance in every +direction. The only natural source of water-supply, a few water +holes and ponds, were visited and found to be of such a nature that +the survival of <i>Simulium</i>, far less its propagation in them, is absolutely +impossible. The nearest stream affording possibilities as a +source of <i>Simulium</i> is 60 miles away, while the average distance of +such possibility is not less than 100 miles."</p> + +<p>"Artificial sources of water-supply were also investigated carefully +and were found to offer no opportunity for the breeding of +<i>Simulium</i>."</p> + +<p>"At Midland the histories of five cases of pellagra were obtained, +which gave clear evidence that this place or its immediate vicinity +was the point of origin. Persons of long residence in the country +were questioned as to the occurrence of such flies as <i>Simulium</i> and +returned negative answers. These included a retired cattle owner, +who is a man of education and a keen observer, an expert veterinarian +stationed in the country who has the cattle of the country under +constant observation, and a practical cattle man, manager of a ranch +and of wide experience. The latter had had experience with 'Buffalo +gnats' in other localities (in the East) and is well acquainted +with them. His close personal supervision of the cattle under his +charge, makes it practically certain that he would have discovered +these gnats had they been present in the country."</p> + +<p>"At the time the study was made, <i>Simulium</i> was breeding and +active in the adult state in the vicinity of Dallas, Texas, in the +eastern part of the state. We have here a region in which cases of +pellagra have originated, yet in which <i>Simulium</i> does not and cannot +breed."</p> + +<p>Other possible insect vectors of pellagra have been studied in +great detail and the available evidence indicates that if <i>any</i> insect +plays a rôle in the spread of the disease, <i>Stomoxys calcitrans</i> most +nearly fills the conditions. This conclusion was announced by +Jennings and King in 1912, and has been supported by their subsequent +work.</p> + +<p>Yet, after all the studies of the past decade, the old belief that +pellagra is essentially of dietary origin is gaining ground. Goldberger, +Waring and Willets (1914) of the United States Public Health<span class="pagenum"><a name="Page_252" id="Page_252">[Pg 252]</a></span> +Service summarize their conclusions in the statement, (1) that it is +dependent on some yet undetermined fault in a diet in which the +animal or leguminous protein component is disproportionately large +and (2) that no pellagra develops in those who <i>consume</i> a mixed, +well-balanced, and varied diet, such, for example, as that furnished +by the Government to the enlisted men of the Army, Navy, and +Marine Corps.</p> + + +<h3><a name="Leprosy" id="Leprosy"></a>Leprosy</h3> + +<p><b>Leprosy</b> is a specific, infectious disease due to <i>Bacillus lepræ</i>, and +characterized by the formation of tubercular nodules, ulcerations, +and disturbances of sensation. In spite of the long time that the +disease has been known and the dread with which it is regarded, +little is known concerning the method of transfer of the causative +organism or the means by which it gains access to the human body.</p> + +<p>It is known that the bacilli are to be found in the tubercles, the +scurf of the skin, nasal secretions, the sputum and, in fact in practically +all the discharges of the leper. Under such conditions it is +quite conceivable that they may be transferred in some instances +from diseased to healthy individuals through the agency of insects +and other arthropods. Many attempts have been made to demonstrate +this method of spread of the disease, but with little success.</p> + +<p>Of the suggested insect carriers none seem to meet the conditions +better than mosquitoes, and there are many suggestions in literature +that these insects play an important rôle in the transmission of +leprosy. The literature has been reviewed and important experimental +evidence presented by Currie (1910). He found that mosquitoes +feeding, under natural conditions, upon cases of nodular leprosy +so rarely, if ever, imbibe the lepra bacillus that they cannot be +regarded as one of the ordinary means of transference of this bacillus +from lepers to the skin of healthy persons. He believes that the +reason that mosquitoes that have fed on lepers do not contain the +lepra bacillus is that when these insects feed they insert their proboscis +directly into a blood vessel and thus obtain bacilli-free blood, +unmixed with lymph.</p> + +<p>The same worker undertook to determine whether flies are able +to transmit leprosy. He experimented with five species found in +Honolulu,—<i>Musca domestica</i>, <i>Sarcophaga pallinervis</i>, <i>Sarcophaga +barbata</i>, <i>Volucella obesa</i> and an undetermined species of <i>Lucilia</i>. +The experiments with <i>Musca domestica</i> were the most detailed.<span class="pagenum"><a name="Page_253" id="Page_253">[Pg 253]</a></span> +From these experiments he concluded, first, that all of the above-named +flies, when given an opportunity to feed upon leprous fluids, +will contain the bacilli in their intestinal tracts and feces for several +days after such feeding. Second, that considering the habits of +these flies, and especially those of <i>Musca domestica</i>, it is certain that, +given an exposed leprous ulcer, these insects will frequently convey +immense numbers of lepra bacilli, directly or indirectly, to the skins, +nasal mucosa, and digestive tracts of healthy persons. Additional +evidence along this line has recently been brought forward by +Honeij and Parker (1914), who incriminate both <i>Musca domestica</i> +and <i>Stomoxys calcitrans</i>. Whether or not such insect-borne bacilli +are capable of infecting persons whose skin and mucosa are thus +contaminated, Currie was unwilling to maintain, but he concludes +that until we have more accurate knowledge on this point, we are +justified in regarding these insects with grave suspicion of being +one of the means of disseminating leprous infection.</p> + +<p>Various students of the subject have suggested that bed-bugs +may be the carriers of leprosy and have determined the presence of +acid-fast bacilli in the intestines of bed-bugs which had fed on leprous +patients. Opposed to this, the careful experiments of Thompson +(1913) and of Skelton and Parkham (1913) have been wholly negative.</p> + +<p>Borrel has recently suggested that <i>Demodex</i>, may play a rôle in +spreading the infection in families. Many other insects and acariens +have been suggested as possible vectors, but the experimental data +are few and in no wise conclusive. The most that can be said is that +it is quite possible that under favorable conditions the infection +might be spread by any of the several blood-sucking forms or by +house-flies.</p> + + +<h3><a name="Verruga_peruviana" id="Verruga_peruviana"></a>Verruga peruviana</h3> + +<p><b>Verruga peruviana</b> is defined by Castellani and Chalmers as "a +chronic, endemic, specific, general disorder of unknown origin, not +contagious, but apparently inoculable, and characterized by an irregular +fever associated with rheumatoid pains, anemia, followed +by granulomatous swellings in the skin, mucous membranes, and +organs of the body." It has been generally believed by medical +men interested that the comparatively benign eruptive verruga is +identical with the so-called Oroya, or Carrion's fever, a malignant +type. This view is not supported by the work of Strong, Tyzzer +and Brues, (1913).<span class="pagenum"><a name="Page_254" id="Page_254">[Pg 254]</a></span></p> + +<p>The disease is confined to South America and to definitely limited +areas of those countries in which it does occur. It is especially +prevalent in some parts of Peru.</p> + +<p>The causative organism and the method of transfer of verruga +are unknown. Castellani and Chalmers pointed out in 1910 that the +study of the distribution of the disease in Peru would impress one +with the similarity to the distribution of the Rocky Mountain fever +and would lead to the conclusion that the ætiological cause must in +some way be associated with some blood-sucking animal, perhaps an +arachnid, and that this is supported by the fact that the persons +most prone to the infection are those who work in the fields.</p> + +<p>More recently, Townsend (1913), in a series of papers, has maintained +that verruga and Carrion's disease are identical, and that they +are transmitted to man by the bites of the Psychodid fly, <i>Phlebotomus +verrucarum</i>. He succeeded in producing the eruptive type of the +disease in experimental animals by injecting a physiological salt +trituration of wild Phlebotomus flies. A cebus monkey was exposed +from October so to November 6, by chaining him to a tree in the +verruga zone, next to a stone wall from which the flies emerged in +large numbers every night. Miliar eruption began to appear on the +orbits November 13 and by November 21, there were a number of +typical eruptions, with exudation on various parts of the body +exactly like miliar eruptive sores commonly seen on legs of human +cases.</p> + +<p>An assistant in the verruga work, George E. Nicholson, contracted +the eruptive type of the disease, apparently as a result of being bitten +by the Phlebotomus flies. He had slept in a verruga zone, under a +tight net. During the night he evidently put his hands in contact +with the net, for in the morning there were fifty-five unmistakable +Phlebotomus bites on the backs of his hands and wrists.</p> + +<p>Townsend believes that in nature, lizards constitute the reservoir +of the disease and that it is from them that the Phlebotomus flies +receive the infection.</p> + + +<h3><a name="Cancer" id="Cancer"></a>Cancer</h3> + +<p>There are not wanting suggestions that this dread disease is +carried, or even caused, by arthropods. Borrel (1909) stated that +he had found mites of the genus <i>Demodex</i> in carcinoma of the face +and of the mammæ. He believed that they acted as carriers of the +virus.<span class="pagenum"><a name="Page_255" id="Page_255">[Pg 255]</a></span></p> + +<p>Saul (1910) and Dahl (1910) go much further, since they attribute +the production of the malignant growth to the presence of mites +which Saul had found in cancers. These Dahl described as belonging +to a new species, which he designated <i>Tarsonemus hominis</i>. These +findings have since been confirmed by several workers. Nevertheless, +the presence of the mite is so rare that it cannot be regarded as +an important factor in the causation of the disease. The theory +that cancer is caused by an external parasite is given little credence +by investigators in this field.</p> + + +<p><b>In conclusion</b>, it should be noted that the medical and entomological +literature of the past few years abounds in suggestions, and in +unsupported direct statements that various other diseases are insect-borne. +Knab (1912) has well said "Since the discovery that certain +blood-sucking insects are the secondary hosts of pathogenic parasites, +nearly every insect that sucks blood, whether habitually or +occasionally, has been suspected or considered a possible transmitter +of disease. No thought seems to have been given to the conditions +and the characteristics of the individual species of blood-sucking +insects, which make disease transmission possible."</p> + +<p>He points out that "in order to be a potential transmitter of human +blood-parasites, an insect must be closely associated with man and +normally have opportunity to suck his blood repeatedly. It is not +sufficient that occasional specimens bite man, as, for example, is the +case with forest mosquitoes. Although a person may be bitten by a +large number of such mosquitoes, the chances that any of these +mosquitoes survive to develop the parasites in question, (assuming +such development to be possible), and then find opportunity to bite +and infect another person, are altogether too remote. Applying +this criterion, not only the majority of mosquitoes but many other +blood-sucking insects, such as Tabanidæ and Simuliidæ, may be +confidently eliminated. Moreover, these insects are mostly in +evidence only during a brief season, so that we have an additional +difficulty of a very long interval during which there could be no propagation +of the disease in question." He makes an exception of +tick-borne diseases, where the parasites are directly transmitted from +the tick host to its offspring and where, for this reason, the insect +remains a potential transmitter for a very long period. He also +cites the trypanosome diseases as possible exceptions, since the causative +organisms apparently thrive in a number of different vertebrate +hosts and may be transmitted from cattle, or wild animals, to man.<span class="pagenum"><a name="Page_256" id="Page_256">[Pg 256]</a></span></p> + +<p>Knab's article should serve a valuable end in checking irresponsible +theorizing on the subject of insect transmission of disease. +Nevertheless, the principles which he laid down cannot be applied +to the cases of accidental carriage of bacterial diseases, or to those +of direct inoculation of pyogenic organisms, or of blood parasites +such as the bacillus of anthrax, or of bubonic plague. Accumulated +evidence has justified the conclusion that certain trypanosomes +pathogenic to man are harbored by wild mammals, and so form an +exception. Townsend believes that lizards constitute the natural +reservoir of verruga; and it seems probable that field mice harbor +the organism of tsutsugamushi disease. Such instances are likely to +accumulate as our knowledge of the relation of arthropods to disease +broadens.<span class="pagenum"><a name="Page_257" id="Page_257">[Pg 257]</a></span></p> + + + + + +<hr style="width: 65%;" /> +<h2><a name="CHAPTER_XII" id="CHAPTER_XII"></a>CHAPTER XII</h2> + +<h2>HOMINOXIOUS ARTHROPODS</h2> + + +<p>The following synoptic tables are presented in the hope that they +may be of service in giving the reader a perspective of the relationships +of the Arthropoda in general and enabling him to identify the +more important species which have been found noxious to man. +Though applicable chiefly to the arthropods found in the United +States, exotic genera and species which are concerned in the transmission +of disease are also included. For this reason the keys to the +genera of the Muscids of the world are given. As will be seen, the +tables embrace a number of groups of species which are not injurious. +This was found necessary in order that the student might not be +lead to an erroneous determination which would result were he to +attempt to identify a species which heretofore had not been considered +noxious, by means of a key containing only the noxious forms. The +names printed in <b>bold faced type</b> indicate the hominoxious arthropods +which have been most commonly mentioned in literature.</p> + + +<h3><a name="CRUSTACEA" id="CRUSTACEA"></a>CRUSTACEA</h3> + +<p>Arthropods having two pairs of antennæ which are sometimes +modified for grasping, and usually with more than five pairs of legs. +With but few exceptions they are aquatic creatures. Representatives +are: Crabs, lobsters, shrimps, crayfish, water-fleas, and woodlice. +To this class belongs the <b>Cyclops</b> (<a href="#Fig_122">fig. 122</a>) a genus of minute aquatic +crustaceans of which at least one species harbors <i>Dracunculus medinensis</i>, +the Guinea worm (<a href="#Fig_121">fig. 121</a>).</p> + + +<h3><a name="MYRIAPODA_1" id="MYRIAPODA_1"></a>MYRIAPODA</h3> + +<p>Elongate, usually vermiform, wingless, terrestrial creatures having +one pair of antennæ, legs attached to each of the many intermediate +body segments. This group is divided into two sections, now usually +given class rank: the <b>Diplopoda</b> or millipedes (<a href="#Fig_13">fig. 13</a>), commonly +known as thousand legs, characterized by having two pairs of legs +attached to each intermediate body segment, and the <b>Chilopoda</b> +or centipedes (<a href="#Fig_14">fig. 14</a>) having only one pair of legs to each body segment.<span class="pagenum"><a name="Page_258" id="Page_258">[Pg 258]</a></span></p> + + +<h3>ARACHNIDA</h3> + +<p>In this class the antennæ are apparently wanting, wings are never +present, and the adults are usually provided with four pairs of legs. +Scorpions, harvest-men, spiders, mites, etc.</p> + + +<h3>HEXAPODA (Insects)</h3> + +<p>True insects have a single pair of antennæ, which is rarely vestigial, +and usually one or two pairs of wings in the adult stage. Familiar +examples are cockroaches, crickets, grasshoppers, bugs, dragon-flies, +butterflies, moths, mosquitoes, flies, beetles, ants, bees and wasps.</p> + + +<h3><a name="ORDERS_OF_THE_ARACHNIDA" id="ORDERS_OF_THE_ARACHNIDA"></a>ORDERS OF THE ARACHNIDA</h3> + +<div class="hanging"> +<p>a. Abdomen distinctly segmented. A group of orders including scorpions, +(<a href="#Fig_11">fig. 11</a>), whip-scorpions (<a href="#Fig_10">fig. 10</a>), pseudo-scorpions, solpugids (<a href="#Fig_12">fig. 12</a>) +harvest-men (daddy-long-legs or harvestmen), etc. <span class="rightalign"><b>Arthrogastra</b></span></p> + +<p>aa. Abdomen unsegmented, though sometimes with numerous annulations +<span class="rightalign"><b>Sphærogastra</b></span></p> + +<div class="hanging"> +<p>b. A constriction between cephalothorax and abdomen (<a href="#Fig_7">fig. 7</a>). True Spiders +<span class="rightalign"><b>Araneida</b></span></p> + +<p>bb. No deep constriction between these parts.</p> + +<div class="hanging"> +<p>c. Legs usually well developed, body more or less depressed (<a href="#Fig_49">fig. 49</a>). Mites +<span class="rightalign"><b>Acarina</b></span></p> + +<p>cc. Legs stumpy or absent, body more or less elongate or vermiform, or if +shorter, the species is aquatic or semi-aquatic in habit.</p> + +<div class="hanging"> +<p>d. Four pairs of short legs; species inhabiting moss or water. Water-bears. +<span class="rightalign"><b>Tardigrada</b></span></p> + +<p>dd. Two pairs of clasping organs near the mouth, instead of legs, in the +adult; worm-like creatures parasitic within the nasal passages, +lungs, etc. of mammals and reptiles (<a href="#Fig_148">fig. 148</a>). Tongue worms. +<span class="rightalign"><b>Linguatulina</b></span></p> +</div></div></div></div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_148" id="Fig_148"></a> +<a href="images/f148-full.png"><img src="images/f148.png" width="500" height="229" alt="148. Linguatula. (a) larva; (enlarged). (b) adult; (natural size)." title="148. Linguatula. (a) larva; (enlarged). (b) adult; (natural size)." /></a> +<span class="caption">148. Linguatula. (<i>a</i>) larva; (enlarged). (<i>b</i>) adult; (natural size).</span> +</div><p><span class="pagenum"><a name="Page_259" id="Page_259">[Pg 259]</a></span></p> + + +<h3><a name="ACARINA_KEY" id="ACARINA_KEY"></a>ACARINA<a name="FNanchor_E_5" id="FNanchor_E_5"></a><a href="#Footnote_E_5" class="fnanchor">[E]</a></h3> + +<div class="hanging"> +<p>a. Abdomen annulate, elongate; very minute forms, often with but four legs +(<a href="#Fig_62">fig. 62</a>). <span class="rightalign"><span class="smcap"><b>Demodicoidea</b></span></span></p> + +<div class="hanging"> +<p>b. With but four legs of five segments each. Living on plants, often forming +galls. <span class="rightalign"><span class="smcap"><b>Eriophyidæ</b></span></span></p> + +<p>bb. With eight legs, of three segments each. Living in the skin of mammals. +<span class="rightalign"><span class="smcap"><b>Demodicidæ</b></span></span></p> + +<div class="hanging"> +<p>To this family belongs the genus <b>Demodex</b> found in the sebaceous glands +and hair follicles of various mammals, including man. <i>D. phylloides</i> +Csokor has been found in Canada on swine, causing white tubercles +on the skin. <i>D. bovis</i> Stiles has been reported from the United States +on cattle, upon the skin of which they form swellings. <b>D. folliculorum</b> +Simon is the species found on man. See <a href="#Page_78">page 78</a>.</p> +</div></div> + +<p>aa. Abdomen not annulate nor prolonged behind; eight legs in the adult stage.</p> + +<div class="hanging"> +<p>b. With a distinct spiracle upon a stigmal plate on each side of the body (usually +ventral) above the third or fourth coxæ or a little behind (<a href="#Fig_50">fig. 50</a>); +palpi free; skin often coriaceous or leathery; tarsi often with a sucker.</p> + +<div class="hanging"> +<p>c. Hypostome large (<a href="#Fig_50">fig. 50</a>), furnished below with many recurved teeth; +venter with furrows, skin leathery; large forms, usually parasitic. +<span class="rightalign"><span class="smcap"><b>Ixodoidea</b></span></span></p> + +<div class="hanging"> +<p>d. Without scutum but covered by a more or less uniform leathery integument; +festoons absent; coxæ unarmed, tarsi without ventral spurs; +pulvilli absent or vestigial in the adults; palpi cylindrical; sexual +dimorphism slight. <span class="rightalign"><span class="smcap"><b>Argasidæ</b></span></span></p> + +<div class="hanging"> +<p>e. Body flattened, oval or rounded, with a distinct flattened margin +differing in structure from the general integument; this margin +gives the body a sharp edge which is not entirely obliterated even +when the tick is full fed. Capitulum (in adults and nymphs) +entirely invisible dorsally, distant in the adult by about its own +length from the anterior border. Eyes absent. <span class="rightalign"><b>Argus</b> Latr.</span></p> + +<div class="hanging"> +<p>f. Body oblong; margin with quadrangular cells; anterior tibiæ and +metatarsi each about three times as long as broad. On poultry, +southwest United States. <span class="rightalign"><b>A. persicus miniatus</b></span></p> + +<div class="hanging"> +<p><i>A. brevipes</i> Banks, a species with proportionately shorter legs has +been recorded from Arizona.</p> +</div> + +<p>ff. With another combination of characters. About six other species +of <i>Argas</i> from various parts of the world, parasitic on birds and +mammals.</p> +</div> + +<p>ee. Body flattened when unfed, but usually becoming very convex on +distention; anterior end more or less pointed and hoodlike; +margin thick and not clearly defined, similar in structure to the +rest of the integument and generally disappearing on distention; +capitulum subterminal, its anterior portions often visible dorsally +in the adult; eyes present in some species.</p> + +<div class="hanging"> +<p>f. Integument pitted, without rounded tubercles; body provided +with many short stiff bristles; eyes absent. On horses, cattle +and man (<a href="#Fig_48">fig. 48</a>). <span class="rightalign"><b>Otiobius</b> Banks.</span></p> + +<div class="hanging"> +<p><b>O. megnini</b>, a widely distributed species, is the type of this genus.<span class="pagenum"><a name="Pg_260" id="Pg_260"></a>[Pg 260]</span></p> +</div> + +<p>ff. Integument with rounded tubercles or granules; body without stiff +bristles. <span class="rightalign"><b>Ornithodoros</b> Koch.</span></p> + +<div class="hanging"> +<p>g. Two pairs of eyes; tarsi IV with a prominent subterminal spur +above; leg I strongly roughened. On cattle and man. +<span class="rightalign"><b>O. coriaceus</b></span></p> + +<p>gg. No eyes; no such spur on the hind tarsi.</p> + +<div class="hanging"> +<p>h. Tarsi I without humps above. <span class="rightalign"><i>O. talaje.</i></span></p> + +<p>hh. Tarsi I with humps above.</p> + +<div class="hanging"> +<p>i. Tarsi IV without distinct humps above. On hogs, cattle +and man. <span class="rightalign"><b>O. turicata</b></span></p> + +<p>ii. Tarsi IV with humps nearly equidistant (<a href="#Fig_142">fig. 142</a>). Africa. +<span class="rightalign"><b>O. moubata</b></span></p> +</div></div></div></div></div> + +<p>dd. With scutum or shield (<a href="#Fig_50">fig. 50</a>); festoons usually present; coxæ +usually armed with spurs, tarsi generally with one or two ventral +spurs; pulvilli present in the adults; sexual dimorphism pronounced. +<span class="rightalign"><span class="smcap"><b>Ixodidæ</b></span></span></p> + +<div class="hanging"> +<p>e. With anal grooves surrounding anus in front; inornate; without eyes; +no posterior marginal festoons; venter of the male with non-salient +plates. Numerous species, 14 from the United States, +among them <b>I. ricinus</b> (fig. <a href="#Fig_49">49</a> and <a href="#Fig_50">50</a>), <b>scapularis</b>, <b>cookei</b>, <i>hexagonus</i>, +<i>bicornis</i>. <span class="rightalign"><b>Ixodes</b> Latr. (including Ceratixodes).</span></p> + +<p>ee. With anal groove contouring anus behind, or groove faint or obsolete.</p> + +<div class="hanging"> +<p>f. With short palpi (<a href="#Fig_149">fig. 149</a>).</p> + +<div class="hanging"> +<p>g. Without eyes, inornate, with posterior marginal festoons; male +without ventral plates. Numerous species. <i>H. chordeilis</i> +and <i>leporis-palustris</i> from the United States. +<span class="rightalign"><i>Hæmaphysalis</i> Koch.</span><span class="pagenum"><a name="Page_262" id="Page_262">[Pg 262]</a></span></p> + +<p>gg. With eyes.</p> + +<div class="hanging"> +<p>h. Anal groove distinct; posterior marginal festoons present.</p> + +<div class="hanging"> +<p>i. Base of the capitulum (<a href="#Fig_150">fig. 150c</a>) rectangular dorsally; +usually ornate. <span class="rightalign"><b>Dermacentor</b> Koch.</span></p> + +<div class="hanging"> +<p>j. Adults with four longitudinal rows of large denticles on +each half of hypostome; stigmal plate nearly circular, +without dorso-lateral prolongation, goblets very large, +attaining 43µ to 115µ in diameter; not over 40 per +plate, each plate surrounded by an elevated row of +regularly arranged supporting cells; white rust wanting; +base of capitulum distinctly broader than long, +its postero-lateral angles prolonged slightly, if at all; +coxæ T with short spurs; trochanter I with small +dorso-terminal blade. Texas, Arizona, etc. <span class="rightalign"><i>D. nitens</i></span></p> + +<p>jj. Adults with three longitudinal rows of large denticles on +each half of hypostome; goblet cells always more +than 40 per plate; whitish rust usually present.</p> + +<div class="hanging"> +<p>k. Dorso-lateral prolongation of stigmal plate small or +absent; plates of the adults distinctly longer than +broad; goblet cells large, usually 30µ to 85µ in +diameter, appearing as very coarse punctations on +untreated specimens, but on specimens treated +with caustic potash they appear very distinct in +outline; base of capitulum distinctly (usually about +twice) broader than long, the postero-lateral angles +distinctly produced caudad; spurs of coxæ I long, +lateral spur slightly longer than median; trochanter +I with dorso-terminal spur. <i>D. albipictus</i>, +(= <i>variegatus</i>), <i>salmoni</i>, <i>nigrolineatus</i>.<span class="pagenum"><a name="Page_263" id="Page_263"></a>[Pg 263]</span></p> + +<p>kk. Dorso-lateral prolongation of stigmal plate distinct.</p> + +<div class="hanging"> +<p>l. Body of plate distinctly longer than broad; goblet +cells of medium size, usually 17.5µ to 35µ or 40µ in +diameter, appearing as medium sized punctuations +on untreated specimens, but on the specimens +treated with caustic potash they appear +very distinct in outline, which is not circular; +base of capitulum usually less than twice as broad +as long, the postero-lateral angles always distinctly +prolonged caudad.</p> + +<div class="hanging"> +<p>m. Trochanter I with distinct dorso-subterminal +retrograde sharp, digitate spur; postero-lateral +angles of capitulum pronouncedly +prolonged caudal, 112µ to 160µ long; goblet +cells attain 13µ to 40µ in diameter; type +locality California. <span class="rightalign"><b>D. occidentalis</b></span></p> + +<p>mm. Trochanter I with dorso-terminal blade; postero-lateral +angles of capitulum with rather short +prolongations.</p> + +<div class="hanging"> +<p>n. Stigmal plate small, goblet cells not exceeding +45 in the male or 100 in the female; scutum +with little rust, coxa I with short spurs, the +inner distinctly shorter than the outer. +<span class="rightalign"><i>D. parumapertus-marginatus</i></span></p> + +<p>nn. Stigmal plate larger; goblet cells over 70 in +the male and over 100 in the female; coxa I +with longer spurs, inner slightly shorter +than the outer; scutum with considerable +rust. <span class="rightalign"><b>D. venustus</b><a name="FNanchor_F_6" id="FNanchor_F_6"></a><a href="#Footnote_F_6" class="fnanchor">[F]</a></span></p> +</div></div> + +<p>ll. Goblet cells small, rarely exceeding 17.6µ, occasionally +reaching 19µ in diameter; on untreated specimens +they appear as very fine granulations, and on +specimens treated with caustic potash they may +be difficult to see, but their large number can +be determined from the prominent stems of the +goblets; surface of outline of the goblets distinctly +circular; base of the capitulum usually less +than twice as broad as long, the postero-lateral +angle distinctly prolonged caudad; spurs of +coxæ I long. +<i>D. reticulatus</i> and <i>electus</i> (= <i>variabilis</i>?)</p> +</div></div></div> + +<p>ii. Base of the capitulum (<a href="#Fig_151">fig. 151</a>) usually hexagonal (except +in the male of <i>puchellus</i>); and usually inornate.<span class="pagenum"><a name="Page_264" id="Page_264">[Pg 264]</a></span></p> + +<div class="hanging"> +<p>j. No ventral plate or shield in either sex (<a href="#Fig_153">fig. 153</a>). <b>R. +bicornis</b> from the United States. <span class="rightalign"><b>Rhipicentor</b> Nuttall</span></p> + +<p>jj. Males with a pair of adanal shields, and usually a pair of +accessory adanal shields. Numerous species, among +them <i>R. sanguineus</i> (<a href="#Fig_154">fig. 154</a>) and <i>texanus</i>, the latter +from the United States. <span class="rightalign"><i>Rhipicephalus</i> Koch</span></p> +</div></div> + +<p>hh. Anal grooves faint or obsolete; no marginal festoons.</p> + +<div class="hanging"> +<p>i. Short palpi; highly chitinized; unfed adults of large size; +coxæ conical; male with a median plate prolonged in two +long spines projecting caudad; segments of leg pair IV +greatly swollen (fig. <a href="#Fig_155">155</a>, <a href="#Fig_156">156</a>). <i>M. winthemi</i> +<span class="rightalign"><i>Margaropus</i> Karsch</span></p> + +<p>ii. Very short palpi, ridged dorsally and laterally; slightly +chitinized; unfed adults of smaller size; coxæ I bifid; +male with adanal and accessory adanal shields (<a href="#Fig_139">fig. 139</a>). +<b>B. annulatus.</b> <span class="rightalign"><b>Boophilus</b> Curtis</span></p> +</div></div></div> + +<p>ff. Palpi longer than broad (<a href="#Fig_157">fig. 157</a>).</p> + +<div class="hanging"> +<p>g. Male with pair of adanal shields, and two posterior abdominal +protrusions capped by chitinized points; festoons present or +absent. Several species, among them <b>H. ægypticum</b> (<a href="#Fig_140">fig. 140</a>) +from the old world. <span class="rightalign"><b>Hyalomma</b> Koch</span></p> + +<p>gg. Male without adanal shields but small ventral plaques are +occasionally present close to the festoons. Many species, a +few from the United States (<a href="#Fig_157">fig. 157</a>). <span class="rightalign"><b>Amblyomma</b> Koch</span></p> + +<div class="hanging"> +<p>h. Coxa I with but one spine, metatarsi (except I) with two +thickened spurs at tips. <span class="rightalign"><i>A. maculatum</i></span></p> + +<p>hh. Coxa I with two spines; metatarsi without stout spurs at +tips, only slender hairs.<span class="pagenum"><a name="Page_265" id="Page_265">[Pg 265]</a></span></p> + +<div class="hanging"> +<p>i. Projections of coxa I blunt and short. Large species on the +gopher tortoise in Florida. <span class="rightalign"><i>A. tuberculatum</i></span></p> + +<p>ii. Projections of coxa I longer, and at least one of them sharp +pointed; second segment of palpus twice as long as the +third; coxa IV of the male with a long spine.</p> + +<div class="hanging"> +<p>j. Porose areas nearly circular; shield of both sexes pale +yellowish, with some silvery streaks and marks, and +some reddish spots; shield of female as broad as long. +<span class="rightalign"><b>A. cajennense</b> (= <b>mixtum</b>).</span></p> + +<p>jj. Porose areas elongate, shield brown, in the female with +an apical silvery mark, in the male with two small +and two or four other silvery spots; shield of the female +longer than broad (<a href="#Fig_158">fig 158 e</a>). <span class="rightalign"><b>A. americanum.</b></span></p> +</div></div></div></div></div></div></div> + +<p>cc. Hypostome small, without teeth, venter without furrows; body often +with coriaceous shields, posterior margin of the body never crenulate +(i.e. without festoons); no eyes. <span class="rightalign">GAMASOIDEA.</span></p> + +<div class="hanging"> +<p>d. Parasitic on vertebrates; mandibles fitted for piercing; body sometimes +constricted. <span class="rightalign"><span class="smcap">Dermanyssidæ.</span></span></p> + +<div class="hanging"> +<p>e. Anal plate present. <span class="rightalign"><span class="smcap">Dermanyssinæ.</span></span></p> + +<div class="hanging"> +<p>f. Body short; legs stout, hind pair reaching much beyond the tip of +the body. On bats. <span class="rightalign"><i>Pteroptus</i> Dufour.</span></p> + +<p>ff. Body long; hind legs not reaching beyond the tip of the body.</p> + +<div class="hanging"> +<p>g. Peritreme on the dorsum, very short; body distinctly constricted. +<span class="rightalign"><i>Ptilonyssus</i> Berl.</span></p> + +<p>gg. Peritreme on the venter, longer; body not distinctly constricted.</p> + +<div class="hanging"> +<p>h. Mandibles in both sexes chelate. Parasitic on bats, mice +and birds (<a href="#Fig_150">fig. 150, h</a>). <span class="rightalign"><b>Liponyssus</b> Kol.</span></p> + +<div class="hanging"> +<p>The species <b>L.</b> (= <b>Leiognathus</b>) <b>sylviarum</b> frequents the +nests of warblers. An instance is on record of these mites +attacking man, causing a pruritis.<span class="pagenum"><a name="Pg_266" id="Pg_266"></a>[Pg 266]</span></p> +</div> + +<p>hh. Mandibles in the male chelate (<span title="refers to the female"><a name="AC_13" id="AC_13"></a><a href="#Fig_158">fig. 158 j</a></span>), in the female long, +styliform. Parasitic on birds. <span class="rightalign"><b>Dermanyssus</b> Dug.</span></p> + +<div class="hanging"> +Two species of importance may be noted, <i>D. hirundinus</i> +and <b>D. gallinæ</b>. The latter (<a href="#Fig_51">fig. 51</a>) is a serious pest +of poultry, sometimes attacking man, causing itching +and soreness. +</div></div></div></div> + +<p>ee. Anal plate absent. In lungs and air passages of some mammals. +<span class="rightalign"><span class="smcap"><b>Halarachninæ.</b></span></span></p> +</div> + +<p>dd. Free or attached to insects, rarely on vertebrates.</p> + +<div class="hanging"> +<p>e. First pair of legs inserted within the same body opening as the oral +tube; genital apertures surrounded by the sternum. On insects. +<span class="rightalign"><span class="smcap">Uropodidæ.</span></span></p> + +<p>ee. First pair of legs inserted at one side of the mouth opening; male +genital aperture usually on the anterior margin of the sternal +plate. <span class="rightalign"><b>Gamasidæ.</b></span></p> + +<div class="hanging"> +<p>This family contains a number of genera, some of which are found +upon mammals, though the majority affect only other arthropods. +One species, <b>Lælaps stabularis</b>, frequents the bedding +in stables, and in one instance at least, has occasioned irritation +and itching, in man.</p> +</div></div></div></div> + +<p>bb. No distinct spiracle in the stigmal plate on each side of the body.</p> + +<div class="hanging"> +<p>c. Body usually coriaceous, with few hairs, with a specialized seta arising +from a pore near each posterior corner of the cephalothorax; no eyes; +mouth parts and palpi very small; ventral openings of the abdomen +large; tarsi without sucker. Not parasitic. <span class="rightalign">ORIBATOIDEA.</span></p> + +<p>cc. Body softer; without such specialized seta.</p> + +<div class="hanging"> +<p>d. Aquatic species. <span class="rightalign">HYDRACHNOIDEA.</span></p> + +<p>dd. Not aquatic.<span class="pagenum"><a name="Page_267" id="Page_267">[Pg 267]</a></span></p> + +<div class="hanging"> +<p>e. Palpi small, three segmented, adhering for some distance to the lip; +ventral suckers at genital opening or near anal opening usually +present; no eyes; tarsi often end in suckers; beneath the skin on +the venter are seen rod-like epimera that support the legs; body +often entire. Adults frequently parasitic. <span class="rightalign"><b>SARCOPTOIDEA.</b></span></p> + +<div class="hanging"> +<p>f. With tracheæ; no ventral suckers; legs ending in claws; body +divided into cephalothorax and abdomen; the female with a +clavate hair between legs I and II. Usually not parasitic +on birds and mammals. <span class="rightalign"><span class="smcap"><b>Tarsonemidæ</b></span></span></p> + +<div class="hanging"> +<p>g. Hind legs of female ending in claw and sucker as in the other +pairs. <span class="rightalign"><span class="smcap"><b>Pediculoidinæ</b></span></span></p> + +<div class="hanging"> +<p>To this sub-family belongs the genus <span class="smcap"><b>Pediculoides</b></span></p> + +<p><b>P. ventricosus</b> is described on <a href="#Page_69">page 69</a>.</p> +</div> + +<p>gg. Hind legs of the female end in long hairs. <span class="rightalign"><span class="smcap"><b>Tarsoneminæ</b></span></span></p> + +<div class="hanging"> +<b>Tarsonemus intectus</b> Karpelles, normally found upon grain, +is said to attack man in Hungary and Russia. Other +species of the genus affect various plants (c.f. <a href="#Fig_150">fig. 150, g</a>). +</div></div> + +<p>ff. Without tracheæ; no such clavate hair.</p> + +<div class="hanging"> +<p>g. Genital suckers usually present; integument usually without +fine parallel lines.</p> + +<div class="hanging"> +<p>h. Legs short, without clavate hair on tarsi I and II. On +insects. <span class="rightalign"><span class="smcap">Canestrinidæ.</span></span></p> + +<p>hh. Legs longer, with a clavate hair on tarsi I and II. Not +normally parasitic except on bees. <span class="rightalign"><span class="smcap"><b>Tyroglyphidæ</b></span></span></p> + +<div class="hanging"> +<p>i. Dorsal integument more or less granulate; claws very weak, +almost invisible; some hairs of the body plainly feathered; +ventral apertures large. <span class="rightalign"><b>Glyciphagus</b> Her.</span></p> + +<div class="hanging"> +<p>This genus occurs in the United States. In Europe the +mites have been found feeding on all sorts of substances. +They are known as sugar mites and cause the disease<span class="pagenum"><a name="Pg_268" id="Pg_268"></a>[Pg 268]</span> +known as grocer's itch. <b>G. domesticus</b> and <b>G. prunorum</b> +are old world species (<a href="#Fig_150">fig. 150, d</a>).</p> +</div> + +<p>ii. Dorsal integument not granulate; claws distinct; no +prominent feathered hairs; ventral aperture small.</p> + +<div class="hanging"> +<p>j. Mandibles not chelate; elongate, and toothed below; +body without long hairs; palpi enlarged at tip and +provided with two divergent bristles. Species feed on +decaying substances. <span class="rightalign"><i>Histiostoma</i> Kram.</span></p> + +<p>jj. Mandibles chelate; palpi not enlarged at the tip, nor +with two bristles.</p> + +<div class="hanging"> +<p>k. No clavate hair on the base of tarsi I and II; no +suture between cephalothorax and abdomen. Live +on bees or in their nests. <span class="rightalign"><i>Trichotarsus</i> Can.</span></p> + +<p>kk. A clavate or thickened hair at the base of tarsi I and II.</p> + +<div class="hanging"> +<p>l. The bristle on the penultimate segment of the legs +arises from near the middle; no suture between the +cephalothorax and abdomen. The species, some +of which occur in the United States, feed on dried +fruit, etc. <span class="rightalign"><i>Carpoglyphus</i> Robin.</span></p> + +<p>ll. The bristle on the penultimate segment of the legs +arise from near the tip; a suture between cephalothorax +and abdomen.</p> + +<div class="hanging"> +<p>m. Cephalothorax with four distinct and long bristles +in a transverse row; tarsi I and II about twice +as long as the preceding segment (<a href="#Fig_150">fig. 150 f</a>). +<span class="rightalign"><b>Tyroglyphus</b> Latr.</span></p> + +<div class="hanging"> +<p>n. Some bristles on tarsi I and II near middle, +distinctly spine-like; the sense hair about its +length from the base of the segment. Several +species in the United States belong to this +group.</p> + +<p>nn. No spine-like bristles near the middle of the +tarsi; sense hair not its length from the base +of the segment.</p> + +<div class="hanging"> +<p>o. Of the terminal abdominal bristles, only two +are about as long as the abdomen; leg I +of the male greatly thickened and with a +spine at apex of the femur below. <span class="rightalign"><b>T. farinæ.</b></span></p> + +<p>oo. Of the terminal abdominal bristles at least +six or more are very long, nearly as long +as the body.</p> + +<div class="hanging"> +<p>p. Bristles of the body distinctly plumose or +pectinate; tarsi very long. <span class="rightalign"><b>T. longior.</b></span></p> + +<p>pp. Bristles of the body not pectinate.</p> + +<div class="hanging"> +<p>q. In mills, stored foods, grains, etc. Third +and fourth joints of hind legs scarcely +twice as long as broad; abdominal +bristles not unusually long; legs I<span class="pagenum"><a name="Page_269" id="Page_269">[Pg 269]</a></span> +and II of the male not unusually +stout. <span class="rightalign"><b>T. americanus.</b></span></p> + +<p>qq. With other characters and habits. +<i>T. lintneri</i> (<a href="#Fig_150">fig. 150</a> f) the mushroom +mite, and several other species.</p> +</div></div></div></div> + +<p>mm. Cephalothorax with but two long distinct +bristles (besides the frontal pair), but sometimes +a very minute intermediate pair; +tarsi I and II unusually short and not twice +as long as the preceding segment.</p> + +<div class="hanging"> +<p>n. Tarsi with some stout spines. <span class="rightalign"><b>Rhizoglyphus</b> Clap.</span></p> + +<div class="hanging"> +<p>The species of this genus are vegetable feeders. +Several occur in the United States. +<b>R. parasiticus</b> and <b>R. spinitarsus</b> have been +recorded from the old world, attacking human +beings who handle affected plants.</p> +</div> + +<p>nn. Tarsi with only fine hairs. <span class="rightalign"><b>Monieziella</b> Berl.</span></p> + +<div class="hanging"> +<p>The species of this genus, as far as known, +are predaceous or feed on recently killed +animal matter. Several species occur +in the United States. <b>M.</b> (= <b>Histiogaster</b>) +<b>entomophaga</b> (<a href="#Fig_152">fig. 152</a>) from the old +world has been recorded as injurious +to man.</p> +</div></div></div></div></div></div></div></div> + +<p>gg. Genital suckers absent; integument with fine parallel lines. +Parasitic on birds and mammals.</p> + +<div class="hanging"> +<p>h. Possessing a specially developed apparatus for clinging to +hairs of mammals. <span class="rightalign"><span class="smcap"><b>Listrophoridæ.</b></span></span></p> + +<p>hh. Without such apparatus.</p> + +<div class="hanging"> +<p>i. Living on the plumage of birds. <span class="rightalign"><span class="smcap">Analgesidæ.</span></span></p> + +<p>ii. In the living tissues of birds and mammals.</p> + +<div class="hanging"> +<p>j. Vulva longitudinal. In the skin and cellular tissues of +birds. <span class="rightalign"><span class="smcap">Cytoleichidæ.</span></span></p> + +<div class="hanging"> +<p>This family contains two species, both occurring in the +United States on the common fowl. <i>Laminosioptes +cysticola</i> occurs on the skin and also bores into the +subcutaneous tissue where it gives rise to a calcareous +cyst. <i>Cytoleichus nudus</i> is most commonly +found in the air passages and air cells.</p> +</div> + +<p>jj. Vulva transverse. In the skin of mammals and birds +<span class="rightalign"><span class="smcap"><b>Sarcoptidæ</b></span></span></p> + +<div class="hanging"> +<p>k. Anal opening on the dorsum.</p> + +<div class="hanging"> +<p>l. Third pair of legs in the male without apical suckers. +On cats and rabbits. <span class="rightalign"><b>Notœdres</b> Rail.</span></p> + +<div class="hanging"> +<p>The itch mite of the cat, <b>N. cati</b> (<a href="#Fig_61">fig. 61</a>) has been +recorded on man.</p> +</div> + +<p>ll. Third leg in the male with suckers. On bats +<span class="rightalign"><i>Prosopodectes</i> Can.</span><span class="pagenum"><a name="Pg_270" id="Pg_270"></a>[Pg 270]</span></p> +</div> + +<p>kk. Anal opening below.</p> + +<div class="hanging"> +<p>l. Pedicel of the suckers jointed; mandibles styliform +and serrate near the tip. <span class="rightalign"><b>Psoroptes</b> Gerv.</span></p> + +<div class="hanging"> +<p><b>P. communis ovis</b> is the cause of sheep scab.</p> +</div> + +<p>ll. Pedicel of the suckers not jointed; mandibles +chelate.</p> + +<div class="hanging"> +<p>m. No suckers on the legs of the females; parasitic +on birds, including chickens. <i>C. mutans</i> is +itch mite of chickens. <span class="rightalign"><i>Cnemidocoptes</i> Fürst.</span></p> + +<p>mm. Suckers at least on legs I and II; parasitic on +mammals.</p> + +<div class="hanging"> +<p>n. Legs very short; in the male the hind pairs +equal in size; body usually short. +<span class="rightalign"><b>Sarcoptes</b> Latr.</span></p> + +<div class="hanging"> +<p><b>S. scabiei</b> is the itch mite of man (<a href="#Fig_56a">fig. 56</a>).</p> +</div> + +<p>nn. Legs more slender; in the male the third pair +is much larger than the fourth; body more +elongate.</p> + +<div class="hanging"> +<p>o. Female with suckers on the fourth pair of +legs. Species do not burrow in the skin, +but produce a scab similar to sheep scab. +They occur in the ox, horse, sheep and goat. +<span class="rightalign"><b>Chorioptes</b> Gerv.</span></p> + +<div class="hanging"> +<p><b>C. symbiotes bovis</b> of the ox has been +recorded a few times on man.</p> +</div> + +<p>oo. Female without suckers to the fourth legs.</p> + +<div class="hanging"> +<p>p. Hind part of the male abdomen with two +lobes. On a few wild animals. +<span class="rightalign"><i>Caparinia</i> Can.</span><span class="pagenum"><a name="Page_271" id="Page_271">[Pg 271]</a></span></p> + +<p>pp. Hind part of the male abdomen without +lobes. Live in ears of dogs and cats. +<span class="rightalign"><i>Otodectes</i> Canestr.</span></p> + +<div class="hanging"> +<p><i>O. cynotis</i> Hering (<a href="#Fig_150">fig. 150 e</a>) has been +taken in the United States.</p> +</div> +</div></div></div></div></div></div></div></div></div></div></div> + +<p>ee. Palpi usually of four or five segments, free; rarely with ventral +suckers near genital or anal openings; eyes often present; tarsi +never end in suckers; body usually divided into cephalothorax +and abdomen; rod-like epimera rarely visible; adults rarely +parasitic.</p> + +<div class="hanging"> +<p>f. Last segment of the palpi never forms a thumb to the preceding +segment; palpi simple, or rarely formed to hold prey; body +with but few hairs. <span class="rightalign"><b>EUPODOIDEA.</b></span></p> + +<div class="hanging"> +<p>g. Palpi often geniculate, or else fitted for grasping prey; mandibles +large and snout like; cephalothorax with four long +bristles above, two in front, two behind; last segment of leg I +longer than the preceding segment, often twice as long. +<span class="rightalign"><span class="smcap">Bdellidæ.</span></span></p> + +<p>gg. Palpi never geniculate (<a href="#Fig_158">fig. 158a</a>), nor fitted for grasping prey: +beak small; cephalothorax with bristles in different arrangement; +last segment of leg I shorter or but little longer than +the preceding joint; eyes when present near posterior +border. <span class="rightalign"><span class="smcap"><b>Eupodidæ</b></span></span></p> + +<div class="hanging"> +<p>Moniez has described a species from Belgium (<b>Tydeus +molestus</b>) which attacks man. It is rose colored; eyeless; +its legs are scarcely as long as its body, the hind +femur is not thickened; the mandibles are small and the +anal opening is on the venter. The female attains a +length of about 0.3 mm.</p> +</div></div> + +<p>ff. Last segment of the palpus forms a thumb to the preceding, which +ends in a claw (with few exceptions); body often with many +hairs (<a href="#Fig_158">fig. 158 k</a>). <span class="rightalign"><b>TROMBIDOIDEA.</b></span></p> + +<div class="hanging"> +<p>g. Legs I and II with processes bearing spines; skin with several +shields; coxæ contiguous. <span class="rightalign"><span class="smcap">Cæculidæ.</span></span></p> + +<p>gg. Legs I and II without such processes; few if any shields.</p> + +<div class="hanging"> +<p>h. Palpi much thickened on the base, moving laterally, last +joint often with two pectinate bristles; no eyes; legs I +ending in several long hairs; adult sometimes parasitic. +<span class="rightalign"><span class="smcap"><b>Cheyletidæ</b></span></span></p> + +<div class="hanging"> +<b>Cheyletus eruditus</b>, which frequents old books, has once +been found in pus discharged from the ear of man. +</div> + +<p>hh. Palpi less thickened, moving vertically; eyes usually present; +leg I not ending in long hairs.</p> + +<div class="hanging"> +<p>i. Coxæ contiguous, radiate; legs slender, bristly; body with +few hairs; no dorsal groove; tarsi not swollen. +<span class="rightalign"><span class="smcap">Erythræidæ.</span></span></p> + +<p>ii. Coxæ more or less in two groups; legs less bristly.<span class="pagenum"><a name="Page_273" id="Page_273">[Pg 273]</a></span></p> + +<div class="hanging"> +<p>j. Body with fewer, longer hairs; often spinning threads; +no dorsal groove; tarsi never swollen; mandibles +styliform (for piercing). <span class="rightalign"><span class="smcap"><b>Tetranychidæ</b></span></span></p> + +<div class="hanging"> +<p>The genus <b>Tetranychus</b> may be distinguished from the +other genera occurring in the United States by the +following characters: No scale-like projections on +the front of the cephalothorax; legs I as long or +longer than the body; palp ends in a distinct thumb; +the body is about 1.5 times as long as broad. <b>T. molestissimus</b> +Weyenb. from South America, and +<b>T. telarius</b> from Europe and America ordinarily +infesting plants, are said also to molest man.</p> +</div> + +<p>jj. Body with many fine hairs or short spines; not spinning +threads; often with dorsal groove; tarsi often +swollen.</p> + +<div class="hanging"> +<p>k. Mandibles styliform for piercing. <span class="rightalign"><span class="smcap">Rhycholophidæ.</span></span></p> + +<p>kk. Mandibles chelate, for biting. <span class="rightalign"><span class="smcap"><b>Trombididæ</b></span></span></p> + +<div class="hanging"> +<p>The genus <b>Trombidium</b> has recently been subdivided +by Berlese into a number of smaller +ones, of which some five or six occur in the +United States. The mature mite is not parasitic +but the larvæ which are very numerous in +certain localities will cause intense itching, +soreness, and even more serious complications. +They burrow beneath the skin and produce +inflamed spots. They have received the +popular name of "<b>red bug</b>." The names <b>Leptus +americanus</b> and <b>L. irritans</b> have been applied to +them, although they are now known to be immature +stages. (<a href="#Fig_44">Fig. 44</a>.)</p> +</div></div></div></div></div></div></div></div></div></div></div></div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_149" id="Fig_149"></a> +<a href="images/f149-full.png"><img src="images/f149.png" width="500" height="276" alt="149. Hæmaphysalis wellingtoni. Note short palpi. After Nuttall and Warburton." title="149. Hæmaphysalis wellingtoni. Note short palpi. After Nuttall and Warburton." /></a> +<span class="caption">149. Hæmaphysalis wellingtoni. Note short palpi. After Nuttall and Warburton.</span> +</div> + +<div class="figcenter" style="width: 450px;"><a name="Fig_150" id="Fig_150"></a> +<a href="images/f150-full.png"><img src="images/f150.png" width="450" height="653" alt="150. Stigmal plate of Dermacentor andersoni; (a) of +male, (b) of female. After Stiles. (c) Dermacentor variabilis, male; +(d) Glyciphagus obesus; (e) Otodectes cynotis; (f) Tyroglyphus +lintneri; (g) Tarsonemus pallidus; (h) anal plate and mandible of +Liponyssus; (c) to (h) after Banks." title="150. Stigmal plate of Dermacentor andersoni; (a) of +male, (b) of female. After Stiles. (c) Dermacentor variabilis, male; +(d) Glyciphagus obesus; (e) Otodectes cynotis; (f) Tyroglyphus +lintneri; (g) Tarsonemus pallidus; (h) anal plate and mandible of +Liponyssus; (c) to (h) after Banks." /></a> +<span class="caption">150. Stigmal plate of Dermacentor andersoni; (<i>a</i>) of +male, (<i>b</i>) of female. After Stiles. (<i>c</i>) Dermacentor variabilis, male; +(<i>d</i>) Glyciphagus obesus; (<i>e</i>) Otodectes cynotis; (<i>f</i>) Tyroglyphus +lintneri; (<i>g</i>) Tarsonemus pallidus; (<i>h</i>) anal plate and mandible of +Liponyssus; (<i>c</i>) to (<i>h</i>) after Banks.</span> +</div> + +<div class="figcenter" style="width: 450px;"><a name="Fig_151" id="Fig_151"></a> +<a href="images/f151-full.png"><img src="images/f151.png" width="450" height="427" alt="151. Rhipicephalus bursa, male. +After Nuttall and Warburton." title="151. Rhipicephalus bursa, male. +After Nuttall and Warburton." /></a> +<span class="caption">151. Rhipicephalus bursa, male. +After Nuttall and Warburton.</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_152" id="Fig_152"></a> +<a href="images/f152-full.png"><img src="images/f152.png" width="500" height="343" alt="152. Monieziella (Histiogaster) emtomophaga-spermatica, ventral aspect, +male and female. After Trouessart." title="152. Monieziella (Histiogaster) emtomophaga-spermatica, ventral aspect, +male and female. After Trouessart." /></a> +<span class="caption">152. Monieziella (Histiogaster) emtomophaga-spermatica, ventral aspect, +male and female. After Trouessart.</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_153" id="Fig_153"></a> +<a href="images/f153-full.png"><img src="images/f153.png" width="500" height="329" alt="153. Rhipicentor bicornis, ventral aspect, male. After Nuttall and +Warburton." title="153. Rhipicentor bicornis, ventral aspect, male. After Nuttall and +Warburton." /></a> +<span class="caption">153. Rhipicentor bicornis, ventral aspect, male. After Nuttall and +Warburton.</span> +</div> + +<div class="figcenter" style="width: 425px;"><a name="Fig_154" id="Fig_154"></a> +<a href="images/f154-full.png"><img src="images/f154.png" width="425" height="407" alt="154. Rhicephalus sanguineus, male. +After Nuttall and Warburton." title="154. Rhicephalus sanguineus, male. +After Nuttall and Warburton." /></a> +<span class="caption">154. Rhicephalus sanguineus, male. +After Nuttall and Warburton.</span> +</div> + +<div class="figcenter" style="width: 450px;"><a name="Fig_155" id="Fig_155"></a> +<a href="images/f155-full.png"><img src="images/f155.png" width="450" height="396" alt="155. Margaropus winthemi, male. After +Nuttall and Warburton." title="155. Margaropus winthemi, male. After +Nuttall and Warburton." /></a> +<span class="caption">155. Margaropus winthemi, male. After +Nuttall and Warburton.</span> +</div> + +<div class="figcenter" style="width: 200px;"><a name="Fig_156" id="Fig_156"></a> +<a href="images/f156-full.png"><img src="images/f156.png" width="200" height="294" alt="156. Margaropus winthemi, +capitulum and scutum. +After Nuttall and Warburton." title="156. Margaropus winthemi, +capitulum and scutum. +After Nuttall and Warburton." /></a> +<span class="caption">156. Margaropus winthemi, +capitulum and scutum. +After Nuttall and Warburton.</span> +</div> + +<div class="figcenter" style="width: 350px;"><a name="Fig_157" id="Fig_157"></a> +<a href="images/f157-full.png"><img src="images/f157.png" width="350" height="400" alt="157. Amblyomma, female. After Nuttall +and Warburton." title="157. Amblyomma, female. After Nuttall +and Warburton." /></a> +<span class="caption">157. Amblyomma, female. After Nuttall +and Warburton.</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_158" id="Fig_158"></a> +<a href="images/f158-full.png"><img src="images/f158.png" width="500" height="706" alt="158. (a) Tydeus, beak and leg from below; (b) Cheyletus pyriformis, beak and palpus; +(c) beak and claw of Pediculoides; (d) leg of Sarcoptes; (e) scutum of +female of Amblyomma americana; (f) leg I and tip of mandible of Histiostoma +americana; (g) Histiogaster malus, mandible and venter; (h) Aleurobius +farinæ, and leg I of male; (i) Otodectes cynotis, tip of abdomen of male, +(j) beak and anal plate of Dermanyssus gallinæ; (k) palpus of Allothrombium. +(a) to (j) after Banks." title="158. (a) Tydeus, beak and leg from below; (b) Cheyletus pyriformis, beak and palpus; +(c) beak and claw of Pediculoides; (d) leg of Sarcoptes; (e) scutum of +female of Amblyomma americana; (f) leg I and tip of mandible of Histiostoma +americana; (g) Histiogaster malus, mandible and venter; (h) Aleurobius +farinæ, and leg I of male; (i) Otodectes cynotis, tip of abdomen of male, +(j) beak and anal plate of Dermanyssus gallinæ; (k) palpus of Allothrombium. +(a) to (j) after Banks." /></a> +<span class="caption">158. (<i>a</i>) Tydeus, beak and leg from below; (<i>b</i>) Cheyletus pyriformis, beak and palpus; +(<i>c</i>) beak and claw of Pediculoides; (<i>d</i>) leg of Sarcoptes; (<i>e</i>) scutum of +female of Amblyomma americana; (<i>f</i>) leg I and tip of mandible of Histiostoma +americana; (<i>g</i>) Histiogaster malus, mandible and venter; (<i>h</i>) Aleurobius +farinæ, <span title="insert "palpus" before "and leg""><a name="AC_14" id="AC_14"></a>and leg I</span> of male; (<i>i</i>) Otodectes cynotis, tip of abdomen of male, +(<i>j</i>) beak and anal plate of Dermanyssus gallinæ; (<i>k</i>) palpus of Allothrombium. +(<i>a</i>) to (<i>j</i>) after Banks.</span> +</div> + + +<h3><a name="HEXAPODA_Insecta" id="HEXAPODA_Insecta"></a>HEXAPODA (Insecta)</h3> + +<p>The Thysanura (springtails and bristletails), the Neuropteroids +(may-flies, stone-flies, dragon-flies, caddis-flies, etc.), Mallophaga +(bird lice), Physopoda (thrips), Orthoptera (grasshoppers, crickets, +roaches), are of no special interest from our viewpoint. The remaining +orders are briefly characterized below.</p> + + +<h3>SIPHUNCULATA (<a href="#Page_275">page 275</a>)</h3> + +<p>Mouth parts suctorial; beak fleshy, not jointed; insect wingless; +parasitic upon mammals. Metamorphosis incomplete. Lice.</p> + + +<h3>HEMIPTERA (<a href="#Page_275">page 275</a>)</h3> + +<p>Mouth parts suctorial; beak or the sheath of the beak jointed; +in the mature state usually with four wings. In external appearance<span class="pagenum"><a name="Page_274" id="Page_274">[Pg 274]</a></span> +the immature insect resembles the adult except that the immature +form (i.e. nymph) never has wings, the successive instars during +the process of growth, therefore, are quite similar; and the metamorphosis +is thus incomplete. To this order belong the true bugs, +the plant lice, leaf hoppers, frog hoppers, cicadas, etc.</p> + + +<h3>LEPIDOPTERA</h3> + +<p>The adult insect has the body covered with scales and (with the +rare exception of the females of a few species) with four wings also +covered with scales. Proboscis, when present, coiled, not segmented, +adapted for sucking. Metamorphosis complete, i.e. the +young which hatches from the egg is quite unlike the adult, and after +undergoing several molts transforms into a quiescent pupa which is +frequently enclosed in a cocoon from which the adult later emerges. +The larvæ are known as caterpillars. Butterflies and moths.</p> + + +<h3>DIPTERA (<a href="#Page_285">page 285</a>)</h3> + +<p>The adult insect is provided with two, usually transparent, +wings, the second pair of wings of other insects being replaced by a +pair of halteres or balancers. In a few rare species the wings, or +halteres, or both, are wanting. The mouth parts, which are not +segmented, are adapted for sucking. The tarsi are five-segmented. +Metamorphosis complete. The larvæ, which are never provided +with jointed legs, are variously known as maggots, or grubs, or +wrigglers. Flies, midges, mosquitoes.</p> + + +<h3>SIPHONAPTERA (<a href="#Page_316">page 316</a>)</h3> + +<p>Mouth parts adapted for sucking; body naked or with bristles +and spines; prothorax well developed; body compressed; tarsi +with five segments; wings absent. Metamorphosis complete. +The larva is a wormlike creature. Fleas.</p> + + +<h3>COLEOPTERA</h3> + +<p>Adult with four wings (rarely wanting), the first pair horny or +leathery, veinless, forming wing covers which meet in a line along +the middle of the back. Mouth parts of both immature stages and +adults adapted for biting and chewing. Metamorphosis complete. +The larvæ of many species are known as grubs. Beetles.<span class="pagenum"><a name="Page_275" id="Page_275">[Pg 275]</a></span></p> + + +<h3>HYMENOPTERA</h3> + +<p>Adult insect with four, usually transparent, wings, wanting in +some species. Mouth parts adapted for biting and sucking; palpi +small; tarsi four or five-segmented. Metamorphosis complete. +Parasitic four-winged flies, ants, bees, and wasps.</p> + + +<h3><a name="SIPHUNCULATA_AND_HEMIPTERA" id="SIPHUNCULATA_AND_HEMIPTERA"></a>SIPHUNCULATA AND HEMIPTERA</h3> + +<div class="hanging"> +<p>a. Legs with claws fitted for clinging to hairs; wings wanting; spiracles of the +abdomen on the dorsal surface. (= <b>ANOPLURA</b> = <b>PARASITICA</b>) +<span class="rightalign"><b>SIPHUNCULATA.</b></span></p> + +<div class="hanging"> +<p>b. Legs not modified into clinging hooks; tibia and tarsus very long and +slender; tibia without thumb-like process; antennæ five-segmented. +<span class="rightalign"><span class="smcap">Hæmatomyzidæ</span> Endr.</span></p> + +<div class="hanging"> +<p><i>Hæmatomyzus elephantis</i> on the elephant.</p> +</div> + +<p>bb. Legs modified into clinging hooks; tibia and tarsus usually short and +stout; tibia with a thumb-like process; head not anteriorly prolonged, +tube-like.</p> + +<div class="hanging"> +<p>c. Body depressed; a pair of stigmata on the mesothorax, and abdominal +segments three to eight; antennæ three to five-segmented.</p> + +<div class="hanging"> +<p>d. Eyes large, projecting, distinctly pigmented; pharynx short and +broad; fulturæ (inner skeleton of head) very strong and broad, +with broad arms; proboscis short, scarcely attaining the thorax. +<span class="rightalign"><span class="smcap"><b>Pediculidæ</b></span></span></p> + +<div class="hanging"> +<p>e. Antennæ three-segmented. A few species occurring upon old +world monkeys. <span class="rightalign"><i>Pedicinis</i> Gerv.</span></p> + +<p>ee. Antennæ five-segmented.</p> + +<div class="hanging"> +<p>f. All legs stout; thumb-like process of the tibia very long and +slender, beset with strong spines, fore legs stouter than the +others; abdomen elongate, segments without lateral processes; +the divided telson with a conical process posteriorly +upon the ventral side. <span class="rightalign"><b>Pediculus</b> L.</span></p> + +<div class="hanging"> +<p>g. Upon man.</p> + +<div class="hanging"> +<p>h. Each abdominal segment dorsally with from one to three +more or less regular transverse rows of small setæ; +antenna about as long as the width of the head. Head +louse (<a href="#Fig_65">fig. 65</a>). <span class="rightalign"><b>P. humanus.</b></span></p> + +<p>hh. "No transverse rows of abdominal setæ; antenna longer +than the width of the head; species larger." Piaget. +Body louse of man. <span class="rightalign"><b>P. corporis.</b></span></p> +</div> + +<p>gg. Upon apes and other mammals. <span class="rightalign"><i>P. pusitatus</i> (?).</span></p> +</div> + +<p>ff. Fore legs delicate, with very long and slender claws; other legs +very stout with short and stout claws; thumb-like process of +the tibia short and stout; abdomen very short and broad; +segment one to five closely crowded, thus the stigmata of segments +three to five apparently lying in one segment; segments +five to eight with lateral processes; telson without lateral +conical appendages (<a href="#Fig_69">fig. 69</a>). Crab louse of man. +<span class="rightalign"><b>Phthirus pubis.</b></span><span class="pagenum"><a name="Page_276" id="Page_276">[Pg 276]</a></span></p> +</div></div> + +<p>dd. Eyes indistinct or wanting; pharynx long and slender, fulturæ very +slender and closely applied to the pharynx; proboscis very long. +Several genera found upon various mammals. <span class="rightalign"><span class="smcap">Hæmatopinidæ.</span></span></p> +</div> + +<p>cc. Body swollen; meso- and metathorax, and abdominal segments two to +eight each with a pair of stigmata; eyes wanting; antennæ four or +five-segmented; body covered with stout spines. Three genera found +upon marine mammals. <span class="rightalign"><span class="smcap">Echinophthiriidæ</span></span></p> +</div></div> + +<p>aa. Legs fitted for walking or jumping; spiracles of abdomen usually ventral; +beak segmented.</p> + +<div class="hanging"> +<p>b. Apex of head usually directed anteriorly; beak arising from its apex; sides +of the face remote from the front coxæ; first pair of wings when present +thickened at base, with thinner margins. <span class="rightalign"><b>HETEROPTERA</b></span></p> + +<div class="hanging"> +<p>c. Front tarsi of one segment, spade-form (palæformes); beak short, at +most two-segmented; intermediate legs long, slender; posterior pair +adapted for swimming. <span class="rightalign"><span class="smcap">Corixidæ</span></span></p> + +<p>cc. Front tarsi rarely one-segmented, never spade-form; beak free, at least +three-segmented.</p> + +<div class="hanging"> +<p>d. Pulvilli wanting.</p> + +<div class="hanging"> +<p>e. Hemelytra usually with a distinct clavus (<a href="#Fig_159">fig. 159</a>), clavus always +ends behind the apex of the scutellum, forming the commissure. +(Species having the wings much reduced or wanting should be +sought for in both sections.)</p> + +<div class="hanging"> +<p>f. Antennæ very short; meso- and metasternum composite; eyes +always present.<span class="pagenum"><a name="Page_277" id="Page_277">[Pg 277]</a></span></p> + +<div class="hanging"> +<p>g. Ocelli present; beak four-segmented. <span class="smcap">Ochteridæ</span> and +<span class="smcap">Nerthridæ</span>.</p> + +<p>gg. Ocelli wanting; antennæ more or less hidden in a groove.</p> + +<div class="hanging"> +<p>h. Anterior coxæ inserted at or near anterior margin of the +prosternum; front legs raptorial; beak three-segmented. +<span class="smcap"><b>Belostomidæ</b></span> (with swimming legs), <span class="smcap">Nepidæ</span>, <span class="smcap">Naucoridæ</span>.</p> + +<div class="hanging"> +<p>i. Metasternum without a median longitudinal keel; antennæ +always four-segmented.</p> + +<div class="hanging"> +<p>j. Beak short, robust, conical; the hairy fleck on the corium +elongate, large, lying in the middle between the inner +angle of the membrane and the outer vein parallel to +the membrane margin; membrane margin S-shaped.</p> + +<div class="hanging"> +<p>k. The thick fore femur with a relatively deep longitudinal +furrow to receive the tibia. Several American +species (<a href="#Fig_19">fig. 19f.</a>). <span class="rightalign"><b>Belostoma</b> (= Lethocerus Mayer)</span></p> + +<p>kk. The less thickened fore femur without such a furrow. +<span class="rightalign"><b>B. griseus.</b> <b>Benacus</b> Stäl.</span></p> +</div> + +<p>jj. Beak slender, cylindrical; the hairy spot on the corium +rounded lying next to the inner angle of the membrane.</p> + +<div class="hanging"> +<p>k. Membrane large, furrow of the embolium broadened. +<i>Z. aurantiacum</i>, <i>fluminea</i>, etc. <span class="rightalign"><i>Zaitha</i></span></p> + +<p>kk. Membrane very short; furrow of embolium not +broadened. Western genus. <span class="rightalign"><i>Pedinocoris</i></span></p> +</div></div> + +<p>ii. Metasternum with a long median longitudinal keel. Southwestern +forms. <span class="rightalign"><i>Abedus ovatus</i> and <i>Deniostoma dilatato</i></span></p> +</div> + +<p>hh. Anterior coxæ inserted at the posterior margin of the +prosternum; legs natatorial. Back swimmers (<a href="#Fig_19">fig. 19 b.</a>). +<span class="rightalign"><span class="smcap"><b>Notonectidæ</b></span></span></p> + +<div class="hanging"> +<p>i. Apices of the hemelytra entire; the three pairs of legs +similar in shape; beak three-segmented; abdomen not +keeled or hairy. <span class="rightalign"><i>Plea</i> Leach</span></p> + +<p>ii. Apices of hemelytra notched; legs dissimilar; beak four-segmented; +abdomen keeled and hairy.</p> + +<div class="hanging"> +<p>j. Hemelytra usually much longer than the abdomen; +fourth segment of the antenna longer than the third +segment; hind tarsi with claws. <span class="rightalign"><i>Bueno</i> Kirk.</span></p> + +<p>jj. Hemelytra but little longer than the abdomen; fourth +segment of the antenna shorter than the third segment; +hind tarsi without claws (<a href="#Fig_19">fig. 19b</a>). <span class="rightalign"><b>Notonecta L.</b></span></p> +</div></div></div></div> + +<p>ff. Antennæ longer than the head; or if shorter, then the eyes and +ocelli absent.</p> + +<div class="hanging"> +<p>g. Eyes, ocelli, and scutellum wanting; beak three-segmented; +head short; hemelytra always short; membrane wanting. +Insects parasitic on bats. <span class="rightalign"><span class="smcap">Polyctenidæ</span></span></p> + +<p>gg. Eyes present.</p> + +<div class="hanging"> +<p>h. First two antennal segments very short, last two long, pilose, +third thickened at the base; ocelli present, veins of the +hemelytra forming cells. <span class="smcap">Dipsocoridæ</span> (= <span class="smcap">Ceratocombidæ</span>) +including <span class="smcap">Schizopteridæ</span>.<span class="pagenum"><a name="Page_278" id="Page_278">[Pg 278]</a></span></p> + +<p>hh. Third segment of the antenna not thickened at the base, +second as long or longer than the third, rarely shorter.</p> + +<div class="hanging"> +<p>i. Posterior coxæ hinged (cardinate), if rarely rotating, the +cuneus is severed, the membrane is one or two-celled, +and the meso- and metasternum are composite.</p> + +<div class="hanging"> +<p>j. Ocelli absent, clypeus dilated toward the apex; hemelytra +always short, membrane wanting. Species parasitic. +Bed bugs, etc. <span class="rightalign"><span class="smcap"><b>Cimicidæ</b></span></span></p> + +<div class="hanging"> +<p>k. Beak short, reaching to about the anterior coxæ; +scutellum acuminate at the apex; lateral margin of +the elytra but little reflexed, apical margin more or +less rounded; intermediate and posterior coxæ +very remote.</p> + +<div class="hanging"> +<p>l. Body covered with short hairs, only the sides of the +pronotum and the hemelytra fringed with longer +hairs; antennæ with the third and fourth segments +very much more slender than the first and +second; pronotum with the anterior margin very +<i>deeply sinuate</i>. <span class="rightalign"><b>Cimex</b> L.</span></p> + +<div class="hanging"> +<p>m. Sides of the pronotum widely dilated, broader +than the breadth of one eye, and densely +fringed with backward curved hairs; apical +margin of the hemelytra nearly straight, rounded +toward the interior or exterior angles.</p> + +<div class="hanging"> +<p>n. Body covered with very short hairs; second +segment of the antenna shorter than the third; +sides of the pronotum feebly reflexed, fringed +with shorter hairs than the breadth of one +eye; hemelytra with the commissural (inner) +margin rounded and shorter than the scutellum, +apical margin rounded towards the +interior angle. The common bed bug (<a href="#Fig_19">fig. 19h</a>). +<span class="rightalign"><b>C. lectularius</b> Linn</span></p> + +<p>nn. Body covered with longer hairs; second and +third segments of the antenna of equal +length; side of the pronotum narrowly, but +distinctly, reflexed, fringed with longer +hairs than the breadth of one eye; hemelytra +with the commissural margin straight and +longer than the scutellum, apical margin +rounded towards the exterior angle. Species +found on bats in various parts of the United +States. <span class="rightalign"><i>C. pillosellus</i> Hov.</span></p> +</div> + +<p>mm. Sides of the pronotum neither dilated, nor +reflexed, fringed with less dense and nearly +straight hairs; hemelytra with the apical +margin distinctly rounded. Parasitic on +man, birds and bats. Occurs in the old +world, Brazil and the West Indies. +<span class="rightalign"><b>C. hemipterus</b> Fabr. (= rotundatus)</span><span class="pagenum"><a name="Page_279" id="Page_279">[Pg 279]</a></span></p> +</div> + +<p>ll. Body clothed with rather longer silky hairs; third +and fourth segments of the antenna somewhat +more slender than the first and second; anterior +margin of the pronotum <i>very slightly sinuate</i> or +nearly straight in the middle, produced at the +lateral angles. This is the species which in American +collections is known as <i>C. hirundinis</i>, the +latter being an old world form. It is found in +swallows nests. <b>O. vicarius</b>. <span class="rightalign"><b>Oeciacus</b> Stäl</span></p> +</div> + +<p>kk. Beak long, reaching to the posterior coxæ; scutellum +rounded at the apex; lateral margins of the elytra +strongly reflexed, apical margin slightly sinuate +toward the middle; intermediate and posterior +coxæ sub-contiguous. This species infests poultry +in southwest United States and in Mexico. <b>H. +inodorus.</b> <span class="rightalign"><b>Hæmatosiphon</b> Champ.</span></p> +</div> + +<p>jj. Ocelli present, if rarely absent in the female, then the +tarsus has two segments; or if with three tarsal segments, +the wing membrane with one or two cells.</p> + +<div class="hanging"> +<p>k. Beak four-segmented, or with two-segmented tarsi. +<span class="smcap">Isometopidæ</span>, <span class="smcap">Microphysidæ</span>, and some <span class="smcap">Capsidæ</span>.</p> + +<p>kk. Beak three-segmented.</p> + +<div class="hanging"> +<p>l. Hemelytra with embolium; head horizontal, more +or less conical; membrane with one to four veins, +rarely wanting. <span class="rightalign"><span class="smcap"><b>Anthocoridæ</b></span></span></p> + +<div class="hanging"> +<p>Several species of this family affecting man have +been noted, <b>Anthocoris kingi</b> and <b>congolense</b>, +from Africa and <b>Lyctocoris campestris</b> from +various parts of the world. <b>Lyctocoris fitchii</b> +Reuter (<a href="#Fig_19">fig. 19 j</a>), later considered by Reuter as +a variety of <b>L. campestris</b>, occurs in the United +States.</p> +</div> + +<p>ll. Hemelytra without embolium. Superfamily <span class="smcap">Acanthioidea</span> +(= <span class="smcap">Saldæ</span> Fieber and <span class="smcap">Leptopodæ</span> +Fieber)<span class="pagenum"><a name="Page_280" id="Page_280">[Pg 280]</a></span></p> +</div></div></div> + +<p>ii. Posterior coxæ rotating.</p> + +<div class="hanging"> +<p>j. Claws preapical; aquatic forms. <span class="rightalign"><span class="smcap">Gerridæ</span> and <span class="smcap">Veliadæ</span></span></p> + +<p>jj. Claws apical.</p> + +<div class="hanging"> +<p>k. Prosternum without stridulatory sulcus (notch for +beak).</p> + +<div class="hanging"> +<p>l. Tarsus with three segments; membrane with two or +three longitudinal cells from which veins radiate; +rarely with free longitudinal veins (Arachnocoris) +or veins nearly obsolete (Arbela); clavus and +corium coriaceous; ocelli rarely absent. <span class="rightalign"><span class="smcap"><b>Nabidæ</b></span></span></p> + +<div class="hanging"> +<p><b>Reduviolus</b> (= <b>Coriscus</b>) <b>subcoleoptratus</b> (<a href="#Fig_19">fig. 19 g</a>), +a species belonging to this family, occurring in +the United States, has been accused of biting +man. This insect is flat, of a jet black color, +bordered with yellow on the sides of the abdomen, +and with yellowish legs. It is predaceous, +feeding on other insects.</p> +</div> + +<p>ll. With other combinations of characters. <span class="smcap">Hydrometridæ</span>, +<span class="smcap">Henicocephalidæ</span>, <span class="smcap">Næogeidæ</span>, <span class="smcap">Mesoveliadæ</span>, +<span class="smcap">Joppeicidæ</span></p> +</div> + +<p>kk. Prosternum with stridulatory sulcus (notch for beak); +with three segments, short, strong.</p> + +<p>l. Antennæ filiform or sometimes more slender apically, +geniculate; wing membrane with two or three +large basal cells; scutellum small or moderate +<span class="rightalign"><span class="smcap"><b>Reduviidæ</b></span></span></p> + +<div class="hanging"> +<p>For a key to the genera and species see <a href="#Reduviidae_of_the_United_States">next page</a>.</p> +</div> + +<p>ll. Last antennal segment clavate or fusiform; win +membrane with the veins often forked and anastomosing; +scutellum large; tarsi each with two +segments; fore legs strong. (= <span class="smcap">Phymatidæ</span>) +<span class="rightalign"><span class="smcap">Macrocephalidæ</span></span></p> +</div></div></div></div></div></div> + +<p>ee. Clavus noticeably narrowed towards the apex, never extending +beyond the scutellum, the two not meeting to form a commissure; +head horizontal, much prolonged between the antennæ, on each +side with an antennal tubercle, sometimes acute; ocelli absent; +meso- and metasternum simple; tarsi each with two segments; +body flattened (<a href="#Fig_19">fig. 19c</a>). <span class="smcap">Aradidæ</span>, including <span class="smcap">Dysodiidæ</span>.</p> +</div> + +<p>dd. Pulvilli present (absent in one Australian family <span class="smcap">Thaumatocoridæ</span> +in which case there is a membranous appendage at the tip of the +tibia). <span class="smcap"><b>Capsidæ</b></span> (= <span class="smcap"><b>Miridæ</b></span>),<a name="FNanchor_G_7" id="FNanchor_G_7"></a><a href="#Footnote_G_7" class="fnanchor">[G]</a> <i>Eotrechus</i> (in family <span class="smcap">Gerridæ</span>), +<span class="smcap">Næogaidæ</span>, <span class="smcap">Tingitidæ</span>, <span class="smcap">Piesmidæ</span>, <span class="smcap">Myodochidæ</span>, <span class="smcap">Corizidæ</span>, +<span class="smcap">Coreidæ</span>, <span class="smcap">Alydidæ</span>, <span class="smcap">Pentatomidæ</span>, <span class="smcap">Scutelleridæ</span>, etc.</p> +</div></div> + +<p>bb. Apex of head directed ventrally, beak arising from the hinder part of the +lower side of the head; sides of face contiguous to the front coxæ; first<span class="pagenum"><a name="Page_281" id="Page_281">[Pg 281]</a></span> +pair of wings, when present, of uniform thickness. Cicadas, scale +insects, plant lice (Aphids), spittle-insects, leaf hoppers, etc. +<span class="rightalign">HOMOPTERA</span></p> +</div></div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_159" id="Fig_159"></a> +<a href="images/f159-full.png"><img src="images/f159.png" width="500" height="358" alt="159. Taxonomic details of Hemiptera-Heteroptera. (a) Dorsal aspect; (b) seta from +bedbug; (c) wing of Heteropteron; (d) leg; (e) wing of Sinea." title="159. Taxonomic details of Hemiptera-Heteroptera. (a) Dorsal aspect; (b) seta from +bedbug; (c) wing of Heteropteron; (d) leg; (e) wing of Sinea." /></a> +<span class="caption">159. Taxonomic details of Hemiptera-Heteroptera. (<i>a</i>) Dorsal aspect; (<i>b</i>) seta from +bedbug; (<i>c</i>) wing of Heteropteron; (<i>d</i>) leg; (<i>e</i>) wing of Sinea.</span> +</div> + +<div class="figcenter" style="width: 250px;"><a name="Fig_160" id="Fig_160"></a> +<a href="images/f160-full.png"><img src="images/f160.png" width="250" height="285" alt="160. Pselliopsis (Milyas) +cinctus (×2). After +C. V. Riley." title="160. Pselliopsis (Milyas) +cinctus (×2). After +C. V. Riley." /></a> +<span class="caption">160. Pselliopsis (Milyas) +cinctus (×2). After +C. V. Riley.</span> +</div> + + +<h3><a name="Reduviidae_of_the_United_States" id="Reduviidae_of_the_United_States"></a><span class="smcap">Reduviidæ of the United States</span></h3> + +<p class="center">(Adapted from a key given by Fracker).</p> + +<div class="hanging"> +<p>a. Ocelli none; wings and hemelytra always present in the adults; no <span title="for discodial read discoidal"><a name="AC_15" id="AC_15"></a>discodial</span> +areole in the corium near the apex of the clavus. <i>Orthometrops decorata</i>, +<i>Oncerotrachelus acuminatus</i>, etc., Pennsylvania and south. <span class="rightalign"><i>Sarcinæ</i></span></p> + +<p>aa. Ocelli present in the winged individuals; anterior coxæ not as long as the +femora.</p> + +<div class="hanging"> +<p>b. Hemelytra without a quadrangular or discoidal areole in the corium near +the apex of the clavus.</p> + +<div class="hanging"> +<p>c. Ocelli not farther cephalad than the caudal margins of the eyes; segment +two of the antenna single.</p> + +<div class="hanging"> +<p>d. Thorax usually constricted caudad of the middle; anterior coxæ externally +flat or concave. <span class="rightalign"><span class="smcap"><b>Piratinæ</b></span></span></p> + +<div class="hanging"> +<p>e. Middle tibiæ without spongy fossa, head long, no lateral tubercle +on neck. <i>S. stria</i>, Carolina, Ill., Cal. <span class="rightalign"><i>Sirthenia</i> Spinola</span></p> + +<p>ee. Middle tibiæ with spongy fossa; fore tibiæ convex above; neck +with a small tubercle on each side.</p> + +<div class="hanging"> +<p>f. Apical portion of anterior tibiæ angularly dilated beneath, the +spongy fossa being preceded by a small prominence. +<span class="rightalign"><b>Melanolestes</b> Stäl</span></p> + +<div class="hanging"> +<p>g. Black, with piceous legs and antennæ. N. E. States (<a href="#Fig_19">fig. 19a</a>) +<span class="rightalign"><b>M. picipes</b></span></p> + +<p>gg. Sides, and sometimes the whole dorsal surface of the abdomen +red. Ill., and southward. <span class="rightalign"><b>M. abdominalis</b></span></p> +</div> + +<p>ff. Tibiæ not dilated as in "f"; spongy fossa elongate; metapleural +sulci close to the margin. <b>R. biguttatus</b> (<a href="#Fig_22">fig. 22</a>). South. +<span class="rightalign"><b>Rasahus</b> A. and S.</span></p> +</div></div> + +<p>dd. Thorax constricted in the middle or cephalad of the middle; anterior +tarsi each three-segmented.</p> + +<div class="hanging"> +<p>e. Apex of the scutellum narrow, without spines or with a single spine +<span class="rightalign"><span class="smcap"><b>Reduviinæ</b></span></span></p> + +<div class="hanging"> +<p>f. Antennæ inserted in the lateral or dorso-lateral margins of the head; +antenniferous tubercles slightly projecting from the sides of the +head; head produced strongly cephalad; ocelli at least as far +apart as the eyes.</p> + +<div class="hanging"> +<p>g. Antennæ inserted very near the apex of the head; segments +one and three of the beak short, segment two nearly four +times as long as segment one. <b>R. prolixus.</b> W. I. +<span class="rightalign"><b>Rhodnius</b> Stäl</span></p> + +<p>gg. Antennæ inserted remote from the vertex of the head.</p> + +<div class="hanging"> +<p>h. Body slightly hairy; pronotum distinctly constricted; angles +distinct; anterior lobe four-tuberculate, with the middle +tubercles large and conical. <i>M. phyllosoma</i>, large species +<span title="insert "from" before "the""><a name="AC_16" id="AC_16"></a>the</span> southwest. <span class="rightalign"><i>Meccus</i> Stäl</span><span class="pagenum"><a name="Page_282" id="Page_282">[Pg 282]</a></span></p> + +<p>hh. Body smooth, margin of the pronotum sinuous, scarcely +constricted; anterior lobe lined with little tubercles. +<span class="rightalign"><b>Conorhinus</b> Lap.</span></p> + +<div class="hanging"> +<p>i. Surface of the pronotum and prosternum more or less +granular.</p> + +<div class="hanging"> +<p>j. Eyes small, head black; body very narrow, a fifth as +wide as long; beak reaches the middle of the prosternum. +California. <span class="rightalign"><b>C. protractus</b></span></p> + +<p>jj. Eyes large, head fuscous; body at least a fourth as wide +as long. Southern species. <span class="rightalign"><i>C. rubrofasciatus</i></span></p> +</div> + +<p>ii. Pronotum and prosternum destitute of granules.</p> + +<div class="hanging"> +<p>j. Border of abdomen entirely black except for a narrow +yellowish spot at the apex of one segment. Texas +<span class="rightalign"><i>C. gerstaeckeri</i></span></p> + +<p>jj. Border of abdomen otherwise marked.</p> + +<div class="hanging"> +<p>k. Beak slender, joints one and two slightly pilose, two +more than twice as long as one; tubercles at the +apical angles of the pronotum slightly acute, conical. +Md. to Ill. and south. The masked bed bug hunter +(<a href="#Fig_71">fig. 71</a>). <span class="rightalign"><b>C. sanguisugus</b></span></p> + +<p>kk. Beak entirely pilose, joint two a third longer than +joint one; joint one much longer than three; +tubercles at the apical angles of pronotum slightly +elevated, obtuse. Ga., Ill., Tex., Cal. <span class="rightalign"><i>C. variegatus</i></span></p> +</div></div></div></div></div> + +<p>ff. Antenna inserted on top of the head between margins, close to the +eyes; antenniferous tubercles not projecting from the side of the +head.</p> + +<div class="hanging"> +<p>g. Anterior lobe of the pronotum with a bispinous or bituberculate +disc; femora unarmed. <i>S. arizonica</i>, <i>S. bicolor</i>. Southwestern +species. <span class="rightalign"><i>Spiniger</i> Burm.</span></p> + +<p>gg. Disc of pronotum unarmed; apex of scutellum produced into +a spine; ocelli close to the eyes; eyes large and close together. +<span class="rightalign"><b>Reduvius</b> Lamarck</span></p> + +<div class="hanging"> +<p>h. Color piceous. Widely distributed in the United States. +(<a href="#Fig_20">Fig. 20</a>). <span class="rightalign"><b>R. personatus</b></span></p> + +<p>hh. More or less testaceous in color. Southwestern states +<span class="rightalign"><b>R. senilis</b></span></p> +</div></div></div> + +<p>ee. Apex of scutellum broad, with two or three spines. <span class="rightalign"><span class="smcap">Ectrichodiinæ</span></span></p> + +<div class="hanging"> +<p>f. First segment of the antenna about as long as the head. <span class="rightalign"><i>E. cruciata</i></span></p> + +<div class="hanging"> +<p>Pa. and south; <i>E. cinctiventris</i>, Tex. and Mex. +<span class="rightalign"><i>Ectrichodia</i> L. et S.</span></p> +</div> + +<p>ff. First segment of the antennæ short. <i>P. æneo-nitens</i>. South +<span class="rightalign"><i>Pothea</i> A. et S.</span></p> +</div></div></div> + +<p>cc. Ocelli cephalad of the hind margins of the eyes; first segment of the +antennæ stout, second segment divided into many smaller segments. +South and west. <i>Homalocoris maculicollis</i>, and <i>Hammatocerus +purcis</i>. <span class="rightalign"><span class="smcap">Hammatocerinæ</span></span><span class="pagenum"><a name="Page_283" id="Page_283">[Pg 283]</a></span></p> +</div> + +<p>bb. Hemelytra with a quadrangular or discoidal areole in the corium near the +apex of the clavus (<a href="#Fig_159">fig. 159e</a>).</p> + +<div class="hanging"> +<p>c. Anal areole of the membrane not extending as far proximad as the costal +areole; basal segment of the antenna thickened, porrect; the other +segments slender, folding back beneath the head and the first segment. +<span class="rightalign"><span class="smcap">Stenopodinæ</span></span></p> + +<div class="hanging"> +<p>d. Head armed with a ramous or furcate spine below each side, caudad +of the eyes.</p> + +<div class="hanging"> +<p>e. First segment of the antenna thickened, apex produced in a spine +beyond the insertion of the second segment. Species from Va., +Ill. and south. <span class="rightalign"><i>Pnirontis</i> Stäl.</span></p> + +<p>ee. First segment of the antenna not produced beyond the insertion +of the second segment. <i>Pygolampis</i>, N. E. states and south; +<i>Gnathobleda</i>, S. W. and Mex.</p> +</div> + +<p>dd. Head unarmed below or armed with a simple spine; rarely with a +subfurcate spine at the side of the base. Carolina, Missouri and +south. <i>Stenopoda</i>, <i>Schumannia</i>, <i>Diaditus</i>, <i>Narvesus</i>, <i>Oncocephalus</i></p> +</div> + +<p>cc. Anal areole of membrane extending farther proximad than the costal +areole.</p> + +<div class="hanging"> +<p>d. Ocelli farther apart than the eyes. <i>A. crassipes</i>, widely distributed +in the United States; other species occur in the southwest. +<span class="rightalign"><i>Apiomerus</i> Hahn.</span></p> + +<p>dd. Ocelli not so far apart as the eyes. <span class="rightalign"><span class="smcap">Zelinæ</span></span></p> + +<div class="hanging"> +<p>e. Sides of mesosternum without a tubercle or fold in front.</p> + +<div class="hanging"> +<p>f. Fore femur as long as or longer than the hind femur; first segment +of the beak much shorter than the second. <i>Z. audax</i>, in the +north eastern states; other species south and west. <span class="rightalign"><i>Zelus</i> Fabr.</span></p> + +<p>ff. Fore femur shorter than the hind femur, rarely of equal length, +in this case the first segment of the beak as long or longer than +the second.</p> + +<div class="hanging"> +<p>g. First segment of the beak shorter than the second; fore femur +a little shorter than the hind femur; the first segment of the +beak distinctly longer than the head before the eyes. <i>P. +cinctus</i> a widely distributed species (<a href="#Fig_160">fig. 160</a>). <i>P. punctipes</i>, +<i>P. spinicollis</i>, Cal., Mex. <span class="rightalign">(= <i>Milyas</i>) <i>Pselliopus</i> Berg.</span></p> + +<p>gg. First segment of the beak as long or longer than the second.</p> + +<div class="hanging"> +<p>h. Pronotum armed with spines on the disc.</p> + +<div class="hanging"> +<p>i. Juga distinctly prominent at the apex and often acute or +subacute; fore femur distinctly thickened; hemelytra +usually not reaching the apex of the abdomen. <i>Fitchia +aptera</i>, N. Y., south and west; <i>F. spinosula</i>, South; +<i>Rocconata annulicornis</i>, Texas, etc.</p> + +<p>ii. Juga when prominent, obtuse at apex; eyes full width of +the head; fore femur not thickened; pronotum with four +spines on posterior lobe. <i>R. taurus</i>, Pa., south and west +<span class="rightalign"><i>Repipta</i> Stäl.</span></p> +</div> + +<p>hh. Pronotum unarmed on the disc.<span class="pagenum"><a name="Page_284" id="Page_284">[Pg 284]</a></span></p> + +<div class="hanging"> +<p>i. Spines on each apical angle of the penultimate abdominal +segment. <i>A. cinereus</i>, Pa., and south. <span class="rightalign"><i>Atrachelus</i> A. et S.</span></p> + +<p>ii. Apical angle of the penultimate abdominal segment unarmed. +<i>Fitchia</i> (in part); <i>Castolus ferox</i>, Arizona.</p> +</div></div></div></div> + +<p>ee. Sides of the mesosternum with a <span title="for "tubercle of" read "tubercle or""><a name="AC_17" id="AC_17"></a>tubercle of</span> fold in front at the hind +angles of the prosternum; first segment of the beak longer than +the part of the head cephalad of the eyes.</p> + +<div class="hanging"> +<p>f. Fore femur thickened, densely granulated; hind femur unarmed.</p> + +<div class="hanging"> +<p>g. Fore tibiæ each with three long spines on the ventral side. +<i>S. diadema</i> (<a href="#Fig_159">fig. 159e</a>), a widely distributed species; and +several southwestern species. <span class="rightalign"><i>Sinea</i> A. et S.</span></p> + +<p>gg. Fore tibiæ unarmed. <i>A. multispinosa</i>, widely distributed; +<i>A. tabida</i>, Cal. <span class="rightalign"><i>Acholla</i> Stäl.</span></p> +</div> + +<p>ff. Fore femur unarmed, rarely a little thickened, a little granulated.</p> + +<div class="hanging"> +<p>g. Pronotum produced caudad over the scutellum, with a high +mesal tuberculate ridge (<a href="#Fig_19">fig. 19e</a>). <b>A. cristatus.</b> N. Y. to +Cal. and south. <span class="rightalign"><b>Arilus</b> Hahn.</span></p> + +<p>gg. Caudal lobe of the pronotum six sided, neither elevated nor +produced caudad. <i>H. americanus</i>, Southwest; also several +W. I. and Mexican genera. <span class="rightalign"><b>Harpactor</b></span></p> +</div></div></div></div></div></div></div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_161" id="Fig_161"></a> +<a href="images/f161-full.png"><img src="images/f161.png" width="500" height="357" alt="161. Taxonomic details of Diptera. (a) Ventral aspect of abdomen of Cynomyia; +(b) antenna of Tabanus; (c) ventral aspect of abdomen of Chortophila; (d) +ventral aspect of abdomen of Stomoxys; (e) claw of Aedes (Culex) sylvestris, +male; (f) claw of Hippoboscid; (g) foot of dipterous insect showing +empodium developed pulvilliform; (h) hind tarsal segment of Simulium +vittatum, female; (i) foot of dipterous insect showing bristle-like empodium." title="161. Taxonomic details of Diptera. (a) Ventral aspect of abdomen of Cynomyia; +(b) antenna of Tabanus; (c) ventral aspect of abdomen of Chortophila; (d) +ventral aspect of abdomen of Stomoxys; (e) claw of Aedes (Culex) sylvestris, +male; (f) claw of Hippoboscid; (g) foot of dipterous insect showing +empodium developed pulvilliform; (h) hind tarsal segment of Simulium +vittatum, female; (i) foot of dipterous insect showing bristle-like empodium." /></a> +<span class="caption">161. Taxonomic details of Diptera. (<i>a</i>) Ventral aspect of abdomen of Cynomyia; +(<i>b</i>) antenna of Tabanus; (<i>c</i>) ventral aspect of abdomen of Chortophila; +(<i>d</i>) ventral aspect of abdomen of Stomoxys; (<i>e</i>) claw of Aedes (Culex) sylvestris, +male; (<i>f</i>) claw of Hippoboscid; (<i>g</i>) foot of dipterous insect showing +empodium developed pulvilliform; (<i>h</i>) hind tarsal segment of Simulium +vittatum, female; (<i>i</i>) foot of dipterous insect showing bristle-like empodium.</span> +<p><span class="pagenum"><a name="Page_285" id="Page_285">[Pg 285]</a></span></p> +</div> + + +<h3><a name="DIPTERA_Mosquitoes_Midges_Flies" id="DIPTERA_Mosquitoes_Midges_Flies"></a>DIPTERA (Mosquitoes, Midges, Flies)</h3> + +<div class="hanging"> +<p>a. Integument leathery, abdominal segments indistinct; wings often wanting; +parasitic forms. <span class="rightalign"><b>PUPIPARA</b></span></p> + +<div class="hanging"> +<p>b. Head folding back on the dorsum of the thorax; wingless flies parasitic +on bats. Genus <i>Nycteribia</i>. <span class="rightalign"><span class="smcap">Nycteribiidæ</span></span></p> + +<p>bb. Head not folding back upon the dorsum of the thorax; flies either winged +or wingless; parasitic on birds and on bats and other mammals.</p> + +<div class="hanging"> +<p>c. Antennæ reduced, wings when present, with distinct parallel veins and +outer crossveins; claws simple; palpi leaf-like, projecting in front of +the head. Flies chiefly found on bats. Several genera occur in North +America. <span class="rightalign"><span class="smcap">Streblidæ</span></span></p> + +<p>cc. Antennæ more elongate, segments more or less distinctly separated; +head sunk into an emargination of the thorax; wings when present +with the veins crowded toward the anterior margin; palpi not leaf-like. +<span class="rightalign"><span class="smcap"><b>Hippoboscidæ</b></span></span></p> + +<div class="hanging"> +<p>d. Wings absent or reduced and not adapted for flight.</p> + +<div class="hanging"> +<p>e. Wings and halteres (balancers) absent. <i>M. ovinus</i>, the sheep tick. +<span class="rightalign"><i>Melophagus</i> Latr.</span></p> + +<p>ee. Wing reduced (or cast off), halteres present.</p> + +<div class="hanging"> +<p>f. Claw bidentate; ocelli present. On deer after the wings are cast +off. <i>L. depressa.</i> <span class="rightalign"><i>Lipoptena</i> Nitsch</span></p> + +<p>ff. Claw tridentate (<a href="#Fig_161">fig. 161 f</a>). On <i>Macropis</i>. <i>B. femorata.</i> +<span class="rightalign"><i>Brachypteromyia</i> Will.</span></p> +</div></div> + +<p>dd. Wings present and adapted for flight.</p> + +<div class="hanging"> +<p>e. Claws bidentate.</p> + +<div class="hanging"> +<p>f. Ocelli present; head flat; wings frequently cast off. On birds +before casting of the wing. <span class="rightalign"><i>Lipoptena</i> Nitsch.</span></p> + +<p>ff. Ocelli absent; head round; wings present. The horse tick +<b>H. equina</b> may attack man (<a href="#Fig_162">fig. 162</a>). <span class="rightalign"><b>Hippobosca</b> L.</span></p> +</div> + +<p>ee. Claws tridentate (<a href="#Fig_161">fig. 161 f.</a>).</p> + +<div class="hanging"> +<p>f. Anal cell closed at apical margin by the anal crossvein.</p> + +<div class="hanging"> +<p>g. Ocelli absent. <span class="rightalign"><i>Stilbometopa</i> Coq.</span></p> + +<p>gg. Ocelli present.<span class="pagenum"><a name="Page_286" id="Page_286">[Pg 286]</a></span></p> + +<div class="hanging"> +<p>h. R<sub>4+5</sub> does not form an angle at the crossvein. On birds. +There is a record of one species of this genus attacking man. +<span class="rightalign"><b>Ornithomyia</b> Latr.</span></p> + +<p>hh. R<sub>4+5</sub> makes an angle at the crossvein. <i>O. confluens.</i> +<span class="rightalign"><i>Ornithoica</i> Rdi.</span></p> +</div></div> + +<p>ff. Anal cell not closed by an anal crossvein. <i>Lynchia</i>, <i>Pseudolfersia</i>, +and <i>Olfersia</i> are chiefly bird parasites. The first mentioned +genus is said to be the intermediate host of <i>Hæmoproteus columbæ</i>.</p> +</div></div></div></div></div> + +<p>aa. Abdominal segments chitinous; not parasitic in the adult stage.</p> + +<div class="hanging"> +<p>b. Antennæ with six or more segments and empodium not developed pulvilliform; +palpi often with four segments.</p> + +<div class="hanging"> +<p>c. Ocelli present. <span class="smcap"><b>Blepharoceridæ</b></span>, <span class="smcap">Rhyphidæ</span>, <span class="smcap">Bibionidæ</span>, <span class="smcap">Mycetophilidæ</span>, +besides some isolated genera of other families.</p> + +<p>cc. Ocelli absent.</p> + +<div class="hanging"> +<p>d. Dorsum of the thorax with a V-shaped suture; wings usually with +numerous veins; legs often very long and slender. Crane flies. +<span class="rightalign"><span class="smcap">Tipulidæ</span></span></p> + +<p>dd. Dorsum of the thorax without a V-shaped suture.</p> + +<div class="hanging"> +<p>e. Not more than four longitudinal veins ending in the wing margin; +wing usually hairy: antennæ slender; coxæ not long; tibiæ: without +spurs, legs long and slender. Small, delicate flies often called +Gall gnats. <span class="rightalign"><span class="smcap">Cecidomyiidæ.</span></span></p> + +<p>ee. More than four longitudinal veins ending in the wing margin.</p> + +<div class="hanging"> +<p>f. The costal vein is not produced beyond the tip of the wing; radius +with not more than three branches.</p> + +<div class="hanging"> +<p>g. Antennæ short, composed of ten or eleven closely united segments; +legs stout; body stout; abdomen oval; anterior +veins stout, posterior ones weak (<a href="#Fig_163">fig. 163 b</a>); eyes of the male +contiguous over the antennæ. Black flies, buffalo flies, +turkey gnats. Many North American species, several of +them notorious for their blood sucking propensities. +<span class="rightalign"><span class="smcap"><b>Simuliidæ</b></span></span></p> + +<div class="hanging"> +<p>h. Second joint of the hind tarsus with basal scale-like process and +dorsal excision (<a href="#Fig_161">fig. 161 h</a>); radial sector not forked; no +small cell at the base of the wing. <i>S. forbesi</i>, <i>jenningsi</i>, +<i>johannseni</i>, <i>meridionale</i>, <i>piscicidium</i>, <b>venustum</b>, <b>vittatum</b>, +etc. Widely distributed species. +<span class="rightalign">(= <b>Eusimulium</b>) <b>Simulium</b> Latr.</span></p> + +<p>hh. No basal scale-like process on the second joint of the hind +tarsus; radial sector usually forked (<a href="#Fig_163">fig. 163 b</a>).</p> + +<div class="hanging"> +<p>i. Face broad, small basal cell of the wing present. <i>P. fulvum</i>, +<b>hirtipes</b>, <i>mutatum</i>, <b>pecuarum</b>, <i>pleurale</i>. <span class="rightalign"><b>Prosimulium</b> Roub.</span></p> + +<p>ii. Face linear; small basal cell of the wing absent. One +species, <i>P. furcatum</i>, from California. +<span class="rightalign"><i>Parasimulium</i> Malloch</span></p> +</div></div> + +<p>gg. Flies of a different structure.</p> + +<div class="hanging"> +<p>h. Antennæ composed of apparently two segments and a terminal +arista formed of a number of closely united segments. +Rare flies with aquatic larvæ. <span class="rightalign"><span class="smcap">Orphnephilidæ</span></span><span class="pagenum"><a name="Page_287" id="Page_287">[Pg 287]</a></span></p> + +<p>hh. Antennæ of six to fifteen segments, those of the male usually +plumose; legs frequently slender and wings narrow +<span class="rightalign"><span class="smcap"><b>Chironomidæ</b></span></span></p> + +<div class="hanging"> +<p>i. Media forked (except in the European genus <i>Brachypogon</i>); +thorax without longitudinal fissure and not produced over +the head (except in four exotic genera); antennæ usually +fourteen-jointed in both sexes; fore tibia with a simple +comb of setulæ, hind tibia with two unequal combs, +middle tibia without comb. <span class="rightalign"><span class="smcap"><b>Ceratopogoninæ</b></span></span></p> + +<div class="hanging"> +<p>j. Thorax produced cap-like over the head, wing narrow +and very long. <i>Jenkinsia</i>, <i>Macroptilum</i> and <i>Calyptopogon</i>, +eastern hemisphere; <i>Paryphoconus</i>, Brazil.</p> + +<p>jj. Thorax not produced over the head.</p> + +<div class="hanging"> +<p>k. Eyes pubescent, empodium well developed, or if short +then R<sub>2+3</sub> distinct and crossvein-like or the +branches of R coalescent; r-m crossvein present; +fore femora not thickened; wing either with appressed +hairs or with microscopic erect setulæ. +<span class="rightalign"><i>Dasyhelea</i> Kieff.</span></p> + +<p>kk. Eyes bare, or otherwise differing from the foregoing.</p> + +<div class="hanging"> +<p>l. Empodium well developed, nearly as long as the +claws and with long hairs at the base; femora and +fifth tarsal segments unarmed, i.e. without spines +or stout setæ; fourth tarsal segment cylindrical.</p> + +<div class="hanging"> +<p>m. Wing with erect and microscopic setulæ. Widely +distributed. +<span class="rightalign">(= Atrichopogon) <i>Ceratopogon</i> Meig.</span></p> + +<p>mm. Wing with long and depressed hairs. Widely +distributed. <span class="rightalign"><i>Forcipomyia</i></span></p> + +<div class="hanging"> +<p>n. Hind metatarsus shorter or not longer than the +following (i.e. the second tarsal) segment +<span class="rightalign">Subgenus <i>Prohelea</i> Kieff</span></p> + +<p>nn. Hind metatarsus longer than the following +segment. <span class="rightalign">Subgenus <i>Forcipomyia</i> Meig.</span></p> +</div></div> + +<p>ll. Empodium short, scarcely reaching the middle of +the claws, or vestigial.</p> + +<div class="hanging"> +<p>m. R-m crossvein wanting.</p> + +<div class="hanging"> +<p>n. Palpi four segmented; inferior fork of the media +obliterated at the base. Australia. +<span class="rightalign"><i>Leptoconops</i> Skuse</span></p> + +<p>nn. Palpi three-segmented.</p> + +<div class="hanging"> +<p>o. Legs spinulose, tarsal claws of the female +with a basal tooth or strong bristle, those +of the male unequal, the anterior with a +long sinuous tooth, the posterior with a +short arcuate tooth. Italy. +<span class="rightalign"><b>Mycterotypus</b> Noé</span><span class="pagenum"><a name="Page_288" id="Page_288">[Pg 288]</a></span></p> + +<p>oo. Legs unarmed; no crossvein between the +branches of the radius (<a href="#Fig_163">fig. 163e</a>). New +Mexico. <span class="rightalign"><b>Tersesthes</b> Townsend</span></p> +</div></div> + +<p>mm. R-m crossvein present.</p> + +<div class="hanging"> +<p>n. Fore femora very much swollen, armed with +spines below, fore tibia arcuate and applied +closely to the inferior margin of the femur.</p> + +<div class="hanging"> +<p>o. R<sub>2+3</sub> present, therefore cell R<sub>1</sub> and R<sub>2</sub> both +present; wing usually fasciate. United +States. <span class="rightalign"><i>Heteromyia</i> Say.</span></p> + +<p>oo. R<sub>2+3</sub> not distinct from R<sub>4+5</sub>, hence cell +R<sub>3</sub> obliterated. South America. +<span class="rightalign"><i>Pachyleptus</i> Arrib. (Walker)</span></p> +</div> + +<p>nn. Fore femur not distinctly swollen.</p> + +<div class="hanging"> +<p>o. R<sub>2+3</sub> present therefore cells R<sub>1</sub> and R<sub>3</sub> +both present, or if not, then the branches +of the radius more or less coalescent, +obliterating the cells.</p> + +<div class="hanging"> +<p>p. At least the tip of the wing with erect +setulæ; tip of R<sub>4+5</sub> scarcely attaining +the middle of the wing, empodium rather +indistinct, not reaching the middle of the +claws, the claws not toothed, equal, with +long basal bristle; legs without stout +setæ. Widely distributed. +<span class="rightalign"><b>Culicoides</b> Latr.</span></p> + +<div class="hanging"> +<p><b>Hæmatomyidium</b> and <b>Oecacta</b> are probable +synonyms of this.</p> +</div> + +<p>pp. Wings bare, if rarely with hair, then the +radius reaches beyond two-thirds the +length of the wing, or the femur or +fifth tarsal segment with stout black +spines.</p> + +<div class="hanging"> +<p>q. Media unbranched. Europe. +<span class="rightalign"><i>Brachypogon</i> Kieff</span></p> + +<p>qq. Media branched.</p> + +<div class="hanging"> +<p>r. Hind femur much swollen and spined. +America and Europe. <span class="rightalign"><i>Serromyia</i> Meg.</span></p> + +<p>rr. Hind femur not distinctly swollen.</p> + +<div class="hanging"> +<p>s. Cell R<sub>1</sub> not longer than high; fork +of the media distad of the crossvein; +wing with microscopic setulæ. +<span class="rightalign"><i>Stilobezzia</i> Kieff</span></p> + +<p>ss. Cell R<sub>1</sub> elongate.</p> + +<div class="hanging"> +<p>t. Femora unarmed. Widely distributed. +(= Sphaeromias Kieff. +1913 not Curtis?). +<span class="rightalign"><b>Johannseniella</b> Will.</span><span class="pagenum"><a name="Page_289" id="Page_289">[Pg 289]</a></span></p> + +<p>tt. Femora, at least in part, with +strong black spines. Widely +distributed. <span class="rightalign"><i>Palpomyia</i> Megerle</span></p> +</div></div></div></div></div> + +<p>oo. R<sub>2+3</sub> coalescent with R<sub>4+5</sub> hence cell R<sub>3</sub> +is obliterated.</p> + +<div class="hanging"> +<p>p. In the female the lower branch of the +media with an elbow near its base projecting +proximad, the petiole of the +media coalescent with the basal section +of the radius, wing long and narrow, +radial sector ending near the tip of the +wing; venation of the male as in <i>Bezzia</i>; +front concave. United States. +<span class="rightalign"><i>Stenoxenus</i> Coq.</span></p> + +<p>pp. Venation otherwise, front not concave.</p> + +<div class="hanging"> +<p>q. Subcosta and R<sub>1</sub> more or less coalescent +with the costa; wing pointed at the +apex, much longer than the body; +antennæ fourteen segmented, not plumose. +India. <span class="rightalign"><i>Haasiella</i> Kieff.</span></p> + +<p>qq. Subcosta and radius distinct from the +costa.</p> + +<div class="hanging"> +<p>r. Abdomen petiolate. <span class="rightalign"><i>Dibezzia</i> Kieff.</span></p> + +<p>rr. Abdomen not petiolate.</p> + +<div class="hanging"> +<p>s. Head semi-globose; hind tarsi unusually +elongate in the female; +antennæ of the male not plumose. +Europe. <span class="rightalign"><i>Macropeza</i> Meigen.</span></p> + +<p>ss. Head not globose, more or less +flattened in front; antennæ of +the male plumose. Widely distributed. +<span class="rightalign"><i>Bezzia</i> Kieff.</span></p> + +<div class="hanging"> +<p>t. Fore femora, at least, armed with +stout spines below. +<span class="rightalign">Subgenus <i>Bezzia</i> Kieff.</span></p> + +<p>tt. Femora unarmed. +<span class="rightalign">Subgenus <i>Probezzia</i> Kieff.</span></p> +</div></div></div></div></div></div></div></div></div></div></div> + +<p>ii. Media of the wing simple, and otherwise not as in "i". To +this group belong numerous Chironomid genera, none of +which are known to be noxious to man.</p> +</div></div></div> + +<p>ff. The costal vein apparently is continued around the hind margin of +the wing; radius with at least four branches.</p> + +<div class="hanging"> +<p>g. Wing ovate pointed, with numerous veins; crossveins, if evident, +before the basal third of the wing; veins very hairy; +very small moth-like flies. <span class="rightalign"><span class="smcap"><b>Psychodidæ</b></span></span></p> + +<div class="hanging"> +<p>h. With elongate biting proboscis; the petiole of the anterior +forked cell of the wing (R<sub>2</sub>) arises at or beyond the middle of +the wing (<a href="#Fig_163">fig. 163d</a>). <span class="rightalign"><b>Phlebotomus</b> Rdi.</span><span class="pagenum"><a name="Page_291" id="Page_291">[Pg 291]</a></span></p> + +<p>hh. With shorter proboscis; the petiole of the anterior forked +cell arises near the base of the wing. +<span class="rightalign"><i>Psychoda</i>, <i>Pericoma</i>, etc.</span></p> +</div> + +<p>gg. The r-m crossvein placed at or beyond the center of the wing; +wings not folded roof-like over the abdomen.</p> + +<div class="hanging"> +<p>h. Proboscis short, not adapted for piercing; wings bare (<span class="smcap">Dixidæ</span>); +or wings scaled (<span class="smcap">Culicidæ</span>, Subf. <span class="smcap">Corethrinæ</span>).</p> + +<p>hh. Proboscis elongate, adapted for piercing; wings scaled, +fringed on the hind margin; antennæ of the male bushy +plumose. Mosquitoes. +<span class="smcap"><b>Culicidæ</b></span> (exclusive of <span class="smcap">Corethrinæ</span>)</p> + +<div class="hanging"> +<p>i. Metanotum without setæ.</p> + +<div class="hanging"> +<p>j. Proboscis strongly decurved; body with broad, appressed, +metalescent scales; cell R<sub>2</sub> less than half as +long as its petiole; claws of female simple, some of the +claws of the male toothed. Several large southern +species believed to feed only on nectar of flowers +<span class="rightalign"><i>Megarhinus</i> R. D.</span></p> + +<p>jj. Proboscis straight or nearly so, or otherwise different.</p> + +<div class="hanging"> +<p>k. Scutellum evenly rounded, not lobed; claws simple in +both sexes. <span class="rightalign"><b>Anopheles</b> Meig.</span></p> + +<div class="hanging"> +<p>l. Abdomen with clusters of broad outstanding scales +along the sides; outstanding scales on the veins of +the wing rather narrow, lanceolate; upper side of +the thorax and scutellum bearing many appressed +lanceolate scales. Florida and southward (<b>Cellia</b>).</p> + +<div class="hanging"> +<p>m. Hind feet from the middle of the second segment +largely or wholly snow white.</p> + +<div class="hanging"> +<p>n. With a black band at the base of the last segment +of each hind foot. +<span class="rightalign"><b>A. albimanus</b>*<a name="FNanchor_H_8" id="FNanchor_H_8"></a><a href="#Footnote_H_8" class="fnanchor">[H]</a> and <b>tarsimaculata</b>*</span></p> + +<p>nn. Without such a band <span class="rightalign"><b>A. argyritarsis</b>*</span></p> +</div> + +<p>mm. Hind feet black, mottled with whitish and with +bands of the same color at the sutures of the +segments. W. I. <span class="rightalign"><b>A. maculipes</b></span></p> +</div> + +<p>ll. Abdomen without such a cluster of scales; outstanding +scales of the wing veins rather narrow, lanceolate; +tarsi wholly black.</p> + +<div class="hanging"> +<p>m. Deep black, thorax obscurely lined with violaceous, +especially posteriorly; head, abdomen and +legs black; no markings on the pleura; abdomen +without trace of lighter bindings; +wing scales outstanding, uniform, not forming +spots, though little thicker at the usual points +indicating the spottings. Florida. <span class="rightalign"><b>A. atropus</b></span><span class="pagenum"><a name="Page_292" id="Page_292">[Pg 292]</a></span></p> + +<p>mm. Otherwise marked when the wings are unspotted.</p> + +<div class="hanging"> +<p>n. Wings unspotted.</p> + +<div class="hanging"> +<p>o. Petiole of the first forked cell (R<sub>2</sub>) more than a +third the length of the cell. Mississippi +valley. <span class="rightalign"><b>A. walkeri</b></span></p> + +<p>oo. Petiole of the first forked cell a third the +length of the cell. Md. <span class="rightalign"><b>A. barberi</b></span></p> +</div> + +<p>nn. Wings spotted.</p> + +<div class="hanging"> +<p>o. Front margin of the wings with a patch of +whitish and yellow scales at a point about +two-thirds or three-fourths of the way from +base to apex of wing.</p> + +<div class="hanging"> +<p>p. Veins of the wing with many broad obovate +outstanding scales; thorax with a black +dot near the middle of each side. W. I. +<span class="rightalign"><b>A. grabhami</b>*</span></p> + +<p>pp. The outstanding scales of the wings rather +narrow, lanceolate.</p> + +<div class="hanging"> +<p>q. Scales of the last vein of the wings white, +those at each end black; R<sub>4+5</sub> black +scaled, the extreme apex white scaled. +Widely distributed north and south +(<a href="#Fig_131">fig. 131</a>). <span class="rightalign"><b>A. punctipennis</b></span></p> + +<div class="hanging"> +<p>A dark variety from Pennsylvania has +been named <b>A. perplexens</b>.</p> +</div> + +<p>qq. Scales of the last vein of the wing white, +those at its apex black; R<sub>4+5</sub> white +scaled and with two patches of +black scales. South and the tropics. +<b>A. franciscanus</b> and <b>pseudopunctipennis</b>*</p> +</div></div> + +<p>oo. Front margin of the wings wholly black +scaled.</p> + +<div class="hanging"> +<p>p. Last (anal) vein of the wings white scaled +with three patches of black scales (<a href="#Fig_132">fig. 132</a>). +New Jersey to Texas. <span class="rightalign"><b>A. crucians</b>*</span></p> + +<p>pp. Last vein of the wings wholly black +scaled.</p> + +<div class="hanging"> +<p>q. Widely distributed north and south +(<a href="#Fig_130">fig. 130</a>), (= <b>maculipennis</b>). +<span class="rightalign"><b>A. quadrimaculatus</b>*</span></p> + +<p>qq. Distributed from Rocky Mountains +westward. <span class="rightalign"><b>A. occidentalis</b></span></p> +</div></div></div></div></div></div> + +<p>kk. Scutellum distinctly trilobed.</p> + +<div class="hanging"> +<p>l. Cell R<sub>2</sub> less than half as long as its petiole; thorax +with metallic blue scales; median lobe of the +scutellum not tuberculate; few small species which +are not common. <span class="rightalign"><b>Uranotænia</b> Arrib.</span><span class="pagenum"><a name="Page_293" id="Page_293">[Pg 293]</a></span></p> + +<p>ll. Cell R<sub>2</sub> nearly or quite as long as its petiole, or +otherwise distinct.</p> + +<div class="hanging"> +<p>m. Femora with erect outstanding scales; occiput +broad and exposed. Large species. <b>P. ciliata.</b> +<b>P. howardi.</b> <span class="rightalign"><b>Psorophora</b> R. D.</span></p> + +<p>mm. Femora without erect scales.</p> + +<div class="hanging"> +<p>n. Clypeus bearing several scales or hairs, scutellum +with broad scales only; back of head +with broad scales; scales along the sides of the +mesonotum narrow; some or the claws +toothed; thorax marked with a pair of +silvery scaled curved stripes; legs black +with white bands at the bases of some of the +segments (<a href="#Fig_134">fig. 134</a>). Yellow Fever mosquito +<span class="rightalign"><b>Aedes</b> (= <b>Stegomyia</b>) <b>calopus</b>.</span></p> + +<p>nn. With another combination of characters. +Numerous species of mosquitoes belonging +to several closely related genera, widely +distributed over the country. (<i>Culex</i>, <i>Aedes</i>, +<i>Ochlerotatus</i>, etc.). <b>Culex</b> in the wide sense.</p> +</div></div></div></div></div> + +<p>ii. Metanotum with setæ. <i>Wyeomyia</i> (found in the United +States); and related tropic genera.</p> +</div></div></div></div></div></div></div> + +<p>bb. Antennæ composed of three segments with a differentiated style or bristle; +third segment sometimes complex or annulate, in which case the empodium +is usually developed like the pulvilli, i.e., pad-like (<a href="#Fig_161">fig. 161 g</a>).</p> + +<div class="hanging"> +<p>c. Empodium developed pad-like (pulvilliform) i.e., three nearly equal +membranous appendages on the underside of the claw (<a href="#Fig_161">fig. 161g</a>).</p> + +<div class="hanging"> +<p>d. Squamæ, head, and eyes large; occiput flattened or concave; third +segment of the antennæ with four to eight annuli or segments, +proboscis adapted for piercing; body with fine hairs, never with +bristles; middle tibia with two spurs; wing venation as figured +(<a href="#Fig_163">fig. 163f</a>); marginal vein encompasses the entire wing. Horse +flies, greenheads, deer flies, gad flies. <span class="rightalign"><span class="smcap"><b>Tabanidæ</b></span><a name="FNanchor_I_9" id="FNanchor_I_9"></a><a href="#Footnote_I_9" class="fnanchor">[I]</a></span></p> + +<div class="hanging"> +<p>e. Hind tibia with spurs at tip; ocelli usually present. (<span class="smcap"><b>Pangoninæ</b></span>)</p> + +<div class="hanging"> +<p>f. Third joint of the antennæ with seven or eight segments; proboscis +usually prolonged.</p> + +<div class="hanging"> +<p>g. Each section the third antennal segment branched. Central +American species, <i>P. festæ</i>. <span class="rightalign"><i>Pityocera</i> G. T.</span></p> + +<p>gg. Sections of the third antennal segment not branched.</p> + +<div class="hanging"> +<p>h. Upper corner of the eyes in the female terminating in an acute +angle; wings of both sexes dark anteriorly. <i>G. chrysocoma</i>, +a species from the eastern states. <span class="rightalign"><i>Goniops</i> Ald.</span></p> + +<p>hh. Upper corner of the eye in the female not so terminating; +wings nearly uniform in color, or hyaline.</p> + +<div class="hanging"> +<p>i. Proboscis scarcely extending beyond the palpi; front of the +female wide; much wider below than above. S. W. +States. <span class="rightalign"><i>Apatolestes</i> Will.</span><span class="pagenum"><a name="Page_294" id="Page_294">[Pg 294]</a></span></p> + +<p>ii. Proboscis extending beyond the palpi.</p> + +<div class="hanging"> +<p>j. Wing with cell M<sub>3</sub> closed. Tropic America. +<span class="rightalign">(= <i>Diclisa</i>) <i>Scione</i> Wlk.</span></p> + +<p>jj. Cell M<sub>3</sub> open; ocelli present or absent. Two or three +eastern species; many south and west. <span class="rightalign"><b>Pangonia</b> Rdi.</span></p> +</div></div></div></div> + +<p>ff. Third segment of the antenna with five divisions; ocelli present.</p> + +<div class="hanging"> +<p>g. First and second segments of the antenna short, the second only +half as long as the first, three western species. <span class="rightalign"><b>Silvius</b> Rdi.</span></p> + +<p>gg. First and second segments of the antenna long, the second +distinctly over half as long as the first. Deer flies. Many +species, widely distributed. <span class="rightalign"><b>Chrysops</b> Meig.</span></p> +</div></div> + +<p>ee. Hind tibia without spurs; ocelli absent.</p> + +<div class="hanging"> +<p>f. Third segment of antenna with four divisions, no tooth or angulation; +wings marked with rings and circles of darker coloring; +front of the female very wide. Widely distributed. <i>H. americana</i>, +<i>H. punctulata</i>. <span class="rightalign"><b>Hæmatopota</b> Meig.</span></p> + +<p>ff. Third segment of the antenna with five divisions (<a href="#Fig_161">fig. 161b</a>).</p> + +<div class="hanging"> +<p>g. Third segment of the antenna not furnished with a tooth or +distinct angular projection.</p> + +<div class="hanging"> +<p>h. Body covered with metallic scales; front of female of normal +width; front and middle tibiæ greatly dilated. <i>L. +lepidota</i>. <span class="rightalign"><i>Lepidoselaga</i> Macq.</span></p> + +<p>hh. Body without metallic scales; antennæ not very long, the +third segment not cylindrical, not situated on a projecting +tubercle; front of the female narrow. South. <i>D. ferrugatus.</i> +<span class="rightalign">(= <i>Diabasis</i>) <i>Diachlorus</i> O. S.</span></p> +</div> + +<p>gg. Third segment of the antenna furnished with a tooth or a +distinct angular projection.</p> + +<div class="hanging"> +<p>h. Hind tibiæ ciliate with long hairs. S. W. and tropics. +<span class="rightalign"><i>Snowiella</i> and <i>Stibasoma</i>.</span></p> + +<p>hh. Hind tibiæ not ciliate.</p> + +<div class="hanging"> +<p>i. Species of slender build, usually with a banded thorax and +abdomen; third segment of the antenna slender, the +basal prominence long; wings mostly with brownish +markings. Tropic America. <span class="rightalign"><i>Dichelacera</i> Macq.</span></p> + +<p>ii. Species of a stouter build; third segment of the antenna +stout, its basal process short (<a href="#Fig_161">fig. 161b</a>). Many species, +widely distributed <span class="rightalign"><b>Tabanus</b> L.</span></p> +</div></div></div></div></div> + +<p>dd. With another group of characters.</p> + +<div class="hanging"> +<p>e. Squamæ small, antennæ variable, thinly pilose or nearly bare species, +without distinct bristles; wing veins not crowded anteriorly, R<sub>4</sub> and +R<sub>5</sub> both present, basal cells large; middle tibiæ at least with spurs. +<span class="rightalign"><span class="smcap"><b>Leptidæ</b></span></span></p> + +<div class="hanging"> +<p>f. Flagellum of the antenna more or less elongated, composed of +numerous more or less distinct divisions. +<span class="rightalign"><span class="smcap">Xylophaginæ</span> and <span class="smcap">Arthroceratinæ</span>.</span></p> + +<p>ff. Antennæ short, third segment simple, with arista or style; face +small, proboscis short <span class="rightalign"><span class="smcap"><b>Leptinæ</b></span></span><span class="pagenum"><a name="Page_295" id="Page_295">[Pg 295]</a></span></p> + +<div class="hanging"> +<p>g. Front tibiæ each with one or two spurs, or if absent, then no +discal cell. <i>Triptotricha</i>, <i>Pheneus</i>, <i>Dialysis</i>, <i>Hilarimorpha</i>.</p> + +<p>gg. Front tibiæ without terminal spurs, discal cell present.</p> + +<div class="hanging"> +<p>h. Hind tibiæ each with a single spur.</p> + +<div class="hanging"> +<p>i. Anal cell open (<a href="#Fig_163">fig. 163g</a>); third antennal segment kidney-shaped +with dorsal or subdorsal arista; first antennal +segment elongate and thickened. About a dozen species +have been described from the United States, of which at +least one (<b>S. pachyceras</b>) is known to be a vicious blood +sucker. <span class="rightalign"><b>Symphoromyia</b> Frauenf.</span></p> + +<p>ii. Anal cell closed; third antennal segment not kidney-shaped. +<i>Chrysopila</i>, <i>Ptiolina</i>, <i>Spania</i>.</p> +</div> + +<p>hh. Hind tibiæ each with two spurs.</p> + +<div class="hanging"> +<p>i. Third segment kidney-shaped, the arista subdorsal; anal +cell closed. <span class="rightalign"><i>Atherix</i> Meig.</span></p> + +<p>ii. Third segment of the antenna short and with terminal +arista; anal cell open. <span class="rightalign"><i>Leptis</i> Fabr.</span></p> + +<div class="hanging"> +<p>Two European species of this genus have been accused of +blood sucking habits, but the record seems to have +been based upon error in observation.</p> +</div></div></div></div></div> + +<p>ee. With another combination of characters. +<span class="rightalign"><span class="smcap">Stratiomyiidæ</span>, <span class="smcap">Cyrtidæ</span>, etc.</span></p> +</div></div> + +<p>cc. Empodium bristlelike or absent.</p> + +<div class="hanging"> +<p>d. Antennæ apparently two segmented, with three-segmented arista, +wings (rarely wanting) with several stout veins anteriorly, the +weaker ones running obliquely across the wing (<a href="#Fig_163">fig. 163h</a>); small, +quick running, bristly, humpbacked flies. Several genera; <b>Aphiochæta</b>, +<b>Phora</b>, <b>Trineura</b>, etc. <span class="rightalign"><span class="smcap"><b>Phoridæ</b></span></span></p> + +<p>dd. Flies with other characters.</p> + +<div class="hanging"> +<p>e. No frontal lunule above the base of the antennæ; both R<sub>4</sub> and R<sub>5</sub> +often present; third segment of the antenna often with a terminal +bristle. <span class="smcap"><b>Asilidæ</b></span>, <span class="smcap">Mydaidæ</span>, <span class="smcap">Apioceridæ</span>, <span class="smcap">Therevidæ</span>, <span class="smcap">Scenopinidæ</span>, +<span class="smcap">Bombyliidæ</span>, <span class="smcap">Empididæ</span>, <span class="smcap">Dolichopodidæ</span>, <span class="smcap">Lonchopteridæ</span>.</p> + +<p>ee. A frontal lunule above the base of the antennæ; third segment of the +antenna always simple, i.e., not ringed, usually with a dorsal +arista; R<sub>4</sub> and R<sub>5</sub> coalesced into a simple vein.</p> + +<div class="hanging"> +<p>f. A spurious vein or fold between the radius and the media, rarely +absent; the cell R<sub>4+5</sub> closed at the apex by vein M<sub>1</sub>; few or no +bristles on the body, none on the head; flies frequently with +yellow markings. <b>Eristalis</b> (<a href="#Fig_163">fig. 163i</a>), <b>Helophilus</b>, and many +other genera. <span class="rightalign"><span class="smcap"><b>Syrphidæ</b></span></span></p> + +<p>ff. No spurious vein present.</p> + +<div class="hanging"> +<p>g. Body without bristles; proboscis elongate and slender, often +folding; front of both male and female broad. <span class="rightalign"><span class="smcap">Conopidæ</span></span></p> + +<p>gg. Bristles almost always present on head, thorax, abdomen and +legs.<span class="pagenum"><a name="Page_296" id="Page_296">[Pg 296]</a></span></p> + +<div class="hanging"> +<p>h. Arista terminal; hind metatarsus enlarged, sometimes ornamented, +hind tarsus more or less flattened beneath. +<span class="rightalign"><span class="smcap">Platypezidæ</span></span></p> + +<p>hh. Flies having a different combination of characters.</p> + +<div class="hanging"> +<p>i. Head large, eyes occupying nearly the entire head; cell +R<sub>4+5</sub> narrowed in the margin; small flies. <span class="rightalign"><span class="smcap">Pipunculidæ</span></span></p> + +<p>ii. Head and eyes not unusually large.</p> + +<div class="hanging"> +<p>j. Squamæ (tegulæ, or calyptræ, or alulæ) not large, often +quite small, the lower one lacking, or at most barely +projecting from below the upper one (antisquama); +front of both male and female broad, the eyes therefore +widely separated; posthumeral and intraalar macrochæta +not simultaneously present; thorax usually +without a complete transverse suture; postalar callus +usually absent; the connectiva adjoining the ventral +sclerites always visible; hypopleural macrochætæ +absent; last section of R<sub>4+5</sub> and M<sub>1+2</sub> with but few +exceptions nearly parallel; subcostal vein often wanting +or vestigial or closely approximated to R<sub>1</sub>; the latter +often short, basal cells small, the posterior ones often +indistinct or wanting; vibrissæ present or absent +<span class="rightalign"><span class="smcap"><b>Acalyptrate Muscoidea</b></span></span></p> + +<div class="hanging"> +<p>k. Subcosta present, distinctly separated from R<sub>1</sub> at the +tip; R<sub>1</sub> usually ends distad of the middle of the +wing; the small basal cells of the wing distinct.</p> + +<div class="hanging"> +<p>l. A bristle (vibrissa) on each side of the face near the +margin of the mouth. <span class="smcap"><b>Cordyluridæ</b></span>, <span class="smcap"><b>Sepsidæ</b></span>, +<span class="smcap">Phycodromidæ</span>, <span class="smcap">Heteroneuridæ</span>, <span class="smcap">Helomyzidæ</span>.</p> + +<p>ll. No vibrissæ present.</p> + +<div class="hanging"> +<p>m. Head nearly spherical, cheeks broad and retreating; +proboscis short; the cell R<sub>5</sub> closed or +narrowed in the margin; legs very long; tarsi +shorter than the tibiæ. <b>Calobata</b> and other +genera. <span class="rightalign"><span class="smcap"><b>Micropezidæ</b></span></span></p> + +<p>mm. Flies with another combination of characters. +<span class="smcap">Rhopalomeridæ</span>, <span class="smcap">Trypetidæ</span>, <span class="smcap">Ortalidæ</span>, +<span class="smcap">Sciomyzidæ</span>.</p> +</div></div> + +<p>kk. Subcosta absent or vestigial, or if present, then +apparently ending in the costa at the point where +R<sub>1</sub> joins it; R<sub>1</sub> usually ends in the costa at or before +the middle of the wing.</p> + +<div class="hanging"> +<p>l. Arista long plumose, or pectinate above; oral vibrissæ +present; anal cell complete; costa broken at +the apex of R<sub>1</sub>. <b>Drosophila</b>, <b>Phortica</b>, and other +genera. <span class="rightalign"><span class="smcap"><b>Drosophilidæ</b></span></span></p> + +<p>ll. With another combination of characters.</p> + +<div class="hanging"> +<p>m. The cell M and first M<sub>2</sub> not separated by a crossvein; +anal cell absent; front bare or only<span class="pagenum"><a name="Page_297" id="Page_297">[Pg 297]</a></span> +bristly above; usually light colored flies. +<b>Hippelates</b>, <b>Oscinus</b>, and other genera. (See +also m m m.) <span class="rightalign"><span class="smcap"><b>Oscinidæ</b></span></span></p> + +<p>mm. Cell M and cell first M<sub>2</sub> often separated by a +crossvein; anal cell present, complete, though +frequently small; scutellum without spines +or protuberances; oral vibrissæ present; +arista bare or short plumose; front bristly at +vertex only; small dark flies. <b>Piophila</b> +(<a href="#Fig_99">fig. 99</a>), <b>Sepsis</b> and other genera. <span class="rightalign"><span class="smcap"><b>Sepsidæ</b></span></span></p> + +<p>mmm. The <span class="smcap">Geomyzidæ</span>, <span class="smcap">Agromyzidæ</span>, <span class="smcap">Psilidæ</span>, +<span class="smcap">Trypetidæ</span>, <span class="smcap">Rhopalomeridæ</span>, <span class="smcap">Borboridæ</span> +and <span class="smcap">Diopsidæ</span> differ in various particulars +from either the <span class="smcap"><b>Oscinidæ</b></span> and the <span class="smcap"><b>Sepsidæ</b></span> +noted above.</p> +</div></div></div> + +<p>jj. Squamæ well developed, usually large, the lower one +frequently projecting from below the upper one; both +posthumeral and intraalar macrochætæ present; +thorax with a complete transverse suture; postalar +callus present and separated by a distinct suture from +the dorsum of the thorax; front of the female broad, +of the male frequently narrow, the eyes then nearly or +quite contiguous; the connectiva adjoining the ventral +sclerites either visible or not; hypopleural macrochætæ +present or absent; subcosta always distinct in +its whole course, R<sub>1</sub> never short. <span class="rightalign"><span class="smcap"><b>Calyptrate Muscoidea</b></span><a name="FNanchor_J_10" id="FNanchor_J_10"></a><a href="#Footnote_J_10" class="fnanchor">[J]</a></span></p> +</div> + +<p>k. Oral opening small, mouth parts usually much reduced +or vestigial. This family is undoubtedly of polyphyletic +origin but for convenience it is here considered +as a single family. <span class="rightalign"><span class="smcap"><b>Oestridæ.</b></span></span></p> + +<div class="hanging"> +<p>l. The costal vein ends at the tip of R<sub>4+5</sub>, M<sub>1+2</sub> +straight, not reaching the wing margin, hence +cell R<sub>5</sub> wide open (<a href="#Fig_163">fig. 163j</a>); squamæ small; +arista bare; ovipositor of the female elongate. +Larvæ in the alimentary canal of horses, etc. <span class="rightalign"><b>Gastrophilus</b></span></p> + +<div class="hanging"> +<p>m. Posterior crossvein (m-cu) wanting; wings +smoky or with clouds. Europe. <span class="rightalign"><b>G. pecorum</b></span></p> + +<p>mm. Posterior crossvein (m-cu) present, at least in +part.<span class="pagenum"><a name="Page_298" id="Page_298">[Pg 298]</a></span></p> + +<div class="hanging"> +<p>n. Wing hyaline with smoky median cross band, +and two or three spots; posterior trochanters +with hook in the male and a prominence in +the female. World wide distribution. <span class="rightalign"><b>G. equi.</b></span></p> + +<p>nn. Wings without spots.</p> + +<div class="hanging"> +<p>o. Posterior crossvein (m-cu) distad of the +anterior crossvein (r-m); legs, particularly +the femora, blackish brown. Europe and +North America. <span class="rightalign"><b>G. hæmorrhoidalis</b></span></p> + +<p>oo. Posterior crossvein opposite or proximad of +the anterior crossvein. Europe and North +America. <span class="rightalign"><b>G. nasalis</b></span></p> +</div></div></div> + +<p>ll. The costal vein ends at the tip of M<sub>1+2</sub>, M<sub>1+2</sub> with a +bend, the cell R<sub>5</sub> hence much narrowed in the +margin, or closed.</p> + +<div class="hanging"> +<p>m. Proboscis geniculate, inserted in a deep slit; +female without extricate ovipositor; arista +either bare or plumose; squamæ large; facial +grooves approximated below.</p> + +<div class="hanging"> +<p>n. Arista bare, short. Larvæ in rodents. Tropic +America. <i>B. princeps</i>. <span class="rightalign"><i>Bogeria</i> Austen</span></p> + +<p>nn. Arista pectinate above.</p> + +<div class="hanging"> +<p>o. Tarsi broadened and flattened, hairy, anal +lobe of the wing large. Larvæ in rodents. +A number of American species. <span class="rightalign"><i>Cuterebra.</i></span></p> + +<p>oo. Tarsi slender, not hairy; anal lobe of the +wing moderate. Larvæ in man and other +mammals. Tropic America. <b>D. cyaniventris.</b> +<span class="rightalign"><b>Dermatobia</b> Br.</span></p> +</div></div> + +<p>mm. Mouth parts very small, vestigial; arista bare.</p> + +<div class="hanging"> +<p>n. Facial grooves approximated below, leaving a +narrow median depression or groove.</p> + +<div class="hanging"> +<p>o. Cell R<sub>5</sub> closed and petiolate, body nearly +bare. Larvæ in the nasal cavities of the +smaller Ungulates. The sheep bot fly. +<b>O. ovis.</b> Widely distributed <span class="rightalign"><b>Oestrus</b> L.</span></p> + +<p>oo. Cell R<sub>5</sub> narrowly open, body hairy. Larvæ +parasitic on deer. Europe and America. <span class="rightalign"><i>Cephenomyia</i> Latr.</span></p> +</div> + +<p>nn. Facial grooves far apart, enclosing between +them a broad shield-shaped surface; squamæ +large; female with elongate ovipositor. +Larvæ hypodermatic on Ungulates. <span class="rightalign"><b>Hypoderma</b> Clark</span></p> + +<div class="hanging"> +<p>o. Palpi wanting; tibiæ thickened in the middle.</p> + +<div class="hanging"> +<p>p. Hair at apex of the abdomen yellow; legs +including femora yellowish brown. <span class="rightalign"><b>H. diana</b></span><span class="pagenum"><a name="Page_299" id="Page_299">[Pg 299]</a></span></p> + +<p>pp. Hair at the apex of the abdomen reddish +yellow. Europe and America.</p> + +<div class="hanging"> +<p>q. Tibiæ and tarsi yellow; femora black. <span class="rightalign"><b>H. lineata</b></span></p> + +<p>qq. Legs black with black hair; tips of +hind tibia and tarsi yellowish brown. <span class="rightalign"><b>H. bovis</b></span></p> +</div></div> + +<p>oo. Palpi small, globular; tibiæ cylindrical, +straight. On reindeer. <i>O. tarandi</i> +<span class="rightalign"><i>Oedemagena</i> Latr.</span></p> +</div></div></div></div> + +<p>kk. Oral opening of the usual size; mouth parts not +vestigial.</p> + +<div class="hanging"> +<p>l. Hypopleurals wanting; if three sternopleurals are +present the arrangement is 1:2; conjunctiva +(<a href="#Fig_161">fig. 161c</a>) of the venter usually present; if the +terminal section of M<sub>1+2</sub> is bent it has neither fold +nor appendage. (<span class="smcap"><b>Anthomyiidæ</b></span> of Girschner).</p> + +<div class="hanging"> +<p>m. Sternopleurals wanting; M<sub>1+2</sub> straight toward +the apex, costa ends at or slightly beyond the +tip of R<sub>4+5</sub>; mouth parts vestigial. +<span class="rightalign"><span class="smcap"><b>Gastrophilinæ.</b></span> See <span class="smcap"><b>Oestridæ</b></span></span></p> + +<p>mm. Sternopleurals present, if rarely absent then +differing in other characters.</p> + +<div class="hanging"> +<p>n. Caudal margin of the fifth ventral abdominal +sclerite of the male deeply notched on the +median line usually to beyond the middle; +abdomen often cylindrical or linear; abdomen +often with four to eight spots; eyes of the +male usually widely separated; sternopleurals +three, arranged in an equilateral +triangle; subapical seta of the hind tibia +placed very low; M<sub>1+2</sub> straight; anal vein +abbreviated; wings not rilled. <i>Cænosia</i>, +<i>Caricea</i>, <i>Dexiopsis</i>, <i>Hoplogaster</i>, <i>Schœnomyia</i>, +etc. (<span class="smcap">Cœnosinæ</span>)<a name="FNanchor_K_11" id="FNanchor_K_11"></a><a href="#Footnote_K_11" class="fnanchor">[K]</a>. +<span class="rightalign"><span class="smcap">Anthomyiidæ</span> in part</span></p> + +<p>nn. Caudal margin of the fifth ventral abdominal +sclerite of the male incurved, rarely deeply +cleft, rarely entire, in a few genera +deeply two or three notched; M<sub>1+2</sub> straight<span class="pagenum"><a name="Page_300" id="Page_300">[Pg 300]</a></span> +or curved; abdomen usually short or elongate +oval; sternopleurals, if three are present, +arranged in the order 1:2 in a right triangle. +<span class="rightalign">(<span class="smcap">Muscinæ-Anthomyiinæ</span> of Girschner)</span></p> + +<div class="hanging"> +<p>o. M<sub>1+2</sub> straight, hence cell R<sub>5</sub> not narrowed in +the margin. <span class="rightalign"><span class="smcap"><b>Anthomyiidæ</b></span> in part</span></p> + +<div class="hanging"> +<p>p. Underside of the scutellum more or less +sparsely covered with fine hairs; anal +vein nearly always reaches the hind +margin of the wing; extensor surface of +the hind tibiæ with a number of stout +setæ; squamæ often small and equal. +<b>Anthomyia</b>, <i>Chortophila</i>, <i>Eustalomyia</i>, +<i>Hammomyia</i>, <i>Hylemyia</i>, <i>Prosalpia</i>, <i>Pegomyia</i>, +etc. <span class="rightalign"><span class="smcap"><b>Hylemyinæ-Pegomyinæ</b></span></span></p> + +<p>pp. Underside of the scutellum bare; anal +vein does not reach the wing margin.</p> + +<div class="hanging"> +<p>q. First anal vein short, second anal suddenly +flexed upwards; hind tibiæ each +with one or two strong setæ on the +extensor surface. <b>Fannia</b> (= <b>Homalomyia</b>), +<i>Cœlomyia</i>, <i>Choristoma</i>, <i>Euryomma</i>, +<i>Azelia</i>, etc. <span class="rightalign"><span class="smcap"><b>Fanninæ-Azelinæ</b></span></span></p> + +<p>qq. Anal veins parallel or divergent.</p> + +<div class="hanging"> +<p>r. Setæ on the exterior surface of the hind +tibiæ wanting (except in <i>Limnaricia</i> +and <i>Cœnosites</i>), lower squama not +broadened to the margin of the +scutellum. <i>Leucomelina</i>, <i>Limnophora</i>, +<i>Limnospila</i>, <i>Lispa</i>, <i>Mydæa</i>, +<i>Spilogaster</i>, etc. <span class="rightalign"><span class="smcap">Mydæinæ-Limnophorinæ</span></span></p> + +<p>rr. One (rarely more) seta on the extensor +surface of the hind tibia; squamæ +usually large and unequal. <b>Hydrotaea</b>, +<i>Aricia</i>, <i>Drymeia</i>, <i>Ophyra</i>, +<i>Phaonia</i> (= <i>Hyetodesia</i>), <i>Pogonomyia</i>, +<i>Trichophthicus</i>, etc. <span class="rightalign"><span class="smcap"><b>Aricinæ</b></span></span></p> +</div></div></div> + +<p>oo. M<sub>1+2</sub> curved or bent, hence the cell R<sub>5</sub> more +or less narrowed in the margin. +(<span class="smcap"><b>Muscinæ</b></span>). <span class="smcap"><b>Muscidæ</b></span> in part. See +<a href="#Page_303">page 303</a> for generic synopsis.</p> +</div></div></div> + +<p>ll. Hypopleurals present; when three sternopleurals +are present the arrangement is 2:1 or 1:1:1. +<span class="rightalign">(<span class="smcap">Tachinidæ</span> of Girschner)</span></p> + +<div class="hanging"> +<p>m. Conjunctiva of the ventral sclerites of the abdomen +present, frequently well developed, +surrounding the sclerites.<span class="pagenum"><a name="Page_301" id="Page_301">[Pg 301]</a></span></p> + +<div class="hanging"> +<p>n. Mouth parts vestigial. <span class="smcap"><b>Oestridæ.</b></span> See <a href="#Page_297">page +297</a> for generic synopsis.</p> + +<p>nn. Mouth parts well developed.</p> + +<div class="hanging"> +<p>o. M<sub>1+2</sub> straight, hence cell R<sub>5</sub> wide open in +the margin; costa ending at the tip of R<sub>5</sub>; +three sternopleurals present; antennal +arista plumose. <i>Syllegoptera</i>. Europe. +(<span class="smcap">Syllegopterinæ</span>). <span class="rightalign"><span class="smcap">Dexiidæ</span> in part</span></p> + +<p>oo. M<sub>1+2</sub> bent, hence cell R<sub>5</sub> narrowed in the +margin; sternopleurals rarely wanting, +usually 1:1 or 0:1; facial plate strongly +produced below vibrissal angle like the +bridge of the nose; antennal arista bare. +Parasitic on Hemiptera and Coleoptera. +<i>Allophora</i>, <i>Cistogaster</i>, <i>Clytia</i>, <i>Phasia</i>, +etc. (<span class="smcap">Phasiinæ</span>). <span class="rightalign"><span class="smcap">Tachinidæ</span> in part.</span></p> +</div></div> + +<p>mm. Conjunctiva of the ventral sclerites invisible +(<a href="#Fig_161">fig. 161a</a>).</p> + +<div class="hanging"> +<p>n. Second ventral sclerite of the abdomen lying +with its edges either upon or in contact with +the ventral edges of the corresponding +dorsal sclerite.</p> + +<div class="hanging"> +<p>o. Outermost posthumeral almost always lower +(more ventrad) in position than the presutural +macrochæta; fifth ventral abdominal +sclerite of the male cleft beyond the +middle, often strongly developed; body +color very frequently metallic green or +blue, or yellow; arista plumose. (<span class="smcap"><b>Calliphorinæ</b></span>) +<span class="rightalign"><span class="smcap"><b>Muscidæ</b></span> in part.</span></p> + +<div class="hanging"> +<p>See <a href="#Page_303">page 303</a> for generic synopsis.</p> +</div> + +<p>oo. Outermost posthumeral macrochæta on +level or higher (more dorsad) than the +presutural macrochæta; arista bare, pubescent, +or plumose only on the basal two-thirds; +body coloring usually grayish +(<a href="#Fig_106">fig. 106</a>). <span class="rightalign"><span class="smcap"><b>Sarcophagidæ</b></span></span></p> + +<div class="hanging"> +<p>p. Fifth ventral sclerite of the male either +wanting or with the caudal margin +straight; presutural intraalar rarely +present. <span class="rightalign">(<span class="smcap"><b>Sarcophaginæ</b></span>)</span></p> + +<div class="hanging"> +<p>q. Fifth ventral abdominal sclerite of the +male much reduced, the remaining +segments with straight posterior margin, +overlapping scale-like; in the +female only segment one and two scale-like, +the others wholly or in part +covered; sternopleurals usually three +or more. <b>Sarcophaga</b> and related +genera.<span class="pagenum"><a name="Page_302" id="Page_302">[Pg 302]</a></span></p> + +<p>qq. Fifth ventral sclerite of the male plainly +visible; sternopleurals usually two. +<b>Sarcophila</b>, <b>Wohlfahrtia</b>, <i>Brachycoma</i>, +<i>Hilarella</i>, <i>Miltogramma</i>, <i>Metopia</i>, +<i>Macronychia</i>, <i>Nyctia</i>, <i>Paramacronychia</i>, +<i>Pachyphthalmus</i>, etc.</p> +</div> + +<p>pp. Fifth ventral abdominal sclerite of the +male cleft to beyond the middle; ventral +sclerites usually visible, shield-like. +<i>Rhinophora</i>, <i>Phyto</i>, <i>Melanophora</i>. +<span class="rightalign"><span class="smcap">Rhinophorinæ</span></span></p> +</div></div> + +<p>nn. Second ventral abdominal sclerite as well as +the others more or less covered, sometimes +wholly, by the edges of the dorsal sclerite.</p> + +<div class="hanging"> +<p>o. The presutural intraalar wanting; ventral +sclerites two to five nearly or quite covered +by the edges of the corresponding dorsal +sclerites; base of the antennæ usually at or +below the middle of the eye; arista usually +plumose; legs usually elongate; abdominal +segments with marginal and often +discal macrochætæ. <span class="rightalign"><span class="smcap"><b>Dexiidæ</b></span></span></p> + +<p>oo. Presutural intraalar present, if absent, then +the ventral sclerites broadly exposed +or the fifth ventral sclerite vestigial;<span class="pagenum"><a name="Page_303" id="Page_303">[Pg 303]</a></span> +base of the antennæ usually above the +middle of the eye; arista bare; at least +two posthumerals and three posterior +intraalars present. Parasitic on caterpillars, +etc. <span class="rightalign"><span class="smcap">Tachinidæ</span></span></p> +</div></div></div></div></div></div></div></div></div></div></div></div></div> + +<div class="figcenter" style="width: 450px;"><a name="Fig_162" id="Fig_162"></a> +<a href="images/f162-full.png"><img src="images/f162.png" width="450" height="285" alt="162. Hippobosca equina, ×4. After Osborn." title="162. Hippobosca equina, ×4. After Osborn." /></a> +<span class="caption">162. Hippobosca equina, ×4. After Osborn.</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_163" id="Fig_163"></a> +<a href="images/f163-full.png"><img src="images/f163.png" width="500" height="707" alt="163. Wings of Diptera. (a) Anopheles; (b) Prosimulium; (c) Johannseniella; (d) Phlebotomus +(After Doerr and Russ); (e) Tersesthes (after Townsend); (f) Tabanus; +(g) Symphoromyia; (h) Aphiochæta; (i) Eristalis; (j) Gastrophilus; +(k) Fannia; (l) Musca." title="163. Wings of Diptera. (a) Anopheles; (b) Prosimulium; (c) Johannseniella; (d) Phlebotomus +(After Doerr and Russ); (e) Tersesthes (after Townsend); (f) Tabanus; +(g) Symphoromyia; (h) Aphiochæta; (i) Eristalis; (j) Gastrophilus; +(k) Fannia; (l) Musca." /></a> +<span class="caption">163. Wings of Diptera. (<i>a</i>) Anopheles; (<i>b</i>) Prosimulium; (<i>c</i>) Johannseniella; (<i>d</i>) Phlebotomus +(After Doerr and Russ); (<i>e</i>) Tersesthes (after Townsend); (<i>f</i>) Tabanus; +(<i>g</i>) Symphoromyia; (<i>h</i>) Aphiochæta; (<i>i</i>) Eristalis; (<i>j</i>) Gastrophilus; +(<i>k</i>) Fannia; (<i>l</i>) Musca.</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_164" id="Fig_164"></a> +<a href="images/f164-full.png"><img src="images/f164.png" width="500" height="427" alt="164. Glossina palpalis. (×4.) After Austen." title="164. Glossina palpalis. (×4.) After Austen." /></a> +<span class="caption">164. Glossina palpalis. (×4.) After Austen.</span> +</div> + + +<h3>SYNOPSIS OF THE PRINCIPAL GENERA OF THE MUSCIDÆ OF THE WORLD</h3> + +<div class="hanging"> +<p>a. Proboscis long, directed forward, adapted for piercing, or oral margin much +produced, snout-like.</p> + +<div class="hanging"> +<p>b. Oral margin produced snout-like; vibrissa placed high above the oral +margin; antennal arista either pectinate or more or less plumose.</p> + +<div class="hanging"> +<p>c. Antennal arista short or long-plumose; neither sex with distinct +orbital bristles.</p> + +<div class="hanging"> +<p>d. No facial carina between the antennæ. <span class="rightalign"><span class="smcap">Rhynchomyiinæ</span></span></p> + +<div class="hanging"> +<p>e. Arista short-plumose. <i>R. speciosa.</i> Europe. <span class="rightalign"><i>Rhynchomyia</i> R. D.</span></p> + +<p>ee. Arista long-plumose. <i>I. phasina</i>. Europe and Egypt. <span class="rightalign"><i>Idiopsis.</i> B. B.</span></p> +</div> + +<p>dd. With flattened carina, the bases of the antennæ separated; no abdominal +macrochætæ. <span class="rightalign"><span class="smcap">Cosmininæ</span></span></p> + +<div class="hanging"> +<p><i>C. fuscipennis</i>. South Africa. <span class="rightalign"><i>Cosmina</i></span></p> +</div></div> + +<p>cc. Antennal arista pectinate; bases of the antennæ separated by a flattened +carina. <span class="rightalign"><span class="smcap">Rhiniinæ</span> R. D.</span></p> + +<div class="hanging"> +<p>d. Cell R<sub>5</sub> open, or closed at the margin.</p> + +<div class="hanging"> +<p>e. Third segment of the antenna twice as long as the second; claws of +both sexes short; cell R<sub>5</sub> open. <i>I. lunata.</i> Eastern Hemisphere. +<span class="rightalign"><i>Idia</i> Meigen</span></p> + +<p>ee. Third segment of the antenna three times as long as the second; +cell R<sub>5</sub> open or closed; claws of the male long and slender, of the +female shorter than the last tarsal joint. <i>I. mandarina</i>, China. +<span class="rightalign"><i>Idiella</i> B. B.</span></p> +</div> + +<p>dd. Cell R<sub>5</sub> petiolate. <span class="rightalign"><i>Rhinia</i>; and <i>Beccarimyia</i> Rdi.</span></p> +</div></div> + +<p>bb. Proboscis long, directed forward, adapted for piercing. <span class="rightalign"><span class="smcap"><b>Stomoxinæ</b></span></span></p> + +<div class="hanging"> +<p>c. Arista flat, pectinate above with plumose rays; sternopleurals 1:2; +bases of the veins R<sub>1</sub> and R<sub>4+5</sub> without setæ; base of the media bowed +down; apical cell opens before the apex of the wing. African species +<span class="rightalign"><b>Glossina</b> Wied.</span></p> + +<div class="hanging"> +<p>d. Species measuring over twelve mm. in length. <i>G. longipennis</i> and <b>fusca</b>.</p> + +<p>dd. Species less than twelve mm. in length.</p> + +<div class="hanging"> +<p>e. All segments of the hind tarsi black.</p> + +<div class="hanging"> +<p>f. The fourth and fifth segments of the fore tarsi black; antennæ +black (<a href="#Fig_164">fig. 164</a>). <span class="rightalign"><b>G. palpalis</b> R. D.</span></p> + +<p>ff. Otherwise marked. <span class="rightalign"><i>G. bocagei</i>, <i>tachinoides</i>, <i>pallicera</i>.</span></p> +</div> + +<p>ee. First three segments of the hind tarsi are yellow, the fourth and +fifth segments are black.</p> + +<div class="hanging"> +<p>f. Fourth and fifth segments of the first and second pair of tarsi are +black.</p> + +<div class="hanging"> +<p>g. The yellow bands of the abdominal segments occupy a third of +the segment (<a href="#Fig_165">fig. 165</a>). <span class="rightalign"><b>G. morsitans</b> Westw.</span></p> + +<p>gg. The yellow band on each segment of the abdomen occupies a +sixth of the segment. <span class="rightalign"><b>G. longipalpis</b> Wied.</span><span class="pagenum"><a name="Page_304" id="Page_304">[Pg 304]</a></span></p> +</div> + +<p>ff. Tarsi of the first and second pairs of legs wholly yellow. +<span class="rightalign"><i>G. pallidipes</i> Austen</span></p> +</div></div></div> + +<p>cc. Rays of the arista not plumose; only one or two sternopleurals; base of +the media not strongly bowed down; apical cell opens at or very near +the apex of the wing.</p> + +<div class="hanging"> +<p>d. Vein R<sub>4+5</sub> without setæ at the base; palpi about as long as the proboscis.</p> + +<div class="hanging"> +<p>e. Arista pectinate (i. e. rays on one side only), the rays often undulate; +two yellow sternopleurals often difficult to detect; vein M<sub>1+2</sub> +only slightly bent, the apical cell hence wide open. The horn fly, +<b>H. irritans</b> (= <i>Lyperosia serrata</i>) and related species. Widely distributed +(figs. <a href="#Fig_167">167</a>, <a href="#Fig_168">168</a>). <span class="rightalign"><b>Hæmatobia</b> R. D. not B. B.</span></p> + +<p>ee. Arista also with rays below; vein M<sub>1+2</sub> more strongly bent, the +apical cell hence less widely open.</p> + +<div class="hanging"> +<p>f. Palpi strongly spatulate at the tips, lower rays of the arista about +six in number, <b>B. sanguinolentus</b>. South Asia. +<span class="rightalign"><b>Bdellolarynx</b> Austen</span></p> + +<p>ff. Palpi feebly spatulate; apical cell of the wing narrowly open +slightly before the tip; sternopleurals black, anterior bristle +sometimes absent. <b>H. atripalpis</b>. Europe. +<span class="rightalign"><b>Hæmatobosca</b> Bezzi</span></p> +</div></div> + +<p>dd. Vein R<sub>4+5</sub> with setæ at the base.<a name="FNanchor_L_12" id="FNanchor_L_12"></a><a href="#Footnote_L_12" class="fnanchor">[L]</a><span class="pagenum"><a name="Page_305" id="Page_305">[Pg 305]</a></span></p> + +<div class="hanging"> +<p>e. Veins R<sub>1</sub> and R<sub>4+5</sub> with setæ at the base; two equally prominent +sternopleural macrochætæ; arista with rays both above and below; +palpi as long as the proboscis; apical cell of the wing wide +open. <b>L. tibialis.</b> (<i>Hæmatobia</i> B. B. not R. D.). +<span class="rightalign"><b>Lyperosiops</b> Town.</span></p> + +<p>ee. Only vein R<sub>4+5</sub> with basal setæ; anterior sternopleural macrochæta +wanting; arista pectinate.</p> + +<div class="hanging"> +<p>f. Palpi as long as the proboscis, the latter stout, with fleshy terminal +labellæ; apical cell narrowly open; sternopleural macrochætæ +black. <b>S. maculosa</b> from Africa and related species +from Asia. <span class="rightalign"><b>Stygeromyia</b> Austen</span></p> + +<p>ff. Palpi much shorter than the proboscis, the latter pointed at the +apex, without fleshy labellæ; apical cell of the wing wide open. +<b>S. calcitrans</b>, the stable fly and related species. Widely distributed +in both hemispheres (<a href="#Fig_110">fig. 110</a>). <span class="rightalign"><b>Stomoxys</b> Geof.</span></p> +</div></div></div></div></div> + +<p>aa. Proboscis neither slender nor elongate, the labellæ fleshy and not adapted for +piercing.</p> + +<div class="hanging"> +<p>b. Hypopleuræ without a vertical row of macrochætæ. <span class="rightalign"><span class="smcap"><b>Muscinæ</b></span></span></p> + +<div class="hanging"> +<p>c. Arista bare; distal portion of <span title="for R_{4+5} read M_{1+2}"><a name="AC_18a" id="AC_18a"></a>R<sub>4+5</sub></span> broadly curved at the end; hypopleuræ +with a sparse cluster of fine hairs. <i>S. braziliana</i>, Southern +States and southward. <span class="rightalign"><i>Synthesiomyia</i> B. B.</span></p> + +<p>cc. Arista pectinate or plumose.</p> + +<div class="hanging"> +<p>d. Arista pectinate. <i>H. vittigera</i>, with the posterior half of the abdomen +metallic blue. Mexico. <span class="rightalign"><i>Hemichlora</i> V. d. W.</span></p> + +<p>dd. Arista plumose.</p> + +<div class="hanging"> +<p>e. Middle tibia with one or more prominent setæ on the inner (flexor) +surface beyond the middle, or inner surface very hairy.</p> + +<div class="hanging"> +<p>f. R<sub>1</sub> ends distad of the m-cu crossvein; <span title="for R_{4+5} read M_{1+2}"><a name="AC_18b" id="AC_18b"></a>R<sub>4+5</sub></span> with a broad curve +near its apical end. (= <i>Neomesembrina</i> Schnabl. = <i>Metamesembrina</i> Town). +<i>M. meridiana.</i> Europe. <span class="rightalign"><i>Mesembrina</i> Meigen</span></p> + +<p>ff. R<sub>1</sub> ends proximad of the m-cu crossvein.</p> + +<div class="hanging"> +<p>g. Eyes pilose, sometimes sparsely in the female.</p> + +<div class="hanging"> +<p>h. Female with two or three stout orbital setæ; the hind metatarsus +of the male thickened below at the base and penicillate. +<i>D. pratorum.</i> Europe. <span class="rightalign"><i>Dasyphora</i> R. D.<a name="FNanchor_M_13" id="FNanchor_M_13"></a><a href="#Footnote_M_13" class="fnanchor">[M]</a></span></p> + +<p>hh. Neither sex with orbital setæ.</p> + +<div class="hanging"> +<p>i. Abdomen without macrochætæ; arista plumose. <i>C. asiatica.</i> +Eastern Hemisphere. <span class="rightalign"><i>Cryptolucilia</i> B. B.</span></p> + +<p>ii. Abdomen with strong macrochætæ; arista very short-plumose, +nearly bare. <i>B. tachinina.</i> Brazil. <span class="rightalign"><i>Reinwardtia</i> B. B.</span></p> +</div></div> + +<p>gg. Eyes bare.</p> + +<div class="hanging"> +<p>h. Body densely pilose; thoracic macrochætæ wanting; middle +tibiæ much elongate and bent; last section of <span title="for R_{4+5} read M_{1+2}"><a name="AC_18c" id="AC_18c"></a>R<sub>4+5</sub></span> with a +gentle curve. H. (<i>Mesembrina</i>) <i>mystacea, et al.</i>, Europe +and <i>H. solitaria</i>, N. America. <span class="rightalign"><i>Hypodermodes</i> Town.</span></p> + +<p>hh. Body not densely pilose.<span class="pagenum"><a name="Page_306" id="Page_306">[Pg 306]</a></span></p> + +<div class="hanging"> +<p>i. Dorsocentrals six; last section of <span title="for R_{4+5} read M_{1+2}"><a name="AC_18d" id="AC_18d"></a>R<sub>4+5</sub></span> with a gentle curve.</p> + +<div class="hanging"> +<p>j. Inner dorsocentrals ("acrostichals") wanting; sternopleurals +arranged 1:3. <i>P. cyanicolor</i>, <i>cadaverina</i>, etc. +Europe and America. <span class="rightalign"><i>Pyrellia</i> R. D.</span></p> + +<p>jj. Inner dorsocentrals ("acrostichals") present; sternopleurals +arranged 1:2. E. <i>latreillii.</i> North America. +<span class="rightalign"><i>Eumesembrina</i> Town.</span></p> +</div> + +<p>ii. Dorsocentrals five; inner dorsocentrals present; last +section of <span title="for R_{4+5} read M_{1+2}"><a name="AC_18e" id="AC_18e"></a>R<sub>4+5</sub></span> with a rounded angle; sternopleurals +arranged 1:2. <i>P. cornicina</i> Europe and America. +(<i>Pseudopyrellia</i> Girsch.) <span class="rightalign"><i>Orthellia</i> R. D.</span></p> +</div></div></div></div> + +<p>ee. Middle tibia without a prominent bristle on the inner surface beyond +the middle.</p> + +<div class="hanging"> +<p>f. Squamula thoracalis broadened mesad and caudad as far as the +scutellum.</p> + +<div class="hanging"> +<p>g. Sternopleural macrochætæ arranged in an equilateral triangle; +front of both sexes broad; genæ bare; dorsocentrals six, +small; wing not rilled. (To <span class="smcap">Coenosinæ</span>). <span class="rightalign"><i>Atherigona</i> Rdi.</span></p> + +<p>gg. Sternopleural macrochætæ when three are present, arranged +in a right triangle.</p> + +<div class="hanging"> +<p>h. Last section of <span title="for R_{4+5} read M_{1+2}"><a name="AC_18f" id="AC_18f"></a>R<sub>4+5</sub></span> with a more or less rounded angle +(<a href="#Fig_163">fig. 163l</a>).</p> + +<div class="hanging"> +<p>i. Eyes of the male pilose or pubescent, of the female nearly +bare; m-cu crossvein usually at or proximad of the mid-distance +between the r-m crossvein and the bend of +<span title="for R_{4+5} read M_{1+2}"><a name="AC_18g" id="AC_18g"></a>R<sub>4+5</sub></span>. P. (= <i>Placomyia</i> R. D.) <i>vitripennis</i>. +<span class="rightalign"><i>Plaxemyia</i> R. D.</span></p> + +<p>ii. Eyes bare; the m-cu crossvein always nearer to the bend of +<span title="for R_{4+5} read M_{1+2}"><a name="AC_18h" id="AC_18h"></a>R<sub>4+5</sub></span> than to the r-m crossvein.</p> + +<div class="hanging"> +<p>j. Apex of the proboscis when extended reveals a circlet of +stout chitinous teeth. <b>P. insignis</b> Austen, of India, +bites both man and animals. (= <i>Pristirhynchomyia</i>) +<span class="rightalign"><b>Philæmatomyia</b> Austen</span><span class="pagenum"><a name="Page_307" id="Page_307">[Pg 307]</a></span></p> + +<p>jj. Apex of the proboscis without black teeth.</p> + +<div class="hanging"> +<p>k. Eyes of male separated by a distance equal to a fourth +the width of the head. House or typhoid fly. +<b>M. domestica</b> L. Widely distributed. <span class="rightalign"><b>Musca</b> L.</span></p> + +<p>kk. Eyes of the male contiguous. <b>E. corvina.</b> Europe. +<span class="rightalign"><b>Eumusca</b> Town</span></p> +</div></div></div> + +<p>hh. Last section of <span title="for R_{4+5} read M_{1+2}"><a name="AC_18i" id="AC_18i"></a>R<sub>4+5</sub></span> with a gentle curve (<a href="#Fig_102">fig. 102</a>).</p> + +<div class="hanging"> +<p>i. Eyes pilose.</p> + +<div class="hanging"> +<p>j. Claws in the male somewhat elongated; no orbitals in +either sex; antennæ separated at the base by a flat +carina; abdomen marked with red or yellow. <i>G. maculata.</i> +Europe and America. <span class="rightalign"><i>Graphomyia</i> R. D.</span></p> + +<p>jj. Claws short and equal in the two sexes; two or three +stout orbital macrochætæ in the female; R<sub>1</sub> scarcely +produced beyond the r-m crossvein; eyes contiguous +in the male. <i>P. obsoleta.</i> Brazil. <span class="rightalign"><i>Phasiophana</i> Br.</span></p> +</div> + +<p>ii. Eyes bare; fronto-orbital macrochætæ in a double row, +antennæ contiguous at the base.</p> + +<div class="hanging"> +<p>j. One or more pairs of well developed anterior inner dorsocentral +(acrostichal) macrochætæ; seta on extensor +surface of hind tibia. <b>M. assimilis</b>, <b>stabulans</b>, etc. +Europe and America. <span class="rightalign"><b>Muscina</b> R. D.</span></p> + +<p>jj. Anterior inner dorsocentrals and the setæ; on the extensor +surface of the hind tibia wanting. <i>M. micans</i>, +etc. Europe and North America. <span class="rightalign"><i>Morellia</i> R. D.</span></p> +</div></div></div></div> + +<p>ff. Squamula thoracalis not broadened mesad and caudad, not +reaching the margin of the scutellum; macrochætæ on extensor +surface of the hind tibia wanting.</p> + +<div class="hanging"> +<p>g. Eyes pubescent. <i>M. meditabunda.</i> Europe and America. +<span class="rightalign"><i>Myiospila</i> Rdi.</span></p> + +<p>gg. Eyes bare; R<sub>1</sub> ends before the middle of the wing. A number +of species from the tropics of both hemispheres. +<span class="rightalign"><i>Clinopera</i> V. d. W.</span></p> +</div></div></div></div></div> + +<p>bb. Hypopleuræ with a vertical row of macrochætæ.</p> + +<div class="hanging"> +<p>c. Eyes pubescent.</p> + +<div class="hanging"> +<p>d. R<sub>1</sub> ends about opposite the r-m crossvein; basal section of R<sub>4+5</sub> bristly +nearly to the crossvein; <i>S. enigmatica</i>. Africa. <span class="rightalign"><i>Somalia</i> Hough</span></p> + +<p>dd. R<sub>1</sub> ends distad of the r-m crossvein.</p> + +<div class="hanging"> +<p>e. Eastern hemisphere. Australasia. <i>N. ochracea</i>, <i>dasypthalma</i>. +<span class="rightalign"><i>Neocalliphora</i> Br.</span></p> + +<p>ee. Western Hemisphere. <i>T. muscinum.</i> Mexico. <span class="rightalign"><i>Tyreomma</i> V. d. W.</span></p> +</div></div> + +<p>cc. Eyes bare.</p> + +<div class="hanging"> +<p>d. The vibrissal angle situated at a noticeable distance above the level of +the margin of the mouth.</p> + +<div class="hanging"> +<p>e. Sternopleural macrochætæ arranged in the order 1:1.</p> + +<div class="hanging"> +<p>f. Genæ with microchætæ.</p> + +<div class="hanging"> +<p>g. Body grayish, with depressed yellow woolly hair among the +macrochætæ; wings folded longitudinally over the body when<span class="pagenum"><a name="Page_308" id="Page_308">[Pg 308]</a></span> +at rest. Cluster flies. <i>P. rudis</i> and related species, widely +distributed. <span class="rightalign"><i>Pollenia</i> R. D.<a name="FNanchor_N_14" id="FNanchor_N_14"></a><a href="#Footnote_N_14" class="fnanchor">[N]</a></span></p> + +<p>gg. Body metallic blue or green. Eastern Hemisphere.</p> + +<div class="hanging"> +<p>h. Vibrissal angle placed very high above the oral margin; a +carina between the antennæ; outer posthumeral wanting; +anterior intraalar present. <i>T. viridaurea.</i> Java. +<span class="rightalign"><i>Thelychæta</i> Br.</span></p> + +<p>hh. Vibrissal angle moderately high above the oral margin; +carina small or wanting; no post humeral macrochæta; +lower squamæ hairy above. (= <i>Paracompsomyia</i> +Hough) (<a href="#Fig_166">fig. 166</a>). <span class="rightalign"><i>Pycnosoma</i> Br.</span></p> +</div></div> + +<p>ff. Genæ bare. <i>S. terminata</i>. Eastern Hemisphere. +<span class="rightalign"><i>Strongyloneura</i> Bigot</span></p> +</div> + +<p>ee. Sternopleurals arranged 2:1.</p> + +<div class="hanging"> +<p>f. Body metallic green or blue, with gray stripes; genæ hairy to the +lower margin; post humerals often wanting; lower squamæ bare +above. (= <i>Compsomyia</i> Rdi.). <span class="rightalign"><b>Chrysomyia</b> R. D.</span></p> + +<div class="hanging"> +<p>g. With one or two orbitals; height of bucca less than half the +height of the eye. South and east U. S. (<a href="#Fig_107">fig. 107</a>). +<span class="rightalign"><b>C. marcellaria</b></span></p> + +<p>gg. No orbitals; height of bucca about a third less than height of +eye. West U. S. <span class="rightalign"><i>C. wheeleri</i> Hough</span><span class="pagenum"><a name="Page_309" id="Page_309">[Pg 309]</a></span></p> +</div> + +<p>ff. Body black or sordidly metallic greenish gray, usually yellow pollinose +or variegate; genæ at most hairy above. <i>N. stygia.</i> +Eastern Hemisphere. <span class="rightalign"><i>Neopollenia</i> Br.</span></p> +</div></div> + +<p>dd. Vibrissal angle situated nearly on a level of the oral margin.</p> + +<div class="hanging"> +<p>e. Species wholly blackish, bluish, or greenish metallic in color.</p> + +<div class="hanging"> +<p>f. First section of <span title="for R_{4+5} read M_{1+2}"><a name="AC_18j" id="AC_18j"></a>R<sub>4+5</sub></span> with at most three or four small bristles at +the immediate base.</p> + +<div class="hanging"> +<p>g. The bend of <span title="for R_{4+5} read M_{1+2}"><a name="AC_18k" id="AC_18k"></a>R<sub>4+5</sub></span> a gentle curve; costal spine present; cell R<sub>5</sub> +closed, ending before the apex of the wing. <i>S. cuprina.</i> +Java. <span class="rightalign"><i>Synamphoneura</i> Bigot</span></p> + +<p>gg. Bend of R<sub>4+5</sub> angular; or the insect differs in other characters; +dorsal surface of the squamula thoracalis hairy (except in +<i>Melinda</i>); arista plumose only on the basal two-thirds +(except usually in <i>Calliphora</i> and <i>Eucalliphora</i>).</p> + +<div class="hanging"> +<p>h. Arista plumose only on the basal two-thirds.</p> + +<div class="hanging"> +<p>i. Base of the antennæ ventrad of the middle of the eye; eyes +of the male nearly contiguous; genæ hairy; second +abdominal segment with median marginal macrochætæ; +two, rarely three, postsutural intraalar macrochætæ.</p> + +<div class="hanging"> +<p>j. Squamula thoracalis dorsally with long black hairs; male +hypopygium two-segmented, large, projecting; claws +and pulvilli of the male elongate; three strong sternopleural +macrochætæ; genæ at least half the width of the +eye; buccæ (cheeks) half the height of the eyes; oviviparous. +<i>O. sepulcralis.</i> Europe. <span class="rightalign"><i>Onesia</i> R. D.</span></p> + +<p>jj. Dorsal surface of the squamula thoracalis bare; male +hypopygium small, scarcely projecting below; claws +and pulvilli not elongate; two stout sternopleural +macrochætæ, sometimes with a delicate one below the +anterior; genæ nearly linear in the male; buccæ about +a third of the eye height; oviparous. <i>M. cærulea.</i> +Europe. <span class="rightalign"><i>Melinda</i>. R. D.</span><span class="pagenum"><a name="Page_311" id="Page_311">[Pg 311]</a></span></p> +</div> + +<p>ii. Base of the antennæ dorsad of the middle of the eye; eyes +of both sexes distinctly separated; dorsal surface of +the squamula thoracalis with black hairs; two postsutural +intraalar macrochætæ.</p> + +<div class="hanging"> +<p>j. Hypopygium of the male large, with a pair of slightly +curved forceps whose ends are concealed in a longitudinal +slit in the fifth ventral sclerite; third posterior +inner dorso-central (acrostichal) macrochætæ absent; +anterior intraalar rarely present; abdomen usually not +pollinose; the second segment without median marginal +macrochætæ; face yellow. <i>C. mortuorum</i>, <i>cadaverina</i>, +and related species. Both hemispheres. +<span class="rightalign"><i>Cynomyia</i> R. D.<a name="FNanchor_O_15" id="FNanchor_O_15"></a><a href="#Footnote_O_15" class="fnanchor">[O]</a></span></p> + +<p>jj. Three pairs of posterior inner dorsocentrals (acrostichals) +present; second abdominal segment with a row of +marginal macrochætæ; genæ hairy, at least above.</p> + +<div class="hanging"> +<p>k. Hypopygium of the male with a projecting style. +<i>S. stylifera</i>. Europe. <span class="rightalign"><i>Steringomyia</i> Pok.</span><span class="pagenum"><a name="Page_312" id="Page_312">[Pg 312]</a></span></p> + +<p>kk. Hypopygium of the male without style. <i>A. stelviana</i> +B. B. <span class="rightalign"><i>Acrophaga</i> B. B.</span></p> +</div></div></div> + +<p>hh. Arista usually plumose nearly to the tip; posterior dorsocentrals +and inner dorsocentrals (acrostichals) well +developed; dorsal surface of the squamula thoracalis +hairy; abdomen metallic and usually pollinose; genæ +hairy.</p> + +<div class="hanging"> +<p>i. With one pair of ocellar macrochætæ. <b>C. vomitoria</b>, +<b>erythrocephala</b>, <b>viridescens</b>, and related species. Both +hemispheres. <span class="rightalign"><b>Calliphora</b> R. D.</span></p> + +<p>ii. With two strong pairs of ocellar macrochætæ. <i>E. latifrons.</i> +Pacific slope of the U. S. <span class="rightalign"><i>Eucalliphora</i> Town.</span></p> +</div></div></div> + +<p>ff. First section of R<sub>4+5</sub> bristly near or quite half way to the small +crossvein; dorsal surface of the squamula thoracalis is bare; +the hypopygium of the male is inconspicuous.</p> + +<div class="hanging"> +<p>g. Genæ bare; posterior inner and outer dorsocentrals distinct +and well developed. <i>L. cæsar</i>, <i>sericata</i>, <i>sylvarum</i>, and +related species. Widely distributed in both hemispheres +(<a href="#Fig_103">fig. 103</a>). <span class="rightalign"><b>Lucilia</b> R. D.</span></p> + +<p>gg. Genæ with microchætæ, at least down to the level of the base +of the arista.</p> + +<div class="hanging"> +<p>h. Mesonotum flattened behind the transverse suture.</p> + +<div class="hanging"> +<p>i. Posterior dorsocentrals inconstant and unequally developed; +one pair of posterior inner dorsocentrals. <i>P. terrænovæ.</i> +North America. <span class="rightalign"><i>Protophormia</i> Town.</span></p> + +<p>ii. Posterior dorsocentrals well developed, the inner dorsocentrals +(acrostichals) unequally developed. <i>P. azurea</i>, +<i>chrysorrhœa</i>, etc. Europe and America. +<span class="rightalign"><i>Protocalliphora</i> Hough</span></p> +</div> + +<p>hh. Mesonotum not flattened behind the transverse suture; +posterior inner and outer dorsocentrals inconstant +and unequally developed. <i>P. regina.</i> Europe and +America. <span class="rightalign"><i>Phormia</i> R. D.</span></p> +</div></div></div> + +<p>ee. Species more or less rufous or yellow in color.</p> + +<div class="hanging"> +<p>f. Anterior dorsocentrals wanting; first section of the R<sub>4+5</sub> at most +only bristly at the base, bend <span title="for "near apex" read "of M_{1+2}""><a name="AC_20" id="AC_20"></a>near apex</span> rectangular, R<sub>1</sub> ends over +the crossvein; fronto-orbital macrochæta absent; eyes of the +male contiguous. <i>C. semiviridis.</i> Mexico. <span class="rightalign"><i>Chloroprocta</i> V. d. W.</span></p> + +<p>ff. With another combination of characters.</p> + +<div class="hanging"> +<p>g. Body robust, of large size, abdomen elongate, not round; genæ +with several ranges of microchætæ; vibrissal ridges strongly +convergent; abdomen with well developed macrochætæ; +costal spine usually absent; eyes of the male widely separated.</p> + +<div class="hanging"> +<p>h. Peristome broad, pteropleural macrochætæ distinct; one or +two sternopleurals; in the female a single orbital macrochæta; +last abdominal segment without discal macrochætæ; +hypopygial processes of the male with a long +stylet; second abdominal segment of the female sometimes<span class="pagenum"><a name="Page_314" id="Page_314">[Pg 314]</a></span> +much elongate. <b>A. luteola</b> (<a href="#Fig_86">fig. 86</a>). Africa. The sub-genus +<i>Chœromyia</i> Roub. is included here. <span class="rightalign"><b>Auchmeromyia</b> B. B.</span></p> + +<p>hh. Peristome narrow; no pteropleurals, two sternopleurals; +two orbitals in the female; second segment not elongate; +the fourth with two well developed discal macrochætæ. +<b>B. depressa.</b> Africa. <span class="rightalign"><b>Bengalia</b> R. D</span></p> +</div> + +<p>gg. With another combination of characters.</p> + +<div class="hanging"> +<p>h. Costal spine present; body in part black; antennæ noticeably +shorter than the epistome, inserted above the middle +of the eye and separated from each other by a carina; +abdominal segments with marginal macrochætæ; sternopleurals +2:1 or 1:1. <span class="rightalign"><i>Paratricyclea</i> Villen.</span></p> + +<p>hh. Costal spine not distinct, or if present, insect otherwise +different.</p> + +<div class="hanging"> +<p>i. Genæ with several ranges of microchætæ; vibrissal ridges +strongly converging; peristome broad; arista moderately +plumose; sternopleurals usually 1:1; color entirely +testaceous. <b>C. anthropophaga</b> (<a href="#Fig_87">fig. 87</a>) and <b>grunbergi</b>. +Africa. <span class="rightalign"><b>Cordylobia</b> Grünb.</span></p> + +<p>ii. Genæ bare or with but one range of setæ; vibrissal ridges +less converging; peristome narrow; arista long plumose.</p> + +<div class="hanging"> +<p>j. Genæ with a single row of microchætæ.</p> + +<div class="hanging"> +<p>k. Sternopleurals 2:1; color entirely testaceous. +<span class="rightalign"><i>Ochromyia</i> Macq.<a name="FNanchor_P_16" id="FNanchor_P_16"></a><a href="#Footnote_P_16" class="fnanchor">[P]</a></span></p> + +<p>kk. Sternopleurals 1:1. <i>P. varia</i> Hough. Africa. +<span class="rightalign"><i>Parochromyia</i> Hough</span></p> +</div> + +<p>jj. Genæ bare.</p> + +<div class="hanging"> +<p>k. Basal section of R<sub>4+5</sub> bristly only at the immediate +base, <span title="for "distal section" read "distally M_{1+2}""><a name="AC_22" id="AC_22"></a>distal section</span> with a broad curve; distal +portion of the abdomen metallic; sternopleurals +usually 1:1, rarely 2:1. <i>M. æneiventris</i> Wd. Tropic +America. <span class="rightalign"><i>Mesembrinella.</i> G. T.</span></p> + +<p>kk. R<sub>4+5</sub> bristly at least nearly half way to the small +crossvein; sternopleurals 1:1.</p> + +<div class="hanging"> +<p>l. Macrochætæ of the abdomen marginal; neither sex +with orbitals; no carina between the base of the +antennæ; three pairs of presutural inner dorsocentrals. +Eastern hemisphere. <i>T. ferruginea.</i> +<i>Tricyclea V. d. W.</i> (= <i>Zonochroa</i> B. B. according +to Villeneuve 1914).</p> + +<p>ll. Abdomen without macrochætæ; wing usually with +a marginal streak and gray markings. Brazil +<span class="rightalign">Hemilucilia B. B.</span></p> +</div></div></div></div></div></div></div></div></div></div></div></div> + +<div class="figcenter" style="width: 450px;"><a name="Fig_165" id="Fig_165"></a> +<a href="images/f165-full.png"><img src="images/f165.png" width="450" height="365" alt="165. Glossina morsitans. (×4.) After Austen." title="165. Glossina morsitans. (×4.) After Austen." /></a> +<span class="caption">165. Glossina morsitans. (×4.) After Austen.</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_166" id="Fig_166"></a> +<a href="images/f166-full.png"><img src="images/f166.png" width="500" height="316" alt="166. Pycnosoma marginale. (×4.) After Graham-Smith." title="166. Pycnosoma marginale. (×4.) After Graham-Smith." /></a> +<span class="caption">166. Pycnosoma marginale. (×4.) After Graham-Smith.</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_167" id="Fig_167"></a> +<a href="images/f167-full.png"><img src="images/f167.png" width="500" height="339" alt="167. Horn fly. (a) egg; (b) larva; (c) puparium; (d) adult. (×4). Bureau of Entomology" title="167. Horn fly. (a) egg; (b) larva; (c) puparium; (d) adult. (×4). Bureau of Entomology" /></a> +<span class="caption">167. Horn fly. (<i>a</i>) egg; (<i>b</i>) larva; (<i>c</i>) puparium; (<i>d</i>) adult. (×4). Bureau of Entomology</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_168" id="Fig_168"></a> +<a href="images/f168-full.png"><img src="images/f168.png" width="500" height="247" alt="168. Head of horn-fly (Lyperosia irritans); (a) female; (b) male; (c) lateral aspect of female." title="168. Head of horn-fly (Lyperosia irritans); (a) female; (b) male; (c) lateral aspect of female." /></a> +<span class="caption"><span title="add Bureau of Entomology">168. Head of horn-fly (Lyperosia irritans); (<i>a</i>) female; (<i>b</i>) male; (<i>c</i>) lateral aspect of female.</span></span> +</div> + +<div class="figcenter" style="width: 500px;"> +<a href="images/f169-full.png"><img src="images/f169.png" width="500" height="792" alt="169. Lateral and dorsal aspects of the thorax, and frontal aspect of the head of a muscoidean +fly, with designations of the parts commonly used in taxonomic work." title="169. Lateral and dorsal aspects of the thorax, and frontal aspect of the head of a muscoidean +fly, with designations of the parts commonly used in taxonomic work." /></a> +<span class="caption">169. Lateral and dorsal aspects of the thorax, and frontal aspect of the head of a muscoidean +fly, with designations of the parts commonly used in taxonomic work.</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_170" id="Fig_170"></a> +<a href="images/f170-full.png"><img src="images/f170.png" width="500" height="386" alt="170. Sepsis violacea; puparium and adult. (See page 297.) After Howard." title="170. Sepsis violacea; puparium and adult. (See page 297.) After Howard." /></a> +<span class="caption">170. Sepsis violacea; puparium and adult. (See <a href="#Page_297">page 297</a>.) After Howard.</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_171" id="Fig_171"></a> +<a href="images/f171-full.png"><img src="images/f171.png" width="500" height="326" alt="171. Stigmata of the larvæ of Muscoidea. Third instar. (a) Cynomyia cadaverina; (b) Phormia regina; (c) Chrysomyia macellaria; +(d) Musca domestica; (e) Sarcophaga sp.; (f) Oestris ovis; (g) Gastrophilus equi; (h) Sarcophaga sp.; (i) Pegomyia +vicina; (j) Protocalliphora azurea; (k) Hypoderma lineata; (l) Muscina stabulans. Magnification for f, g, and k, ×25; +all others, ×50." title="171. Stigmata of the larvæ of Muscoidea. Third instar. (a) Cynomyia cadaverina; (b) Phormia regina; (c) Chrysomyia macellaria; +(d) Musca domestica; (e) Sarcophaga sp; (f) Oestris ovis; (g) Gastrophilus equi; (h) Sarcophaga sp; (i) Pegomyia +vicina; (j) Protocalliphora azurea; (k) Hypoderma lineata; (l) Muscina stabulans. Magnification for f, g, and k, ×25; +all others, ×50." /></a> +<span class="caption">171. Stigmata of the larvæ of Muscoidea. Third instar. (<i>a</i>) Cynomyia cadaverina; (<i>b</i>) Phormia regina; (<i>c</i>) Chrysomyia macellaria; +(<i>d</i>) Musca domestica; (<i>e</i>) Sarcophaga sp; (<i>f</i>) Oestris ovis; (<i>g</i>) Gastrophilus equi; (<i>h</i>) Sarcophaga sp; (<i>i</i>) Pegomyia +vicina; (<i>j</i>) Protocalliphora azurea; (<i>k</i>) Hypoderma lineata; (<i>l</i>) Muscina stabulans. Magnification for f, g, and k, ×25; +all others, ×50.</span> +</div> + +<div class="figcenter" style="width: 500px;"><a name="Fig_172" id="Fig_172"></a> +<a href="images/f172-full.png"><img src="images/f172.png" width="500" height="698" alt="172. Left hand stigmata of the larvæ of muscoidea. Third instar. (a) Lucilia cæsar; +(b) Calliphora vomitoria; (c) Stomoxys calcitrans; (d) Pseudopyrellia cornicina; +(e) Pyrellia cadavarina; (f) Lyperosia irritans; (g) Mesembrina mystacea; (h) +Mesembrina meridiana; (i) Myospila meditabunda; (j) Mydæa umbana; (k) +Polietes albolineata; (l) Polietes lardaria; (m) Morellia hortorum; (n) Hydrotæa +dentipes; (o) Hebecnema umbratica; (p) H. vespertina; (q) Limnophora septemnotata; +(r) Muscina stabulans. (a and b) after MacGregor; (d) after Banks; +all others after Portchinsky. Magnification varies. The relative distance to the +median line is indicated in each figure." title="172. Left hand stigmata of the larvæ of muscoidea. Third instar. (a) Lucilia cæsar; +(b) Calliphora vomitoria; (c) Stomoxys calcitrans; (d) Pseudopyrellia cornicina; +(e) Pyrellia cadavarina; (f) Lyperosia irritans; (g) Mesembrina mystacea; (h) +Mesembrina meridiana; (i) Myospila meditabunda; (j) Mydæa umbana; (k) +Polietes albolineata; (l) Polietes lardaria; (m) Morellia hortorum; (n) Hydrotæa +dentipes; (o) Hebecnema umbratica; (p) H. vespertina; (q) Limnophora septemnotata; +(r) Muscina stabulans. (a and b) after MacGregor; (d) after Banks; +all others after Portchinsky. Magnification varies. The relative distance to the +median line is indicated in each figure." /></a> +<span class="caption">172. Left hand stigmata of the larvæ of muscoidea. Third instar. (<i>a</i>) Lucilia cæsar; +(<i>b</i>) Calliphora vomitoria; (<i>c</i>) Stomoxys calcitrans; (<i>d</i>) <span title="for Pseudopyrellia read Orthellia">Pseudopyrellia</span> cornicina; +(<i>e</i>) Pyrellia cadavarina; (<i>f</i>) <span title="for Lyperosia read Hæmatobia">Lyperosia</span> irritans; (<i>g</i>) Mesembrina mystacea; +(<i>h</i>) Mesembrina meridiana; (<i>i</i>) Myospila meditabunda; (<i>j</i>) Mydæa <span title="for Umbana read urbana">umbana</span>; +(<i>k</i>) Polietes albolineata; (<i>l</i>) Polietes lardaria; (<i>m</i>) Morellia hortorum; (<i>n</i>) Hydrotæa +dentipes; (<i>o</i>) Hebecnema umbratica; (<i>p</i>) H. vespertina; (<i>q</i>) Limnophora septemnotata; +(<i>r</i>) Muscina stabulans. (<i>a</i> and <i>b</i>) after MacGregor; (<i>d</i>) after Banks; +all others after Portchinsky. Magnification varies. The relative distance to the +median line is indicated in each figure.</span> +</div><p><span class="pagenum"><a name="Page_316" id="Page_316">[Pg 316]</a></span></p> + + +<h3><a name="SIPHONAPTERA_Fleas" id="SIPHONAPTERA_Fleas"></a>SIPHONAPTERA. Fleas</h3> + +<p class="center">Adapted from a table published by Oudemans.</p> + +<div class="hanging"> +<p>a. Elongated fleas, with jointed (articulated) head, with combs (ctenidia) on +head and thorax; with long, oval, free-jointed flagellum of the antenna +(<a href="#Fig_92">fig. 92d</a>). <span class="rightalign">Suborder FRACTICIPATA</span></p> + +<div class="hanging"> +<p>b. With ctenidia in front of the antennæ and on the genæ (cheeks); maxillæ +with acute apices; labial palpi five-segmented, symmetrical; eyes poorly +developed or wanting. On rodents. <span class="rightalign"><span class="smcap">Hystrichopsyllidæ</span></span></p> + +<div class="hanging"> +<p>c. Abdominal segments without ctenidia.</p> + +<div class="hanging"> +<p>d. Post-tibial spines in pairs and not in a very close set row; head with +ctenidia. <span class="rightalign"><i>Ctenophthalmus</i> Kol.</span></p> + +<p>dd. Post-tibial spines mostly single and in a close set row. <i>Ctenopsyllus</i> +and <i>Leptopsyllus</i>. The last genus has recently been erected for +<i>L. musculi</i>, a widely distributed species occurring on rats and mice.</p> +</div> + +<p>cc. Abdominal segments with one or more ctenidia; post-tibial spines in +numerous, short, close-set transverse rows on posterior border with +about four spines in each row. <i>H. americana.</i> <span class="rightalign"><i>Hystrichopsylla</i> Taschenb.</span></p> +</div> + +<p>bb. With only two pairs of subfrontal ctenidia; labial palpi five-segmented, +symmetrical; eyes vestigial or wanting. On bats. (= <span class="smcap">Ischnopsyllidæ</span>). +<span class="rightalign"><span class="smcap">Nycteridipsyllidæ</span></span></p> + +<div class="hanging"> +<p>With more or less blunt maxilla; all tibiæ with notch; a single antepygidial +bristle; metepimeron without ctenidium. <i>N. crosbyi</i> from +Missouri was found on bats. Rothschild suggests that this is probably +the same as <i>N. insignis</i>. +<span class="rightalign">(= <i>Ischnopsyllus</i> = <i>Ceratopsyllus</i>), <i>Nycteridiphilus</i></span></p> +</div></div> + +<p>aa. Head not jointed, i.e. the segments coalescent, traces of the segmentation +still being visible in the presence of the vertex tubercle, the falx (sickle-shaped +process), and a suture. <span class="rightalign">Suborder INTEGRICIPITA</span></p> + +<div class="hanging"> +<p>b. Flagellum of the antennæ long and oval.</p> + +<div class="hanging"> +<p>c. Usually elongate fleas, with a free-segmented flagellum of the antenna; +thorax not shorter than the head, longer than the first tergite.</p> + +<div class="hanging"> +<p>d. Genæ of the head and the pronotum with ctenidia. <span class="rightalign"><span class="smcap">Neopsyllidæ</span></span></p> + +<div class="hanging"> +<p>e. Labial palpi four or five-segmented; symmetrical; hind coxæ with +patch of spines inside; row of six spatulate spines on each side in +front of the antennæ. <i>C. ornate</i> found on a California mole. +<span class="rightalign"><i>Corypsylla</i></span></p> + +<p>ee. Labial palpi two-segmented, transparent, membranous. On +hares. <span class="rightalign"><i>Spilopsyllus</i> Baker</span></p> +</div> + +<p>dd. No ctenidium on the head.</p> + +<div class="hanging"> +<p>e. Pronotum with ctenidium. <span class="rightalign"><span class="smcap"><b>Dolichopsyllidæ</b></span></span></p> + +<div class="hanging"> +<p>f. Labial palpi five-segmented, symmetrical.</p> + +<div class="hanging"> +<p>g. Antepygidial bristles one to three; eyes present.</p> + +<div class="hanging"> +<p>h. Inner side of hind coxæ distally with a comb of minute teeth; +falx present. On rodents and carnivores. +<span class="rightalign"><i>Odontopsyllus</i> Baker</span></p> + +<p>hh. Inner side of hind coxæ without comb or teeth. Many +North American species on rodents. +<span class="rightalign"><b>Ceratophyllus</b> Curtis</span><span class="pagenum"><a name="Page_317" id="Page_317">[Pg 317]</a></span></p> +</div> + +<p>gg. Antepygidial bristles five on each side; eyes absent; suture +white. <i>D. stylosus</i> on rodents. <span class="rightalign"><i>Dolichopsyllus</i> Baker</span></p> +</div> + +<p>ff. Labial palpi four or five-segmented; asymmetrical (membranous +behind), apex acute. <i>Hoplopsyllus anomalus</i> found on Spermophiles +in Colorado. <span class="rightalign"><span class="smcap">Hoplopsyllidæ</span></span></p> +</div> + +<p>ee. Pronotum without ctenidium. <i>Anomiopsyllus californicus</i> and +<i>nudatus</i> on rodents. <span class="rightalign"><span class="smcap">Anomiopsyllidæ</span></span></p> +</div></div> + +<p>cc. Very short fleas; flagellum of the antenna with pseudo-segments coalescent; +thorax much shorter than the head and than the first tergite +<span class="rightalign"><span class="smcap"><b>Hectopsyllidæ</b></span></span></p> + +<div class="hanging"> +<p>Flagellum of the antenna with six coalescent pseudo-segments; maxilla +blunt. The chigger on man (<a href="#Fig_93">fig. 93</a>). <b>D. penetrans.</b> +<span class="rightalign">(= <b>Rhynchoprion</b> = <b>Sarcopsylla</b>) <b>Dermatophilus</b> Guérin</span></p> +</div></div> + +<p>bb. Flagellum short, round, free portion of the first segment shaped like a +mandolin.</p> + +<div class="hanging"> +<p>c. Thorax not shorter than the head, longer than the first tergite; flagellum +either with free segments or in part with the segments coalescent.</p> + +<div class="hanging"> +<p>d. Head and pronotum with ctenidium; labial palpi asymmetrical. <span class="rightalign"><span class="smcap"><b>Archæopsyllidæ</b></span></span></p> + +<div class="hanging"> +<p>With four subfrontal, four genal, and one angular ctenidia. Widely +distributed. <span class="rightalign"><b>Ctenocephalus</b> Kol</span>.</p> + +<p>e. Head rounded in front (<a href="#Fig_92">fig. 92a</a>). Dog flea. <span class="rightalign"><b>C. canis</b></span></p> + +<p>ee. Head long and flat (<a href="#Fig_92">fig. 92b</a>). Cat flea. <span class="rightalign"><b>C. felis</b></span></p> +</div> + +<p>dd. Neither head nor pronotum with ctenidium. Labial palpi asymmetrical, +membranous behind. <span class="rightalign"><span class="smcap"><b>Pulicidæ</b></span></span></p> + +<div class="hanging"> +<p>e. Mesosternite narrow, without internal rod-like thickening from the +insertion of the coxæ upwards. Human flea, etc. <span class="rightalign"><b>Pulex</b> L.</span></p> + +<p>ee. Mesosternite broad with a rod-like internal thickening from the +insertion of the coxæ upwards (<a href="#Fig_89">fig. 89</a>). <b>X. (Lœmopsylla) cheopis</b>, +plague or rat flea. <span class="rightalign"><b>Xenopsylla</b></span></p> +</div></div> + +<p>cc. Thorax much shorter than the head and than the first tergite. <b>Echidnophagidæ.</b> +<b>E. gallinacea</b>, the hen flea also attacks man (<a href="#Fig_96">fig. 96</a>). +<span class="rightalign">(= <b>Argopsylla</b> = <b>Xestopsylla</b>) <b>Echidnophaga</b> Olliff.</span></p> +</div></div></div> + + +<div class="footnotes"><h3>FOOTNOTES:</h3> + +<div class="footnote"><p><a name="Footnote_E_5" id="Footnote_E_5"></a><a href="#FNanchor_E_5"><span class="label">[E]</span></a> Adapted from Banks, Nuttall, Warburton, Stiles, <i>et al.</i></p></div> + +<div class="footnote"><p><a name="Footnote_F_6" id="Footnote_F_6"></a><a href="#FNanchor_F_6"><span class="label">[F]</span></a> Dr. C. W. Stiles considers the species which is responsible for spotted fever distinct from the +<i>venustus</i> of Banks, separating it as follows: +</p><p> +Goblet cells about 75 in the male or 105 in the female. Texas. <i>D. venustus.</i> +</p><p> +Goblet cells 157 in the male, or 120 in the female; stigmal plate shaped as shown in the figure +(<a href="#Fig_150">figs. 150 a, b</a>). Montana, etc. <b>D. andersoni.</b></p></div> + +<div class="footnote"><p><a name="Footnote_G_7" id="Footnote_G_7"></a><a href="#FNanchor_G_7"><span class="label">[G]</span></a> Professor C. R. Crosby who has been working upon certain capsids states that he and his +assistant have been bitten by <b>Lygus pratensis</b>, the tarnished plant bug, by <b>Chlamydatus associatus</b> +and by <b>Orthotylus flavosparsus</b>, though without serious results.</p></div> + +<div class="footnote"><p><a name="Footnote_H_8" id="Footnote_H_8"></a><a href="#FNanchor_H_8"><span class="label">[H]</span></a> Species marked with an * are known to transmit malaria. Species found only in tropical +North America and not known to carry malaria have been omitted from this table, but all found +in the United States are included.</p></div> + +<div class="footnote"><p><a name="Footnote_I_9" id="Footnote_I_9"></a><a href="#FNanchor_I_9"><span class="label">[I]</span></a> This table to the North American genera of the Tabanidæ is adapted from one given by +Miss Ricardo.</p></div> + +<div class="footnote"><p><a name="Footnote_J_10" id="Footnote_J_10"></a><a href="#FNanchor_J_10"><span class="label">[J]</span></a> The classification of the Muscoidea as set forth by Schiner and other earlier writers has +long been followed, although it is not satisfactory, being admittedly more or less artificial. Within +the last two or three decades several schemes have been advanced, that of Brauer and Bergenstamm +and of Girschner, with the modifications of Schnabl and Dziedzicki having obtained most +favor in Europe. Townsend, in 1908, proposed a system which differs from Girschner's in some +respects, but unfortunately it has not yet been published in sufficient detail to permit us to adopt +it. From considerations of expediency we use here the arrangement given in Aldrich's Catalogue +of North American Diptera, though we have drawn very freely upon Girschner's most excellent +paper for taxonomic characters to separate the various groups.</p> + +<p>It may sometimes be found that a species does not agree in all the characters with the synopsis; +in this case it must be placed in the group with which it has the most characters in common.</p></div> + +<div class="footnote"><p><a name="Footnote_K_11" id="Footnote_K_11"></a><a href="#FNanchor_K_11"><span class="label">[K]</span></a> There are several genera of flies of the family <i>Cordyluridæ</i>; (i.e. <i>Acalyptratæ</i>) which might be +placed with the <i>Anthomyiidæ</i> (i.e. <i>Calyptratæ</i>), owing to the relatively large size of their squamæ. +As there is no single character which will satisfactorily separate all doubtful genera of these two +groups we must arbitrarily fix the limits. In general those forms on the border line having a +costal spine, or lower squama larger than the upper, or the lower surface of the scutellum more +or less pubescent, or the eyes of the male nearly or quite contiguous, or the eyes hairy, or the +frontal setæ decussate in the female; or any combination of these characters may at once be +placed with the <i>Anthomyiidæ</i>. Those forms which lack these characteristics and have at least +six abdominal segments (the first and second segments usually being more or less coalescent) +are placed with the Acalyptrates. There are other acalyptrates with squamæ of moderate size +which have either no vibrissæ, or have the subcosta either wholly lacking or coalescent in large +part with R<sub>1</sub> or have spotted wings; they, therefore will not be confused with the calyptrates.</p></div> + +<div class="footnote"><p><a name="Footnote_L_12" id="Footnote_L_12"></a><a href="#FNanchor_L_12"><span class="label">[L]</span></a> <i>Pachymyia</i> Macq. is closely related to <i>Stomoxys</i>. It differs in having the arista rayed both +above and below. <i>P. vexans</i>, Brazil.</p></div> + +<div class="footnote"><p><a name="Footnote_M_13" id="Footnote_M_13"></a><a href="#FNanchor_M_13"><span class="label">[M]</span></a> The genus <i>Eudasyphora</i> Town. has recently been erected to contain <i>D. lasiophthalma</i>.</p></div> + +<div class="footnote"><p><a name="Footnote_N_14" id="Footnote_N_14"></a><a href="#FNanchor_N_14"><span class="label">[N]</span></a> <i>Nitellia</i>, usually included in this genus has the apical cell petiolate. <i>Apollenia</i> Bezzi, has +recently been separated from <i>Pollenia</i> to contain the species <i>P. nudiuscula</i>. Both genera belong +to the Eastern hemisphere.</p></div> + +<div class="footnote"><p><a name="Footnote_O_15" id="Footnote_O_15"></a><a href="#FNanchor_O_15"><span class="label">[O]</span></a> The following three genera are not sufficiently well defined to place in this synopsis. In +color and structural characters they are closely related to <i>Cynomyia</i> from which they may be +distinguished as follows. <i>Catapicephala</i> Macq., represented by the species <i>C. splendens</i> from +Java, has the setæ on the facial ridges rising to the base of the antennæ and has median marginal +macrochætæ on the abdominal segments two to four: <i>Blepharicnema</i> Macq., represented by +<i>B. splendens</i> from Venezuela has bare genæ, oral setæ not ascending; tibiæ villose; claws short +in both sexes; <i>Sarconesia</i> Bigot with the species <i>S. chlorogaster</i> from Chile, setose genæ; legs +slender, not villose; claws of the male elongate.</p></div> + +<div class="footnote"><p><a name="Footnote_P_16" id="Footnote_P_16"></a><a href="#FNanchor_P_16"><span class="label">[P]</span></a> <i>Plinthomyia</i> Rdi. and <i>Hemigymnochæta</i> Corti are related to <i>Ochromyia</i>, though too briefly +described to place in the key.</p></div></div> +<p><span class="pagenum"><a name="Page_318" id="Page_318">[Pg 318]</a></span></p> + + + + +<h2><a name="APPENDIX" id="APPENDIX"></a>APPENDIX</h2> + + +<h3><a name="HYDROCYANIC_ACID_GAS_AGAINST_HOUSEHOLD_INSECTS" id="HYDROCYANIC_ACID_GAS_AGAINST_HOUSEHOLD_INSECTS"></a>HYDROCYANIC ACID GAS AGAINST HOUSEHOLD INSECTS</h3> + +<p>The following directions for fumigating with hydrocyanic acid +gas are taken from Professor Herrick's circular published by the +Cornell Reading Course:</p> + +<p>Hydrocyanic acid gas has been used successfully against household +insects and will probably be used more and more in the future. +It is particularly effective against bed-bugs, and cockroaches, but +because <i>it is such a deadly poison it must be used very carefully</i>.</p> + +<p>The gas is generated from the salt potassium cyanid, by treating +it with sulfuric acid diluted with water. Potassium cyanid is a +most poisonous substance and the gas emanating from it is also +deadly to most, if not all, forms of animal life. The greatest care +must always be exercised in fumigating houses or rooms in buildings +that are occupied. Before fumigation a house should be vacated. +It is not necessary to move furniture or belongings except brass or +nickel objects, which may be somewhat tarnished, and butter, milk, +and other larder supplies that are likely to absorb gas. If the nickel +and brass fixtures or objects are carefully covered with blankets +they will usually be sufficiently protected.</p> + +<p>There may be danger in fumigating one house in a solid row of +houses if there is a crack in the walls through which the gas may find +its way. It also follows that the fumigation of one room in a house +may endanger the occupants of an adjoining room if the walls between +the two rooms are not perfectly tight. It is necessary to keep +all these points in mind and to do the work deliberately and thoughtfully. +The writer has fumigated a large college dormitory of 253 +rooms, once a year for several years, without the slightest accident +of any kind. In order to fumigate this building about 340 pounds +of cyanid and the same amount of sulfuric acid were used each time. +In addition to this, the writer has fumigated single rooms and smaller +houses with the gas. In one instance the generating jars were too +small; the liquid boiled over and injured the floors and the rugs. +Such an accident should be avoided by the use of large jars and by +placing old rugs or a quantity of newspapers beneath the jars.<span class="pagenum"><a name="Page_319" id="Page_319">[Pg 319]</a></span></p> + + +<h3><a name="The_Proportions_of_Ingredients" id="The_Proportions_of_Ingredients"></a><span class="smcap">The Proportions of Ingredients</span></h3> + +<p>Experiments and experience have shown that the potassium +cyanid should be ninety-eight per cent pure in order to give satisfactory +results. The purchaser should insist on the cyanid being of +at least that purity, and it should be procurable at not more than +forty cents per pound. The crude form of sulfuric acid may be used. +It is a thickish, brown liquid and should not cost more than four or +five cents a pound. If a room is made tight, one ounce of cyanid for +every one hundred cubic feet of space has been shown to be sufficient. +It is combined with the acid and water in the following proportions:</p> + + +<div class="center"> +<table border="0" cellpadding="4" cellspacing="0" summary=""> +<tr><td align="left">Potassium cyanid</td><td align="left">1 ounce</td></tr> +<tr><td align="left">Commercial sulfuric acid</td><td align="left">1 fluid ounce</td></tr> +<tr><td align="left">Water</td><td align="left">3 fluid ounces</td></tr> +</table></div> + + +<h3><a name="A_Single_Room_As_an_Example" id="A_Single_Room_As_an_Example"></a><span class="smcap">A Single Room As an Example</span></h3> + +<p>Suppose a room to be 12 by 15 by 8 feet. It will contain +12 × 15 × 8, or 1440 cubic feet. For convenience the writer always +works on the basis of complete hundreds; in this case he would +work on the basis of 1500 cubic feet, and thus be sure to have enough. +The foregoing room, then, would require 15 ounces of cyanid, 15 +ounces of sulfuric acid, and 45 ounces of water. The room should +be made as tight as possible by stopping all the larger openings, +such as fireplaces and chimney flues, with old rags or blankets. +Cracks about windows or in other places should be sealed with narrow +strips of newspaper well soaked in water. Strips of newspaper two +or three inches wide that have been thoroughly soaked in water may +be applied quickly and effectively over the cracks around the window +sash and elsewhere. Such strips will stick closely for several hours +and may be easily removed at the conclusion of the work.</p> + +<p>While the room is being made tight, the ingredients should be +measured according to the formula already given. The water should +be measured and <i>poured first</i> into a stone jar for holding at least two +gallons. The jar should be placed in the middle of the room, with +an old rug or several newspapers under it in order to protect the floor.</p> + +<p>The required amount of sulfuric acid should then be poured +rather slowly into the water. <i>This process must never be reversed; +that is, the acid must never be poured into the jar first.</i> The cyanid +should be weighed and put into a paper bag beside the jar. All hats, +coats, or other articles that will be needed before the work is over<span class="pagenum"><a name="Page_320" id="Page_320">[Pg 320]</a></span> +should be removed from the room. When everything is ready the +operator should drop the bag of cyanid gently into the jar, holding +his breath, and should walk quickly out of the room. The steam-like +gas does not rise immediately under these conditions, and ample +time is given for the operator to walk out and shut the door. If +preferred, however, the paper bag may be suspended by a string +passing through a screw eye in the ceiling and then through the keyhole +of the door. In this case the bag may be lowered from the outside +after the operator has left the room and closed the door.</p> + +<p>The writer has most often started the fumigation toward evening +and left it going all night, opening the doors in the morning. The +work can be done, however, at any time during the day and should +extend over a period of five or six hours at least. It is said that better +results will be obtained in a temperature of 70° F., or above, than +at a lower degree.</p> + +<p>At the close of the operation the windows and doors may be opened +from the outside. In the course of two or three hours the gas should +be dissipated enough to allow a person to enter the room without +danger. The odor of the gas is like that of peach kernels and is easily +recognized. The room should not be occupied until the odor has +disappeared.</p> + + +<h3><a name="Fumigating_a_Large_House" id="Fumigating_a_Large_House"></a><span class="smcap">Fumigating a Large House</span></h3> + +<p>The fumigation of a large house is merely a repetition, in each room +and hall, of the operations already described for a single room. All +the rooms should be made tight, and the proper quantities of water +and sulfuric acid should be measured and poured into jars placed +in each room with the cyanid in bags besides the jars. When all +is ready, the operator should <i>go to the top floor and work downward</i> +because the gas is lighter than air and tends to rise.</p> + + +<h3><a name="Precautions" id="Precautions"></a><span class="smcap">Precautions</span></h3> + +<p>The cyanid should be broken up into small pieces not larger than +small eggs. This can best be done on a cement or brick pavement. +It would be advantageous to wear gloves in order to protect the hands, +although the writer has broken many pounds of cyanid without any +protection on the hands. Wash the hands thoroughly at frequent +intervals in order to remove the cyanid.</p> + +<p>The operations of the work must be carried out according to +directions.<span class="pagenum"><a name="Page_321" id="Page_321">[Pg 321]</a></span></p> + +<p>The work should be done by a calm, thoughtful and careful +person—best by one who has had some experience.</p> + +<p>Conspicuous notices of what has been done should be placed on +the doors, and the doors should be locked so that no one can stray +into the rooms.</p> + +<p>The gas is lighter than air, therefore one should always begin in the +rooms at the top of the house and work down.</p> + +<p>After fumigation is over the contents of the jar should be emptied +into the sewer or some other safe place. The jars should be washed +thoroughly before they are used again.</p> + +<p><i>It must be remembered that cyanid is a deadly poison</i>; but it is +very efficient against household insects, if carefully used, and is not +particularly dangerous when properly handled.</p> + + +<h3><a name="LESIONS_PRODUCED_BY_THE_BITE_OF_THE_BLACK-FLY" id="LESIONS_PRODUCED_BY_THE_BITE_OF_THE_BLACK-FLY"></a>LESIONS PRODUCED BY THE BITE OF THE BLACK-FLY</h3> + +<p>While this text was in press there came to hand an important paper +presenting a phase of the subject of black fly injury so different from +others heretofore given that we deem it expedient to reproduce here +the author's summary. The paper was published in <i>The Journal +of Cutaneous Diseases</i>, for November and December, 1914, under the +title of "A Clinical, Pathological and Experimental Study of the +Lesions Produced by the Bite of the Black Fly (<i>Simulium venustum</i>)," +by Dr. John Hinchman Stokes, of the University of Michigan.</p> + + +<h3><span class="smcap">Resume and Discussion of Experimental Findings</span></h3> + +<p>The principal positive result of the work has been the experimental +reproduction of the lesion produced by the black-fly in characteristic +histological detail by the use of preserved flies. The experimental +lesions not only reproduced the pathological pictures, but followed +a clinical course, which in local symptomatology especially, tallied +closely with that of the bite. This the writer interprets as satisfactory +evidence that the lesion is not produced by any living infective +agent. The experiments performed do not identify the nature +of the toxic agent. Tentatively they seem to bring out, however, +the following characteristics.</p> + +<p>1. The product of alcoholic extraction of flies do not contain +the toxic agent.</p> + +<p>2. The toxic agent is not inactivated by alcohol.</p> + +<p>3. The toxic agent is not destroyed by drying fixed flies.</p> + +<p>4. The toxic agent is not affected by glycerin, but is, if anything, +more active in pastes made from the ground fly and glycerin, than +in the ground flies as such.<span class="pagenum"><a name="Page_322" id="Page_322">[Pg 322]</a></span></p> + +<p>5. The toxic agent is rendered inactive or destroyed by hydrochloric +acid in a concentration of 0.25%.</p> + +<p>6. The toxic agent is most abundant in the region of the anatomical +structures connected with the biting and salivary apparatus +(head and thorax).</p> + +<p>7. The toxic agent is not affected by a 0.5% solution of sodium +bicarbonate.</p> + +<p>8. The toxic agent is not affected by exposure to dry heat at +100° C. for two hours.</p> + +<p>9. The toxic agent is destroyed or rendered inactive in alkaline +solution by a typical hydrolytic ferment, pancreatin.</p> + +<p>10. Incomplete experimental evidence suggests that the activity +of the toxic agent may be heightened by a possible lytic action of +the blood serum of a sensitive individual, and that the sensitive serum +itself may contain the toxic agent in solution.</p> + +<p>These results, as far as they go (omitting No. 10), accord with +Langer's except on the point of alcoholic solubility and the effect +of acids. The actual nature of the toxic agent in the black-fly is +left a matter of speculation.</p> + +<p>The following working theories have suggested themselves to +the writer. First, the toxin may be, as Langer believes in the case +of the bee, an alkaloidal base, toxic as such, and neutralized after +injection by antibodies produced for the occasion by the body. In +such a case the view that a partial local fixation of the toxin occurs, +which prevents its immediate diffusion, is acceptable. Through +chemotactic action, special cells capable of breaking up the toxin +into harmless elements are attracted to the scene. Their function +may be, on the other hand, to neutralize directly, not by lysis. +This would explain the rôle of the eosinophiles in the black-fly lesion. +If their activities be essential to the destruction or neutralization +of the toxin, one would expect them to be most numerous where +there was least reaction. This would be at the site of a bite in an +immune individual. A point of special interest for further investigation, +would be the study of such a lesion.</p> + +<p>Second, it is conceivable that the injected saliva of the fly does +not contain an agent toxic as such. It is possible, that like many +foreign proteins, it only becomes toxic when broken down. The +completeness and rapidity of the breaking down depends on the +number of eosinophiles present. In such a case immunity should +again be marked by intense eosinophilia.<span class="pagenum"><a name="Page_323" id="Page_323">[Pg 323]</a></span></p> + +<div class="figcenter" style="width: 400px;"><a name="Fig_173" id="Fig_173"></a> +<a href="images/f173-full.png"><img src="images/f173.png" width="400" height="299" alt="173. Fifth day mature lesion. Lower power drawing showing papillary œdema and infiltrate in the region +of the puncture." title="173. Fifth day mature lesion. Lower power drawing showing papillary œdema and infiltrate in the region +of the puncture." /></a> +<span class="caption"><span title="add "After Dr. J. H. Stokes"">173. Fifth day mature lesion. Lower power drawing showing papillary œdema and infiltrate in the region +of the puncture.</span></span> +</div><p><span class="pagenum"><a name="Page_324" id="Page_324">[Pg 324]</a></span></p> + +<p>Third, lytic agents in the blood serum may play the chief rôle +in the liberation of the toxic agent from its non-toxic combination. +An immune individual would then be one whose immunity was not +the positive one of antibody formation, but the negative immunity +of failure to metabolize. An immune lesion in such a case might +be conceived as presenting no eosinophilia, since no toxin is liberated. +If the liberation of the toxin is dependent upon lytic agents present +in the serum rather than in any cellular elements, a rational explanation +would be available for the apparent results (subject to confirmation) +of the experiment with sensitive and immune sera. In +this experiment it will be recalled that the sensitive serum seemed to +bring out the toxicity of the ground flies, and the serum itself seemed +even to contain some of the dissolved or liberated toxin. The +slowness with which a lesion develops in the case of the black-fly +bite supports the view of the initial lack of toxicity of the injected +material. The entire absence of early subjective symptoms, such +as pain, burning, etc., is further evidence for this view. It would +appear as if no reaction occurred until lysis of an originally non-toxic +substance had begun. Regarding the toxin itself as the chemotactic +agent which attracts eosinophiles, its liberation in the lytic +process and diffusion through the blood stream attracts the cells +in question to the point at which it is being liberated. Arriving +upon the scene, these cells assist in its neutralization.</p> + +<p>The last view presented is the one to which the author inclines +as the one which most adequately explains the phenomena.</p> + +<p>A fourth view is that the initial injection of a foreign protein by +the fly (i.e., with the first bite) sensitizes the body to that protein. +Its subsequent injection at any point in the skin gives rise to a +local expression of systematic sensitization. Such local sensitization +reactions have been described by Arthus and Breton, by Hamburger +and Pollack and by Cowie. The description of such a lesion +given by the first named authors, in the rabbit, however, does not +suggest, histopathologically at least, a strong resemblance to that +of the black-fly. Such an explanation of many insect urticariæ +deserves further investigation, however, and may align them under +cutaneous expressions of anaphylaxis to a foreign protein injected +by the insect. Depending on the chemical nature of the protein +injected, a specific chemotactic reaction like eosinophilia may or +may not occur. Viewed in this light the development of immunity +to insect bites assumes a place in the larger problem of anaphylaxis.<span class="pagenum"><a name="Page_325" id="Page_325">[Pg 325]</a></span></p> + +<div class="figcenter" style="width: 375px;"><a name="Fig_174" id="Fig_174"></a> +<a href="images/f174-full.png"><img src="images/f174.png" width="375" height="477" alt="174. Experimental lesion produced from alcohol-fixed flies, dried and ground into a +paste with glycerin." title="174. Experimental lesion produced from alcohol-fixed flies, dried and ground into a +paste with glycerin." /></a> +<span class="caption"><span title="add "After Dr. J. H. Stokes"">174. Experimental lesion produced from alcohol-fixed flies, dried and ground into a +paste with glycerin.</span></span> +</div><p><span class="pagenum"><a name="Page_326" id="Page_326">[Pg 326]</a></span></p> + + +<h3><span class="smcap">Summary</span></h3> + +<p>In order to bring the results of the foregoing studies together, +the author appends the following résumé of the clinical data presented +in the first paper.</p> + +<p>The black-fly, <i>Simulium venustum</i>, inflicts a painless bite, with +ecchymosis and hæmorrhage at the site of puncture. A papulo-vesicular +lesion upon an urticarial base slowly develops, the full +course of the lesion occupying several days to several weeks. Marked +differences in individual reaction occur, but the typical course involves +four stages. These are, in chronological order, the papular +stage, the vesicular or pseudovesicular, the mature vesico-papular or +weeping papular stage and the stage of involution terminating in a +scar. The papule develops in from 3 to 24 hours. The early pseudovesicle +develops in 24 to 48 hours. The mature vesico-papular lesion +develops by the third to fifth day and may last from a few days to +three weeks. Involution is marked by cessation of oozing, subsidence +of the papule and scar-like changes at the site of the lesion. The +symptoms accompanying this cycle consist of severe localized or +diffused pruritus, with some heat and burning in the earlier stages +if the œdema is marked. The pruritus appears with the pseudovesicular +stage and exhibits extraordinary persistence and a marked +tendency to periodic spontaneous exacerbation. The flies tend to +group their bites and confluence of the developing lesions in such +cases may result in extensive œdema with the formation of oozing +and crusted plaques. A special tendency on the part of the flies +to attack the skin about the cheeks, eyes and the neck along the +hair line and behind the ears, is noted. In these sites inflammation +and œdema may be extreme.</p> + +<p>A distinctive satellite adenopathy of the cervical glands develops +in the majority of susceptible persons within 48 hours after being +bitten in the typical sites. This adenopathy is marked, discrete +and painful, the glands often exquisitely tender on pressure. It +subsides without suppuration.</p> + +<p>Immunity may be developed to all except the earliest manifestations, +by repeated exposures. Such an immunity in natives of an +infested locality is usually highly developed. There are also apparently +seasonal variations in the virulence of the fly and variations +in the reaction of the same individual to different bites.</p> + +<p>Constitutional effects were not observed but have been reported.<span class="pagenum"><a name="Page_327" id="Page_327">[Pg 327]</a></span></p> + + + +<hr style="width: 65%;" /> +<h2><a name="BIBLIOGRAPHY" id="BIBLIOGRAPHY"></a>BIBLIOGRAPHY</h2> + + +<div class="hanging"><p><b>Aldrich, J. M.</b> 1905. A catalogue of North American Diptera. Washington, +D. C. 1-680.</p> + +<p><b>Alessandri, G.</b> 1910. Studii ed esperienze sulle larve della Piophila casei. +Arch. Parasit. xiii, p. 337-387.</p> + +<p><b>Anderson, J. F.</b> and <b>Frost, W. H.</b> 1912. Transmission of poliomyelitis by +means of the stable-fly (Stomoxys calcitrans). Public Health Reports. +Washington. xxvii, p. 1733-1735.</p> + +<p>---- 1913. Further attempts to transmit the disease through the agency of +the stable-fly (Stomoxys calcitrans). Public Health Repts., Washington. +xxviii, p. 833-837.</p> + +<p><b>Anderson, J. F.</b> and <b>Goldberger, J.</b> 1910. On the infectivity of tabardillo or +Mexican typhus for monkeys, and studies on its mode of transmission. Public +Health Repts., Washington. xxv, p. 177.</p> + +<p><b>Annandale, N.</b> 1910. The Indian species of papataci fly (Phlebotomus). +Records of Indian Mus. iv, p. 35-52, pls. iv-vi.</p> + +<p><b>Austen, E. E.</b> 1903. 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The relapsing fever of tropical Africa. A Review. Parasitology, +iv, p. 183-203.</p> + +<p>---- 1911 b. On the life cycle of <i>Spirochæta gallinarum</i>. ibid., iv. p. 463-477.</p> + +<p><b>Hindle, E.</b> and <b>Merriman, G.</b> 1914. The range of flight of <i>Musca domestica</i>. +Journ. of Hygiene, xiv. p. 23-45.<span class="pagenum"><a name="Page_332" id="Page_332">[Pg 332]</a></span></p> + +<p><b>Hine, J. S.</b> 1903. Tabanidæ of Ohio. Papers Ohio Acad. Sci. No. 5, 55 p.</p> + +<p>---- 1906. Habits and life-histories of some flies of the family Tabanidæ. +U. S. Dept. Agric. Bur. Ent. tech. bul. 12, p. 19-38.</p> + +<p>---- 1907. Second report upon the horse-flies of Louisiana. La. Stat. Exp. +Bul. 93, p. 1-59.</p> + +<p><b>Hodge, C. F.</b> 1910. A practical point in the study of the typhoid, or filth-fly. +Nature Study Review, vi, p. 195-199.</p> + +<p>---- 1913. The distance house-flies, blue-bottles, and stable flies may +travel over water. Science, n. s. xxxviii, p. 513.</p> + +<p><b>Honeij, J. A.</b> and <b>Parker, R. R.</b> 1914. Leprosy: flies in relation to the transmission +of the disease. Journ. Med. Research, Boston, xxx, p. 127-130.</p> + +<p><b>Hooker, W. A.</b> 1908 a. Life history, habits, and methods of study of the +Ixodoidea. Jour. Econ. Ent. i, p. 34-51.</p> + +<p>---- 1908 b. A review of the present knowledge of the rôle of ticks in the +transmission of disease. ibid., i, p. 65-76.</p> + +<p><b>Hope, F. W.</b> 1837. On insects and their larvæ occasionally found in the human +body. Trans. Ent. Soc., London, ii, p. 256-271.</p> + +<p><b>Hough, G. de N.</b> 1899 a. Synopsis of the Calliphorinæ of the United States. +Zoological Bulletin, ii, p. 283-290.</p> + +<p>---- 1899 b. Some Muscinæ of North America, Biological Bulletin i, +p. 19-33.</p> + +<p>---- 1899 c. Some North American Genera of Calliphorinæ. Entom. +News, x, p. 62-66.</p> + +<p><b>Hovarth, G.</b> 1912. Revision of the American Cimicidæ. Ann. Mus. Nat. +Hungarici, x, p. 257-262.</p> + +<p><b>Howard, C. W.</b> 1908. A list of the ticks of South Africa, with descriptions and +keys to all the forms known. Ann. Transvaal Mus. 1, p. 73-170.</p> + +<p><b>Howard, C. W.</b> and <b>Clark, P. F.</b> 1912. Experiments on insect transmission +of the virus of poliomyelitis. Journ. Exper. Med. xvi, p. 805-859.</p> + +<p><b>Howard, L. O.</b> 1899. Spider bites and kissing bugs. Pop. Sci. Mo. lv, p. 31-42.</p> + +<p>---- 1900. A contribution to the study of the insect fauna of human excrement. +Proc. Wash. Acad. Sci. ii, p. 541-604.</p> + +<p>---- 1901. Mosquitoes, how they live, how they carry disease, how they are +classified, how they may be destroyed. 8vo. New York, Doubleday, +Page & Co. (xv + 241 p.)</p> + +<p>---- 1909. Economic loss to the people of the United States, through insects +that carry disease. U. S. Dept. Agric. Bur. of Ent. Bul. 78, p. 1-40.</p> + +<p><b>Howard, L. O.</b>, <b>Dyar, H. G.</b> and <b>Knab, F.</b> 1913-. The mosquitoes of North +and Central America and the West Indies. Vol. I. A general consideration +of mosquitoes, their habits, and their relations to the human species. 4<sup>o</sup>. +Carnegie Institution of Washington (vii + 520 p.).</p> + +<p><b>Howard, L. O.</b> and <b>Marlatt, C. L.</b> 1902. The principal household insects of +the United States. U. S. Dept. Agric., Bur. Ent. Bul. 4.</p> + +<p><b>Huebner, W.</b> 1907. Ueber das Pfeilgift der Kalahari. Arch. exper. Path. +und Pharm., lvii, p. 358-366.</p> + +<p><b>Hunter, S. J.</b> 1913. Pellagra and the sand-fly. Jour. econ. Ent. vi, p. 96-99.</p> + +<p><b>Hunter, W. D.</b> 1913. American interest in medical entomology. Jour. econ. +Ent. vi, p. 27-39.</p> + +<p><b>Hunter, W. D.</b> and <b>Bishopp, F. C.</b> 1910. Some of the more important ticks of +the United States. U. S. Dept. Agric. Yearbook 1910, p. 219-230, pls. xv-xvi.</p> + +<p>---- 1911. The Rocky Mountain spotted fever tick. With special reference +to the problem of its control in the Bitter Root Valley in Montana. +U. S. Dept. Agric., Bur. Ent. Bul. 105, p. 1-47.</p> + +<p><b>Hutchison, R. H.</b> 1914. The migratory habit of housefly larvæ as indicating +a favorable remedial measure. An account of progress. U. S. Dept. Agric., +Bul. 14, p. 1-11.</p> + +<p><b>Jennings, A. H.</b> 1914. Summary of two years' study of insects in relation to +pellagra. Journ. of Parasitology, i, p. 10-21.<span class="pagenum"><a name="Page_333" id="Page_333">[Pg 333]</a></span></p> + +<p><b>Jennings, A. H.</b> and <b>King, W. V.</b> 1913. One of the possible factors in the +causation of pellagra. Journ. Amer. Med. Assoc., lx, p. 271-274.</p> + +<p><b>Jepson, F. P.</b> 1909. Notes on colouring flies for purposes of identification. +Rep't to the Local Gov't Board on Publ. Health, n. s. 16, p. 4-9.</p> + +<p><b>Johannsen, O. A.</b> 1903. Aquatic Nematocerous Diptera. N. Y. State Mus. +Bul., 68, p. 328-448, pls. 32-50.</p> + +<p>---- 1905. Aquatic Nematocerous Diptera II. (Chironomidæ). ibid. 86, +p. 76-330, pls. 16-37.</p> + +<p>---- 1908. North America Chironomidæ. ibid., 124, p. 264-285.</p> + +<p>---- 1911. The typhoid fly and its allies. Maine Agric. Exp. Sta. Bul., +401, p. 1-7.</p> + +<p>---- 1911. Simulium and pellagra. Insect Notes for 1910. Maine Agr. +Exper. Station. Bul, 187, p. 4.</p> + +<p><b>Kellogg, V. L.</b> 1915. Spider poison. Jour. of Parasitology, i, p. 107+</p> + +<p><b>Kelly, H. A.</b> 1907. Walter Reed and yellow fever. 8vo. New York, McClure, +Phillips & Co. (xix + 310 p.).</p> + +<p><b>Kephart, Cornelia F.</b> 1914. The poison glands of the larva of the browntail +moth (Euproctis chrysorrhoea Linn.). Journ. Parasit., i, p.</p> + +<p><b>Kieffer, J. J.</b> 1906. Chironomidæ. Genera Insectorum. Fasc. 42, p. 1-78.</p> + +<p>---- 1913. Nouv. étude sur les Chironomides de l'Indien Museum de +Calcutta. Records of the Indian Mus., ix, p. 119-197.</p> + +<p><b>King, A. F. A.</b> 1883. Insects and disease—mosquitoes and malaria. Pop. +Sci. Mo. xxiii, p. 644-658.</p> + +<p><b>Kirkland, A. H.</b> 1907. Second annual report of the Superintendent for suppressing +the gypsy and browntail moths. 8vo. Boston. 170 p.</p> + +<p><b>Kleine, E.</b> 1909. Postive Infektionsversuche mit <i>Trypanosoma brucei</i> durch +<i>Glossina palpalis.</i> Deutsche med. Wochenschr., xxxv, p. 469-470.</p> + +<div class="hanging"><p>Weitere wissenschaftliche Beobachtungen über die Entwicklung von Trypanosomen +in Glossinen. ibid. p. 924-925.</p> + +<p>Weitere Untersuchungen über die Ætiologie der Schlafkrankheit. ibid., p. 1257-1260.</p> + +<p>Weitere Beobachtungen über Tsetsefliegen und Trypanosomen. ibid., p. 1956-1958.</p></div> + +<p><b>Kling, C.</b> and <b>Levaditi, C.</b> 1913. Études sur la poliomyélite aiguë épidémique. +Ann. Inst. Pasteur, xxvii, p. 718-749, 739-855.</p> + +<p><b>Knab, F.</b> 1912. Unconsidered factors in disease-transmission by blood-sucking +insects. Journ. Econ. Ent., v, p. 196-200.</p> + +<p>---- 1913 a. The species of Anopheles that transmit human malaria. +Amer. Journ. Trop. Dis. and Preventive Med., i, p. 24-43.</p> + +<p>---- 1913 b. Anopheles and malaria. ibid., i, p. 217.</p> + +<p>---- 1913 c. The life history of <i>Dermatobia hominis</i>. ibid., i, p. 464-468.</p> + +<p><b>Knab, F.</b> See Howard, Dyar, and Knab.</p> + +<p><b>Kobert, R.</b> 1893. Lehrbuch der Intoxikationen. 4<sup>o</sup>. Stuttgart, Enke. +(xxii + 816 p.). 2d ed. in 2 vols., 1906.</p> + +<p>---- 1901. Beiträge zur Kenntniss der Giftspinnen. 8<sup>o</sup>. Stuttgart, Enke. +(viii + 191 p.).</p> + +<p><b>Kolbe, H. J.</b> 1894. Der Pfeilgiftkäfer der Kalahari-Wüste, <i>Diamphidia simplex</i>. +Stett. Ent. Zeitg., iv, p. 79-86.</p> + +<p><b>Krause, M.</b> 1907. Untersuchungen über Pfeilgifte aus unseren africanischen +Kolonien. Verhand. deutsche Kolonien kong. 1905. p. 264-288.</p> + +<p><b>Lallier, P.</b> 1897. Étude sur la myase du tube digestif chez l'homme. Thesis, +Paris, 8<sup>o</sup>. 120 p.</p> + +<p><b>Langer, J.</b> 1897. Ueber das Gift unserer Honigbiene. Archiv. exper. Path. +und Pharm., xxxviii, p. 381-396.</p> + +<p><b>Lavinder, C. H.</b> 1911. Pellagra: a précis. U. S. Publ. Health Service Bul. +48, 37 p.</p> + +<p><b>Leidy, J.</b> 1847. History and anatomy of the hemipterous genus <i>Belostoma</i>. +Journ. Acad. Philad. (2), i, p. 57-67.</p> + +<p><b>Leiper, R. T.</b> 1907. The etiology and prophylaxis of dracontiasis. British +Med. Journ. 1907, p. 129-132.<span class="pagenum"><a name="Page_334" id="Page_334">[Pg 334]</a></span></p> + +<p><b>Leishman, W. B.</b> 1910 a. Observations on the mechanism of infection in tick +fever and on the hereditary transmission of <i>Spirochæta duttoni</i> in the tick. +Trans. Soc. Trop. Med. Hyg., iii, p. 77-95. Abstr. in Bul. Inst. Pasteur, +viii, p. 312-313.</p> + +<p>---- 1910 b. On the hereditary transmission and mechanism of infection +in tick fever and on the hereditary transmission of <i>Spirochæta duttoni</i> in +the tick. Lancet., clxxvii, p. 11.</p> + +<p><b>Linnell, R. McC.</b> 1914. Notes on a case of death following the sting of a scorpion. +Lancet, 1914, p. 1608-1609.</p> + +<p><b>Livingstone, D.</b> 1857. Missionary travels and researches in South Africa.</p> + +<p><b>Lucas, H.</b> 1843. (note) <i>Latrodectus malmignatus</i> Bul. Soc. Ent., France, +1843, p. viii.</p> + +<p><b>Ludlow, C. S.</b> 1914. Disease bearing mosquitoes of North and Central America, +the West Indies and the Philippine Islands. War Dept., Office of Surgeon +General. Bul. No. 4, 1-96.</p> + +<p><b>Lugger</b>, 1896. Insects injurious in 1896. Agr. Exp. Sta. Bul. 48. p. 33to 270.</p> + +<p><b>MacCallum, W. C.</b> 1898. On the hæmatozoan infection of birds. Journ. +Exp. Med. iii, p. 117.</p> + +<p><b>MacGregor, M. E.</b> 1914. The posterior stigmata of dipterous larvæ as a diagnostic +character. Parasitology, vii, p. 176-188.</p> + +<p><b>Macloskie, G.</b> 1888. The poison apparatus of the mosquito. Amer. Naturalist, +xxii, p. 884-888.</p> + +<p><b>Malloch, J. P.</b> 1913. American black-flies or Buffalo gnats. U. S. Dept. +Agric. Bur. Ent. Tech. Bul. 26, p. 1-72.</p> + +<p>---- 1914. Notes on North American Diptera. Bul. Illinois State Lab. +Nat. Hist., x, p. 213-243.</p> + +<p><b>Manson, P.</b> 1911. Tropical diseases: a manual of the diseases of warm climates. +8<sup>o</sup>. London, Cassell & Co. (xx + 876 p.). 4 ed. (1907). Reprinted.</p> + +<p><b>Marchoux, E.</b> and <b>Couvy, L.</b> 1913. Argas et spirochætes (1 mémoire). Les +granules de Leishman. Ann. Inst. Pasteur, xxvii, p. 450-480. 2 mémoire. +Le virus chez l'acarien. ibid. p. 620-643.</p> + +<p><b>Marchoux, E.</b> and <b>Selimbeni, A.</b> 1903. La spirillose des poules. Ann. Inst. +Pasteur, xvii, p. 569-580.</p> + +<p><b>Marchoux, E.</b> and <b>Simond, P. L.</b> 1905. Études sur la fièvre jaune. Ann. Inst. +Pasteur, xx, pp. 16-40, 104-148, 161-205.</p> + +<p><b>Marlatt, C. L.</b> 1902. (See Howard, L. O. and Marlatt, C. L.)</p> + +<p>---- 1907. The bed-bug (<i>Cimex lectularius</i> L.) U. S. Dept. Agric., Bur. +Ent., Circ. No. 47, revised ed., 8 pp.</p> + +<p><b>Martin, G. Leboeuf</b>, and <b>Roubaud</b>. 1909. Rapport de la mission d'études de la +maladie du sommeil au Congo francais. 4<sup>o</sup>. Paris, Masson & Cie. (vi + +722 p., 8 pls. and map.).</p> + +<p><b>Maver, Maria B.</b> 1911. Transmission of spotted fever by other than Montana +and Idaho ticks. Journ. Infec. Dis., viii, p. 322-326.</p> + +<p><b>McClintic, T. B.</b> 1912. Investigations of and tick eradications in Rocky Mountain +spotted fever. Publ. Health Repts., Washington, xxvii, p. 732-760.</p> + +<p><b>Meckel, H.</b> 1847. Uber schwarzes Pigment in der Milz und im Blute einer +Geisteskranken. Allgem. Zeitschr. f. Psychiatrie, iv, p. 198-226.</p> + +<p><b>Megni, P.</b> 1906. Les insectes buveurs de sang. 12mo. Paris, Rudeval. +(150 p.).</p> + +<p><b>Melnikoff, N.</b> 1869. Ueber die Jugendzustände der <i>Tænia cucumerina</i>. +Arch. f. Naturg., xxxv, p. 62-70.</p> + +<p><b>Mense, C.</b> 1913. Handbuch der Tropenkrankheiten. 1 Band. 4<sup>o</sup>. Leipzig, +Barth (xv + 295 p.) Entomological parts by A. Eysell, and by Doerr and Russ.</p> + +<p><b>Minchin, E. A.</b> 1912. An introduction to the study of the Protozoa, with special +reference to the parasitic forms. 8<sup>o</sup>. London. Arnold (xi + 517 p.).</p> + +<p><b>Mitchell, Evelyn G.</b> 1907. Mosquito life. 8vo. New York, Putmans. +(xxii + 281 p.).</p> + +<p><b>Mitzmain, M. B.</b> 1910. General observations on the bionomics of the rodent +and human flies. U. S. Publ. Health Service. Bul., 38, p. 1-34.<span class="pagenum"><a name="Page_335" id="Page_335">[Pg 335]</a></span></p> + +<p>---- 1912. The rôle of <i>Stomoxys calcitrans</i> in the transmission of <i>Trypanosoma +evansi</i>. Philippine Journ. Sci., vii, p. 475-519, 5 pls.</p> + +<p>---- 1913 a. The biology of <i>Tabanus striatus</i> Fabricius, the horsefly of the +Philippines. ibid., vii, B. p. 197-221.</p> + +<p>---- 1913 b. The mechanical transmission of surra. ibid., viii, sec. B., +p. 223-229.</p> + +<p>---- 1914 a. Experimental insect transmission of anthrax. U. S. Public +Health Repts. xxix, p. 75-77.</p> + +<p>---- 1914 b. I. Collected studies on the insect transmission of <i>Trypanosoma +evansi</i>. II. Summary of experiments in the transmission of anthrax +by biting flies. U. S. Pub. Health Service, Hyg. Lab. Bul., 94, p. 1-48.</p> + +<p><b>Miyake, H.</b> and <b>Scriba, J.</b> 1893. Vorläufige Mitteilung über einen neuen +Parasit des Menschen. Berl. klin. Wochenschr., xxx, p. 374.</p> + +<p><b>Mollers, B.</b> 1907. Experimentelle Studien über die Uebertragung des Rückfallfiebers +durch Zecken. Zeitschr. für Hyg. u. Infektionskrankheiten, lviii, +p. 277-286.</p> + +<p><b>Mote, D. C.</b> 1914. The cheese-skipper (<i>Piophila casei</i>). Ohio Naturalist xiv, p. 309-310.</p> + +<p><b>Neiva, A.</b> 1910. Beiträge zur Biologie der <i>Conorhinus megistus</i> Burm. Memorias +de Institute Oswaldo Cruz., ii, p. 206-212.</p> + +<p><b>Neveu-Lemaire, M.</b> 1907. Un nouveau cas de parasitisme accidental d'un +myriapode dans le tube digestif de l'homme, C. R. Soc. der Biol., lxiii +p. 305-308.</p> + +<p>---- 1908. Précis de parasitologie humaine. 8vo. Paris, Rudeval. (v + +712 p.).</p> + +<p><b>Newstead, R.</b> 1911. The papataci flies (Phlebotomus) of the Maltese Islands. +Bul. of Ent. Research, ii, p. 47-78, pls. 1-3.</p> + +<p><b>Nicoll, W.</b> 1911. On the part played by flies in the disposal of the eggs of +parasitic worms. Repts. to the Local Gov't. Board on Publ. Health and Med. +Subjects, n. s. No. 53, p. 13-30.</p> + +<p><b>Nicolle, C.</b> 1910, Recherches expérimentales sur la typhus exanthématique +entreprises à l'Institut Pasteur de Tunis pendant l'année 1909. Ann. Inst. +Pasteur, xxiv, p. 243-275.</p> + +<p>---- 1911. Recherches expérimentales sur la typhus exanthématique +entreprises à l'Institut Pasteur de Tunis pendant l'année 1910. ibid., xxv, +p. 1-55, 97-154.</p> + +<p><b>Nicolle, C.</b>, <b>Blaizot, A.</b>, and <b>Conseil, E.</b> 1912 a. Étiologie de la fièvre récurrente. +Son mode de transmission par le pou. C. R. Acad. Sci., cliv, p. 1636-1638.</p> + +<p>---- 1912 b. Conditions de transmission de la fièvre récurrente par le pou. +ibid., clv., p. 481-484.</p> + +<p><b>Nicolle, C.</b> and <b>Catouillard, G.</b> 1905. Sur le venin d'un scorpion commun de +Tunisie (<i>Heterometrus maurus</i>). C. R. Soc. Biol. lviii: p. 100-102.</p> + +<p><b>Noe, G.</b> 1901. Sul ciclo evolutivo della <i>Filaria bancrofti</i> e delta <i>Filaria immitis</i>. +Ricerche labr. anat. comp. norm. Univ. di Roma., viii, p. 275-353.</p> + +<p><b>Norman, W. W.</b> 1896. The effect of the poison of centipedes. Trans. Texas +Acad. Sci., i, p. 118-119.</p> + +<p><b>Nuttall, G. H. F.</b> 1899. On the rôle of insects, arachnids, and myriapods as +carriers in the spread of bacterial and parasitic diseases of man and animals. +Johns Hopkins Hosp. Repts., viii, 154 p., 3 pls.</p> + +<p>---- 1908 a. On the behavior of Spirochætæ in <i>Acanthia lectularia</i>. Parasitology, +i, p. 143-151.</p> + +<p>---- 1908 b. The transmission of <i>Trypanosoma lewisi</i> by fleas and lice. +ibid., i, p. 296-301.</p> + +<p>---- 1908 c. The Ixodoidea or ticks, spirochætosis in man and animals, +piroplasmosis. Journ. Roy. Inst. Publ. Health, xvi, p. 385-403, 449-464, +513-526.</p> + +<p>---- 1914. Tick paralysis in man and animals. 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Review of work by Pantel and Portchinski on reproductive and +early stage characters of Muscoid flies. Proc. Ent. Soc., Washington, xiii, +p. 151-170.</p> + +<p>---- 1912. Muscoid names. ibid., xiv, p. 45.</p> + +<p>---- 1913 a. Preliminary characterization of the vector of verruga, <i>Phlebotomus +verrucarum</i> sp. nov. Insecutor Inscitiæ Menstruus, Washington, i, +p. 107-109.</p> + +<p>---- 1913 b. The transmission of verruga by <i>Phlebotomus</i>. Journ. Amer. +Med. Assoc., lxi, p. 1717.<span class="pagenum"><a name="Page_340" id="Page_340">[Pg 340]</a></span></p> + +<p>---- 1914 a. The relations between lizards and <i>Phlebotomus verrucarum</i> +as indicating the reservoir of verruga. Science n. s., xl, p. 212-214.</p> + +<p>---- 1914 b. Progress of verruga work with <i>Phlebotomus verrucarum</i> T. +Journ. Econ. Ent., vii. p. 357-367.</p> + +<p><b>Trouessart, E.</b> 1902. 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Manual of the North American Diptera, New Haven, +p. 1-405.</p> + +<p><b>Wilson, G. B.</b> and <b>Chowning, W. M.</b> 1903. Studies in <i>Piroplasmosis hominis</i>. +Journ. Inf. Dis., iv, p. 31-57.</p> + +<p><b>Wilson, W. H.</b> 1904. On the venom of scorpions. Rec. Egyptian Gov't +School of Medicine, Cairo, ii, p. 7-44.</p></div><p><span class="pagenum"><a name="Page_341" id="Page_341">[Pg 341]</a></span></p> + + + +<hr style="width: 65%;" /> +<h2><a name="INDEX" id="INDEX"></a>INDEX</h2> + + +<pre> +Abscess, <a href="#Page_178">178</a> + +Acanthia, <a href="#Page_87">87</a> + +Acariasis, <a href="#Page_58">58</a> + +Acarina, <a href="#Page_23">23</a>, <a href="#Page_58">58</a>, <a href="#Page_131">131</a>, <a href="#Page_259">259</a> + +Acarus dysenteriæ, <a href="#Page_132">132</a> + +Accidental parasites, <a href="#Page_131">131</a>, <a href="#Page_132">132</a>, <a href="#Page_134">134</a> + +Aedes, <a href="#Page_194">194</a>, <a href="#Page_293">293</a> + +Aedes calopus, <a href="#Page_182">182</a>, <a href="#Page_201">201</a>, <a href="#Page_205">205</a>, <a href="#Page_206">206</a>, <a href="#Page_208">208</a> + +Aedes cantator, <a href="#Page_101">101</a> + +Aedes sollicitans, <a href="#Page_101">101</a> + +Aedes tæniorhynchus, <a href="#Page_101">101</a> + +Aerobic bacteria, <a href="#Page_152">152</a> + +Æstivo-autumnal, <a href="#Page_186">186</a> + +African Relapsing Fever, <a href="#Page_230">230</a> + +Akis spinosa, <a href="#Page_177">177</a> + +Alternation of Generations, <a href="#Page_175">175</a> + +Amblyomma, <a href="#Page_264">264</a> + +Amblyomma americanum, <a href="#Page_67">67</a> + +Amblyomma cajennense, <a href="#Page_67">67</a> + +American dog tick, <a href="#Page_228">228</a> + +Amœboid organism, <a href="#Page_189">189</a> + +Anisolabis annulipes, <a href="#Page_177">177</a> + +Anterior poliomyelitis, <a href="#Page_241">241</a> + +Anopheles, <a href="#Page_194">194</a>, <a href="#Page_291">291</a> + +Anopheles crucians, <a href="#Page_199">199</a> + +Anopheles maculipennis, <a href="#Page_182">182</a> + +Anopheles punctipennis, <a href="#Page_198">198</a> + +Anopheles quadrimaculatus, <a href="#Page_197">197</a> + +Anopheline, <a href="#Page_192">192</a> + +Anthocoris, <a href="#Page_279">279</a> + +Anthomyiidæ, <a href="#Page_300">300</a> + +Anthomyia, <a href="#Page_138">138</a> + +Anthrax, <a href="#Page_165">165</a> + +Antipruritic treatment, <a href="#Page_72">72</a> + +Ants, <a href="#Page_42">42</a> + +Aphiochæta, <a href="#Page_295">295</a> + +Apis mellifica, <a href="#Page_36">36</a> + +Arachnida, <a href="#Page_258">258</a> + +Araneida, <a href="#Page_6">6</a> + +Argas, <a href="#Page_64">64</a> + +Argas persicus, <a href="#Page_63">63</a>, <a href="#Page_235">235</a>, <a href="#Page_237">237</a> + +Argasidæ, <a href="#Page_62">62</a> + +Argopsylla, <a href="#Page_317">317</a> + +Argus, <a href="#Page_259">259</a> + +Arilus, <a href="#Page_284">284</a> + +Arthropods, poisonous, <a href="#Page_6">6</a> + +Asopia farinalis, <a href="#Page_177">177</a> + +Assassin-bugs, <a href="#Page_31">31</a>, <a href="#Page_219">219</a> + +Auchmeromyia, <a href="#Page_117">117</a> + +Automeris io, <a href="#Page_47">47</a> + +Avicularoidea, <a href="#Page_12">12</a> + + +Babesia, <a href="#Page_226">226</a> + +Babesia bovis, <a href="#Page_223">223</a> + +Babesia ovis, <a href="#Page_225">225</a> + +Babesiosis, <a href="#Page_221">221-222</a> + +Bacilli, <a href="#Page_170">170</a> + +Bacillus icteroides, <a href="#Page_202">202</a>, <a href="#Page_205">205</a> + +Bacillus pestis, <a href="#Page_166">166</a> + +Bacillus typhosus, <a href="#Page_153">153</a> + +Back swimmers, <a href="#Page_30">30</a> + +Bdellolarynx, <a href="#Page_304">304</a> + +Beauperthuy, Louis Daniel, <a href="#Page_2">2</a> + +Bed-bug, <a href="#Page_86">86</a>, <a href="#Page_88">88</a>, <a href="#Page_90">90</a>, <a href="#Page_173">173</a>, <a href="#Page_219">219-220</a> + +Bed-bug, cone-nosed, <a href="#Page_92">92</a> + +Blister beetles, <a href="#Page_54">54</a> + +Belostoma, <a href="#Page_28">28</a>, <a href="#Page_277">277</a> + +Belostoma americana, <a href="#Page_31">31</a> + +Belostomatidæ, <a href="#Page_30">30</a> + +Bengalia, <a href="#Page_314">314</a> + +Bird-spiders, <a href="#Page_10">10</a> + +Black death, <a href="#Page_1">1</a>, <a href="#Page_166">166</a> + +Black flies, <a href="#Page_33">33</a>, <a href="#Page_104">104</a>, <a href="#Page_247">247</a> + +Black heads, <a href="#Page_80">80</a> + +Blaps mortisaga, <a href="#Page_134">134</a> + +Blepharoceridæ, <a href="#Page_286">286</a> + +Boophilus, <a href="#Page_264">264</a> + +Boophilus annulatus, <a href="#Page_67">67</a>, <a href="#Page_223">223-225</a> + +Bot-flies, <a href="#Page_112">112</a> + +Blue bottle flies, <a href="#Page_140">140</a> + +Brill's disease, <a href="#Page_238">238</a> + +Brown-tailed moth, <a href="#Page_48">48</a> + +Bruck, <a href="#Page_34">34</a> + +Buthus quinquestriatus, <a href="#Page_21">21</a> + + +Cabbage butterfly, <a href="#Page_56">56</a> + +Calliphora, <a href="#Page_136">136</a>, <a href="#Page_140">140</a>, <a href="#Page_312">312</a> + +Calliphora erythrocephala, <a href="#Page_141">141</a><span class="pagenum"><a name="Page_342" id="Page_342">[Pg 342]</a></span> + +Calobata, <a href="#Page_296">296</a> + +Camponotinæ, <a href="#Page_43">43</a> + +Cancer, <a href="#Page_254">254</a> + +Cantharidin, <a href="#Page_54">54</a> + +Cantharidin poison, <a href="#Page_55">55</a> + +Canthariasis, <a href="#Page_134">134</a> + +Capsidæ, <a href="#Page_280">280</a> + +Carriers, simple, <a href="#Page_4">4</a>, <a href="#Page_144">144</a> + +Carriers of disease, <a href="#Page_144">144</a> + +Carrion's fever, <a href="#Page_253">253</a> + +Caterpillar rash, <a href="#Page_45">45</a> + +Cat flea, <a href="#Page_172">172</a> + +Cattle ticks, <a href="#Page_222">222</a> + +Causative organism, <a href="#Page_170">170</a> + +Cellia, <a href="#Page_291">291</a> + +Centipedes, <a href="#Page_25">25</a>, <a href="#Page_257">257</a> + +Ceratophyllus, <a href="#Page_120">120</a>, <a href="#Page_316">316</a> + +Ceratophyllus acutus, <a href="#Page_123">123</a> + +Ceratophyllus fasciatus, <a href="#Page_122">122</a>, <a href="#Page_172">172</a>, <a href="#Page_213">213</a> + +Ceratopogon, <a href="#Page_108">108</a> + +Cheese-fly, <a href="#Page_137">137</a> + +Cheyletus eruditus, <a href="#Page_271">271</a> + +Chigger, <a href="#Page_60">60</a>, <a href="#Page_70">70</a> + +Chigoes, <a href="#Page_126">126</a> + +Chilopoda, <a href="#Page_25">25</a>, <a href="#Page_257">257</a> + +Chiracanthium nutrix, <a href="#Page_18">18</a> + +Chironomidæ, <a href="#Page_107">107</a> + +Chorioptes, <a href="#Pg_270">270</a> + +Chrysomelid, <a href="#Page_55">55</a> + +Chrysomyia, <a href="#Page_136">136</a>, <a href="#Page_308">308</a> + +Chrysomyia macellaria, <a href="#Page_117">117</a>, <a href="#Page_140">140</a> + +Chrysops, <a href="#Page_294">294</a> + +Chylous dropsy, <a href="#Page_179">179</a> + +Chyluria, <a href="#Page_178">178</a> + +Cicadidæ, <a href="#Page_55">55</a> + +Cimex L., <a href="#Page_278">278</a> + +Cimex boueti, <a href="#Page_92">92</a> + +Cimex columbarius, <a href="#Page_92">92</a> + +Cimex hemipterus, <a href="#Page_91">91</a>, <a href="#Page_220">220</a> + +Cimex hirundinis, <a href="#Page_92">92</a> + +Cimex inodorus, <a href="#Page_92">92</a> + +Cimex lectularius, <a href="#Page_87">87</a>, <a href="#Page_219">219</a> + +Citheronia regalis, <a href="#Page_44">44</a> + +Clinocoris, <a href="#Page_87">87</a> + +Coleoptera, <a href="#Page_134">134</a>, <a href="#Page_274">274</a> + +Comedons, <a href="#Page_80">80</a> + +Complete metamorphosis, <a href="#Page_80">80</a> + +Compressor muscle, <a href="#Page_20">20</a> + +Compsomyia, <a href="#Page_117">117</a> + +Cone-nosed bed-bug, <a href="#Page_92">92</a> + +Conjunctivitis, nodular, <a href="#Page_52">52</a> + +Conorhinus, <a href="#Page_282">282</a> + +Conorhinus megistus, <a href="#Page_93">93</a>, <a href="#Page_219">219-220</a> + +Conorhinus rubrofasciatus, <a href="#Page_220">220</a> + +Conorhinus sanguisugus, <a href="#Page_32">32</a>, <a href="#Page_92">92</a> + +Copra itch, <a href="#Page_72">72</a> + +Cordylobia, <a href="#Page_118">118</a> + +Coriscus, <a href="#Page_280">280</a> + +Coriscus subcoleoptratus, <a href="#Page_32">32</a> + +Creeping myasis, <a href="#Page_112">112</a> + +Crustacea, <a href="#Page_257">257</a> + +Cryptocystis, <a href="#Page_176">176</a> + +Cryptotoxic, <a href="#Page_54">54-55</a> + +Cteniza sauvagei, <a href="#Page_13">13</a> + +Ctenocephalus, <a href="#Page_120">120</a>, <a href="#Page_172">172</a>, <a href="#Page_213">213</a>, <a href="#Page_317">317</a> + +Culex, <a href="#Page_194">194</a>, <a href="#Page_201">201</a>, <a href="#Page_293">293</a> + +Culex pipiens, <a href="#Page_35">35</a>, <a href="#Page_98">98</a> + +Culex quinquefasciatus, <a href="#Page_180">180</a> + +Culex sollicitans, <a href="#Page_200">200</a> + +Culex territans, <a href="#Page_101">101</a> + +Culicidæ, <a href="#Page_33">33</a>, <a href="#Page_97">97</a> + +Culicin, <a href="#Page_34">34</a> + +Culicoides, <a href="#Page_109">109</a>, <a href="#Page_288">288</a> + +Cyclops, <a href="#Page_183">183</a>, <a href="#Page_257">257</a> + +Cynomyia, <a href="#Page_136">136</a>, <a href="#Page_311">311</a> + + +Dance, St. Vitus, <a href="#Page_8">8</a> + +Dancing mania, <a href="#Page_8">8</a> + +Deer-flies, <a href="#Page_110">110</a> + +Definitive host, <a href="#Page_192">192</a> + +Demodecidæ, <a href="#Page_78">78</a> + +Demodex, <a href="#Page_259">259</a> + +Demodex folliculorum, <a href="#Page_78">78</a> + +Dermacentor, <a href="#Page_262">262</a> + +Dermacentor andersoni, <a href="#Page_67">67</a>, <a href="#Page_228">228</a> + +Dermacentor occidentalis, <a href="#Page_227">227</a> + +Dermacentor variabilis, <a href="#Page_67">67</a> + +Dermacentor venustus, <a href="#Page_24">24</a>, <a href="#Page_228">228</a> + +Dermanyssidæ, <a href="#Page_68">68</a> + +Dermanyssus, <a href="#Pg_266">266</a> + +Dermanyssus gallinæ, <a href="#Page_68">68</a> + +Dermatitis, <a href="#Page_72">72</a>, <a href="#Page_77">77</a>, <a href="#Page_85">85</a> + +Dermatobia, <a href="#Page_115">115</a>, <a href="#Page_298">298</a> + +Dermatobia cyaniventris, <a href="#Page_163">163</a> + +Dermatophilus, <a href="#Page_317">317</a> + +Dermatophilus penetrans, <a href="#Page_60">60</a>, <a href="#Page_126">126</a><span class="pagenum"><a name="Page_343" id="Page_343">[Pg 343]</a></span> + +Diamphidia simplex, <a href="#Page_55">55</a> + +Dimorphism, <a href="#Page_65">65</a> + +Direct inoculators, <a href="#Page_4">4</a> + +Diplopoda, <a href="#Page_25">25</a>, <a href="#Page_257">257</a> + +Diptera, <a href="#Page_33">33</a>, <a href="#Page_94">94</a>, <a href="#Page_274">274</a> + +Dipterous Larvæ, <a href="#Page_135">135</a> + +Dipylidium, <a href="#Page_175">175</a>, <a href="#Page_221">221</a> + +Dipylidium caninum, <a href="#Page_4">4</a>, <a href="#Page_175">175-176</a> + +Dog flea, <a href="#Page_172">172</a> + +Dracunculus, <a href="#Page_257">257</a> + +Dracunculus medinensis, <a href="#Page_182">182</a> + +Drosophila, <a href="#Page_296">296</a> + +Dum-dum fever, <a href="#Page_220">220</a> + +Dysentery, <a href="#Page_154">154</a> + + +Ear-flies, <a href="#Page_110">110</a> + +Earwig, <a href="#Page_177">177</a> + +Echidnophaga, <a href="#Page_317">317</a> + +Echinorhynchus, <a href="#Page_185">185</a> + +Elephantiasis, <a href="#Page_178">178-179</a> + +Empoasca mali, <a href="#Page_33">33</a> + +Empretia, <a href="#Page_46">46</a> + +English Plague Commission, <a href="#Page_171">171</a> + +Epeira diadema, <a href="#Page_18">18</a> + +Epizootic, <a href="#Page_170">170</a> + +Eristalis, <a href="#Page_137">137</a>, <a href="#Page_295">295</a> + +Essential hosts, <a href="#Page_4">4</a>, <a href="#Page_165">165</a> + +Eumusca, <a href="#Page_307">307</a> + +European Relapsing Fever, <a href="#Page_233">233</a> + +Euproctis chrysorrhœa, <a href="#Page_48">48</a> + +Eusimulium, <a href="#Page_286">286</a> + + +Facultative parasites, <a href="#Page_131">131</a> + +Fannia, <a href="#Page_136">136</a>, <a href="#Page_138">138</a>, <a href="#Page_145">145</a>, <a href="#Page_300">300</a> + +Federal Health Service, <a href="#Page_169">169</a> + +Fever, lenticular, <a href="#Page_237">237</a> + African Relapsing, <a href="#Page_230">230</a>, <a href="#Page_234">234</a> + Carrion's, <a href="#Page_253">253</a> + dum-dum, <a href="#Page_154">154</a> + European Relapsing, <a href="#Page_233">233</a> + pappatici, <a href="#Page_96">96</a> + red water, <a href="#Page_220">220</a> + Rocky Mt. Spotted, <a href="#Page_226">226</a> + three day, <a href="#Page_96">96</a> + Typhus, <a href="#Page_237">237</a> + +Filaria, <a href="#Page_178">178</a>, <a href="#Page_221">221</a> + immitis, <a href="#Page_182">182</a> + +Filariasis, <a href="#Page_178">178</a> + +Flannel-moth larvæ, <a href="#Page_44">44</a> + +Fleas, <a href="#Page_119">119</a>, <a href="#Page_166">166</a>, <a href="#Page_213">213</a> + cat, <a href="#Page_172">172</a> + dog, <a href="#Page_172">172</a> + human, <a href="#Page_172">172</a>, <a href="#Page_176">176</a> + rodent, <a href="#Page_123">123</a>, <a href="#Page_172">172</a> + rat, <a href="#Page_171">171</a> + +Flesope, <a href="#Page_125">125</a> + +Formaldehyde, <a href="#Page_91">91</a> + +Fomites, <a href="#Page_199">199</a>, <a href="#Page_204">204</a> + +Fulgoridæ, <a href="#Page_28">28</a> + +Fumigation, <a href="#Page_320">320</a> + + +Gamasid, <a href="#Page_68">68</a> + +Gangrene, <a href="#Page_129">129</a> + +Gastrophilus, <a href="#Page_113">113</a>, <a href="#Page_297">297</a> + +Giant crab spiders, <a href="#Page_13">13</a> + +Giant water bugs, <a href="#Page_30">30</a> + +Gigantorhynchus, <a href="#Page_185">185</a> + +Glossina, <a href="#Page_117">117</a>, <a href="#Page_297">297</a>, <a href="#Page_303">303</a> + +Glossina morsitans, <a href="#Page_214">214</a>, <a href="#Page_217">217</a> + palpalis, <a href="#Page_215">215</a>, <a href="#Page_217">217</a>, <a href="#Page_218">218</a> + +Glyciphagus, <a href="#Page_267">267</a> + +Grain moth, <a href="#Page_69">69</a> + +Grocer's itch, <a href="#Page_72">72</a> + +Guinea-worm, <a href="#Page_182">182</a> + + +Habronema muscæ, <a href="#Page_156">156</a>, <a href="#Page_183">183</a> + +Hæmatobia, <a href="#Page_166">166</a>, <a href="#Page_304">304</a> + irritans, <a href="#Page_146">146</a> + +Hæmatobosca, <a href="#Page_304">304</a> + +Hæmatomyidium, <a href="#Page_288">288</a> + +Hæmatopinus spinulosus, <a href="#Page_213">213</a> + +Hæmatopota, <a href="#Page_294">294</a> + +Hæmatosiphon, <a href="#Page_279">279</a> + +Hæmoglobinuria, <a href="#Page_222">222</a> + +Hæmozoin, <a href="#Page_189">189</a> + +Harpactor, <a href="#Page_284">284</a> + +Harvest mites, <a href="#Page_60">60</a> + effect of, <a href="#Page_59">59</a> + +Head-louse, <a href="#Page_173">173</a> + +Helminthiasis, <a href="#Page_138">138</a> + +Helophilus, <a href="#Page_295">295</a> + +Hemiptera, <a href="#Page_27">27</a>, <a href="#Page_86">86</a>, <a href="#Page_273">273-275</a> + +Heteropodidæ, <a href="#Page_13">13</a> + +Heuchis sanguinea, <a href="#Page_55">55</a> + +Hexapod larvæ, <a href="#Page_58">58</a> + +Hexapoda, <a href="#Page_27">27</a>, <a href="#Page_80">80</a>, <a href="#Page_258">258</a><span class="pagenum"><a name="Page_344" id="Page_344">[Pg 344]</a></span> + +Hippelates, <a href="#Page_297">297</a> + +Hippobosca, <a href="#Page_285">285</a> + +Histiogaster, <a href="#Page_269">269</a> + spermaticus, <a href="#Page_132">132</a> + +Homalomyia, <a href="#Page_136">136</a>, <a href="#Page_138">138</a>, <a href="#Page_300">300</a> + +Honey bee, <a href="#Page_36">36</a> + poison of, <a href="#Page_37">37</a> + +Hornets, <a href="#Page_43">43</a> + +Horn-fly, <a href="#Page_137">137</a>, <a href="#Page_304">304</a>, <a href="#Page_308">308</a> + +Horse-fly, <a href="#Page_110">110</a>, <a href="#Page_165">165</a> + +House-fly, <a href="#Page_137">137-139</a>, <a href="#Page_144">144</a>, <a href="#Page_183">183</a> + control of, <a href="#Page_156">156</a>, <a href="#Page_160">160</a> + +Human flea, <a href="#Page_124">124</a> + +Host, definitive, <a href="#Page_175">175</a> + intermediate, <a href="#Page_175">175</a> + primary, <a href="#Page_175">175</a> + +Hyalomma, <a href="#Page_264">264</a> + ægypticum, <a href="#Page_224">224-225</a> + +Hydrocyanic Acid Gas, <a href="#Page_318">318</a> + +Hydrotæa, <a href="#Page_300">300</a> + +Hymenolepis diminuta, <a href="#Page_176">176</a> + +Hymenoptera, <a href="#Page_36">36</a>, <a href="#Page_275">275</a> + +Hypoderma, <a href="#Page_113">113</a>, <a href="#Page_298">298</a> + diana, <a href="#Page_113">113</a> + lineata, <a href="#Page_113">113</a> + +Hypopharynx, <a href="#Page_80">80</a> + + +Immunity from stings, <a href="#Page_39">39</a> + +Incomplete metamorphosis, <a href="#Page_80">80</a> + +Infantile paralysis, <a href="#Page_162">162</a>, <a href="#Page_241">241</a> + splenic, <a href="#Page_220">220</a> + Direct inoculation, <a href="#Page_164">164</a> + +Insects, <a href="#Page_258">258</a> + blood-sucking, <a href="#Page_170">170</a> + +Intermediate host, <a href="#Page_192">192</a>, <a href="#Page_203">203</a> + +Intestinal infestation, <a href="#Page_112">112</a>, <a href="#Page_133">133</a> + myasis, <a href="#Page_137">137</a> + +Isosoma, <a href="#Page_69">69</a> + +Itch, <a href="#Page_73">73-74</a> + mite, <a href="#Page_73">73</a> + Norwegian, <a href="#Page_77">77</a> + +Ixodes, <a href="#Pg_260">260</a> + ricinus, <a href="#Page_66">66</a>, <a href="#Page_225">225</a> + scapularis, <a href="#Page_66">66</a> + +Ixodidæ, <a href="#Page_64">64-65</a> + +Ixodoidea, <a href="#Page_62">62</a> + + +Janthinosoma lutzi, <a href="#Page_116">116</a> + +Jigger, <a href="#Page_60">60</a> + +Johannseniella, <a href="#Page_110">110</a>, <a href="#Page_288">288</a> + +Journal of Tropical Medicine and Hygiene, <a href="#Page_36">36</a> + +Julus terrestris, <a href="#Page_25">25</a> + +June bug, <a href="#Page_185">185</a> + + +Kala-azar, <a href="#Page_220">220</a> + +Karakurte, <a href="#Page_14">14</a> + +Katipo, <a href="#Page_14">14</a> + +King, A. F. A., <a href="#Page_3">3</a> + +Kircher, Athanasius, <a href="#Page_1">1</a>, <a href="#Page_8">8</a> + +Kissing-bug, <a href="#Page_31">31</a> + + +Labium, <a href="#Page_29">29</a>, <a href="#Page_80">80</a> + +Labrum, <a href="#Page_28">28</a>, <a href="#Page_80">80</a> + +Lachnosterna, <a href="#Page_185">185</a> + +Lælaps, <a href="#Pg_266">266</a> + +Lagoa crispata, <a href="#Page_45">45</a> + +Lamblia intestinalis, <a href="#Page_154">154</a> + +Langer, Josef, <a href="#Page_37">37</a> + +Larder beetles, <a href="#Page_135">135</a> + +Latrodectus, <a href="#Page_12">12</a>, <a href="#Page_14">14</a>, <a href="#Page_17">17</a> + mactans, <a href="#Page_15">15</a> + +Leishmanioses, <a href="#Page_220">220</a> + +Lenticular fever, <a href="#Page_237">237</a> + +Lepidoptera, <a href="#Page_274">274</a> + +Lepidopterous larvæ, <a href="#Page_134">134</a> + +Leprosy, <a href="#Page_252">252</a> + +Leptidæ, <a href="#Page_112">112</a> + +Leptis, <a href="#Page_295">295</a> + +Leptus, <a href="#Page_60">60</a>, <a href="#Page_273">273</a> + +Lice, <a href="#Page_80">80</a> + +Linguatulina, <a href="#Page_258">258</a> + +Liponyssus, <a href="#Page_265">265</a> + +Lœmopsylla, <a href="#Page_172">172</a>, <a href="#Page_317">317</a> + +Lone star tick, <a href="#Page_228">228</a> + +Louse, body, <a href="#Page_84">84</a> + crab, <a href="#Page_85">85</a> + dog, <a href="#Page_176">176</a> + head, <a href="#Page_82">82</a> + pubic, <a href="#Page_85">85</a> + +Lucilia, <a href="#Page_136">136</a>, <a href="#Page_312">312</a> + +Lycosa tarantula, <a href="#Page_10">10</a> + +Lycosidæ, <a href="#Page_10">10</a> + +Lyctocoris, <a href="#Page_279">279</a> + +Lygus pratensis, <a href="#Page_33">33</a><span class="pagenum"><a name="Page_345" id="Page_345">[Pg 345]</a></span> + +Lymphangitis, <a href="#Page_67">67</a> + +Lymph scrotum, <a href="#Page_178">178</a> + +Lyperosia, <a href="#Page_304">304</a> + +Lyperosiops, <a href="#Page_305">305</a> + + +Macloskie, <a href="#Page_34">34</a> + +Maggots, rat-tail, <a href="#Page_137">137</a> + +Magnes sive de Arte Magnetica, <a href="#Page_8">8</a> + +Malaria, <a href="#Page_186">186</a> + +Malmigniatte, <a href="#Page_14">14</a> + +Mandibles, <a href="#Page_28">28</a>, <a href="#Page_80">80</a> + +Mange, <a href="#Page_73">73-75</a> + +Margaropus, <a href="#Page_237">237</a>, <a href="#Page_264">264</a> + annulatus, <a href="#Page_223">223</a> + +Masked bed-bug hunter, <a href="#Page_32">32</a> + +Mastigoproctus giganteus, <a href="#Page_19">19</a>, <a href="#Page_80">80</a> + +Maxillæ, <a href="#Page_28">28</a> + +Meal infesting species, <a href="#Page_135">135</a> + +Melanin granules, <a href="#Page_189">189</a> + +Melanolestes, <a href="#Page_280">280</a> + picipes, <a href="#Page_32">32</a> + +Mena-vodi, <a href="#Page_14">14</a> + +Mercurialis, <a href="#Page_1">1</a> + +Merozoites, <a href="#Page_190">190</a> + +Metamorphosis, <a href="#Page_80">80</a> + +Miana bug, <a href="#Page_63">63</a> + +Microgametoblast, <a href="#Page_192">192</a> + +Midges, <a href="#Page_107">107</a> + +Migratory ookinete, <a href="#Page_192">192</a> + +Millipedes, <a href="#Page_25">25</a>, <a href="#Page_257">257</a> + +Mites, <a href="#Page_23">23</a>, <a href="#Page_58">58</a> + +Monieziella, <a href="#Page_269">269</a> + +Mosquitoes, <a href="#Page_33">33</a>, <a href="#Page_97">97</a>, <a href="#Page_178">178</a>, <a href="#Page_196">196</a>, <a href="#Page_250">250</a> + treatment for bites of, <a href="#Page_34">34</a>, <a href="#Page_36">36</a>, <a href="#Page_102">102</a> + +Musca, <a href="#Page_137">137</a>, <a href="#Page_307">307</a> + domestica, <a href="#Page_139">139</a>, <a href="#Page_145">145</a>, <a href="#Page_146">146</a>, <a href="#Page_157">157</a>, <a href="#Page_162">162</a> + +Muscidæ, <a href="#Page_117">117</a> + +Muscina, <a href="#Page_137">137</a>, <a href="#Page_146">146</a>, <a href="#Page_307">307</a> + stabulans, <a href="#Page_140">140</a> + +Mutualism, <a href="#Page_57">57</a> + +Myasis, <a href="#Page_112">112</a>, <a href="#Page_135">135</a> + intestinal, <a href="#Page_135">135-140</a> + nasal, <a href="#Page_141">141</a> + +Mycterotypus, <a href="#Page_287">287</a> + +Myiospila, <a href="#Page_146">146</a>, <a href="#Page_307">307</a> + +Myriapoda, <a href="#Page_25">25</a>, <a href="#Page_132">132</a>, <a href="#Page_257">257</a> + + +Nagana, <a href="#Page_165">165</a>, <a href="#Page_214">214</a> + +Nasal infestation, <a href="#Page_114">114</a>, <a href="#Page_133">133</a> + +Necrobia, <a href="#Page_135">135</a> + +Nematode parasite, <a href="#Page_182">182</a> + +Nepa, <a href="#Page_28">28</a> + +Nephrophages sanguinarius, <a href="#Page_132">132</a> + +Nettling insects, <a href="#Page_43">43</a> + larvæ, poison of, <a href="#Page_53">53</a> + +Neurasthenia, <a href="#Page_89">89</a> + +Nits, <a href="#Page_86">86</a> + +North African Relapsing Fever, <a href="#Page_234">234</a> + +Norwegian itch, <a href="#Page_77">77</a> + +No-see-ums, <a href="#Page_109">109</a> + +Notœdres, <a href="#Page_269">269</a> + cati, <a href="#Page_78">78</a> + +Notonecta, <a href="#Page_28">28</a>, <a href="#Page_277">277</a> + +Notonectidæ, <a href="#Page_30">30</a> + +Nott, Dr. Josiah, <a href="#Page_2">2</a> + +Nuttall, <a href="#Page_34">34</a> + + +Occipital headaches, <a href="#Page_138">138</a> + +Oecacta, <a href="#Page_288">288</a> + +Oeciacus, <a href="#Page_279">279</a> + +Œsophageal diverticula, <a href="#Page_35">35</a> + +Oestridæ, <a href="#Page_112">112</a>, <a href="#Page_136">136</a> + +Oestris ovis, <a href="#Page_113">113</a> + +Oestrus, <a href="#Page_298">298</a> + oocyst, <a href="#Page_192">192</a> + ookinete, <a href="#Page_192">192</a> + +Opsicoetes personatus, <a href="#Page_32">32</a> + +Opthalmia, <a href="#Page_155">155</a> + nodosa, <a href="#Page_52">52</a> + +Oriental sore, <a href="#Page_221">221</a> + +Ornithodoros, <a href="#Page_65">65</a>, <a href="#Pg_260">260</a> + moubata, <a href="#Page_220">220</a>, <a href="#Page_230">230</a> + +Orthotylus flavosparsus, <a href="#Page_33">33</a> + +Ornithomyia, <a href="#Page_286">286</a> + +Oroya, <a href="#Page_253">253</a> + +Oscinus, <a href="#Page_297">297</a> + +Otiobius, <a href="#Page_259">259</a> + megnini, <a href="#Page_65">65</a> + +Otodectes, <a href="#Page_271">271</a> + + +Pangonia, <a href="#Page_294">294</a> + +Pappatici fever, <a href="#Page_96">96</a> + +Parasimulium, <a href="#Page_286">286</a> + +Parasite, <a href="#Page_3">3</a>, <a href="#Page_57">57</a>, <a href="#Page_131">131</a>, <a href="#Page_134">134</a>, <a href="#Page_182">182</a> + accidental, <a href="#Page_3">3</a>, <a href="#Page_131">131</a>, <a href="#Page_134">134</a> + facultative, <a href="#Page_3">3</a>, <a href="#Page_57">57</a>, <a href="#Page_131">131</a><span class="pagenum"><a name="Page_346" id="Page_346">[Pg 346]</a></span> + nematode, <a href="#Page_182">182</a> + stationary, <a href="#Page_57">57</a> + temporary, <a href="#Page_57">57</a> + true, <a href="#Page_3">3</a> + +Parasitism, accidental, <a href="#Page_134">134</a> + +Pathogenic bacteria, <a href="#Page_152">152</a> + organisms, <a href="#Page_144">144</a>, <a href="#Page_164">164</a> + +Pawlowsky, <a href="#Page_81">81</a> + +Pediculoides, <a href="#Page_267">267</a> + ventricosus, <a href="#Page_69">69</a>, <a href="#Page_72">72</a> + +Pediculosis, <a href="#Page_81">81</a> + +Pediculus, <a href="#Page_275">275</a> + corporis, <a href="#Page_84">84</a>, <a href="#Page_233">233</a>, <a href="#Page_238">238</a> + humanus, <a href="#Page_82">82</a>, <a href="#Page_173">173</a> + +Pellagra, <a href="#Page_162">162</a>, <a href="#Page_246">246</a> + +Pernicious fever, <a href="#Page_186">186</a> + +Pest, <a href="#Page_166">166</a> + +Phidippus audax, <a href="#Page_19">19</a> + +Philæmatomyia, <a href="#Page_306">306</a> + +Phisalix, <a href="#Page_13">13</a>, <a href="#Page_43">43</a> + +Phlebotomus, <a href="#Page_289">289</a> + papatasii, <a href="#Page_94">94</a> + verrucarum, <a href="#Page_254">254</a> + vexator, <a href="#Page_95">95</a> + +Phora, <a href="#Page_295">295</a> + +Phormia, <a href="#Page_136">136</a> + +Phormictopus carcerides, <a href="#Page_13">13</a> + +Phthirus pubis, <a href="#Page_85">85</a>, <a href="#Page_275">275</a> + +Phortica, <a href="#Page_296">296</a> + +Pieris brassicæ, <a href="#Page_56">56</a> + +Piophila, <a href="#Page_297">297</a> + +Piophila casei, <a href="#Page_136">136</a>, <a href="#Page_137">137</a> + +Piroplasmosis, <a href="#Page_222">222</a> + +Plague, <a href="#Page_166">166</a> + bubonic, <a href="#Page_166">166</a>, <a href="#Page_169">169</a>, <a href="#Page_170">170</a> + pneumonic, <a href="#Page_167">167</a> + +Plasmodium, <a href="#Page_186">186</a> + +Platymetopius acutus, <a href="#Page_33">33</a> + +Plica palonica, <a href="#Page_83">83</a> + +Pneumonic, <a href="#Page_166">166</a> + plague, <a href="#Page_167">167</a>, <a href="#Page_173">173</a> + +Poisoning by nettling larvæ, <a href="#Page_53">53</a> + +Poison of spiders, <a href="#Page_7">7</a> + +Pollenia, <a href="#Page_308">308</a> + rudis, <a href="#Page_146">146</a>, <a href="#Page_147">147</a> + +Primary gland, <a href="#Page_28">28</a> + +Prionurus citrinus, <a href="#Page_20">20</a> + +Prosimulium, <a href="#Page_286">286</a> + +Protocalliphora, <a href="#Page_136">136</a>, <a href="#Page_312">312</a> + +Protozoan blood parasite, <a href="#Page_165">165</a> + +Pseudo-tubercular, <a href="#Page_52">52</a> + +Psorophora, <a href="#Page_293">293</a> + +Psoroptes, <a href="#Pg_270">270</a> + +Psychodidæ, <a href="#Page_94">94</a> + +Pulex, <a href="#Page_120">120</a>, <a href="#Page_124">124</a>, <a href="#Page_126">126</a>, <a href="#Page_172">172</a>, <a href="#Page_317">317</a> + cheopis, <a href="#Page_172">172</a> + irritans, <a href="#Page_124">124</a> + penetrans, <a href="#Page_126">126</a> + serraticeps, <a href="#Page_120">120</a> + +Pulvillus, <a href="#Page_150">150</a> + +Punkies, <a href="#Page_109">109</a> + +Pycnosoma, <a href="#Page_308">308</a> + + +Rasahus, <a href="#Page_280">280</a> + thoracicus, <a href="#Page_32">32</a> + +Rat fleas, <a href="#Page_120">120</a>, <a href="#Page_124">124</a>, <a href="#Page_171">171</a> + +Rat louse, <a href="#Page_213">213</a> + +Red bugs, <a href="#Page_70">70-72</a> + +Reduviidæ, <a href="#Page_31">31</a> + +Reduviolus, <a href="#Page_280">280</a> + +Reduvius, <a href="#Page_282">282</a> + personatus, <a href="#Page_32">32</a> + +Redwater fever, <a href="#Page_222">222</a> + +Relapsing fever, <a href="#Page_230">230</a>, <a href="#Page_233">233</a> + +Rhiœstrus nasalis, <a href="#Page_115">115</a> + +Rhipicentor, <a href="#Page_264">264</a> + +Rhipicephalus, <a href="#Page_264">264</a> + +Rhizoglyphus, <a href="#Page_269">269</a> + +Rhodnius, <a href="#Page_281">281</a> + +Rocky Mountain Spotted Fever, <a href="#Page_226">226</a> + spotted fever tick, <a href="#Page_67">67</a> + +Russian gad-fly, <a href="#Page_115">115</a> + + +St. Vitus's or St. John's dance, <a href="#Page_8">8</a> + +Salivary syringe, <a href="#Page_28">28</a> + +Sand-flies, <a href="#Page_109">109</a>, <a href="#Page_250">250</a> + +Sanguinetti, <a href="#Page_11">11</a> + +Sarcophaga, <a href="#Page_136">136</a>, <a href="#Page_142">142</a>, <a href="#Page_143">143</a> + +Sarcophila, <a href="#Page_302">302</a> + +Sarcopsylla, <a href="#Page_317">317</a> + penetrans, <a href="#Page_126">126</a> + +Sarcoptes, <a href="#Pg_270">270</a> + minor, <a href="#Page_78">78</a> + scabiei, <a href="#Page_73">73</a> + +Sarcoptidæ, <a href="#Page_72">72</a> + +Scabies, <a href="#Page_172">72</a>, <a href="#Page_73">73</a>, <a href="#Page_74">74</a>, <a href="#Page_75">75</a><span class="pagenum"><a name="Page_347" id="Page_347">[Pg 347]</a></span> + +Scaurus striatus, <a href="#Page_177">177</a> + +Schaudinn, <a href="#Page_34">34</a> + +Schizont, <a href="#Page_189">189</a>, <a href="#Page_190">190</a> + +Scholeciasis, <a href="#Page_134">134</a> + +Scolopendra morsitans, <a href="#Page_26">26</a> + +Scorpions, <a href="#Page_20">20</a> + poison of, <a href="#Page_21">21</a> + +Screw worm fly, <a href="#Page_140">140</a> + +Sepsidæ, <a href="#Page_296">296</a> + +Sepsis, <a href="#Page_136">136</a>, <a href="#Page_297">297</a> + +Shipley, <a href="#Page_34">34</a> + +Sibine, <a href="#Page_46">46</a> + +Silvius, <a href="#Page_294">294</a> + +Simple carriers, <a href="#Page_4">4</a>, <a href="#Page_144">144</a> + +Simuliidæ, <a href="#Page_33">33</a>, <a href="#Page_104">104</a> + +Simulium, <a href="#Page_247">247</a>, <a href="#Page_249">249</a>, <a href="#Page_286">286</a>, <a href="#Page_321">321</a> + pictipes, <a href="#Page_104">104</a> + +Siphonaptera, <a href="#Page_119">119</a>, <a href="#Page_274">274</a>, <a href="#Page_316">316</a> + +Siphunculata, <a href="#Page_80">80</a>, <a href="#Page_275">275</a> + +Sitotroga cerealella, <a href="#Page_69">69</a> + +Skippers, <a href="#Page_137">137</a> + +Sleeping sickness, <a href="#Page_166">166</a>, <a href="#Page_215">215</a> + +Snipe-flies, <a href="#Page_112">112</a> + +Solpugida, <a href="#Page_22">22</a> + +Spanish fly, <a href="#Page_54">54</a> + +Spermatozoa, <a href="#Page_192">192</a> + +Spinose ear-tick, <a href="#Page_65">65</a> + +Spirochæta, <a href="#Page_35">35</a> + berberi, <a href="#Page_234">234</a> + duttoni, <a href="#Page_234">234</a> + +Spirochætosis, <a href="#Page_235">235</a> + +Sporozoite, <a href="#Page_189">189</a> + +Spotted fever, <a href="#Page_67">67</a>, <a href="#Page_226">226</a> + +Squirrel flea, <a href="#Page_123">123</a> + +Stable-fly, <a href="#Page_137">137</a>, <a href="#Page_160">160</a>, <a href="#Page_163">163</a>, <a href="#Page_165">165</a> + +Stegomyia, <a href="#Page_182">182</a>, <a href="#Page_293">293</a> + calopus, <a href="#Page_206">206</a> + fasciata, <a href="#Page_206">206</a> + +Stomoxys, <a href="#Page_137">137</a>, <a href="#Page_305">305</a> + calcitrans, <a href="#Page_117">117</a>, <a href="#Page_146">146</a>, <a href="#Page_160">160</a>, <a href="#Page_161">161</a>, <a href="#Page_165">165</a>, <a href="#Page_242">242</a> + +Straw-worm, <a href="#Page_69">69</a> + +Stygeromyia, <a href="#Page_305">305</a> + +Sucking stomach, <a href="#Page_35">35</a> + +Sulphur ointment, <a href="#Page_77">77</a> + +Surra, <a href="#Page_165">165</a> + +Symbiosis, <a href="#Page_57">57</a> + +Symphoromyia, <a href="#Page_112">112</a>, <a href="#Page_295">295</a> + + +Tabanidæ, <a href="#Page_110">110</a> + +Tabanus, <a href="#Page_110">110</a>, <a href="#Page_166">166</a>, <a href="#Page_294">294</a> + striatus, <a href="#Page_165">165</a> + +Taenia, <a href="#Page_175">175</a> + +Tapeworm, <a href="#Page_4">4</a>, <a href="#Page_176">176</a> + +Tarantella, <a href="#Page_8">8</a> + +Tarantism, <a href="#Page_8">8</a> + +Tarantula, <a href="#Page_10">10</a> + +Tarsonemidæ, <a href="#Page_69">69</a> + +Tarsonemus, <a href="#Page_267">267</a> + +Tenebrionid beetles, <a href="#Page_127">127</a> + +Tersesthes, <a href="#Page_110">110</a>, <a href="#Page_288">288</a> + +Tetanus, <a href="#Page_129">129</a> + +Tetranychus, <a href="#Page_273">273</a> + +Texas fever, <a href="#Page_220">220-223</a> + +Three-day fever, <a href="#Page_96">96</a> + +Tick, <a href="#Page_23">23</a>, <a href="#Page_226">226</a> + bites, Treatment of, <a href="#Page_68">68</a> + fever, <a href="#Page_230">230</a> + paralysis, <a href="#Page_67">67</a> + +Treatment, + Bee stings, <a href="#Page_36">36</a>, <a href="#Page_41">41</a> + Bites of, + Bed-bugs, <a href="#Page_90">90</a>, <a href="#Page_93">93</a> + Blackflies, <a href="#Page_107">107</a> + Buffalo flies, <a href="#Page_107">107</a> + Bugs, <a href="#Page_31">31</a>, <a href="#Page_33">33</a> + Centipedes, <a href="#Page_26">26</a>, <a href="#Page_27">27</a> + Chiggers, <a href="#Page_127">127</a> + Chigoes, <a href="#Page_127">127</a> + Fleas, <a href="#Page_127">127</a> + Harvest mites, <a href="#Page_61">61</a> + Jiggers, <a href="#Page_129">129</a> + Lice, <a href="#Page_83">83</a>, <a href="#Page_85">85</a> + Mosquitoes, <a href="#Page_34">34</a>, <a href="#Page_36">36</a>, <a href="#Page_102">102</a> + Phlebotomus flies, <a href="#Page_97">97</a> + Sand flies, <a href="#Page_96">96</a>, <a href="#Page_107">107</a>, <a href="#Page_109">109</a> + Scorpions, <a href="#Page_22">22</a>, <a href="#Page_23">23</a> + Spiders, <a href="#Page_19">19</a> + Ticks, <a href="#Page_61">61</a>, <a href="#Page_68">68</a>, <a href="#Page_72">72</a> + Ticks, ear, <a href="#Page_65">65</a> + Blister beetle poison, <a href="#Page_55">55</a> + Brown-tail moth rash, <a href="#Page_45">45</a> + Cantharidin poison, <a href="#Page_55">55</a> + Caterpillar rash, <a href="#Page_45">45</a> + Ear ticks, <a href="#Page_65">65</a> + House fly control, <a href="#Page_156">156</a>, <a href="#Page_160">160</a> + Itch, <a href="#Page_77">77</a><span class="pagenum"><a name="Page_348" id="Page_348">[Pg 348]</a></span> + Itch, grocer's, <a href="#Page_72">72</a> + Lice, <a href="#Page_85">85</a> + Nasal myasis, <a href="#Page_143">143</a> + Rocky Mt. spotted fever, <a href="#Page_228">228</a>, <a href="#Page_229">229</a> + Rash, caterpillar, <a href="#Page_45">45</a> + Scabies, <a href="#Page_77">77</a> + Sleeping sickness control, <a href="#Page_218">218</a> + Spotted fever, <a href="#Page_228">228</a>, <a href="#Page_229">229</a> + Stings, bee, <a href="#Page_36">36</a>, <a href="#Page_41">41</a> + Typhus fever, prophylaxis, <a href="#Page_239">239</a> + +Trichodectes canis, <a href="#Page_176">176</a> + +Trichoma, <a href="#Page_82">82</a> + +Trineura, <a href="#Page_295">295</a> + +Trochosa singoriensis, <a href="#Page_11">11</a> + +Trombidium, <a href="#Page_60">60</a>, <a href="#Page_273">273</a> + +True insects, <a href="#Page_80">80</a> + +Trypanosoma, <a href="#Page_35">35</a> + +Trypanosoma, brucei, <a href="#Page_165">165</a> + +Trypanosoma cruzi, <a href="#Page_219">219</a> + +Trypanosoma lewisi, <a href="#Page_213">213</a> + +Trypanosomiases, <a href="#Page_212">212</a> + +Trypanosomiasis, <a href="#Page_165">165</a>, <a href="#Page_219">219</a> + +Tsetse flies, <a href="#Page_117">117</a>, <a href="#Page_166">166</a>, <a href="#Page_214">214</a>, <a href="#Page_219">219</a> + +Tsetse flies disease, <a href="#Page_165">165</a> + +Tuberculosis, <a href="#Page_155">155</a> + +Tumbu-fly, <a href="#Page_118">118</a> + +Tydeus, <a href="#Page_271">271</a> + +Typhoid, <a href="#Page_155">155</a> + +Typhoid fever, <a href="#Page_154">154</a> + +Typhus, <a href="#Page_237">237</a> + +Typhus fever, <a href="#Page_237">237</a> + +Tyroglyphus, <a href="#Page_72">72</a>, <a href="#Page_131">131</a>, <a href="#Pg_268">268</a> + +Dr. Tyzzer, <a href="#Page_49">49</a> + + +Uranotænia, <a href="#Page_292">292</a> + + +Vancoho, <a href="#Page_14">14</a> + +Varicose groin glands, <a href="#Page_178">178</a> + +Verruga peruviana, <a href="#Page_253">253</a> + +Vescicating insects, <a href="#Page_54">54</a> + + +Wanzenspritze, <a href="#Page_29">29</a> + +Warble-flies, <a href="#Page_112">112</a> + +Wasps, <a href="#Page_43">43</a> + +Whip-scorpions, <a href="#Page_19">19</a> + +Wohlfahrtia, <a href="#Page_143">143</a>, <a href="#Page_302">302</a> + +Wolf-spiders, <a href="#Page_10">10</a> + +Wyeomyia smithii, <a href="#Page_101">101</a>, <a href="#Page_293">293</a> + + +Xenopsylla, <a href="#Page_172">172</a>, <a href="#Page_317">317</a> + +Xenopsylla cheopis, <a href="#Page_171">171</a>, <a href="#Page_124">124</a> + +Xestopsylla, <a href="#Page_317">317</a> + + +Yaws, <a href="#Page_2">2</a> + +Yellow fever, <a href="#Page_196">196</a>, <a href="#Page_203">203</a>, <a href="#Page_205">205</a> +</pre> + +<div class="bbox" style="padding:1em;"> +<h3>TRANSCRIBERS' NOTES</h3> + +<p>The following discrepancies in the text are as in the original:</p> +<ul> +<li>The inconsistent hyphenation of the following: +<ul> +<li>assassin-bugs/assassin bugs;</li> +<li>bedbug/bed-bug (and bedbugs);</li> +<li>beekeeper/bee-keeper (and beekeepers);</li> +<li>blackflies/black-flies;</li> +<li>blow-flies/blow flies;</li> +<li>bluebottles/blue-bottles;</li> +<li>bot-flies/bot flies;</li> +<li>bristlelike/bristle-like;</li> +<li>browntail/brown-tail;</li> +<li>coextensive/co-extensive;</li> +<li>deer-flies/deer flies;</li> +<li>dorsocentral/dorso-central;</li> +<li>ectoparasites/ecto-parasites;</li> +<li>endoparasites/endo-parasites;</li> +<li>flesh-fly/flesh fly (and flesh flies);</li> +<li>hotbed/hot-bed;</li> +<li>housefly/house-fly (and houseflies);</li> +<li>horsefly/horse-fly (and horse flies);</li> +<li>horse-manure/horse manure;</li> +<li>midsummer/mid-summer;</li> +<li>preeminently/pre-eminently;</li> +<li>sandfly/sand-fly (and sandflies);</li> +<li>screw-worm fly/screw worm fly;</li> +<li>stable-fly/stable fly;</li> +<li>subequal/sub-equal;</li> +<li>subfamily/sub-family;</li> +<li>subtropical/sub-tropical;</li> +<li>tapeworm/tape-worm (and tapeworms);</li> +<li>today/to-day;</li> +<li>tsetse-flies/tsetse flies;</li> +<li>widespread/wide-spread;</li> +<li>wormlike/worm-like.</li> +</ul> +</li> +<li>Inconsistent use of diaeresis in Aëdes/Aedes</li> +<li>Inconsistent spelling of the following: +<ul> +<li>defence/defense;</li> +<li>disc/disk;</li> +<li>hemolysis/hæmolysis;</li> +<li>hemolytic/hæmolytic;</li> +<li>hexapod/hexopod;</li> +<li>Levaditi/Lavaditi;</li> +<li>metalescent/metallescent;</li> +<li>Mitzmain/Mitzman;</li> +<li>Neveau-Lemaire/Neveau-Lamaire;</li> +<li>offence/offense;</li> +<li>Phthirus/Phthirius</li> +<li>Portschinsky/Portchinsky/Portchinski;</li> +<li>travelled/traveled;</li> +<li>ventra-/ventro-;</li> +<li>Villot/Villet;</li> +<li>Wohlfartia/Wohlfahrtia;</li> +</ul> +</li> +<li>Inconsistent use of ligatures in Taenia/Tænia</li> +</ul> +<p>toxine, insiduous, efficaceous, cyanid are spelt as in the original</p> +<p>In the first paragraph of chapter XII, "the student might not be lead" +is as in the original. Lead perhaps should be led</p> +<p>There is no Figure 147 in the original</p> +<p>In the bibliography, the entry for Vaughan without a text specified is +as in the original</p> + +</div> + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of Handbook of Medical Entomology, by +William Albert Riley and Oskar Augustus Johanssen + +*** END OF THIS PROJECT GUTENBERG EBOOK HANDBOOK OF MEDICAL ENTOMOLOGY *** + +***** This file should be named 34279-h.htm or 34279-h.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/4/2/7/34279/ + +Produced by Bryan Ness, Iris Schimandle, Brownfox and the +Online Distributed Proofreading Team at http://www.pgdp.net +(This file was produced from images generously made +available by The Internet Archive/American Libraries.) + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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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: Handbook of Medical Entomology + +Author: William Albert Riley + Oskar Augustus Johanssen + +Release Date: November 11, 2010 [EBook #34279] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK HANDBOOK OF MEDICAL ENTOMOLOGY *** + + + + +Produced by Bryan Ness, Iris Schimandle, Brownfox and the +Online Distributed Proofreading Team at http://www.pgdp.net +(This file was produced from images generously made +available by The Internet Archive/American Libraries.) + + + + + +Transcriber's Note: ae character replaced with ae. Accents have been +removed. The degree symbol has been replaced with ^o. The times symbol +has been replaced with lowercase x. The symbol mu/micro has been +replaced with lowercase u. + + + + +[Illustration: Some early medical entomology. Athanasius Kircher's +illustration of the Italian tarantula and the music prescribed as an +antidote for the poison of its bite. (1643).] + + + + +HANDBOOK OF MEDICAL + +ENTOMOLOGY + +WM. A. RILEY, PH.D. + +Professor of Insect Morphology and Parasitology, Cornell University + +and + +O. A. JOHANNSEN, PH.D. + +Professor of Biology, Cornell University + +[Illustration] + +ITHACA, NEW YORK + +THE COMSTOCK PUBLISHING COMPANY + +1915 + + +COPYRIGHT, 1915 + +BY THE COMSTOCK PUBLISHING COMPANY, + +ITHACA, N. Y. + + +Press of W. F. Humphrey +Geneva, N. Y. + + + + +PREFACE + + +The Handbook of Medical Entomology is the outgrowth of a course of +lectures along the lines of insect transmission and dissemination of +diseases of man given by the senior author in the Department of +Entomology of Cornell University during the past six years. More +specifically it is an illustrated revision and elaboration of his "Notes +on the Relation of Insects to Disease" published January, 1912. + +Its object is to afford a general survey of the field, and primarily to +put the student of medicine and entomology in touch with the discoveries +and theories which underlie some of the most important modern work in +preventive medicine. At the same time the older phases of the +subject--the consideration of poisonous and parasitic forms--have not +been ignored. + +Considering the rapid shifts in viewpoint, and the development of the +subject within recent years, the authors do not indulge in any hopes +that the present text will exactly meet the needs of every one +specializing in the field,--still less do they regard it as complete or +final. The fact that the enormous literature of isolated articles is to +be found principally in foreign periodicals and is therefore difficult +of access to many American workers, has led the authors to hope that a +summary of the important advances, in the form of a reference book may +not prove unwelcome to physicians, sanitarians and working +entomologists, and to teachers as a text supplementing lecture work in +the subject. + +Lengthy as is the bibliography, it covers but a very small fraction of +the important contributions to the subject. It will serve only to put +those interested in touch with original sources and to open up the +field. Of the more general works, special acknowledgment should be made +to those of Banks, Brumpt, Castellani and Chalmers, Comstock, Hewitt, +Howard, Manson, Mense, Neveau-Lemaire, Nuttall, and Stiles. + +To the many who have aided the authors in the years past, by suggestions +and by sending specimens and other materials, sincerest thanks is +tendered. This is especially due to their colleagues in the Department +of Entomology of Cornell University, and to Professor Charles W. Howard, +Dr. John Uri Lloyd, Mr. A. H. Ritchie, Dr. I. M. Unger, and Dr. Luzerne +Coville. + +They wish to express indebtedness to the authors and publishers who have +so willingly given permission to use certain illustrations. Especially +is this acknowledgment due to Professor John Henry Comstock, Dr. L. O. +Howard, Dr. Graham-Smith, and Professor G. H. T. Nuttall. Professor +Comstock not only authorized the use of departmental negatives by the +late Professor M. V. Slingerland (credited as M. V. S.), but generously +put at their disposal the illustrations from the MANUAL FOR THE STUDY OF +INSECTS and from the SPIDER BOOK. Figures 5 and 111 are from Peter's +"Der Arzt und die Heilkunst in der deutschen Vergangenheit." It should +be noted that on examining the original, it is found that Gottfried's +figure relates to an event antedating the typical epidemic of dancing +mania. + + WM. A. RILEY. + O. A. JOHANNSEN. + + CORNELL UNIVERSITY, + January, 1915. + + + + +ADDITIONS AND CORRECTIONS + +vi line 11, for Heilkunft read Heilkunst. + +18 line 2, for tarsi read tarsus. + +32 line 21, and legend under fig. 23, for C. (Conorhinus) abdominalis + read Melanolestes abdominalis. + +47 legend under figure for 33c read 34. + +92 line 22 and 25, for sangiusugus read sanguisugus. + +116 legend under fig. 83, for Graham-Smith read Manson. + +136 line 10, from bottom, insert "ring" after "chitin". + +137 line 3, for meditatunda read meditabunda. + +145 line 7, from bottom, for Rs read R_5. + +158 line 20, for have read has. + +212 after the chapter heading insert "continued". + +219 line 10, from bottom, for Cornohinus read Conorhinus. + +266 line 1, fig. 158j refers to the female. + +272 line 5, insert "palpus" before "and leg". + +281 line 6, for discodial read discoidal. + +281 last line, insert "from" before "the". + +284 line 5, for "tubercle of" read "tubercle or". + +305 lines 19, 28, 44, page 306 lines 1, 9, 22, 27, 30, page 307 line 7, + page 309 lines 8, 11, for R_{4+5} read M_{1+2}. + +309 legend under fig. 168 add Bureau of Entomology. + +312 line 36, for "near apex" read "of M_{1+2}". + +313 running head, for Muscidae read Muscoidea. + +314 line 29, for "distal section" read "distally M_{1+2}". + +315 legend under fig. 172, for Pseudopyrellia read Orthellia, for + Lyperosia read Haematobia, for Umbana read urbana. + +323 and 325 legends under the figures, add "After Dr. J. H. Stokes". + +328 line 7 from bottom for Apiochaeta read Aphiochaeta. + + + + +CONTENTS + + +CHAPTER I + +INTRODUCTION 1-5 + + Early suggestions regarding the transmission of disease by + insects. + The ways in which arthropods may affect the health of man. + + +CHAPTER II + +ARTHROPODS WHICH ARE DIRECTLY POISONOUS 6-56 + + The Araneida, or Spiders. + The tarantulas. Bird spiders. Spiders of the genus + Latrodectus. Other venomous spiders. Summary. + The Pedipalpida, or whip-scorpions. + The Scorpionida, or true scorpions. + The Solpugida, or solpugids. + The Acarina, or mites and ticks. + The Myriapoda, or centipedes and millipedes. + The Hexapoda, or true insects. + Piercing or biting insects poisonous to man. + Hemiptera, or true bugs. + The Notonectidae or back-swimmers. Belostomidae or giant + water-bugs. Reduviidae, or assassin bugs. Other + Hemiptera reported as poisonous to man. + Diptera; the midges, mosquitoes and flies. + Stinging insects. + Apis mellifica, the honey bee. Other stinging forms. + Nettling insects. + Lepidoptera, or butterflies and moths. Relief from + poisoning by nettling larvae. + Vescicating insects and those possessing other poisons + in their blood plasma. The blister beetles. Other + cryptotoxic insects. + + +CHAPTER III + +PARASITIC ARTHROPODS AFFECTING MAN 57-130 + + Acarina, or mites. + The Trombidiidae, or harvest mites. + The Ixodoidea, or ticks. + Argasidae. Ixodidae. Treatment of tick bites. + The mites. + Dermanyssidae. Tarsonemidae. Sarcoptidae, the itch mites. + Demodecidae, the follicle mites. + Hexapoda, or true insects. + Siphunculata, or sucking lice. + Hemiptera. + The bed-bug. Other bed-bugs. + Parasitic Diptera, or flies. + Psychodidae, or moth flies. Phlebotominae. Culicidae, or + mosquitoes. Simuliidae, or black-flies. Chironomidae, or + midges. Tabanidae, or horse-flies. Leptidae or + snipe-flies. Oestridae, or bot-flies. Muscidae, the + stable-fly and others. + Siphonaptera, or fleas. + The fleas affecting man, the dog, cat, and rat. + The true chiggers, or chigoes. + +CHAPTER IV + +ACCIDENTAL OR FACULTATIVE PARASITES 131-143 + + Acarina, or mites. + Myriapoda, or centipedes and millipedes. + Lepidopterous larvae. + Coleoptera, or beetles. + Dipterous larvae causing myiasis. + Piophila casei, the cheese skipper. Chrysomyia macellaria, + the screw-worm fly. Calliphorinae, the bluebottles. + Muscinae, the house or typhoid fly, and others. + Anthomyiidae, the lesser house-fly and others. + Sarcophagidae, the flesh-flies. + + +CHAPTER V + +ARTHROPODS AS SIMPLE CARRIERS OF DISEASE 144-163 + + The house or typhoid fly as a carrier of disease. + Stomoxys calcitrans, the stable-fly. + Other arthropods which may serve as simple carriers of + pathogenic organisms. + + +CHAPTER VI + +ARTHROPODS AS DIRECT INOCULATORS OF DISEASE GERMS 164-174 + + Some illustrations of direct inoculations of disease germs + by arthropods. + The role of fleas in the transmission of the plague. + + +CHAPTER VII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC ORGANISMS 175-185 + + Insects as intermediate hosts of tape-worms. + Arthropods as intermediate hosts of nematode worms. + Filariasis and mosquitoes. + Other nematode parasites of man and animals. + + +CHAPTER VIII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA 186-211 + + Mosquitoes and malaria. + Mosquitoes and yellow fever. + + +CHAPTER IX + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA 212-229 + + Insects and trypanosomiases. + Fleas and lice as carriers of Trypanosoma lewisi. + Tsetse-flies and nagana. + Tsetse-flies and sleeping sickness in man. + South American trypanosomiasis. + Leishmanioses and insects. + Ticks and diseases of man and animals. + Cattle tick and Texas fever. + Ticks and Rocky Mountain Spotted fever of man. + + +CHAPTER X + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA +(CONTINUED) 230-240 + + Arthropods and Spirochaetoses of man and animals. + African relapsing fever of man. + European relapsing fever. + North African relapsing fever of man. + Other types of relapsing fever of man. + Spirochaetosis of fowls. + Other spirochaete diseases of animals. + Typhus fever and lice. + + +CHAPTER XI + +SOME POSSIBLE, BUT IMPERFECTLY KNOWN CASES OF +ARTHROPOD TRANSMISSION OF DISEASE 241-256 + + Infantile paralysis, or acute anterior poliomyelitis. + Pellagra. Leprosy. Verruga peruviana. Cancer. + + +CHAPTER XII + +KEYS TO THE ARTHROPODS NOXIOUS TO MAN 257-317 + + Crustacea. + Myriapoda, or centipedes and millipedes. + Arachnida (Orders of). + Acarina or ticks. + Hexapoda (Insecta). + Siphunculata and Hemiptera (lice and true bugs). + Diptera (mosquitoes, midges, and flies). + Siphonaptera (fleas). + + +APPENDIX + +Hydrocyanic acid gas against household insects 318-320 + Proportion of ingredients. A single room as an example. + Fumigating a large house. Precautions. + +Lesions produced by the bite of the black-fly 321-326 + + +BIBLIOGRAPHY 327-340 + + +INDEX 341-348 + + + + +CHAPTER I. + +INTRODUCTION + +EARLY SUGGESTIONS REGARDING THE TRANSMISSION OF DISEASE BY INSECTS + + +Until very recent years insects and their allies have been considered as +of economic importance merely in so far as they are an annoyance or +direct menace to man, or his flocks and herds, or are injurious to his +crops. It is only within the past fifteen years that there has sprung +into prominence the knowledge that in another and much more insiduous +manner, they may be the enemy of mankind, that they may be among the +most important of the disseminators of disease. In this brief period, +such knowledge has completely revolutionized our methods of control of +certain diseases, and has become an important weapon in the fight for +the conservation of health. + +It is nowhere truer than in the case under consideration that however +abrupt may be their coming into prominence, great movements and great +discoveries do not arise suddenly. Centuries ago there was suggested the +possibility that insects were concerned with the spread of disease, and +from time to time there have appeared keen suggestions and logical +hypotheses along this line, that lead us to marvel that the +establishment of the truths should have been so long delayed. + +One of the earliest of these references is by the Italian physician, +Mercurialis, who lived from 1530 to 1607, during a period when Europe +was being ravaged by the dread "black death", or plague. Concerning its +transmission he wrote: "There can be no doubt that flies feed on the +internal secretions of the diseased and dying, then, flying away, they +deposit their excretions on the food in neighboring dwellings, and +persons who eat of it are thus infected." + +It would be difficult to formulate more clearly this aspect of the facts +as we know them to-day, though it must always be borne in mind that we +are prone to interpret such statements in the light of present-day +knowledge. Mercurialis had no conception of the animate nature of +contagion, and his statement was little more than a lucky guess. + +Much more worthy of consideration is the approval which was given to his +view by the German Jesuit, Athanasius Kircher in 1658. One cannot read +carefully his works without believing that long before Leeuwenhook's +discovery, Kircher had seen the larger species of bacteria. Moreover, he +attributed the production of disease to these organisms and formulated, +vaguely, to be sure, a theory of the animate nature of contagion. It has +taken two and a half centuries to accumulate the facts to prove his +hypothesis. + +The theory of Mercurialis was not wholly lost sight of, for in the +medical literature of the eighteenth century there are scattered +references to flies as carriers of disease. Such a view seems even to +have been more or less popularly accepted, in some cases. Gudger (1910), +has pointed out that, as far back as 1769, Edward Bancroft, in "An Essay +on the Natural History of Guiana in South America," wrote concerning the +contagious skin-disease known as "Yaws": "It is usually believed that +this disorder is communicated by the flies who have been feasting on a +diseased object, to those persons who have sores, or scratches, which +are uncovered; and from many observations, I think this is not +improbable, as none ever receive this disorder whose skins are whole." + +Approaching more closely the present epoch, we find that in 1848, Dr. +Josiah Nott, of Mobile, Alabama, published a remarkable article on the +cause of yellow fever, in which he presented "reasons for supposing its +specific cause to exist in some form of insect life." As a matter of +fact, the bearing of Nott's work on present day ideas of the insect +transmission of disease has been very curiously overrated. The common +interpretation of his theory has been deduced from a few isolated +sentences, but his argument appears quite differently when the entire +article is studied. It must be remembered that he wrote at a period +before the epoch-making discoveries of Pasteur and before the +recognition of micro-organisms as factors in the cause of disease. His +article is a masterly refutation of the theory of "malarial" origin of +"all the fevers of hot climates," but he uses the term "insect" as +applicable to the lower forms of life, and specific references to +"mosquitoes," "aphids," "cotton-worms," and others, are merely in the +way of similes. + +But, while Nott's ideas regarding the relation of insects to yellow +fever were vague and indefinite, it was almost contemporaneously that +the French physician, Louis Daniel Beauperthuy argued in the most +explicit possible manner, that yellow fever and various others are +transmitted by mosquitoes. In the light of the data which were available +when he wrote, in 1853, it is not surprising that he erred by thinking +that the source of the virus was decomposing matter which the mosquito +took up and accidentally inoculated into man. Beauperthuy not only +discussed the role of mosquitoes in the transmission of disease, but he +taught, less clearly, that house-flies scatter pathogenic organisms. It +seems that Boyce (1909) who quotes extensively from this pioneer work, +does not go too far when he says "It is Dr. Beauperthuy whom we must +regard as the father of the doctrine of insect-borne disease." + +In this connection, mention must be made of the scholarly article by the +American physician, A. F. A. King who, in 1883, brought together an all +but conclusive mass of argument in support of his belief that malaria +was caused by mosquitoes. At about the same time, Finley, of Havana, was +forcefully presenting his view that the mosquito played the chief role +in the spread of yellow fever. + +To enter more fully into the general historical discussion is beyond the +scope of this book. We shall have occasion to make more explicit +references in considering various insect-borne diseases. Enough has been +said here to emphasize that the recognition of insects as factors in the +spread of disease was long presaged, and that there were not wanting +keen thinkers who, with a background of present-day conceptions of the +nature of disease, might have been in the front rank of investigators +along these lines. + + +THE WAYS IN WHICH ARTHROPODS MAY AFFECT THE HEALTH OF MAN + +When we consider the ways in which insects and their allies may affect +the health of man, we find that we may treat them under three main +groups: + +A. They may be directly poisonous. Such, for example, are the scorpions, +certain spiders and mites, some of the predaceous bugs, and stinging +insects. Even such forms as the mosquito deserve some consideration from +this viewpoint. + +B. They may be parasitic, living more or less permanently on or in the +body and deriving their sustenance from it. + +Of the parasitic arthropods we may distinguish, first, the _true +parasites_, those which have adopted and become confirmed in the +parasitic habit. Such are the itch mites, the lice, fleas, and the +majority of the forms to be considered as parasitic. + +In addition to these, we may distinguish a group of _accidental_, or +_facultative parasites_, species which are normally free-living, feeding +on decaying substances, but which when accidentally introduced into the +alimentary canal or other cavities of man, may exist there for a greater +or less period. For example, certain fly larvae, or maggots, normally +feeding in putrifying meat, have been known to occur as accidental or +facultative parasites in the stomach of man. + +C. Finally, and most important, arthropods may be transmitters and +disseminators of disease. In this capacity they may function in one of +three ways; as _simple carriers_, as _direct inoculators_, or as +_essential hosts_ of disease germs. + +As simple carriers, they may, in a wholly incidental manner, transport +from the diseased to the healthy, or from filth to food, pathogenic +germs which cling to their bodies or appendages. Such, for instance, is +the relation of the house-fly to the dissemination of typhoid. + +As direct inoculators, biting or piercing species may take up from a +diseased man or animal, germs which, clinging to the mouth parts, are +inoculated directly into the blood of the insect's next victim. It it +thus that horse-flies may occasionally transmit anthrax. Similarly, +species of spiders and other forms which are ordinarily perfectly +harmless, may accidentally convey and inoculate pyogenic bacteria. + +It is as essential hosts of disease germs that arthropods play their +most important role. In such cases an essential part of the life cycle +of the pathogenic organism is undergone in the insect. In other words, +without the arthropod host the disease-producing organism cannot +complete its development. As illustrations may be cited the relation of +the Anopheles mosquito to the malarial parasite, and the relation of the +cattle tick to Texas fever. + +A little consideration will show that this is the most important of the +group. Typhoid fever is carried by water or by contaminated milk, and in +various other ways, as well as by the house-fly. Kill all the +house-flies and typhoid would still exist. On the other hand, malaria is +carried only by the mosquito, because an essential part of the +development of the malarial parasite is undergone in this insect. +Exterminate all of the mosquitoes of certain species and the +dissemination of human malaria is absolutely prevented. + +Once an arthropod becomes an essential host for a given parasite it may +disseminate infection in three different ways: + +1. By infecting man or animals who ingest it. It is thus, for example, +that man, dog, or cat, becomes infected with the double-pored dog +tapeworm, _Dipylidium caninum_. The cysticercoid stage occurs in the +dog louse, or in the dog or cat fleas, and by accidentally ingesting the +infested insect the vertebrate becomes infested. Similarly, _Hymenolepis +diminuta_, a common tapeworm of rats and mice, and occasional in man, +undergoes part of its life cycle in various meal-infesting insects, and +is accidentally taken up by its definitive host. It is very probable +that man becomes infested with _Dracunculus (Filaria) medinensis_ +through swallowing in drinking water, the crustacean, _Cyclops_, +containing the larvae of this worm. + +2. By infecting man or animals on whose skin or mucous membranes the +insect host may be crushed or may deposit its excrement. The pathogenic +organism may then actively penetrate, or may be inoculated by +scratching. The causative organism of typhus fever is thus transmitted +by the body louse. + +3. By direct inoculation by its bite, the insect host may transfer the +parasite which has undergone development within it. The malarial +parasite is thus transferred by mosquitoes; the Texas fever parasite by +cattle ticks. + + + + +CHAPTER II. + +ARTHROPODS WHICH ARE DIRECTLY POISONOUS + + +Of all the myriads of insects and related forms, a very few are of +direct use to man, some few others have forced his approbation on +account of their wonderful beauty, but the great hordes of them are +loathed or regarded as directly dangerous. As a matter of fact, only a +very small number are in the slightest degree poisonous to man or to the +higher animals. The result is that entomologists and lovers of nature, +intent upon dissipating the foolish dread of insects, are sometimes +inclined to go to the extreme of discrediting all statements of serious +injury from the bites or stings of any species. + +Nevertheless, it must not be overlooked that poisonous forms do exist, +and they must receive attention in a consideration of the ways in which +arthropods may affect the health of man. Moreover, it must be recognized +that "what is one man's meat, is another man's poison," and that in +considering the possibilities of injury we must not ignore individual +idiosyncrasies. Just as certain individuals may be poisoned by what, for +others, are common articles of food, so some persons may be abnormally +susceptible to insect poison. Thus, the poison of a bee sting may be of +varying severity, but there are individuals who are made seriously sick +by a single sting, regardless of the point of entry. Some individuals +scarcely notice a mosquito bite, others find it very painful, and so +illustrations of this difference in individuals might be multiplied. + +In considering the poisonous arthropods, we shall take them up by +groups. The reader who is unacquainted with the systematic relationship +of insects and their allies is referred to Chapter XII. No attempt will +be made to make the lists under the various headings exhaustive, but +typical forms will be discussed. + + +ARANEIDA OR SPIDERS + +Of all the arthropods there are none which are more universally feared +than are the spiders. It is commonly supposed that the majority, if not +all the species are poisonous and that they are aggressive enemies of +man and the higher animals, as well as of lower forms. + +That they really secrete a poison may be readily inferred from the +effect of their bite upon insects and other small forms. Moreover, the +presence of definite and well-developed poison glands can easily be +shown. They occur as a pair of pouches (fig. 1) lying within the +cephalothorax and connected by a delicate duct with a pore on the claw +of the chelicera, or so-called "mandible" on the convex surface of the +claw in such a position that it is not plugged and closed by the flesh +of the victim. + +[Illustration: 1. Head of a spider showing poison gland (_c_) and its +relation to the chelicera (_a_).] + +The glands may be demonstrated by slowly and carefully twisting off a +chelicera and pushing aside the stumps of muscles at its base. By +exercising care, the chitinous wall of the chelicera and its claw may be +broken away and the duct traced from the gland to its outlet. The inner +lining of the sac is constituted by a highly developed glandular +epithelium, supported by a basement membrane of connective tissue and +covered by a muscular layer, (fig. 2). The muscles, which are striated, +are spirally arranged (fig. 1), and are doubtless under control of the +spider, so that the amount of poison to be injected into a wound may be +varied. + +[Illustration: 2. Section through a venom gland of Latrodectus +13-guttatus showing the peritoneal, muscular and epithelial layers. +After Bordas.] + +The poison itself, according to Kobert (1901), is a clear, colorless +fluid, of oily consistency, acid reaction, and very bitter taste. After +the spider has bitten two or three times, its supply is exhausted and +therefore, as in the case of snakes, the poison of the bite decreases +quickly with use, until it is null. To what extent the content of the +poison sacs may contain blood serum or, at least, active principles of +serum, in addition to a specific poison formed by the poison glands +themselves, Kobert regards as an open question. He believes that the +acid part of the poison, if really present, is formed by the glands and +that, in the case of some spiders, the ferment-like, or better, active +toxine, comes from the blood. + +[Illustration: 3. Chelicera of a spider.] + +But there is a wide difference between a poison which may kill an insect +and one which is harmful to men. Certain it is that there is no lack of +popular belief and newspaper records of fatal cases, but the evidence +regarding the possibility of fatal or even very serious results for man +is most contradictory. For some years, we have attempted to trace the +more circumstantial newspaper accounts, which have come to our notice, +of injury by North American species. The results have served, mainly, to +emphasize the straits to which reporters are sometimes driven when there +is a dearth of news. The accounts are usually vague and lacking in any +definite clue for locating the supposed victim. In the comparatively few +cases where the patient, or his physician, could be located, there was +either no claim that the injury was due to spider venom, or there was no +evidence to support the belief. Rarely, there was evidence that a +secondary blood poisoning, such as might be brought about by the prick +of a pin, or by any mechanical injury, had followed the bite of a +spider. Such instances have no bearing on the question of the venomous +nature of these forms. + +[Illustration: 4. The Italian tarantula (Lycosa tarantula). After +Kobert.] + +The extreme to which unreasonable fear of the bites of spiders +influenced the popular mind was evidenced by the accepted explanation of +the remarkable dancing mania, or tarantism, of Italy during the Middle +Ages. This was a nervous disorder, supposed to be due to the bite of a +spider, the European tarantula (fig. 4), though it was also, at times, +attributed to the bite of the scorpion. In its typical form, it was +characterized by so great a sensibility to music that under its +influence the victims indulged in the wildest and most frenzied dancing, +until they sank to the ground utterly exhausted and almost lifeless. The +profuse perspiring resulting from these exertions was supposed to be the +only efficacious remedy for the disease. Certain forms of music were +regarded as of especial value in treating this tarantism, and hence the +name of "tarantella" was applied to them. Our frontispiece, taken from +Athanasius Kircher's _Magnes sive de Arte Magnetica_, 1643 ed., +represents the most commonly implicated spider and illustrates some of +what Fabre has aptly designated as "medical choreography." + +The disease was, in reality, a form of hysteria, spreading by sympathy +until whole communities were involved, and was paralleled by the +outbreaks of the so-called St. Vitus's or St. John's dance, which swept +Germany at about the same time (fig. 5). The evidence that the spider +was the cause of the first is about as conclusive as is that of the +demoniacal origin of the latter. The true explanation of the outbreaks +is doubtless to be found in the depleted physical and mental condition +of the people, resulting from the wars and the frightful plagues which +devastated all Europe previous to, and during these times. An +interesting discussion of these aspects of the question is to be found +in Hecker. + +[Illustration: 5. Dancing mania. Illustration from Johann Ludwig +Gottfried's Chronik. 1632.] + +So gross has been the exaggeration and so baseless the popular fear +regarding spiders that entomologists have been inclined to discredit all +accounts of serious injury from their bites. Not only have the most +circumstantial of newspaper accounts proved to be without foundation but +there are on record a number of cases where the bite of many of the +commoner species have been intentionally provoked and where the effect +has been insignificant. Some years ago the senior author personally +experimented with a number of the largest of our northern species, and +with unexpected results. The first surprise was that the spiders were +very unwilling to bite and that it required a considerable effort to get +them to attempt to do so. In the second place, most of those +experimented with were unable to pierce the skin of the palm or the back +of the hand, but had to be applied to the thin skin between the fingers +before they were able to draw blood. Unfortunately, no special attempt +was made to determine, at the time, the species experimented with, but +among them were _Theridion tepidariorum_, _Miranda aurantia_ +(_Argiopa_), _Metargiope trifasciata_, _Marxia stellata_, _Aranea +trifolium_, _Misumena vatia_, and _Agelena naevia_. In no case was the +bite more severe than a pin prick and though in some cases the sensation +seemed to last longer, it was probably due to the fact that the mind was +intent upon the experiment. + +[Illustration: 6. An American tarantula (Eurypelma hentzii). Natural +size. After Comstock.] + +Similar experiments were carried out by Blackwell (1855), who believed +that in the case of insects bitten, death did not result any more +promptly than it would have from a purely mechanical injury of equal +extent. He was inclined to regard all accounts of serious injury to man +as baseless. The question cannot be so summarily dismissed, and we shall +now consider some of the groups which have been more explicitly +implicated. + + +THE TARANTULAS.--In popular usage, the term "tarantula" is loosely +applied to any one of a number of large spiders. The famous tarantulas +of southern Europe, whose bites were supposed to cause the dancing +mania, were Lycosidae, or wolf-spiders. Though various species of this +group were doubtless so designated, the one which seems to have been +most implicated was _Lycosa tarantula_ (L.), (fig. 4). On the other +hand, in this country, though there are many Lycosidae, the term +"tarantula" has been applied to members of the superfamily Avicularoidea +(fig. 6), including the bird-spiders. + +Of the Old World Lycosidae there is no doubt that several species were +implicated as the supposed cause of the tarantism. In fact, as we have +already noted, the blame was sometimes attached to a scorpion. However, +there seems to be no doubt that most of the accounts refer to the spider +known as _Lycosa tarantula_. + +There is no need to enter into further details here regarding the +supposed virulence of these forms, popular and the older medical +literature abound in circumstantial accounts of the terrible effects of +the bite. Fortunately, there is direct experimental evidence which bears +on the question. + +Fabre induced a common south European wolf-spider, _Lycosa narbonensis_, +to bite the leg of a young sparrow, ready to leave the nest. The leg +seemed paralyzed as a result of the bite, and though the bird seemed +lively and clamored for food the next day, on the third day it died. A +mole, bitten on the nose, succumbed after thirty-six hours. From these +experiments Fabre seemed justified in his conclusion that the bite of +this spider is not an accident which man can afford to treat lightly. +Unfortunately, there is nothing in the experiments, or in the symptoms +detailed, to exclude the probability that the death of the animals was +the result of secondary infection. + +As far back as 1693, as we learn from the valuable account of Kobert, +(1901), the Italian physician, Sanguinetti allowed himself to be bitten +on the arm by two tarantulas, in the presence of witnesses. The +sensation was equivalent to that from an ant or a mosquito bite and +there were no other phenomena the first day. On the second day the wound +was inflamed and there was slight ulceration. It is clear that these +later symptoms were due to a secondary infection. These experiments have +been repeated by various observers, among whom may be mentioned Leon +Dufour, Josef Erker and Heinzel, and with the similar conclusion that +the bite of the Italian tarantula ordinarily causes no severe symptoms. +In this conclusion, Kobert, though firmly convinced of the poisonous +nature of some spiders, coincides. He also believes that striking +symptoms may be simulated or artificially induced by patients in order +to attract interest, or because they have been assured that the bite, +under all circumstances, caused tarantism. + +The so-called Russian tarantula, _Trochosa singoriensis_ (fig. 7), is +much larger than the Italian species, and is much feared. Kobert carried +out a series of careful experiments with this species and his results +have such an important bearing on the question of the venomous nature of +the tarantula that we quote his summary. Experimenting first on nearly a +hundred living specimens of _Trochosa singoriensis_ from Crimea he says +that: + +"The tarantulas, no matter how often they were placed on the skin, +handled, and irritated, could not be induced to bite either myself, the +janitor, or the ordinary experimental animals. The objection that the +tarantulas were weak and indifferent cannot stand, for as soon as I +placed two of them on the shaved skin of a rabbit, instead of an attack +on the animal, there began a furious battle between the two spiders, +which did not cease until one of the two was killed." + +[Illustration: 7. Trochosa singoriensis. After Kobert.] + +"Since the spiders would not bite, I carefully ground up the fresh +animals in physiological salt solution, preparing an extract which must +have contained, in solution, all of the poisonous substance of their +bodies. While in the case of _Latrodectus_, as we shall see, less than +one specimen sufficed to yield an active extract, I have injected the +filtered extract of six fresh Russian tarantulas, of which each one was +much heavier than an average _Latrodectus_, subcutaneously and into the +jugular vein of various cats without the animals dying or showing any +special symptoms. On the basis of my experiments I can therefore only +say that the quantity of the poison soluble in physiological salt +solution, even when the spiders are perfectly fresh and well nourished, +is very insignificant. That the poison of the Russian tarantula is not +soluble in physiological salt solution, is exceedingly improbable. +Moreover, I have prepared alcoholic extracts and was unable to find them +active. Since the Russian spider exceeds the Italian in size and in +intensity of the bite, it seems very improbable to me that the +pharmacological test of the Italian tarantula would yield essentially +other results than those from the Russian species." + +To the AVICULAROIDEA belong the largest and most formidable appearing of +the spiders and it is not strange that in the New World they have fallen +heir to the bad reputation, as well as to the name of the tarantula of +Europe. In this country they occur only in the South or in the far West, +but occasionally living specimens are brought to our northern ports in +shipments of bananas and other tropical produce, and are the source of +much alarm. It should be mentioned, however, that the large spider most +frequently found under such circumstances is not a tarantula at all, but +one of the Heteropodidae, or giant crab-spiders, (fig. 8). + +[Illustration: 8. The giant crab-spider or banana spider (Heteropoda +venatoria). Natural size. After Comstock.] + +In spite of their prominence and the fear which they arouse there are +few accurate data regarding these American tarantulas. It has often been +shown experimentally that they can kill small birds and mammals, though +it is doubtful if these form the normal prey of any of the species, as +has been claimed. There is no question but that the mere mechanical +injury which they may inflict, and the consequent chances of secondary +infection, justify, in part, their bad reputation. In addition to the +injury from their bite, it is claimed that the body hairs of several of +the South American species are readily detached and are urticating. + +Recently, Phisalix (1912) has made a study of the physiological effects +of the venom of two Avicularoidea, _Phormictopus carcerides_ Pocock, +from Haiti and _Cteniza sauvagei_ Rossi, from Corsica. The glands were +removed aseptically and ground up with fine, sterilized sand in +distilled water. The resultant liquid was somewhat viscid, colorless, +and feebly alkaline. Injected into sparrows and mice the extract of +_Phormictopus_ proved very actively poisonous, that from a single spider +being sufficient to kill ten sparrows or twenty mice. It manifested +itself first and, above all, as a narcotic, slightly lowering the +temperature and paralyzing the respiration. Muscular and cardiac +weakening, loss of general sensibility, and the disappearance of +reflexes did not occur until near the end. The extract from _Cteniza_ +was less active and, curiously enough, the comparative effect on +sparrows and on mice was just reversed. + + +SPIDERS OF THE GENUS LATRODECTUS.--While most of the popular accounts of +evil effects from the bites of spiders will not stand investigation, it +is a significant fact that, the world over, the best authenticated +records refer to a group of small and comparatively insignificant +spiders belonging to the genus _Latrodectus_, of the family Theridiidae. +The dread "Malmigniatte" of Corsica and South Europe, the "Karakurte" of +southeastern Russia, the "Katipo" of New Zealand, the "Mena-vodi" and +"Vancoho" of Madagascar, and our own _Latrodectus mactans_, all belong +to this genus, and concerning all of these the most circumstantial +accounts of their venomous nature are given. These accounts are not mere +fantastic stories by uneducated natives but in many cases are reports +from thoroughly trained medical men. + +The symptoms produced are general, rather than local. As summarized by +Kobert (1901) from a study of twenty-two cases treated in 1888, in the +Kherson (Russia) Government Hospital and Berislaw (Kherson) District +Hospital the typical case, aside from complications, exhibits the +following symptoms. The victim suddenly feels the bite, like the sting +of a bee. Swelling of the barely reddened spot seldom follows. The +shooting pains, which quickly set in, are not manifested at the point of +injury but localized at the joints of the lower limb and in the region +of the hip. The severity of the pain forces the victim to the hospital, +in spite of the fact that they otherwise have a great abhorrence of it. +The patient is unable to reach the hospital afoot, or, at least, not +without help, for there is usually inability to walk. The patient, even +if he has ridden, reaches the hospital covered with cold sweat and +continues to perspire for a considerable period. His expression +indicates great suffering. The respiration may be somewhat dyspnoeic, +and a feeling of oppression in the region of the heart is common. There +is great aversion to solid food, but increasing thirst for milk and tea. +Retention of urine, and constipation occur. Cathartics and, at night, +strong narcotics are desired. Warm baths give great relief. After three +days, there is marked improvement and usually the patient is dismissed +after the fifth. This summary of symptoms agrees well with other +trustworthy records. + +It would seem, then, that Riley and Howard (1889), who discussed a +number of accounts in the entomological literature, were fully justified +in their statement that "It must be admitted that certain spiders of the +genus _Latrodectus_ have the power to inflict poisonous bites, which may +(probably exceptionally and depending upon exceptional conditions) bring +about the death of a human being." + +And yet, until recently the evidence bearing on the question has been +most conflicting. The eminent arachnologist, Lucas, (1843) states that +he himself, had been repeatedly bitten by the Malmigniatte without any +bad effects. Dr. Marx, in 1890, gave before the Entomological Society of +Washington, an account of a series of experiments to determine whether +the bite of _Latrodectus mactans_ is poisonous or not. He described the +poison glands as remarkably small[A] and stated that he had introduced +the poison in various ways into guinea-pigs and rabbits without +obtaining any satisfactory results. Obviously, carefully conducted +experiments with the supposed venom were needed and fortunately they +have been carried out in the greatest detail by Kobert (1901). + +This investigator pointed out that there were two factors which might +account for the discrepancies in the earlier experiments. In the first +place, the poison of spiders, as of snakes, might be so exhausted after +two or three bites that further bites, following directly, might be +without visible effect. Secondly, the application of the poison by means +of the bite, is exceedingly inexact, since even after the most careful +selection of the point of application, the poison might in one instance +enter a little vein or lymph vessel, and in another case fail to do so. +Besides, there would always remain an incalculable and very large amount +externally, in the nonabsorptive epithelium. While all of these factors +enter into the question of the effect of the bite in specific instances, +they must be as nearly as possible obviated in considering the question +of whether the spiders really secrete a venom harmful to man. + +Kobert therefore sought to prepare extracts which would contain the +active principles of the poison and which could be injected in definite +quantities directly into the blood of the experimental animal. For this +purpose various parts of the spiders were rubbed up in a mortar with +distilled water, or physiological salt solution, allowed to stand for an +hour, filtered, and then carefully washed, by adding water drop by drop +for twenty-four hours. The filtrate and the wash-water were then united, +well mixed and, if necessary, cleared by centrifuging or by exposure to +cold. The mixture was again filtered, measured, and used, in part, for +injection and, in part, for the determination of the organic materials. + +Such an extract was prepared from the cephalothoraces of eight dried +specimens of the Russian _Latrodectus_ and three cubic centimeters of +this, containing 4.29 mg. of organic material, were injected into the +jugular vein of a cat weighing 2450 grams. The previously very active +animal was paralyzed and lay in whatever position it was placed. The +sensibility of the skin of the extremities and the rump was so reduced +that there was no reaction from cutting or sticking. There quickly +followed dyspnoea, convulsions, paralysis of the respiratory muscles +and of the heart. In twenty-eight minutes the cat was dead, after having +exhibited exactly the symptoms observed in severe cases of poisoning of +man from the bite of this spider. + +These experiments were continued on cats, dogs, guinea pigs and various +other animals. Not only extracts from the cephalothorax, but from other +parts of the body, from newly hatched spiders, and from the eggs were +used and all showed a similar virulence. Every effort was made to avoid +sources of error and the experiments, conducted by such a recognized +authority in the field of toxicology, must be accepted as conclusively +showing that this spider and, presumably, other species of the genus +_Latrodectus_ against which the clinical evidence is quite parallel, +possess a poison which paralyzes the heart and central nervous system, +with or without preliminary stimulus of the motor center. If the +quantity of the poison which comes into direct contact with the blood is +large, there may occur haemolysis and thrombosis of the vessels. + +On the other hand, check experiments were carried out, using similar +extracts of many common European spiders of the genera _Tegenaria_, +_Drassus_, _Agelena_, _Eucharia_ and _Argyroneta_, as well as the +Russian tarantula, _Lycosa singoriensis_. In no other case was the +effect on experimental animals comparable to the _Latrodectus_ extract. + +Kobert concludes that in its chemical nature the poison is neither an +alkaloid, nor a glycoside, nor an acid, but a toxalbumen, or poisonous +enzyme which is very similar to certain other animal poisons, notably +that of the scorpion. + +[Illustration: 9. Latrodectus mactans; (_a_) female, x3; (_b_) venter +of female; (_c_) dorsum of male. After Comstock.] + +The genus _Latrodectus_ is represented in the United States by at least +two species, _L. mactans_ and _L. geometricus_. Concerning _L. mactans_ +there are very circumstantial accounts of serious injury and even death +in man[B]. _Latrodectus mactans_ is coal black, marked with red or +yellow or both. It has eight eyes, which are dissimilar in color and +are distinctly in front of the middle of the thorax, the lateral eyes of +each side widely separate. The tarsus of the fourth pair of legs has a +number of curved setae in a single series. It has on the ventral side of +its abdomen an hour-glass shaped spot. The full-grown female is about +half an inch in length. Its globose abdomen is usually marked with one +or more red spots dorsally along the middle line. The male is about half +as long but has in addition to the dorsal spots, four pairs of stripes +along the sides. Immature females resemble the male in coloring (fig. +9). + +Regarding the distribution of _Latrodectus mactans_, Comstock states +that: "Although it is essentially a Southern species, it occurs in +Indiana, Ohio, Pennsylvania, New Hampshire, and doubtless other of the +Northern States." _L. geometricus_ has been reported from California. + + +OTHER VENOMOUS SPIDERS--While conclusive evidence regarding the venomous +nature of spiders is meager and relates almost wholly to that of the +genus _Latrodectus_, the group is a large one and we are not justified +in dismissing arbitrarily, all accounts of injury from their bites. +Several species stand out as especially needing more detailed +investigation. + +_Chiracanthium nutrix_ is a common European species of the family +Clubionidae, concerning which there is much conflicting testimony. Among +the reports are two by distinguished scientists whose accounts of +personal experiences cannot be ignored. A. Forel allowed a spider of +this species to bite him and not only was the pain extreme, but the +general symptoms were so severe that he had to be helped to his house. +The distinguished arachnologist, Bertkau reports that he, himself, was +bitten and that an extreme, burning pain spread almost instantaneously +over the arm and into the breast. There were slight chills the same day +and throbbing pain at the wound lasted for days. While this particular +species is not found in the United States, there are two other +representatives of the genus and it is possible that they possess the +same properties. We are unaware of any direct experimental work on the +poison. + +_Epeira diadema_, of Europe, belongs to a wholly different group, that +of the orb-weavers, but has long been reputed venomous. Kobert was able +to prepare from it an extract whose effects were very similar to that +prepared from _Latrodectus_, though feebler in its action. Under +ordinary circumstances this spider is unable to pierce the skin of man +and though Kobert's results seem conclusive, the spider is little to be +feared. + +_Phidippus audax_ (_P. tripunctatus_) is one of our largest Attids, or +jumping spiders. The late Dr. O. Lugger describes a case of severe +poisoning from the bite of this spider and though details are lacking, +it is quite possible that this and other large species of the same +group, which stalk their prey, may possess a more active poison than +that of web-building species. + + +SUMMARY--It is clearly established that our common spiders are not to be +feared and that the stories regarding their virulence are almost wholly +without foundation. On the other hand, the chances of secondary +infection from the bites of some of the more powerful species are not to +be ignored. + +Probably all species possess a toxin secreted by the poison gland, +virulent for insects and other normal prey of the spiders, but with +little or no effect on man. + +There are a very few species, notably of the genus _Latrodectus_, and +possibly including the European _Chiracanthium nutrix_ and _Epeira +diadema_, which possess, in addition, a toxalbumen derived from the +general body tissue, which is of great virulence and may even cause +death in man and the higher animals. + +[Illustration: 10. A whip-scorpion (Mastigoproctus giganteus). Half +natural size. After Comstock.] + + +THE PEDIPALPIDA OR WHIP-SCORPIONS + +The tailed whip-scorpions, belonging to the family Thelyphonidae, are +represented in the United States by the giant whip-scorpion +_Mastigoproctus giganteus_ (fig. 10), which is common in Florida, Texas +and some other parts of the South. In Florida, it is locally known as +the "grampus" or "mule-killer" and is very greatly feared. There is no +evidence that these fears have any foundation, and Dr. Marx states that +there is neither a poison gland nor a pore in the claw of the +chelicera. + + +THE SCORPIONIDA, OR TRUE SCORPIONS + +The true scorpions are widely distributed throughout warm countries and +everywhere bear an evil reputation. According to Comstock (1912), about +a score of species occur in the Southern United States. These are +comparatively small forms but in the tropics members of this group may +reach a length of seven or eight inches. They are pre-eminently +predaceous forms, which lie hidden during the day and seek their prey by +night. + +[Illustration: 11. A true scorpion. After Comstock.] + +The scorpions (fig. 11) possess large pedipalpi, terminated by strongly +developed claws, or chelae. They may be distinguished from all other +Arachnids by the fact that the distinctly segmented abdomen is divided +into a broad basal region of seven segments and a terminal, slender, +tail-like division of five distinct segments. + +The last segment of the abdomen, or telson, terminates in a +ventrally-directed, sharp spine, and contains a pair of highly developed +poison glands. These glands open by two small pores near the tip of the +spine. Most of the species when running carry the tip of the abdomen +bent upward over the back, and the prey, caught and held by the +pedipalpi, is stung by inserting the spine of the telson and allowing it +to remain for a time in the wound. + +The glands themselves have been studied in _Prionurus citrinus_ by +Wilson (1904). He found that each gland is covered by a sheet of muscle +on its mesal and dorsal aspects, which may be described as the +_compressor muscle_. The muscle of each side is inserted by its edge +along the ventral inner surface of the chitinous wall of the telson, +close to the middle line, and by a broader insertion laterally. A layer +of fine connective tissue completely envelops each gland and forms the +basis upon which the secreting cells rest. The secreting epithelium is +columnar; and apparently of three different types of cells. + +1. The most numerous have the appearance of mucous cells, resembling the +goblet cells of columnar mucous membranes. The nucleus, surrounded by a +small quantity of protoplasm staining with haematoxylin, lies close to +the base of the cell. + +2. Cells present in considerable numbers, the peripheral portions of +which are filled with very numerous fine granules, staining with acid +dyes such as methyl orange. + +3. Cells few in number, filled with very large granules, or irregular +masses of a substance staining with haematoxylin. + +The poison, according to Kobert (1893), is a limpid, acid-reacting +fluid, soluble in water but insoluble in absolute alcohol and ether. +There are few data relative to its chemical nature. Wilson (1901) states +that a common Egyptian species, _Buthus quinquestriatus_, has a specific +gravity of 1.092, and contains 20.3% of solids and 8.4% ash. + +The venom of different species appears to differ not only quantitatively +but qualitatively. The effects of the bite of the smaller species of the +Southern United States may be painful but there is no satisfactory +evidence that it is ever fatal. On the other hand, certain tropical +species are exceedingly virulent and cases of death of man from the bite +are common. + +In the case of _Buthus quinquestriatus_, Wilson (1904) found the +symptoms in animals to be hypersecretion, salivation and lachrymation, +especially marked, convulsions followed by prolonged muscular spasm; +death from asphyxia. The temperature shows a slight, rarely +considerable, rise. Rapid and considerable increase of blood-pressure +(observed in dogs) is followed by a gradual fall with slowing of the +heart-beat. The coagulability of the blood is not affected. + +An interesting phase of Wilson's work was the experiments on desert +mammals. The condition under which these animals exist must frequently +bring them in contact with scorpions, and he found that they possess a +degree of immunity to the venom sufficient at least to protect them from +the fatal effects of the sting. + +As far as concerns its effect on man, Wilson found that much depended +upon the age. As high as 60 per cent of the cases of children under +five, resulted fatally. Caroroz (1865), states that in a Mexican state +of 15,000 inhabitants, the scorpions were so abundant and so much feared +that the authorities offered a bounty for their destruction. A result +was a large number of fatalities, over two hundred per year. Most of the +victims were children who had attempted to collect the scorpions. + +The treatment usually employed in the case of bites by the more +poisonous forms is similar to that for the bite of venomous snakes. +First, a tight ligature is applied above the wound so as to stop the +flow of blood and lymph from that region. The wound is then freely +excised and treated with a strong solution of permanganate of potash, or +with lead and opium lotion. + +In recent years there have been many attempts to prepare an antivenom, +or antiserum comparable to what has been used so effectively in the case +of snake bites. The most promising of these is that of Todd (1909), +produced by the immunization of suitable animals. This antivenom proved +capable of neutralizing the venom when mixed _in vitro_ and also acts +both prophylactically and curatively in animals. Employed curatively in +man, it appears to have a very marked effect on the intense pain +following the sting, and the evidence so far indicates that its prompt +use greatly reduces the chance of fatal results. + + +THE SOLPUGIDA, OR SOLPUGIDS + +The SOLPUGIDA are peculiar spider-like forms which are distinguished +from nearly all other arachnids by the fact that they possess no true +cephalothorax, the last two leg-bearing segments being distinct, +resembling those of the abdomen in this respect. The first pair of legs +is not used in locomotion but seemingly functions as a second pair of +pedipalpi. Figure 12 illustrates the striking peculiarities of the +group. They are primarily desert forms and occur in the warm zones of +all countries. Of the two hundred or more species, Comstock lists twelve +as occurring in our fauna. These occur primarily in the southwest. + +[Illustration: 12. A solpugid (Eremobates cinerea). After Comstock.] + +The Solpugida have long borne a bad reputation and, regarding virulence, +have been classed with the scorpions. Among the effects of their bites +have been described painful swelling, gangrene, loss of speech, cramps, +delirium, unconsciousness and even death. Opposed to the numerous loose +accounts of poisoning, there are a number of careful records by +physicians and zoologists which indicate clearly that the effects are +local and though they may be severe, they show not the slightest symptom +of direct poisoning. + +More important in the consideration of the question is the fact that +there are neither poison glands nor pores in the fangs for the exit of +any poisonous secretion. This is the testimony of a number of prominent +zoologists, among whom is Dr. A. Walter, who wrote to Kobert at length +on the subject and whose conclusions are presented by him. + +However, it should be noted that the fangs are very powerful and are +used in such a manner that they may inflict especially severe wounds. +Thus, there may be more opportunity for secondary infection than is +usual in the case of insect wounds. + +The treatment of the bite of the Solpugida is, therefore, a matter of +preventing infection. The wound should be allowed to bleed freely and +then washed out with a 1:3000 solution of corrosive sublimate, and, if +severe, a wet dressing of this should be applied. If infection takes +place, it should be treated in the usual manner, regardless of its +origin. + + +THE ACARINA, OR MITES AND TICKS + +A number of the parasitic Acarina evidently secrete a specific poison, +presumably carried by the saliva, but in most cases its effect on man is +insignificant. There is an abundant literature dealing with the +poisonous effect of the bite of these forms, especially the ticks, but +until recently it has been confused by failure to recognize that various +species may transmit diseases of man, rather than produce injury through +direct poisoning. We shall therefore discuss the Acarina more especially +in subsequent chapters, dealing with parasitism and with disease +transmission. + +Nevertheless, after the evidence is sifted, there can be no doubt that +the bites of certain ticks may occasionally be followed by a direct +poisoning, which may be either local or general in its effects. Nuttall +(1908) was unable to determine the cause of the toxic effect, for, in +_Argas persicus_, the species most often implicated, he failed to get +the slightest local or general effect on experimental animals, from the +injection of an emulsion prepared by crushing three of the ticks. + +It seems clearly established that the bite of certain ticks may cause a +temporary paralysis, or even complete paralysis, involving the organs of +respiration or the heart, and causing death. In 1912, Dr. I. U. Temple, +of Pendleton, Oregon, reported several cases of what he called "acute +ascending paralysis" associated with the occurrence of ticks on the head +or the back of the neck. A typical severe case was that of a six year +old child, who had retired in her usual normal health. The following +morning upon arising she was unable to stand on her feet. She exhibited +paralysis extending to the knees, slight temperature, no pain, sensory +nerves normal, motor nerves completely paralyzed, reflexes absent. The +following day the paralysis had extended to the upper limbs, and before +night of the third day the nerves of the throat (hypoglossal) were +affected. The thorax and larynx were involved, breathing was labored, +she was unable to swallow liquids, phonation was impossible and she +could only make low, guttural sounds. At this stage, two ticks, fully +distended with blood, were found over the junction of the spinal column +with the occipital bones in the hollow depression. They were removed by +the application of undiluted creoline. Though the child's life was +despaired of, by the following morning she was very much improved. By +evening she was able to speak. The paralysis gradually receded, +remaining longest in the feet, and at the end of one week the patient +was able to go home. + +There was some doubt as to the exact species of tick implicated in the +cases which Dr. Temple reported, although the evidence pointed strongly +to _Dermacentor venustus_.[C] Somewhat later, Hadwen (1913) reported +that "tick paralysis" occurs in British Columbia, where it affects not +only man, but sheep and probably other animals. It is caused by the +bites of _Dermacentor venustus_ and was experimentally produced in lambs +and a dog (Hadwen and Nuttall, 1913). It is only when the tick begins to +engorge or feed rapidly, some days after it has become attached, that +its saliva produces pathogenic effects. + +Ulceration following tick bite is not uncommon. In many of the instances +it is due to the file-like hypostome, with its recurved teeth, being +left in the wound when the tick is forcibly pulled off. + + +THE MYRIAPODA, OR CENTIPEDES AND MILLIPEDES + +The old class, Myriapoda includes the DIPLOPODA, or millipedes, and the +CHILOPODA, or centipedes. The present tendency is to raise these groups +to the rank of classes. + + +The Diplopoda + +The DIPLOPODA, or millipedes (fig. 13), are characterized by the +presence of two pairs of legs to a segment. The largest of our local +myriapods belong to this group. They live in moist places, feeding +primarily on decaying vegetable matter, though a few species +occasionally attack growing plants. + +[Illustration: 13. A millipede. After Comstock.] + +The millipedes are inoffensive and harmless. _Julus terrestris_, and +related species, when irritated pour out over the entire body a +yellowish secretion which escapes from cutaneous glands. It is volatile, +with a pungent odor, and Phisalix (1900) has shown that it is an active +poison when injected into the blood of experimental animals. This, +however, does not entitle them to be considered as poisonous arthropods, +in the sense of this chapter, any more than the toad can be considered +poisonous to man because it secretes a venom from its cutaneous glands. + + +The Chilopoda + +[Illustration: 14. Two common centipedes. + +(_a_) Lithobius forficatus. After Comstock. + +(_b_) Scutigera forceps. Natural size; after Howard.] + +The CHILOPODA, or centipedes (fig. 14), unlike the millipedes, are +predaceous forms, and possess well developed poison glands for killing +their prey. These glands are at the base of the first pair of legs +(fig. 15), which are bent forward so as to be used in holding their +prey. The legs terminate in a powerful claw, at the tip of which is the +outlet of the poison glands. + +The poison is a limpid, homogeneous, slightly acid fluid, which +precipitates in distilled water. Briot (1904) extracted it from the +glands of _Scolopendra morsitans_, a species common in central France, +and found that it was actively venomous for the ordinary experimental +animals. A rabbit of two kilograms weight received an injection of three +cubic centimeters in the vein of the ear and died in a minute. A white +rat, weighing forty-eight grams, received one and a half cubic +centimeters in the hind leg. There was an almost immediate paralysis of +the leg and marked necrosis of the tissues. + +[Illustration: 15. Mandible of Scolopendra cingulata showing venom +gland. After Dubosq.] + +As for the effect on man, there is little foundation for the fear with +which centipedes are regarded. Our native species produce, at most, +local symptoms,--sometimes severe local pain and swelling,--but there is +no authentic record of fatal results. In the tropics, some of the +species attain a large size, _Scolopendra gigantea_ reaching a length of +nearly a foot. These forms are justly feared, and there is good evidence +that death sometimes, though rarely, results from their bite. + +One of the most careful accounts of death from the sting of the scorpion +is that of Linnell, (1914), which relates to a comparatively small +Malayan species, unfortunately undetermined. The patient, a coolie, aged +twenty, was admitted to a hospital after having been stung two days +previously on the left heel. For cure, the other coolies had made him +eat the head of the scorpion. On admission, the patient complained of +"things creeping all over the body". Temp. 102.8^o. On the fourth day he +had paralysis of the legs, and on the fifth day motor paralysis to the +umbilicus, sensation being unaltered. On the sixth day there was +retention of the urine and on the ninth day (first test after third day) +sugar was present. On the thirteenth day the patient became comatose, +but could be roused to eat and drink. The temperature on the following +day fell below 95^o and the patient was still comatose. Death fifteenth +day. + +Examination of the spinal (lumbar) cord showed acute disseminated +myelitis. In one part there was an acute destruction of the anterior +horn and an infiltration of round cells. In another portion Clarke's +column had been destroyed. The perivascular sheaths were crowded with +small round cells and the meninges were congested. Some of the cells of +the anterior horn were swollen and the nuclei eccentric; chromatolysis +had occurred in many of them. + +As for treatment, Castellani and Chalmers (1910), recommend bathing the +part well with a solution of ammonia (one in five, or one in ten). After +bathing, apply a dressing of the same alkali or, if there is much +swelling and redness, an ice-bag. If necessary, hypodermic injections of +morphine may be given to relieve the pain. At a later period +fomentations may be required to reduce the local inflammation. + + +THE HEXAPODA OR TRUE INSECTS + +There are a number of HEXAPODA, or true insects, which are, in one way +or another, poisonous to man. These belong primarily to the orders +Hemiptera, or true bugs; Lepidoptera, or butterflies and moths (larval +forms); Diptera, or flies; Coleoptera, or beetles; and Hymenoptera, or +ants, bees, and wasps. There are various ways in which they may be +poisonous. + +1. _Piercing_ or _biting_ forms may inject an irritating or poisonous +saliva into the wound caused by their mouth-parts. + +2. _Stinging forms_ may inject a poison, from glands at the caudal end +of the abdomen, into wounds produced by a specially modified ovipositer, +the _sting_. + +3. _Nettling_ properties may be possessed by the hairs of the insect. + +4. _Vescicating_, or _poisonous blood plasma_, or _body fluids_ are +known to exist in a large number of species and may, under exceptional +circumstances, affect man. + +For convenience of discussion, we shall consider poisonous insects under +these various headings. In this, as in the preceding discussion, no +attempt will be made to give an exhaustive list of the poisonous forms. +Typical instances will be selected and these will be chosen largely from +North American species. + + +PIERCING OR BITING INSECTS POISONOUS TO MAN + + +HEMIPTERA + +Several families of the true bugs include forms which, while normally +inoffensive, are capable of inflicting painful wounds on man. In these, +as in all of the Hemiptera, the mouth-parts are modified to form an +organ for piercing and sucking. This is well shown by the accompanying +illustration (fig. 16). + +The upper lip, or _labrum_, is much reduced and immovable, the lower +lip, or _labium_, is elongated to form a jointed sheath, within which +the lance-like mandibles and maxillae are enclosed. The mandibles are +more or less deeply serrate, depending on the species concerned. + +[Illustration: 16. Beak of hemipteron.] + +The poison is elaborated by the salivary glands, excepting, possibly, in +_Belostoma_ where Locy is inclined to believe that it is secreted by the +maxillary glands. The salivary glands of the Hemiptera have been the +subject of much study but the most recent, comprehensive work has been +done by Bugnion and Popoff, (1908 and 1910) to whose text the reader is +referred for details. + +The Hemiptera have two pairs of salivary glands: the _primary gland_, of +which the efferent duct leads to the salivary syringe, and the +_accessory gland_, of which the very long and flexuous duct empties into +the primary duct at its point of insertion. Thus, when one observes the +isolated primary gland it appears as though it had efferent ducts +inserted at the same point. In _Nepa_ and the _Fulgoridae_ there are two +accessory glands and therefore apparently three ducts at the same point +on the primary gland. The _ensemble_ differs greatly in appearance in +different species but we shall show here Bugnion and Popoff's figure of +the apparatus of _Notonecta maculata_, a species capable of inflicting a +painful bite on man (fig. 17). + +[Illustration: 17. Salivary glands of Notonecta maculata. After Bugnion +and Popoff.] + +[Illustration: 18. Pharyngeal syringe or salivary pump of Fulgora +maculata. After Bugnion and Popoff.] + +[Illustration: 19. Heteroptera, (_a_) Melanolestes picipes; (_b_) +Notonecta undulata; (_c_, _d_) Aradus robustus (_c_) adult, (_d_) nymph, +much enlarged; (_e_) Arilus cristatus; (_f_) Belostoma americana; (_g_) +Nabis (Coriscus) subcoleoptratus, enlarged; (_h_) Cimex lectularius, +(_i_) Oeciacus vicarius, much enlarged; (_j_) Lyctocoris fitchii, much +enlarged. After Lugger.] + +Accessory to the salivary apparatus there is on the ventral side of the +head, underneath the pharynx, a peculiar organ which the Germans have +called the "Wanzenspritze," or syringe. The accompanying figure of the +structure in _Fulgora maculata_ (fig. 18) shows its relation to the +ducts of the salivary glands and to the beak. It is made up of a +dilatation forming the body of the pump, in which there is a chitinous +piston. Attached to the piston is a strong retractor muscle. The +function of the salivary pump is to suck up the saliva from the salivary +ducts and to force it out through the beak. + +Of the Hemiptera reported as attacking man, we shall consider briefly +the forms most frequently noted. + + +The NOTONECTIDAE, or _back swimmers_, (fig. 19_b_) are small, aquatic +bugs that differ from all others in that they always swim on their +backs. They are predaceous; feeding on insects and other small forms. +When handled carelessly they are able to inflict a painful bite, which +is sometimes as severe as the sting of a bee. In fact, they are known in +Germany as "Wasserbienen." + + +The BELOSTOMATIDAE, or _giant water bugs_, (fig. 19_f_) include the +largest living Hemiptera. They are attracted to lights and on account of +the large numbers which swarm about the electric street lamps in some +localities they have received the popular name "electric light bugs." +Our largest representatives in the northern United States belong to the +two genera _Belostoma_ and _Banacus_, distinguished from each other by +the fact that _Belostoma_ has a groove on the under side of the femur of +the front leg, for the reception of the tibia. + +The salivary glands of Belostoma were figured by Leidy (1847) and later +were studied in more detail by Locy (1884). There are two pairs of the +glands, those of one pair being long and extending back as far as the +beginning of the abdomen, while the others are about one-fourth as long. +They lie on either side of the oesophagus. On each side of the +oesophagus there is a slender tube with a sigmoid swelling which may +serve as a poison reservoir. In addition to this salivary system, there +is a pair of very prominent glands on the ventral side of the head, +opening just above the base of the beak. These Locy has called the +"cephalic glands" and he suggests that they are the source of the +poison. They are the homologues of the maxillary glands described for +other Hemiptera, and it is by no means clear that they are concerned +with the production of venom. It seems more probable that in +_Belostoma_, as in other Hemiptera, it is produced by the salivary +glands, though the question is an open one. + +The Belostomatidae feed not only on insects, but on small frogs, fish, +salamanders and the like. Matheson (1907) has recorded the killing of a +good-sized bird by _Belostoma americana_. A woodpecker, or flicker, was +heard to utter cries of distress, and fluttered and fell from a tree. On +examination it was found that a bug of this species had inserted its +beak into the back part of the skull and was apparently busily engaged +in sucking the blood or brains of the bird. Various species of +_Belostoma_ have been cited as causing painful bites in man. We can +testify from personal experience that the bite of _Belostoma americana_ +may almost immediately cause severe, shooting pains that may extend +throughout the arm and that they may be felt for several days. + +[Illustration: 20. Reduvius (Opsicoetus) personatus. (x2).] + +Relief from the pain may be obtained by the use of dilute ammonia, or a +menthol ointment. In the not uncommon case of secondary infection the +usual treatment for that should be adopted. + +[Illustration: 21. (_a_) Reduvius personatus, nymph. Photograph by M. V. +S.] + +The REDUVIIDAE, or _assassin-bugs_ are capable of inflicting very painful +wounds, as most collectors of Hemiptera know to their sorrow. Some +species are frequently to be found in houses and outhouses and Dr. +Howard suggests that many of the stories of painful spider bites relate +to the attack of these forms. + +[Illustration: 21. (_b_) Reduvius personatus, adult (x2) Photograph by +M. V. S.] + +An interesting psychological study was afforded in the summer of 1899, +by the "kissing-bug" scare which swept over the country. It was reported +in the daily papers that a new and deadly bug had made its appearance, +which had the unpleasant habit of choosing the lips or cheeks for its +point of attack on man. So widespread were the stories regarding this +supposedly new insect that station entomologists all over the country +began to receive suspected specimens for identification. At Cornell +there were received, among others, specimens of stone-flies, may-flies +and even small moths, with inquiries as to whether they were +"kissing-bugs." + +[Illustration: 22. Rasahus biguttatus. (x2). After Howard.] + +Dr. L. O. Howard has shown that the scare had its origin in newspaper +reports of some instances of bites by either _Melanolestes picipes_ +(fig. 19a) or _Opsicoetes personatus_ (fig. 20), in the vicinity of +Washington, D. C. He then discusses in considerable detail the more +prominent of the Reduviidae which, with greater or less frequency pierce +the skin of human beings. These are _Opsicoetes personatus_, +_Melanolestes picipes_, _Coriscus subcoleoptratus_ (fig. 19_g_), +_Rasahus thoracicus_, _Rasahus biguttatus_ (fig. 22), _Conorhinus +sanguisugus_ (fig. 71), and _Melanolestes abdominalis_ (fig. 23). + +[Illustration: 23. Melanolestes abdominalis (x2). After Marlatt.] + +One of the most interesting of these species is _Reduvius personatus_, +(= _Opsicoetus personatus_), which is popularly known as the "masked +bed-bug hunter." It owes this name to the fact that the immature nymphs +(fig. 21) have their bodies and legs completely covered by dust and +lint, and that they are supposed to prey upon bed-bugs. LeConte is +quoted by Howard as stating that "This species is remarkable for the +intense pain caused by its bite. I do not know whether it ever willingly +plunges its rostrum into any person, but when caught, or unskilfully +handled it always stings. In this case the pain is almost equal to the +bite of a snake, and the swelling and irritation which result from it +will sometimes last for a week." + +A species which very commonly attacks man is _Conorhinus sanguisugus_, +the so-called "big bed-bug" of the south and southern United States. It +is frequently found in houses and is known to inflict an exceedingly +painful bite. As in the case of a number of other predaceous Hemiptera, +the salivary glands of these forms are highly developed. The effect of +the bite on their prey and, as Marlatt has pointed out, the constant and +uniform character of the symptoms in nearly all cases of bites in man, +clearly indicate that their saliva contains a specific substance. No +satisfactory studies of the secretions have been made. On the other +hand, Dr. Howard is doubtless right in maintaining that the very serious +results which sometimes follow the bite are due to the introduction of +extraneous poison germs. This is borne out by the symptoms of most of +the cases cited in literature and also by the fact that treatment with +corrosive sublimate, locally applied to the wound, has yielded favorable +results. + + +OTHER HEMIPTERA REPORTED AS POISONOUS TO MAN--A large number of other +Hemiptera have been reported as attacking man. Of these, there are +several species of Lygaeidae, Coreidae, and Capsidae. Of the +latter, _Lygus pratensis_, the tarnished plant-bug, is reported by +Professor Crosby as sucking blood. _Orthotylus flavosparsus_ is another +Capsid which has been implicated. _Empoasca mali_ and _Platymetopius +acutus_ of the Jassidae have also been reported as having similar +habits. + +Whenever the periodical cicada or "seventeen-year locust" becomes +abundant, the newspapers contain accounts of serious results from its +bites. The senior author has made scores of attempts to induce this +species to bite and only once successfully. At that time the bite was in +no wise more severe than a pin-prick. A student in our department +reports a similar experience. There is no case on record which bears +evidence of being worthy of any credence, whatsoever. + +Under the heading of poisonous Hemiptera we might consider the bed-bugs +and the lice. These will be discussed later, as parasites and as +carriers of disease, and therefore need only be mentioned here. + + +DIPTERA + +Several species of blood-sucking Diptera undoubtedly secrete a saliva +possessing poisonous properties. Chief among these are the Culicidae, or +mosquitoes, and the Simuliidae, or black-flies. As we shall consider +these forms in detail under the heading of parasitic species and +insects transmitting disease, we shall discuss here only the poison of +the mosquitoes. + +It is well known that mosquitoes, when they bite, inject into the wound +a minute quantity of poison. The effect of this varies according to the +species of mosquito and also depends very much on the susceptibility of +the individual. Soon after the bite a sensation of itching is noticed +and often a wheal, or eminence, is produced on the skin, which may +increase to a considerable swelling. The scratching which is induced may +cause a secondary infection and thus lead to serious results. Some +people seem to acquire an immunity against the poison. + +The purpose of this irritating fluid may be, as Reaumur suggested, to +prevent the coagulation of the blood and thus not only to cause it to +flow freely when the insect bites but to prevent its rapid coagulation +in the stomach. Obviously, it is not developed as a protective fluid, +and its presence subjects the group to the additional handicap of the +vengeance of man. + +[Illustration: 24. Diagram of a longitudinal section of a mosquito.] + +As to the origin of the poison, there has been little question, until +recent years, that it was a secretion from the salivary glands. +Macloskie (1888) showed that each gland is subdivided into three lobes, +the middle of which differs from the others in having evenly granulated +contents and staining more deeply than the others (fig. 24). This middle +lobe he regarded as the source of the poison. Bruck, (1911), by the use +of water, glycerine, chloroform, and other fluids, extracted from the +bodies of a large number of mosquitoes a toxine which he calls +_culicin_. This he assumes comes from the salivary glands. Animal +experimentation showed that this extract possessed hemolytic powers. +Inoculated into the experimenter's own skin it produced lesions which +behaved exactly as do those of mosquito bites. + +Similarly, most writers on the subject have concurred with the view that +the salivary glands are the source of the poison. However, recent work, +especially that of Nuttall and Shipley (1903), and Schaudinn (1904), has +shown that the evidence is by no means conclusive. Nuttall dissected out +six sets (thirty-six acini) of glands from freshly killed _Culex +pipiens_ and placed them in a drop of salt solution. The drop was +allowed to dry, it being thought that the salt crystals would facilitate +the grinding up of the glands with the end of a small glass rod, this +being done under microscopic control. After grinding up, a small drop of +water was added of the size of the original drop of saline, and an equal +volume of human blood taken from the clean finger-tip was quickly mixed +therewith, and the whole drawn up into a capillary tube. Clotting was +not prevented and no hemolysis occurred. Salivary gland emulsion added +to a dilute suspension of corpuscles did not lead to hemolysis. This +experiment was repeated a number of times, with slight modification, but +with similar results. The data obtained from the series "do not support +the hypothesis that the salivary glands, at any rate in _Culex pipiens_, +contain a substance which prevents coagulation." + +Much more detailed, and the more important experiments made along this +line, are those of Schaudinn (1904). The results of these experiments +were published in connection with a technical paper on the alternation +of generations and of hosts in _Trypanosoma_ and _Spirochaeta_, and for +this reason seem to have largely escaped the notice of entomologists. +They are so suggestive that we shall refer to them in some detail. + +Schaudinn observed that the three oesophageal diverticula (commonly, +but incorrectly, known as the "sucking stomach") (fig. 24) usually +contain large bubbles of gas and in addition, he always found yeast +cells. On the ground of numerous observations, Schaudinn was convinced +that these yeast plants are normal and constant commensals of the +insect. He regarded them as the cause of the gas bubbles to be found in +diverticula. It was found that as the insect fed, from time to time the +abdomen underwent convulsive contractions which resulted in the emptying +of the oesophageal diverticula and the salivary glands through blood +pressure. + +In order to test the supposed toxic action of the salivary glands, +Schaudinn repeatedly introduced them under his skin and that of his +assistant, in a drop of salt solution, and never obtained a suggestion +of the irritation following a bite of the insect, even though the glands +were carefully rubbed to fragments after their implantation. Like +Nuttall, he failed to get satisfactory evidence that the secretion of +the salivary glands retarded coagulation of the blood. + +He then carefully removed the oesophageal diverticula with their +content of yeast and introduced them into an opening in the skin of the +hand. Within a few seconds there was noticeable the characteristic +itching irritation of the mosquito bite; and in a short time there +appeared reddening and typical swelling. This was usually much more +severe than after the usual mosquito bite, and the swelling persisted +and itched longer. This was because by the ordinary bite of the mosquito +most of the yeast cells are again sucked up, while in these experiments +they remained in the wound. These experiments were repeated a number of +times on himself, his assistant and others, and always with the same +result. From them Schaudinn decided that the poisonous action of the +mosquito bite is caused by an enzyme from a commensal fungus. These +conclusions have not, as yet, been satisfactorily tested. + +Relief from the effect of the mosquito bite may be obtained by bathing +the swellings with weak ammonia or, according to Howard, by using moist +soap. The latter is to be rubbed gently on the puncture and is said to +speedily allay the irritation. Howard also quotes from the _Journal of +Tropical Medicine and Hygiene_ to the effect that a few drops of a +solution of thirty to forty grains of iodine to an ounce of saponated +petroleum rubbed into the mosquito bite, or wasp sting, allay the pain +instantaneously. + +Methods of mosquito control will be discussed later, in considering +these insects as parasites and as carriers of disease. + + +STINGING INSECTS + +The stinging insects all belong to the order HYMENOPTERA. In a number of +families of this group the ovipositor is modified to form a sting and is +connected with poison-secreting glands. We shall consider the apparatus +of the honey-bee and then make briefer reference to that of other forms. + + +APIS MELLIFICA, THE HONEY BEE--The sting of the worker honey-bee is +situated within a so-called sting chamber at the end of the abdomen. +This chamber is produced by the infolding of the greatly reduced and +modified eighth, ninth and tenth abdominal segments into the seventh.[D] +From it the dart-like sting can be quickly exserted. + +The sting (fig. 25) is made up of a central shaft, ventro-laterad of +which are the paired _lancets_, or darts, which are provided with sharp, +recurved teeth. Still further laterad lie the paired whitish, +finger-like _sting palpi_. Comparative morphological as well as +embryological studies have clearly established that these three parts +correspond to the three pairs of gonopophyses of the ovipositor of more +generalized insects. + +[Illustration: 25. Sting of a honey bee. _Psn Sc_, base of acid poison +gland; _B Gl_, alkaline poison gland; _Stn Plp_, sting palpi; _Sh B_, +bulb of sting; _Sh A_, basal arm; _Lct_, lancets or darts; _Sh s_, shaft +of sting. Modified from Snodgrass.] + +[Illustration: 26. Poison apparatus of a honey bee. Modified from +Snodgrass.] + +An examination of the internal structures (fig. 26) reveals two distinct +types of poison glands, the acid-secreting and the alkaline-secreting +glands, and a prominent poison reservoir. In addition, there is a small +pair of accessory structures which have been called lubricating glands, +on account of the supposed function of their product. The acid-secreting +gland empties into the distal end of the poison reservoir which in turn +pours the secretion into the muscular bulb-like enlargement at the base +of the shaft. The alkaline secreting gland empties into the bulb ventrad +of the narrow neck of the reservoir. + +The poison is usually referred to as formic acid. That it is not so +easily explained has been repeatedly shown and is evidenced by the +presence of the two types of glands. Carlet maintains that the product +of either gland is in itself innocent,--it is only when they are +combined that the toxic properties appear. + +The most detailed study of the poison of the honey-bee is that of Josef +Langer (1897), who in the course of his work used some 25,000 bees. +Various methods of obtaining the active poison for experimental purposes +were used. For obtaining the pure secretion, bees were held in the +fingers and compressed until the sting was exserted, when a clear drop +of the poison was visible at its tip. This was then taken up in a +capillary tube or dilute solutions obtained by dipping the tip of the +sting into a definite amount of distilled water. + +An aqueous solution of the poison was more readily obtained by pulling +out the sting and poison sacs by means of forceps, and grinding them up +in water. The somewhat clouded fluid was then filtered one or more +times. For obtaining still greater quantities, advantage was taken of +the fact that while alcohol coagulates the poison, the active principle +remains soluble in water. Hence the stings with the annexed glands where +collected in 96 per cent alcohol, after filtering off of the alcohol +were dried at 40^o C., then rubbed to a fine powder and this was +repeatedly extracted with water. Through filtering of this aqueous +extract there was obtained a yellowish-brown fluid which produced the +typical reactions, according to concentration of the poison. + +The freshly expelled drop of poison is limpid, of distinct acid +reaction, tastes bitter and has a delicate aromatic odor. On +evaporation, it leaves a sticky residue, which at 100 degrees becomes +fissured, and suggests dried gum arabic. The poison is readily soluble +in water and possesses a specific gravity of 1.1313. On drying at room +temperature, it leaves a residue of 30 per cent, which has not lost in +poisonous action or in solubility. In spite of extended experiments, +Langer was unable to determine the nature of the active principle. He +showed that it was not, as had been supposed, an albuminous body, but +rather an organic base. + +The pure poison, or the two per cent aqueous solution, placed on the +uninjured skin showed absolutely no irritating effect, though it +produced a marked reaction on the mucus membrane of the nose or eye. A +single drop of one-tenth per cent aqueous solution of the poison brought +about a typical irritation in the conjunctiva of the rabbit's eye. On +the other hand, the application of a drop of the poison, or its +solution, to the slightest break in the skin, or by means of a needle +piercing the skin, produced typical effects. There is produced a local +necrosis, in the neighborhood of which there is infiltration of +lymphocytes, oedema, and hyperaemia. + +The effect of the sting on man (fig. 27) is usually transitory but there +are some individuals who are made sick for hours, by a single sting. +Much depends, too, on the place struck. It is a common experience that +an angry bee will attempt to reach the eye of its victim and a sting on +the lid may result in severe and prolonged swelling. In the case of a +man stung on the cheek, Legiehn observed complete aphonia and a breaking +out of red blotches all over the body. A sting on the tongue has been +known to cause such collateral oedema as to endanger life through +suffocation. Cases of death of man from the attacks of bees are rare but +are not unknown. Such results are usually from a number of stings but, +rarely, death has been known to follow a single sting, entering a blood +vessel of a particularly susceptible individual. + +[Illustration: Effect of bee stings. After Root.] + +It is clearly established that partial immunity from the effects of the +poison may be acquired. By repeated injections of the venom, mice have +been rendered capable of bearing doses that certainly would have killed +them at first. It is a well-known fact that most bee-keepers become +gradually hardened to the stings, so that the irritation and the +swelling become less and less. Some individuals have found this immunity +a temporary one, to be reacquired each season. A striking case of +acquired immunity is related by the Roots in their "A B C and X Y Z of +Bee Culture." The evidence in the case is so clear that it should be +made more widely available and hence we quote it here. + +A young man who was determined to become a bee-keeper, was so +susceptible to the poison that he was most seriously affected by a +single sting, his body breaking out with red blotches, breathing growing +difficult, and his heart action being painfully accelerated. "We finally +suggested taking a live bee and pressing it on the back of his hand +until it merely pierced his skin with the sting, then immediately +brushing off both bee and sting. This was done and since no serious +effect followed, it was repeated inside of four or five days. This was +continued for some three or four weeks, when the patient began to have a +sort of itching sensation all over his body. The hypodermic injections +of bee-sting poison were then discontinued. At the end of a month they +were repeated at intervals of four or five days. Again, after two or +three weeks the itching sensation came on, but it was less pronounced. +The patient was given a rest of about a month, when the doses were +repeated as before." By this course of treatment the young man became so +thoroughly immunized that neither unpleasant results nor swelling +followed the attacks of the insects and he is able to handle bees with +the same freedom that any experienced bee-keeper does. + +In an interesting article in the _Entomological News_ for November, +1914, J. H. Lovell calls attention to the fact that "There has been a +widespread belief among apiarists that a beekeeper will receive more +stings when dressed in black than when wearing white clothing. A large +amount of evidence has been published in the various bee journals +showing beyond question that honey-bees under certain conditions +discriminate against black. A few instances may be cited in +illustration. Of a flock of twelve chickens running in a bee-yard seven +black ones were stung to death, while five light colored ones escaped +uninjured. A white dog ran among the bee-hives without attracting much +attention, while at the same time a black dog was furiously assailed by +the bees. Mr. J. D. Byer, a prominent Canadian beekeeper, relates that a +black and white cow, tethered about forty feet from an apiary, was one +afternoon attacked and badly stung by bees. On examination it was found +that the black spots had five or six stings to one on the white. All +noticed this fact, although no one was able to offer any explanation. A +white horse is in much less danger of being stung, when driven near an +apiary, than a black one. It has, indeed, been observed repeatedly that +domestic animals of all kinds, if wholly or partially black, are much +more liable to be attacked by bees, if they wander among the hives, than +those which are entirely white." + +In order to test the matter experimentally, the following series of +experiments was performed. In the language of the investigator: + +"On a clear, warm day in August I dressed wholly in white with the +exception of a black veil. Midway on the sleeve of my right arm there +was sewed a band of black cloth ten inches wide. I then entered the +bee-yard and, removing the cover from one of the hives, lifted a piece +of comb with both hands and gently shook it. Instantly many of the bees +flew to the black band, which they continued to attack as long as they +were disturbed. Not a single bee attempted to sting the left sleeve, +which was of course entirely white, and very few even alighted upon it." + +"This experiment was repeated a second, third and fourth time; in each +instance with similar results. I estimated the number of bees on the +band of black cloth at various moments was from thirty to forty; it was +evident from their behavior that they were extremely irritable. To the +left white sleeve and other portions of my clothing they paid very +little attention; but the black veil was very frequently attacked." + +"A few days later the experiments were repeated, but the band of black +cloth, ten inches wide, was sewed around my left arm instead of around +the right arm as before. When the bees were disturbed, after the hive +cover had been removed, they fiercely attacked the band of black cloth +as in the previous experiences; but the right white sleeve and the white +suit were scarcely noticed. At one time a part of the black cloth was +almost literally covered with furiously stinging bees, and the black +veil was assailed by hundreds. The bees behaved in a similar manner when +a second hive on the opposite side of the apiary was opened." + +"A white veil which had been procured for this purpose, was next +substituted for the black veil. The result was most surprising, for, +whereas in the previous experiments hundreds of bees had attacked the +black veil, so few flew against the white veil as to cause me no +inconvenience. Undoubtedly beekeepers will find it greatly to their +advantage to wear white clothing when working among their colonies of +bees and manipulating the frames of the hives." + +When a honey-bee stings, the tip of the abdomen, with the entire sting +apparatus, is torn off and remains in the wound. Here the muscles +continue to contract, for some minutes, forcing the barbs deeper and +deeper into the skin, and forcing out additional poison from the +reservoir. + +Treatment, therefore, first consists in removing the sting without +squeezing out additional poison. This is accomplished by lifting and +scraping it out with a knife-blade or the fingernail instead of grasping +and pulling it out. Local application of alkalines, such as weak +ammonia, are often recommended on the assumption that the poison is an +acid to be neutralized on this manner, but these are of little or no +avail. They should certainly not be rubbed in, as that would only +accelerate the absorption of the poison. The use of cloths wrung out in +hot water and applied as hot as can be borne, affords much relief in the +case of severe stings. The application of wet clay, or of the end of a +freshly cut potato is sometimes helpful. + +In extreme cases, where there is great susceptibility, or where there +may have been many stings, a physician should be called. He may find +strychnine injections or other treatment necessary, if general symptoms +develop. + +[Illustration: 28. The poison apparatus of Formica. Wheeler, after +Forel.] + + +OTHER STINGING FORMS--Of the five thousand, or more, species of bees, +most possess a sting and poison apparatus and some of the larger forms +are capable of inflicting a much more painful sting than that of the +common honey-bee. In fact, some, like the bumble bees, possess the +advantage that they do not lose the sting from once using it, but are +capable of driving it in repeatedly. In the tropics there are found many +species of stingless bees but these are noted for their united efforts +to drive away intruders by biting. Certain species possess a very +irritating saliva which they inject into the wounds. + +The ants are not ordinarily regarded as worthy of consideration under +the heading of "stinging insects" but as a matter of fact, most of them +possess well developed stings and some of them, especially in the +tropics, are very justly feared. Even those which lack the sting possess +well-developed poison glands and the parts of the entire stinging +apparatus, in so far as it is developed in the various species, may +readily be homologized with those of the honey-bee. + +The ants lacking a sting are those of the subfamily CAMPONOTINAE, which +includes the largest of our local species. It is an interesting fact +that some of these species possess the largest poison glands and +reservoir (fig. 28) and it is found that when they attack an enemy they +bring the tip of the abdomen forward and spray the poison in such a way +that it is introduced into the wound made by the powerful mandibles. + +[Illustration: 29. A harmless, but much feared larva, the "tomato worm." +Natural size. Photograph by M. V. S.] + +More feared than any of the other Hymenoptera are the hornets and wasps. +Of these there are many species, some of which attain a large size and +are truly formidable. Phisalix (1897), has made a study of the venom of +the common hornet and finds that, like the poison of the honey-bee, it +is neither an albuminoid nor an alkaloid. Its toxic properties are +destroyed at 120^o C. Phisalix also says that the venom is soluble in +alcohol. If this be true, it differs in this respect from that of the +bee. An interesting phase of the work of Phisalix is that several of her +experiments go to show that the venom of hornets acts as a vaccine +against that of vipers. + + +NETTLING INSECTS + +So far, we have considered insects which possess poison glands connected +with the mouth-parts or a special sting and which actively inject their +poison into man. There remain to be considered those insects which +possess poisonous hairs or body fluids which, under favorable +circumstances, may act as poisons. To the first of these belong +primarily the larvae of certain Lepidoptera. + + +LEPIDOPTERA + +[Illustration: 30. Another innocent but much maligned caterpillar, the +larva of the Regal moth. Photograph by M. V. S.] + +When we consider the reputedly poisonous larvae of moths and butterflies, +one of the first things to impress us is that we cannot judge by mere +appearance. Various species of Sphingid, or hawk-moth larvae, bear at the +end of the body a chitinous horn, which is often referred to as a +"sting" and regarded as capable of inflicting dangerous wounds. It would +seem unnecessary to refer to this absurd belief if it were not that each +summer the newspapers contain supposed accounts of injury from the +"tomato worm" (fig. 29) and others of this group. The grotesque, spiny +larva (fig. 30) of one of our largest moths, _Citheronia regalis_ is +much feared though perfectly harmless, and similar instances could be +multiplied. + +[Illustration: 31. The flannel moth (Lagoa crispata). (_a_) Poisonous +larva.] + +[Illustration: 31. (_b_) Adult. Enlarged. Photographs by M. V. S.] + +But if the larvae are often misjudged on account of their ferocious +appearance, the reverse may be true. A group of most innocent looking +and attractive caterpillars is that of the flannel-moth larvae, of which +_Lagoa crispata_ may be taken as an example. Its larva (fig. 31) has a +very short and thick body, which is fleshy and completely covered and +hidden by long silken hairs of a tawny or brown color, giving a convex +form to the upper side. Interspersed among these long hairs are +numerous short spines connected with underlying hypodermal poison +glands. These hairs are capable of producing a marked nettling effect +when they come in contact with the skin. This species is found in our +Atlantic and Southern States. Satisfactory studies of its poisonous +hairs and their glands have not yet been made. + +[Illustration: 32 The poisonous saddle back caterpillar. Empretia +(Sibine) stimulea. Photograph by M. V. S.] + +[Illustration: 33_a_. Io moth larvae on willow. Photograph by M. V. S.] + +_Sibine stimulea (Empretia stimulea)_, or the saddle-back caterpillar +(fig. 32), is another which possesses nettling hairs. This species +belongs to the group of Eucleidae, or slug caterpillars. It can be +readily recognized by its flattened form, lateral, bristling spines and +by the large green patch on the back resembling a saddle-cloth, while +the saddle is represented by an oval, purplish-brown spot. The small +spines are venomous and affect some persons very painfully. The larva +feeds on the leaves of a large variety of forest trees and also on +cherry, plum, and even corn leaves. It is to be found throughout the +Eastern and Southern United States. + +[Illustration: 33_b_. Io moth. Full grown larva. Photograph by M. V. S.] + +[Illustration: 34. Io moth. Adult. Photograph by M. V. S.] + +_Automeris io_ is the best known of the nettling caterpillars. It is the +larva of the Io moth, one of the Saturniidae. The mature caterpillar, +(fig. 33), which reaches a length of two and one-half inches, is of a +beautiful pale green with sublateral stripes of cream and red color and +a few black spines among the green ones. The green radiating spines give +the body a mossy appearance. They are tipped with a slender chitinous +hair whose tip is readily broken off in the skin and whose poisonous +content causes great irritation. Some individuals are very susceptible +to the poison, while others are able to handle the larvae freely without +any discomfort. The larvae feed on a wide range of food plants. They are +most commonly encountered on corn and on willow, because of the +opportunities for coming in contact with them. + +[Illustration: 35. Larva of brown-tail moth. (Natural size). Photograph +by M. V. S.] + +The larvae of the brown-tail moth (_Euproctis chrysorrhoea_) (fig. 35 +and 36), where they occur in this country, are, on account of their +great numbers, the most serious of all poisonous caterpillars. It is not +necessary here, to go into details regarding the introduction of this +species from Europe into the New England States. This is all available +in the literature from the United States Bureau of Entomology and from +that of the various states which are fighting the species. Suffice to +say, there is every prospect that the pest will continue to spread +throughout the Eastern United States and Canada and that wherever it +goes it will prove a direct pest to man as well as to his plants. + +Very soon after the introduction of the species there occurred in the +region where it had gained a foothold, a mysterious dermatitis of man. +The breaking out which usually occurred on the neck or other exposed +part of the body was always accompanied by an intense itching. It was +soon found that this dermatitis was caused by certain short, barbed +hairs of the brown-tail caterpillars and that not only the caterpillars +but their cocoons and even the adult female moths might harbor these +nettling hairs and thus give rise to the irritation. In many cases the +hairs were wafted to clothing on the line and when this was worn it +might cause the same trouble. Still worse, it was found that very +serious internal injury was often caused by breathing or swallowing the +poisonous hairs. + +[Illustration: 36. Browntail moths. One male and two females. Photograph +by M. V. S.] + +The earlier studies seemed to indicate that the irritation was purely +mechanical in origin, the result of the minute barbed hairs working into +the skin in large numbers. Subsequently, however, Dr. Tyzzer (1907) +demonstrated beyond question that the trouble was due to a poison +contained in the hairs. In the first place, it is only the peculiar +short barbed hairs which will produce the dermatitis when rubbed on the +skin, although most of the other hairs are sharply barbed. Moreover, it +was found that in various ways the nettling properties could be +destroyed without modifying the structure of the hairs. This was +accomplished by baking for one hour at 110^o C, by warming to 60^o C in +distilled water, or by soaking in one per cent. or in one-tenth per +cent. of potassium hydrate or sodium hydrate. The most significant part +of his work was the demonstration of the fact that if the nettling +hairs are mingled with blood, they immediately produce a change in the +red corpuscles. These at once become coarsely crenated, and the roleaux +are broken up in the vicinity of the hair (fig. 37_b_). The corpuscles +decrease in size, the coarse crenations are transformed into slender +spines which rapidly disappear, leaving the corpuscles in the form of +spheres, the light refraction of which contrasts them sharply with the +normal corpuscles. The reaction always begins at the basal sharp point +of the hair. It could not be produced by purely mechanical means, such +as the mingling of minute particles of glass wool, the barbed hairs of a +tussock moth, or the other coarser hairs of the brown-tail, with the +blood. + +[Illustration: 37. (_a_) Ordinary hairs and three poison hairs of +subdorsal and lateral tubercles of the larva of the browntail moth. +Drawing by Miss Kephart.] + +The question of the source of the poison has been studied in our +laboratory by Miss Cornelia Kephart. She first confirmed Dr. Tyzzer's +general results and then studied carefully fixed specimens of the larvae +to determine the distribution of the hairs and their relation to the +underlying tissues. + +[Illustration: 37. (_b_) Effect of the poison on the blood corpuscles of +man. After Tyzzer.] + +The poison hairs (fig. 37), are found on the subdorsal and lateral +tubercles (fig. 38), in bunches of from three to twelve on the minute +papillae with which the tubercles are thickly covered. The underlying +hypodermis is very greatly thickened, the cells being three or four +times the length of the ordinary hypodermal cells and being closely +crowded together. Instead of a pore canal through the cuticula for each +individual hair, there is a single pore for each papillae on a tubercle, +all the hairs of the papilla being connected with the underlying cells +through the same pore canal, (figs. 39 and 40). + +[Illustration: 38. Cross section of the larva of the browntail moth +showing the tubercles bearing the poison hairs. Drawing by Miss +Kephart.] + +The hypodermis of this region is of two distinct types of cells. First, +there is a group of slender fusiform cells, one for each poison hair on +the papilla, which are the trichogen, or hair-formative cells. They are +crowded to one side and towards the basement membrane by a series of +much larger, and more prominent cells (fig. 40), of which there is a +single one for each papilla. These larger cells have a granular +protoplasm with large nuclei and are obviously actively secreting. They +are so characteristic in appearance as to leave no question but that +they are the true poison glands. + +[Illustration: 39. Epithelium underlying poison hairs of the larva of +the browntail moth. Drawing by Miss Kephart.] + +Poisonous larvae of many other species have been reported from Europe and +especially from the tropics but the above-mentioned species are the more +important of those occurring in the United States and will serve as +types. It should be noted in this connection that through some curious +misunderstanding Goeldi (1913) has featured the larva of _Orgyia +leucostigma_, the white-marked tussock moth, as the most important of +the poisonous caterpillars of this country. Though there are occasional +reports of irritation from its hairs such cases are rare and there is no +evidence that there is any poison present. Indeed, subcutaneous +implantation of the hairs leads to no poisoning, but merely to temporary +irritation. + +[Illustration: 40. Same as figure 39, on larger scale.] + +Occasionally, the hairs of certain species of caterpillars find +lodgement in the conjunctiva, cornea, or iris of the eye of man and give +rise to the condition known as _opthalmia nodosa_. The essential feature +of this trouble is a nodular conjunctivitis which simulates tuberculosis +of the conjunctiva and hence has been called _pseudo-tubercular_. It may +be distinguished microscopically by the presence of the hairs. + +[Illustration: 41. (_a_) Nodular conjunctivitis in the eye of a child. +De Schweinitz and Shumway.] + +Numerous cases of opthalmia nodosa are on record. Of those from this +country, one of the most interesting is reported by de Schweinitz and +Shumway (1904). It is that of a child of fifteen years whose eye had +become inflamed owing to the presence of some foreign body. Downward and +inward on the bulbar conjunctiva were a number of flattened, +grayish-yellow nodules, between which was a marked congestion of the +conjunctival and episcleral vessels (fig. 41_a_). Twenty-seven nodules +could be differentiated, those directly in the center of the collection +being somewhat confluent and assuming a crescentic and circular +appearance. The nodules were excised and, on sectioning, were found to +be composed of a layer of spindle cells and round cells, outside of +which the tissue was condensed into a capsule. The interior consisted of +epithelioid cells, between which was a considerable intercellular +substance. Directly in the center of a certain number of nodules was +found the section of a hair (fig. 41_b_). The evidence indicated that +the injury had resulted from playing with caterpillars of one of the +Arctiid moths, _Spilosoma virginica_. Other reported cases have been +caused by the hairs of larvae of _Lasiocampa rubi_, _L. pini_, +_Porthetria dispar_, _Psilura monacha_ and _Cnethocampa processionea_. + +[Illustration: 41_b_. Section through one of the nodules showing the +caterpillar hair. De Schweinitz and Shumway.] + + +RELIEF FROM POISONING BY NETTLING LARVAE--The irritation from nettling +larvae is often severe and, especially in regions where the brown-tail +abounds, inquiries as to treatment arise. In general, it may be said +that cooling lotions afford relief, and that scratching, with the +possibilities of secondary infection, should be avoided, in so far as +possible. + +Among the remedies usually at hand, weak solutions of ammonia, or a +paste of ordinary baking soda are helpful. Castellani and Chalmers +recommend cleaning away the hairs by bathing the region with an alkaline +lotion, such as two per cent solution of bicarbonate of soda, and then +applying an ointment of ichthyol (10%). + +In the brown-tail district, there are many proprietary remedies of which +the best ones are essentially the following, as recommended by Kirkland +(1907): + + Carbolic acid 1/2 drachm. + Zinc oxide 1/2 oz. + Lime water 8 oz. + +Shake thoroughly and rub well into the affected parts. + +In some cases, and especially where there is danger of secondary +infection, the use of a weak solution of creoline (one teaspoonful to a +quart of water), is to be advised. + + +VESCICATING INSECTS AND THOSE POSSESSING OTHER POISONS IN THEIR BLOOD +PLASMA + +We have seen that certain forms, for example, the poisonous spiders, not +only secrete a toxine in their poison glands, but that such a substance +may be extracted from other parts of their body, or even their eggs. +There are many insects which likewise possess a poisonous blood plasma. +Such forms have been well designated by Taschenberg as _cryptotoxic_ +([Greek: kryptos] = hidden). We shall consider a few representative +forms. + +[Illustration: 42_a_. Blister beetle.] + +[Illustration: 42_b_. An American blister beetle. Meloe angusticollis. +Photograph by M. V. S.] + + +THE BLISTER BEETLES--Foremost among the cryptotoxic insects are the +_Meloidae_ or "blister beetles," to which the well-known "Spanish fly" +(fig. 42_a_), formerly very generally used in medical practice, belongs. +The vescicating property is due to the presence in the blood plasma of a +peculiar, volatile, crystalline substance known as _cantharidin_, which +is especially abundant in the reproductive organs of the beetle. +According to Kobert, the amount of this varies in different species from +.4 or .5% to 2.57% of the dry weight of the beetle. + +While blister beetles have been especially used for external +application, they are also at times used internally as a stimulant and a +diuretic. The powder or extract was formerly much in vogue as an +aphrodisiac, and formed the essential constituent of various philters, +or "love powders". It is now known that its effects on the reproductive +organs appear primarily after the kidneys have been affected to such an +extent as to endanger life, and that many cases of fatal poison have +been due to its ignorant use. + +There are many cases on record of poisoning and death due to internal +use, and in some instances from merely external application. There are +not rarely cases of poisoning of cattle from feeding on herbage bearing +a large number of the beetles and authentic cases are known of human +beings who have been poisoned by eating the flesh of such cattle. Kobert +states that the beetles are not poisonous to birds but that the flesh of +birds which have fed on them is poisonous to man, and that if the flesh +of chickens or frogs which have fed on the cantharidin be fed to cats it +causes in them the same symptoms as does the cantharidin. + +Treatment of cases of cantharidin poison is a matter for a skilled +physician. Until he can be obtained, emetics should be administered and +these should be followed by white of egg in water. Oils should be +avoided, as they hasten the absorption of the poison. + + +OTHER CRYPTOTOXIC INSECTS--Though the blister beetles are the best known +of the insects with poisonous blood plasma, various others have been +reported and we shall refer to a few of the best authenticated. + +One of the most famous is the Chrysomelid beetle, _Diamphidia simplex_, +the body fluids of whose larvae are used by certain South African bushmen +as an arrow poison. Its action is due to the presence of a toxalbumin +which exerts a haemolytic action on the blood, and produces inflammation +of the subcutaneous connective tissue and mucous membranes. Death +results from general paralysis. Krause (1907) has surmised that the +active principle may be a bacterial toxin arising from decomposition of +the tissues of the larva, but he presents no support of this view and it +is opposed by all the available evidence. + +In China, a bug, _Heuchis sanguinea_, belonging to the family Cicadidae, +is used like the Meloidae, to produce blistering, and often causes +poisoning. It has been assumed that its vescicating properties are due +to cantharidin, but the presence of this substance has not been +demonstrated. + +Certain Aphididae contain a strongly irritating substance which produces, +not merely on mucous membranes but on outer skin, a characteristic +inflammation. + +It has been frequently reported that the larvae of the European cabbage +butterfly, _Pieris brassicae_, accidentally eaten by cows, horses, ducks, +and other domestic animals, cause severe colic, attempts to vomit, +paralysis of the hind legs, salivation, and stomatitis. On _postmortem_ +there are to be found haemorrhagic gastro-enteritis, splenitis, and +nephritis. Kobert has recently investigated the subject and has found a +poisonous substance in the blood of not only the larvae but also the +pupae. + + +FOOTNOTES: + +[A] This is diametrically opposed to the findings of Bordas (1905) in +the case of the European _Latrodectus 13-guttatus_, whose glands are +"much larger than those of other spiders." From a considerable +comparative study, we should also unhesitatingly make this statement +regarding the glands of our American species, _L. mactans_. + +[B] Dr. E. H. Coleman (Kellogg, 1915) has demonstrated its virulence by +a series of experiments comparable with those of Kobert. + +[C] According to Stiles, the species occurring in the Northwest which is +commonly identified as _D. venustus_ should be called _D. andersoni_ +(see footnote, chapter 12). + +[D] It should be remembered that in all the higher Hymenoptera the first +abdominal segment is fused with the thorax and that what is apparently +the sixth segment is, in reality, the seventh. + + + + +CHAPTER III + +PARASITIC ARTHROPODA AFFECTING MAN + + +The relation of insects to man as simple parasites has long been +studied, and until very recent years the bulk of the literature of +medical entomology referred to this phase of the subject. This is now +completely overshadowed by the fact that so many of these parasitic +forms are more than simple parasites, they are transmitters of other +microscopic parasites which are pathogenic to man. Yet the importance of +insects as parasites still remains and must be considered in a +discussion of the relation of insects to the health of man. In taking up +the subject we shall first consider some general features of the +phenomenon of animal parasitism. + +Parasitism is an adaptation which has originated very often among living +organisms and in widely separated groups. It would seem simple to define +what is meant by a "parasite" but, in reality, the term is not easily +limited. It is often stated that a parasite is "An organism which lives +at the expense of another," but this definition is applicable to a +predatory species or, in its broadest sense, to all organisms. For our +purpose we may say with Braun: "A parasite is an organism which, for the +purpose of obtaining food, takes up its abode, temporarily or +permanently, on or within another living organism". + +Thus, parasitism is a phase of the broad biological phenomenon of +_symbiosis_, or living together of organisms. It is distinguished from +_mutualism_, or symbiosis in the narrow sense, by the fact that only one +party to the arrangement obtains any advantage, while the other is to a +greater or less extent injured. + +Of parasites we may distinguish on the basis of their location on or in +the host, _ecto-parasites_, which live outside of the body; and +_endo-parasites_, which live within the body. On account of their method +of breathing the parasitic arthropods belong almost exclusively to the +first of these groups. + +On the basis of relation to their host, we find _temporary parasites_, +those which seek the host only occasionally, to obtain food; and the +_stationary_ or _permanent_ parasites which, at least during certain +stages, do not leave their host. + +_Facultative parasites_ are forms which are not normally parasitic, but +which, when accidentally ingested, or otherwise brought into the body, +are able to exist for a greater or less period of time in their unusual +environment. These are generally called in the medical literature +"pseudoparasites" but the term is an unfortunate one. + +We shall now take up the different groups of arthropods, discussing the +more important of the parasitic forms attacking man. The systematic +relationship of these forms, and key for determining important species +will be found in Chapter XII. + + +ACARINA OR MITES + +The ACARINA, or _mites_, form a fairly natural group of arachnids, +characterized, in general, by a sac-like, unsegmented body which is +generally fused with the cephalothorax. The mouth-parts have been united +to form a beak or rostrum. + +The representatives of this group undergo a marked metamorphosis. +Commonly, the larvae on hatching from the egg, possess but three pairs of +legs, and hence are called _hexapod larvae_. After a molt, they transform +into nymphs which, like the adult, have four pairs of legs and are +called _octopod nymphs_. These after a period of growth, molt one or +more times and, acquiring external sexual organs, become adult. + +Most of the mites are free-living, but there are many parasitic species +and as these have originated in widely separated families, the Acarina +form an especially favorable group for study of the origin of +parasitism. Such a study has been made by Ewing (1911), who has reached +the following conclusions: + +"We have strong evidence indicating that the parasitic habit has +originated independently at least eleven times in the phylogeny of the +Ararina. Among the zoophagous parasites, the parasitic habit has been +developed from three different types of free-living Acarina: (a) +predaceous forms, (b) scavengers, (c) forms living upon the juices of +plants." + +Ewing also showed that among the living forms of Acarina we can trace +out all the stages of advancing parasitism, semiparasitism, facultative +parasitism, even to the fixed and permanent type, and finally to +endoparasitism. + +Of the many parasitic forms, there are several species which are serious +parasites of man and we shall consider the more important of these. +Infestation by mites is technically known as _acariasis_. + +[Illustration: 43. Effect of the harvest mites on the skin of man. +Photograph by J. C. Bradley.] + + +The Trombidiidae, or Harvest Mites + +In many parts of this country it is impossible for a visitor to go into +the fields and, particularly, into berry patches and among tall weeds +and grass in the summer or early fall without being affected by an +intolerable itching, which is followed, later, by a breaking out of +wheals, or papules, surrounded by a bright red or violaceous aureola, +(fig. 43). It is often regarded as a urticaria or eczema, produced by +change of climate, an error in diet, or some condition of general +health. + +Sooner or later, the victim finds that it is due to none of these, but +to the attacks of an almost microscopic red mite, usually called +"jigger" or "chigger" in this country. As the term "chigger" is applied +to one of the true fleas, _Dermatophilus penetrans_, of the tropics, +these forms are more correctly known as "harvest mites." Natives of an +infested region may be so immune or accustomed to its attacks as to be +unaware of its presence, though such immunity is by no means possessed +by all who have been long exposed to the annoyance. + +[Illustration: 44. Harvest mites. (Larvae of Trombidium). After C. V. +Riley.] + +The harvest mites, or chiggers, attacking man are larval forms, +possessing three pairs of legs (fig. 44). Their systematic position was +at first unknown and they were classed under a special genus _Leptus_, a +name which is very commonly still retained in the medical literature. It +is now known that they are the larval forms of various species of the +genus _Trombidium_, a group of predaceous forms, the adults of which +feed primarily on insects and their eggs. In this country the species +best known are those to be found late in summer, as larvae at the base of +the wings of houseflies or grasshoppers. + +There is much uncertainty as to the species of the larvae attacking man +but it is clear that several are implicated. Bruyant has shown that in +France the larvae _Trombidium inapinatum_ and _Trombidium holosericeum_ +are those most frequently found. The habit of attacking man is abnormal +and the larvae die after entering the skin. Normally they are parasitic +on various insects. + +Most recent writers agree that, on man, they do not bore into the skin, +as is generally supposed, but enter a hair follicle or sebaceous gland +and from the bottom of this, pierce the cutis with their elongate +hypopharynx. According to Braun, there arises about the inserted +hypopharynx a fibrous secretion--the so-called "beak" which is, in +reality, a product of the host. Dr. J. C. Bradley, however, has made +careful observations on their method of attack, and he assures us that +the mite ordinarily remains for a long time feeding on the surface of +the skin, where it produces the erythema above described. During this +time it is not buried in the skin but is able to retreat rapidly into it +through a hair follicle or sweat gland. The irritation from the mites +ceases after a few days, but not infrequently the intolerable itching +leads to so much scratching that secondary infection follows. + +Relief from the irritation may be afforded by taking a warm salt bath as +soon as possible after exposure or by killing the mites by application +of benzine, sulphur ointment or carbolized vaseline. When they are few +in number, they can be picked out with a sterile needle. + +Much may be done in the way of warding off their attacks by wearing +gaiters or close-woven stockings extending from ankle to the knee. Still +more efficacious is the sprinkling of flowers of sulphur in the +stockings and the underclothes from a little above the knee, down. The +writers have known this to make it possible for persons who were +especially susceptible to work with perfect comfort in badly infested +regions. Powdered naphthalene is successfully used in the same way and +as Chittenden (1906) points out, is a safeguard against various forms of +man-infesting tropical insect pests. + +The question of the destruction of the mites in the field is sometimes +an important one, and under some conditions, is feasible. Chittenden +states that much can be accomplished by keeping the grass, weeds, and +useless herbage mowed closely, so as to expose the mites to the sun. He +believes that in some cases good may be done by dusting the grass and +other plants, after cutting, with flowers of sulphur or by spraying with +dilute kerosene emulsion in which sulphur has been mixed. More recently +(1914) he calls attention to the value of cattle, and more especially +sheep, in destroying the pests by tramping on them and by keeping the +grass and herbage closely cropped. + + +IXODOIDEA OR TICKS + +Until recently, the ticks attracted comparatively little attention from +entomologists. Since their importance as carriers of disease has been +established, interest in the group has been enormously stimulated and +now they rank second only to the mosquitoes in the amount of detailed +study that has been devoted to them. + +[Illustration: 45_a_. Argus persicus. Capitulum of male. After Nuttall +and Warburton.] + +The ticks are the largest of the Acarina. They are characterized by the +fact that the hypostome, or "tongue" (fig. 45) is large and file-like, +roughened by sharp teeth. They possess a breathing pore on each side of +the body, above the third or fourth coxae (fig. 45_b_). + +[Illustration: 45_b_. Left spiracle of nymph of _Argas persicus_. After +Nuttall and Warburton.] + +There are two distinct families of the IXODOIDEA, differing greatly in +structure, life-history and habits. These are the ARGASIDAE and the +IXODIDAE. We shall follow Nuttall (1908) in characterizing these two +families and in pointing out their biological differences, and shall +discuss briefly the more important species which attack man. The +consideration of the ticks as carriers of disease will be reserved for a +later chapter. + + +Argasidae + +In the ticks belonging to the family ARGASIDAE, there is comparatively +little sexual dimorphism, while this is very marked in the Ixodidae. The +capitulum, or so-called "head" is ventral, instead of terminal; the +palpi are leg-like, with the segments subequal; the scutum, or dorsal +shield, is absent; eyes, when present, are lateral, on supracoxal folds. +The spiracles are very small; coxae unarmed; tarsi without ventral +spurs, and the pulvilli are absent or rudimentary. + +In habits and life history the Argasidae present striking +characteristics. In the first place, they are long-lived, a factor which +counts for much in the maintenance of the species. They are intermittent +feeders, being comparable with the bed-bug in this respect. There are +two or more nymphal stages, and they may molt after attaining maturity. +The female lays comparatively few eggs in several small batches. + +Nuttall (1911) concludes that "The Argasidae represent the relatively +primitive type of ticks because they are less constantly parasitic than +are the Ixodidae. Their nymphs and adults are rapid feeders and chiefly +infest the habitat of their hosts. * * * Owing to the Argasidae infesting +the habitats of their hosts, their resistance to prolonged starvation +and their rapid feeding habits, they do not need to bring forth a large +progeny, because there is less loss of life in the various stages, as +compared with the Ixodidae, prior to their attaining maturity." + +[Illustration: 46. Argus persicus. Dorsal and ventral aspects. (x4). +After Hassell.] + +Of the Argasidae, we have in the United States, several species which +have been reported as attacking man. + +_Argas persicus_, the famous "Miana bug" (fig. 46), is a very widely +distributed species, being reported from Europe, Asia, Africa, and +Australia. It is everywhere preeminently a parasite of fowls. According +to Nuttall it is specifically identical with _Argas americanus_ Packard +or _Argas miniatus_ Koch, which is commonly found on fowls in the United +States, in the South and Southwest. Its habits are comparable to those +of the bed-bug. It feeds intermittently, primarily at night, and instead +of remaining on its host, it then retreats to cracks and crevices. +Hunter and Hooker (1908) record that they have found the larva to remain +attached for five or eight days before dropping. Unlike the Ixodidae, the +adults oviposit frequently. + +[Illustration: 47. Otiobius (Ornithodoros) megnini, head of nymph. After +Stiles.] + +[Illustration: 48. Otiobius (Ornithodoros) megnini, male. (_a_) dorsal, +(_b_) ventral aspect. After Nuttall and Warburton.] + +The most remarkable feature of the biology of this species is the great +longevity, especially of the adult. Hunter and Hooker report keeping +larvae confined in summer in pill boxes immediately after hatching for +about two months while under similar conditions those of the Ixodid, +_Boophilus annulatus_ lived for but two or three days. Many writers have +recorded keeping adults for long periods without food. We have kept +specimens in a tin box for over a year and a half and at the end of that +time a number were still alive. Laboulliene kept unfed adults for over +three years. In view of the effectiveness of sulphur in warding off the +attacks of Trombidiidae, it is astonishing to find that Lounsbury has +kept adults of _Argas persicus_ for three months in a box nearly filled +with flowers of sulphur, with no apparent effect on them. + +We have already called attention to the occasional serious effects of +the bites of this species. While such reports have been frequently +discredited there can be no doubt that they have foundation in fact. The +readiness with which this tick attacks man, and the extent to which old +huts may be infested makes it especially troublesome. + +_Otiobius (Ornithodoros) megnini_, the "spinose ear-tick" (figs. 47, +48), first described from Mexico, as occurring in the ears of horses, is +a common species in our Southwestern States and is recorded by Banks as +occurring as far north as Iowa. + +The species is remarkable for the great difference between the spiny +nymph stage and the adult. The life history has been worked out by +Hooker (1908). Seed ticks, having gained entrance to the ear, attach +deeply down in the folds, engorge, and in about five days, molt; as +nymphs with their spinose body they appear entirely unlike the larvae. As +nymphs they continue feeding sometimes for months. Finally the nymph +leaves the host, molts to form the unspined adult, and without further +feeding is fertilized and commences oviposition. + +The common name is due to the fact that in the young stage the ticks +occur in the ear of their hosts, usually horses or cattle. Not +uncommonly it has been reported as occurring in the ear of man and +causing very severe pain. Stiles recommends that it be removed by +pouring some bland oil into the ear. + +Banks (1908) reports three species of _Ornithodoros_--_O. turicata_, +_coriaceus_ and _talaje_--as occurring in the United States. All of these +attack man and are capable of inflicting very painful bites. + + +Ixodidae + +The ticks belonging to the family IXODIDAE (figs. 49 and 50) exhibit a +marked sexual dimorphism. The capitulum is anterior, terminal, instead +of ventral as in the Argasidae; the palpi are relatively rigid (except in +the subfamily Ixodinae), with rudimentary fourth segment; scutum present; +eyes, when present, dorsal, on side of scutum. The spiracles are +generally large, situated well behind the fourth coxae; coxae generally +with spurs; pulvilli always present. + +In habits and life history the typical Ixodidae differ greatly from +the Argasidae. They are relatively short-lived, though some recent +work indicates that their longevity has been considerably +under-estimated. Typically, they are permanent feeders, remaining on the +host, or hosts, during the greater part of their life. They molt twice +only, on leaving the larval and the nymphal stages. The adult female +deposits a single, large batch of eggs. Contrasting the habits of the +Ixodidae to those of the Argasidae, Nuttall (1911) emphasizes that +the Ixodidae are more highly specialized parasites. "The majority are +parasitic on hosts having no fixed habitat and consequently all stages, +as a rule, occur upon the host." + +[Illustration: 49. Ixodes ricinus; male, ventral aspect. After Braun and +Luehe.] + +As mere parasites of man, apart from their power to transmit disease, +the Ixodidae are much less important than the Argasidae. Many are +reported as occasionally attacking man and of these the following native +species may be mentioned. + +[Illustration: 50. Ixodes ricinus, var. scapularis, female. Capitulum +and scutum; ventral aspect of capitulum; coxae; tarsus 4; spiracle; +genital and anal grooves. After Nuttall and Warburton.] + +_Ixodes ricinus_, the European castor bean tick (figs. 49, 50), is a +species which has been often reported from this country but Banks (1908) +has shown that, though it does occur, practically all of the records +apply to _Ixodes scapularis_ or _Ixodes cookei_. In Europe, _Ixodes +ricinus_ is very abundant and very commonly attacks man. At the point +of penetration of the hypostome there is more or less inflammation but +serious injury does not occur unless there have been introduced +pathogenic bacteria or, unless the tick has been abruptly removed, +leaving the capitulum in the wound. Under the latter circumstances, +there may be an abscess formed about the foreign body and occasionally, +serious results have followed. Under certain conditions the tick, in +various stages, may penetrate under the skin and produce a tumor, within +which it may survive for a considerable period of time. + +_Ixodes cookei_ is given by Banks as "common on mammals in the Eastern +States as far west as the Rockies." It is said to affect man severely. + +_Amblyomma americanum_, (fig. 158c), the "lone star tick," is widely +distributed in the United States. Its common name is derived from the +single silvery spot on the scutum of the female. Hunter and Hooker +regard this species as, next to _Boophilus annulatus_, the most +important tick in the United States. Though more common on cattle, it +appears to attack mammals generally, and "in portions of Louisiana and +Texas it becomes a pest of considerable importance to moss gatherers and +other persons who spend much time in the forests." + +_Amblyomma cajennense_, noted as a pest of man in central and tropical +America, is reported from various places in the south and southwestern +United States. + +_Dermacentor variabilis_ is a common dog tick of the eastern United +States. It frequently attacks man, but the direct effects of its bite +are negligible. + +The "Rocky Mountain spotted fever tick" (_Dermacentor andersoni_ +according to Stiles, _D. venustus_ according to Banks) is, from the +viewpoint of its effects on man, the most important of the ticks of the +United States. This is because, as has been clearly established, it +transmits the so-called "spotted fever" of man in our northwestern +states. This phase of the subject will be discussed later and it need +merely be mentioned here, that this species has been reported as causing +painful injuries by its bites. Dr. Stiles states that he has seen cases +of rather severe lymphangitis and various sores and swellings developing +from this cause. In one case, of an individual bitten near the elbow, +the arm became very much swollen and the patient was confined in bed for +several days. The so-called tick paralysis produced by this species is +discussed in a preceding chapter. + +There are many other records of various species of ticks attacking man, +but the above-mentioned will serve as typical and it is not necessary to +enter into greater detail. + + +TREATMENT OF TICK BITES--When a tick attaches to man the first thing to +be done is to remove it without leaving the hypostome in the wound to +fester and bring about secondary effects. This is best accomplished by +applying to the tick's body some substance which will cause it to more +readily loosen its hold. Gasoline or petroleum, oil or vaseline will +serve. For removing the spinose ear-tick, Stiles recommends pouring some +bland oil into the ear. Others have used effectively a pledget of cotton +soaked in chloroform. + +In general, the treatment recommended by Wellman for the bites of +_Ornithodoros moubata_ will prove helpful. It consists of prolonged +bathing in very hot water, followed by the application of a strong +solution of bicarbonate of soda, which is allowed to dry upon the skin. +He states that this treatment is comforting. For severe itching he +advises smearing the bites with vaseline, which is slightly impregnated +with camphor or menthol. Medical aid should be sought when complications +arise. + +The DERMANYSSIDAE are Gamasid mites which differ from others of the group +in that they are parasitic on vertebrates. None of the species normally +attack man, but certain of them, especially the poultry mite, may be +accidental annoyances. + +[Illustration: 51. Dermanyssus gallinae, female. After Delafond.] + +_Dermanyssus gallinae_ (fig. 51), the red mite of poultry, is an +exceedingly common and widespread parasite of fowls. During the day it +lives in cracks and crevices of poultry houses, under supports of +roosts, and in litter of the food and nests, coming out at night to +feed. They often attack people working in poultry houses or handling +and plucking infested fowls. They may cause an intense pruritis, but +they do not produce a true dermatosis, for they do not find conditions +favorable for multiplication on the skin of man. + + +Tarsonemidae + +The representatives of the family TARSONEMIDAE are minute mites, with the +body divided into cephalothorax and abdomen. There is marked sexual +dimorphism. The females possess stigmata at the anterior part of the +body, at the base of the rostrum, and differ from all other mites in +having on each side, a prominent clavate organ between the first and +second legs. The larva, when it exists, is hexapodous and resembles the +adult. A number of the species are true parasites on insects, while +others attack plants. Several of them may be accidental parasites of +man. + +[Illustration: 52. Pediculoides ventricosus, female. After Webster.] + +[Illustration: 53. Pediculoides ventricosus, gravid female. (x80). +After Webster.] + +_Pediculoides ventricosus_ (fig. 52 and 53) is, of all the +Tarsonemidae reported, the one which has proved most troublesome to +man. It is a predaceous species which attacks a large number of insects +but which has most commonly been met with by man through its fondness +for certain grain-infesting insects, notably the Angoumois grain moth, +_Sitotroga cerealella_, and the wheat straw-worm, _Isosoma grande_. In +recent years it has attracted much attention in the United States and +its distribution and habits have been the object of detailed study by +Webster (1901). + +[Illustration: 54. Pediculoides ventricosus, male. After Braun.] + +There is a very striking sexual dimorphism in this species. The +non-gravid female is elongate, about 200u by 70u (fig. 52), with the +abdomen slightly striated longitudinally. The gravid female (fig. 53) +has the abdomen enormously swollen, so that it is from twenty to a +hundred times greater than the rest of the body. The species is +viviparous and the larvae undergo their entire growth in the body of the +mother. They emerge as sexually mature males and females which soon +pair. The male (fig. 54) is much smaller, reaching a length of only 320u +but is relatively broad, 80u, and angular. Its abdomen is very greatly +reduced. + +As far back as 1850 it was noted as causing serious outbreaks of +peculiar dermatitis among men handling infested grain. For some time the +true source of the difficulty was unknown and it was even believed that +the grain had been poisoned. Webster has shown that in this country (and +probably in Europe as well) its attacks have been mistaken for those of +the red bugs or "chiggers" (larval Trombiidae). More recently a number of +outbreaks of a mysterious "skin disease" were traced to the use of straw +mattresses, which were found to be swarming with these almost +microscopic forms which had turned their attentions to the occupants of +the beds. Other cases cited were those of farmers running wheat through +a fanning mill, and of thrashers engaged in feeding unthrashed grain +into the cylinder of the machine. + +[Illustration: 55. Lesions produced by the attacks of Pediculoides +ventricosus. After Webster.] + +The medical aspects of the question have been studied especially by +Schamberg and Goldberger and from the latter's summary (1910) we derive +the following data. Within twelve to sixteen hours after exposure, +itching appears and in severe cases, especially where exposure is +continued night after night by sleeping on an infested bed, the itching +may become almost intolerable. Simultaneously, there appears an eruption +which characteristically consists of wheals surrounded by a vesicle +(fig. 55). The vesicle as a rule does not exceed a pin head in size but +may become as large as a pea. Its contents rapidly become turbid and in +a few hours it is converted into a pustule. The eruption is most +abundant on the trunk, slight on the face and extremities and almost +absent on the feet and hands. In severe cases there may be +constitutional disturbances marked, at the outset, by chilliness, +nausea, and vomiting, followed for a few days by a slight elevation of +temperature, with the appearance of albumin in the urine. In some cases +the eruption may simulate that of chicken-pox or small-pox. + +Treatment for the purpose of killing the mites is hardly necessary as +they attach feebly to the surface and are readily brushed off by +friction of the clothes. "Antipruritic treatment is always called for; +warm, mildly alkaline baths or some soothing ointment, such as zinc +oxide will be found to fulfil this indication." Of course, reinfestation +must be guarded against, by discarding, or thoroughly fumigating +infested mattresses, or by avoiding other sources. Goldberger suggests +that farm laborers who must work with infested wheat or straw might +protect themselves by anointing the body freely with some bland oil or +grease, followed by a change of clothes and bath as soon as their work +is done. We are not aware of any experiments to determine the effect of +flowers of sulphur, but their efficiency in the case of "red bugs" +suggests that they are worth a trial against _Pediculoides_. + +Various species of TYROGLYPHIDAE (fig. 150_f_) may abound on dried fruits +and other products and attacking persons handling them, may cause a +severe dermatitis, comparable to that described above for _Pediculoides +ventricosus_. Many instances of their occurrence as such temporary +ectoparasites are on record. Thus, workers who handle vanilla pods are +subject to a severe dermatitis, known as vanillism, which is due to the +attacks of _Tyroglyphus siro_, or a closely related species. The +so-called "grocer's itch" is similarly caused by mites infesting various +products. Castellani has shown that in Ceylon, workers employed in the +copra mills, where dried cocoanut is ground up for export, are much +annoyed by mites, which produce the so-called "copra itch." The skin of +the hands, arms and legs, and sometimes of the whole body, except the +face, is covered by fairly numerous, very pruriginous papules, often +covered by small, bloody crusts due to scratching. The condition is +readily mistaken for scabies. It is due to the attacks of _Tyroglyphus +longior castellanii_ which occur in enormous numbers in some samples of +the copra. + + +Sarcoptidae + +The SARCOPTIDAE are minute whitish mites, semi-globular in shape, with a +delicate transversely striated cuticula. They lack eyes and tracheae. The +mouth-parts are fused at the base to form a cone which is usually +designated as the head. The legs are short and stout, and composed of +five segments. The tarsi may or may not possess a claw and may terminate +in a pedunculated sucker, or simple long bristle, or both. The presence +or absence of these structures and their distribution are much used in +classification. The mites live on or under the skin of mammals and +birds, where they produce the disease known as scabies, mange, or itch. +Several species of the Sarcoptidae attack man but the most important of +these, and the one pre-eminent as the "itch mite" is _Sarcoptes +scabiei_. + +The female of _Sarcoptes scabiei_, of man, is oval and yellowish white; +the male more rounded and of a somewhat reddish tinge, and much smaller. +The body is marked by transverse striae which are partly interrupted on +the back. There are transverse rows of scales, or pointed spines, and +scattered bristles on the dorsum. + +[Illustration: 56_a_. Sarcoptes scabiei, male. (x100). After +Furstenberg.] + +The male (fig. 56) which is from 200-240u in length, and 150-200u in +breadth, possesses pedunculated suckers on each pair of legs except the +third, which bears, instead, a long bristle. The female (fig. 56) +300-450u in length and 250-350u in breadth, has the pedunculated suckers +on the first and second pairs of legs, only, the third and fourth +terminating in bristles. + +[Illustration: 56_b_. Sarcoptes scabiei, female. (x100.) After +Furstenberg.] + +The mite lives in irregular galleries from a few millimeters to several +centimeters in length, which it excavates in the epidermis (fig. 57). It +works especially where the skin is thin, such as between the fingers, in +the bend of the elbows and knees, and in the groin, but it is by no +means restricted to these localities. The female, alone, tunnels into +the skin; the males remain under the superficial epidermal scales, and +seldom are found, as they die soon after mating. + +As she burrows into the skin the female deposits her eggs, which measure +about 150 x 100u. Furstenberg says that each deposits an average of +twenty-two to twenty-four eggs, though Gudden reports a single burrow as +containing fifty-one. From these there develop after about seven days, +the hexapod larvae. These molt on the sixteenth day to form an octopod +nymph, which molts again the twenty-first day. At the end of the fourth +week the nymphs molt to form the sexually mature males and the so-called +pubescent females. These pair, the males die, and the females again cast +their skin, and become the oviparous females. Thus the life cycle is +completed in about twenty-eight days. + +The external temperature exercises a great influence on the development +of the mites and thus, during the winter, the areas of infestation not +only do not spread, but they become restricted. As soon as the +temperature rises, the mites increase and the infestation becomes much +more extensive. + +[Illustration: 57. Sarcoptes scabiei. Diagrammatic representation of the +course in the skin of man.] + +In considering the possible sources of infestation, and the chances of +reinfestation after treatment, the question of the ability of the mite +to live apart from its host is a very important one. Unfortunately there +are few reliable data on this subject. Gerlach found that, exposed in +the dry, warm air of a room they became very inactive within twenty-four +hours, that after two days they showed only slight movement, and that +after three or four days they could not be revived by moisture and +warming. The important fact was brought out that in moist air, in folded +soiled underwear, they survived as long as ten days. Bourguignon found +that under the most favorable conditions the mites of _Sarcoptes scabiei +equi_ would live for sixteen days. + +The disease designated the "itch" or "scabies," in man has been known +from time immemorial, but until within less than a hundred years it was +almost universally attributed to malnutrition, errors of diet, or "bad +blood." This was in spite of the fact that the mite was known to Mouffet +and that Bonomo had figured both the adult and the egg and had declared +the mite the sole cause of the disease. In 1834 the Corsican medical +student, Francis Renucci, demonstrated the mite before a clinic in Saint +Louis Hospital in Paris and soon thereafter there followed detailed +studies of the life history of the various itch mites of man and +animals. + +[Illustration: 58. Scabies on the hand. From portfolio of Dermochromes +by permission of Rebman & Co., of New York. Publishers.] + +The disease is a cosmopolitan one, being exceedingly abundant in some +localities. Its spread is much favored where large numbers of people are +crowded together under insanitary conditions and hence it increases +greatly during wars and is widely disseminated and abundant immediately +afterwards. Though more commonly to be met with among the lower classes, +it not infrequently appears among those of the most cleanly, careful +habits, and it is such cases that are most liable to wrong diagnosis by +the physician. + +Infection occurs solely through the passage, direct or indirect, of the +young fertilized females to the skin of a healthy individual. The adult, +oviparous females do not quit their galleries and hence do not serve to +spread the disease. The young females move about more or less at night +and thus the principal source of infestation is through sleeping in the +same bed with an infested person, or indirectly through bedclothes, or +even towels or clothing. Diurnal infestation through contact or clothing +is exceptional. Many cases are known of the disease being contracted +from animals suffering from scabies, or mange. + +When a person is exposed to infestation, the trouble manifests itself +after eight or ten days, though there usually elapses a period of twenty +to thirty days before there is a suspicion of anything serious. The +first symptom is an intense itching which increases when the patient is +in bed. When the point of irritation is examined the galleries may +usually be seen as characteristic sinuous lines, at first whitish in +color but soon becoming blackish because of the contained eggs and +excrement. The galleries, which may not be very distinct in some cases, +may measure as much as four centimeters in length. Little vesicles, of +the size of a pin head are produced by the secretions of the feeding +mite; they are firm, and projecting, and contain a limpid fluid. Figures +58 and 59 show the typical appearance of scabies on the hands, while +figure 60 shows a severe general infestation. The intolerable itching +induces scratching and through this various complications may arise. The +lesions are not normally found on the face and scalp, and are rare on +the back. + +[Illustration: 59. Scabies on the hand. After Duhring.] + +[Illustration: 60. Generalized infection of Scabies. After Morrow.] + +Formerly, scabies was considered a very serious disease, for its cause +and method of treatment were unknown, and potentially it may continue +indefinitely. Generation after generation of the mites may develop and +finally their number become so great that the general health of the +individual is seriously affected. Now that the true cause of the disease +is known, it is easily controlled. + +Treatment usually consists in softening the skin by friction with soap +and warm water, followed by a warm bath, and then applying some +substance to kill the mites. Stiles gives the following directions, +modified from Bourguignon's, as "a rather radical guide, to be modified +according to facilities and according to the delicacy of the skin or +condition of the patient": + +1. The patient, stripped naked, is energetically rubbed all over (except +the head) for twenty minutes, with green soap and warm water. 2. He is +then placed in a warm bath for thirty minutes, during which time the +rubbing is continued. 3. The parasiticide is next rubbed in for twenty +minutes and is allowed to remain on the body for four or five hours; in +the meantime the patient's clothes are sterilized, to kill the eggs or +mites attached to them. 4. A final bath is taken to remove the +parasiticide. + +The parasiticide usually relied on is the officinal sulphur ointment of +the United States pharmacopoeia. When infestation is severe it is +necessary to repeat treatment after three or four days in order to kill +mites which have hatched from the eggs. + +The above treatment is too severe for some individuals and may, of +itself, produce a troublesome dermatitis. We have seen cases where the +treatment was persisted in and aggravated the condition because it was +supposed to be due to the parasite. For delicate-skinned patients the +use of balsam of Peru is very satisfactory, and usually causes no +irritation whatever. Of course, sources of reinfection should be +carefully guarded against. + +_Sarcoptes scabiei crustosae_, which is a distinct variety, if not +species, of the human itch mite, is the cause of so-called Norwegian +itch. This disease is very contagious, and is much more resistant than +the ordinary scabies. Unlike the latter, it may occur on the face and +scalp. + +_Sarcoptes scabiei_ not only attacks man but also occurs on a large +number of mammals. Many species, based on choice of host, and minute +differences in size and secondary characters, have been established, but +most students of the subject relegate these to varietal rank. Many of +them readily attack man, but they have become sufficiently adapted to +their normal host so that they are usually less persistent on man. + +_Notoedres cati_ (usually known as _Sarcoptes minor_) is a species of +itch mites which produce an often fatal disease of cats. The body is +rounded and it is considerably smaller than _Sarcoptes scabiei_, the +female (fig. 61) measuring 215-230u long and 165-175u wide; the males +145-150u by 120-125u. The most important character separating +_Notoedres_ from _Sarcoptes_ is the position of the anus, which is +dorsal instead of terminal. The mite readily transfers to man but does +not persist, the infestation usually disappearing spontaneously in about +two weeks. Infested cats are very difficult to cure, unless treatment is +begun at the very inception of the outbreak, and under ordinary +circumstances it is better to kill them promptly, to avoid spread of the +disease to children and others who may be exposed. + +[Illustration: 61. Notoedres cati, male and female. After Railliet.] + + +Demodecidae + +The DEMODECIDAE are small, elongate, vermiform mites which live in the +hair follicles of mammals. The family characteristics will be brought +out in the discussion of the species infesting man, _Demodex +folliculorum_. + +[Illustration: 62. Demodex folliculorum. (x200) After Blanchard.] + +_Demodex folliculorum_ (fig. 62) is to be found very commonly in the +hair follicles and sebaceous glands of man. It is vermiform in +appearance, and with the elongate abdomen transversely striated so as to +give it the appearance of segmentation. The female is 380-400u long by +45u; the male 300u by 40u. The three-jointed legs, eight in number, are +reduced to mere stubs in the adult. The larval form is hexopod. These +mites thus show in their form a striking adaptation to their +environment. In the sebaceous glands and hair follicles they lie with +their heads down (fig. 63). Usually there are only a few in a gland, but +Gruby has counted as many as two hundred. + +[Illustration: 63. Demodex folliculorum. Section through skin showing +the mites in situ. Magnification of Nos. 1, 2, 6 and 7, x150; Nos, 3, +4, 5, x450. After Megnin.] + +The frequency with which they occur in man is surprising. According to +European statistics they are found in 50 per cent to 60 per cent or even +more. Gruby found them in forty out of sixty persons examined. These +figures are very commonly quoted, but reliable data for the United +States seem to be lacking. Our studies indicate that it is very much +less common in this country than is generally assumed. + +The Demodex in man does not, as a rule, cause the slightest +inconvenience to its host. It is often stated that they give rise to +comedons or "black-heads" but there is no clear evidence that they are +ever implicated. Certain it is that they are not the usual cause. A +variety of the same, or a very closely related species of _Demodex_, on +the dog gives rise to the very resistant and often fatal follicular +mange. + + +HEXAPODA OR TRUE INSECTS + +The HEXAPODA, or true insects, are characterized by the fact that the +adult possesses three pairs of legs. The body is distinctly segmented +and is divided into head, thorax, and abdomen. + +The mouth-parts in a generalized form, consist of an upper lip, or +_labrum_, which is a part of the head capsule, and a central unpaired +_hypopharynx_, two _mandibles_, two _maxillae_ and a lower lip, or +_labium_, made up of the fused pair of second maxillae. These parts may +be greatly modified, dependent upon whether they are used for biting, +sucking, piercing and sucking, or a combination of biting and sucking. + +Roughly speaking, insects may be grouped into those which undergo +_complete metamorphosis_ and those which have _incomplete +metamorphosis_. They are said to undergo complete metamorphosis when the +young form, as it leaves the egg, bears no resemblance to the adult. For +example, the maggot changes to a quiescent pupa and from this emerges +the winged active fly. They undergo incomplete metamorphosis, when the +young insect, as it leaves the egg, resembles the adult to a greater or +less extent, and after undergoing a certain number of molts becomes +sexually mature. + +Representatives of several orders have been reported as accidental or +faculative parasites of man, but the true parasites are restricted to +four orders. These are the Siphunculata; the Hemiptera, the Diptera and +the Siphonaptera. + + +SIPHUNCULATA + +The order SIPHUNCULATA was established by Meinert to include the true +sucking lice. These are small wingless insects, with reduced +mouth-parts, adapted for sucking; thorax apparently a single piece due +to indistinct separation of its three segments: the compound eyes +reduced to a single ommatidium on each side. The short, powerful legs +are terminated by a single long claw. Metamorphosis incomplete. + +There has been a great deal of discussion regarding the structure of the +mouth-parts, and the relationships of the sucking lice, and the +questions cannot yet be regarded as settled. The conflicting views are +well represented by Cholodkovsky (1904 and 1905) and by Enderlein +(1904). + +[Illustration: 64. Pediculus showing the blind sac (_b_) containing the +mouth parts (_a_) beneath the alimentary canal (_p_). After Pawlowsky.] + +Following Graber, it is generally stated that the mouth-parts consist of +a short tube furnished with hooks in front, which constitutes the lower +lip, and that within this is a delicate sucking tube derived from the +fusion of the labrum and the mandibles. Opposed to this, Cholodkovsky +and, more recently, Pawlowsky, (1906), have shown that the piercing +apparatus lies in a blind sac under the pharynx and opening into the +mouth cavity (fig. 64). It does not form a true tube but a furrow with +its open surface uppermost. Eysell has shown that, in addition, there is +a pair of chitinous rods which he regards as the homologues of the +maxillae. + +When the louse feeds, it everts the anterior part of the mouth cavity, +with its circle of hooks. The latter serve for anchoring the bug, and +the piercing apparatus is then pushed out. + +[Illustration: 65. Pediculus humanus, ventral aspect of male. (x10)] + +Most writers have classed the sucking lice as a sub-order of the +Hemiptera, but the more recent anatomical and developmental studies +render this grouping untenable. An important fact, bearing on the +question, is that, as shown by Gross, (1905), the structure of the +ovaries is radically different from that of the Hemiptera. + +Lice infestation and its effects are known medically as _pediculosis_. +Though their continued presence is the result of the grossest neglect +and filthiness, the original infestation may be innocently obtained and +by people of the most careful habits. + +Three species commonly attack man. Strangely enough, there are very few +accurate data regarding their life history. + +_Pediculus humanus_ (fig. 65), the head louse, is the most widely +distributed. It is usually referred to in medical literature as +_Pediculus capitis_, but the Linnean specific name has priority. In +color it is of a pale gray, blackish on the margins. It is claimed by +some authors that the color varies according to the color of the skin of +the host. The abdomen is composed of seven distinct segments, bearing +spiracles laterally. There is considerable variation in size. The males +average 1.8 mm. and the females 2.7 mm. in length. + +The eggs, fifty to sixty in number, stick firmly to the hairs of the +host and are known as nits. They are large and conspicuous, especially +on dark hair and are provided with an operculum, or cap, at the free +end, where the nymphs emerge. They hatch in about six days and about the +eighteenth day the young lice are sexually mature. + +[Illustration: 66. Pediculosis of the head. The illustration shows the +characteristic indications of the presence of lice, viz: the occipital +eczema gluing the hairs together, the swollen cervical glands, and the +porrigo, or eruption of contagious pustules upon the neck. After Fox.] + +The head lice live by preference on the scalp of their host but +occasionally they are found on the eyelashes and beard, or in the pubic +region. They may also occur elsewhere on the body. The penetration of +the rostrum into the skin and the discharge of an irritating saliva +produce a severe itching, accompanied by the formation of an eczema-like +eruption (fig. 66). When the infestation is severe, the discharge from +the pustules mats down the hair, and scabs are formed, under which the +insects swarm. "If allowed to run, a regular carapace may form, called +_trichoma_, and the head exudes a foetid odor. Various low plants may +grow in the trichoma, the whole being known as _plica +palonica_."--Stiles. + +Sources of infestation are various. School children may obtain the lice +from seatmates, by wearing the hats or caps of infested mates, or by the +use, in common, of brushes and combs. They may be obtained from infested +beds or sleeper berths. Stiles reports an instance in which a large +number of girls in a fashionable boarding school developed lousiness a +short time after traveling in a sleeping car. + +Treatment is simple, for the parasites may readily be controlled by +cleanliness and washing the head with a two per cent solution of +carbolic acid or even kerosene. The latter is better used mixed with +equal parts of olive oil, to avoid irritation. The treatment should be +applied at night and followed the next morning by a shampoo with soap +and warm water. It is necessary to repeat the operation in a few days. +Xylol, used pure, or with the addition of five per cent of vaseline, is +also very efficacious. Of course, the patient must be cautioned to stay +away from a lighted lamp or fire while using either the kerosene or +xylol. While these treatments will kill the eggs or nits, they will not +remove them from the hairs. Pusey recommends repeated washings with +vinegar or 25 per cent of acetic acid in water, for the purpose of +loosening and removing the nits. + +Treatment of severe infestations in females is often troublesome on +account of long hair. For such cases the following method recommended by +Whitfield (1912) is especially applicable: + +The patient is laid on her back on the bed with her head over the edge, +and beneath the head is placed a basin on a chair so that the hair lies +in the basin. A solution of 1 in 40 carbolic acid is then poured over +the hair into the basin and sluiced backwards and forwards until the +whole of the hair is thoroughly soaked with it. It is especially +necessary that care should be taken to secure thorough saturation of the +hair over the ears and at the nape of the neck, since these parts are +not only the sites of predilection of the parasites but they are apt to +escape the solution. This sluicing is carried out for ten minutes by the +clock. At the end of ten minutes the hair is lifted from the basin and +allowed to drain, but is not dried or even thoroughly wrung out. The +whole head is then swathed with a thick towel or better, a large piece +of common house flannel, which is fastened up to form a sort of turban, +and is allowed to remain thus for an hour. It can then be washed or +simply allowed to dry, as the carbolic quickly disperses. At the end of +this period every pediculus and what is better, every ovum is dead and +no relapse will occur unless there is exposure to fresh contagion. +Whitfield states that there seem to be no disadvantages in this method, +which he has used for years. He has never seen carboluria result from +it, but would advise first cutting the hair of children under five years +of age. + +_Pediculus corporis_ (= _P. vestimenti_) the body louse, is larger than +the preceding species, the female measuring 3.3 mm., and the male 3 mm. +in length. The color is a dirty white, or grayish. _P. corporis_ has +been regarded by some authorities as merely a variety of _P. humanus_ +but Piaget maintains there are good characters separating the two +species. + +The body louse lives in the folds and seams of the clothing of its host, +passing to the skin only when it wishes to feed. Brumpt states that he +has found enormous numbers of them in the collars of glass-ware or +grains worn by certain naked tribes in Africa. + +Exact data regarding the life-history of this species have been +supplied, in part, by the work of Warburton (1910), cited by Nuttall. He +found that _Pediculus corporis_ lives longer than _P. humanus_ under +adverse conditions. This is doubtless due to its living habitually on +the clothing, whereas _humanus_ lives upon the head, where it has more +frequent opportunities of feeding. He reared a single female upon his +own person, keeping the louse enclosed in a cotton-plugged tube with a +particle of cloth to which it could cling. The tube was kept next to his +body, thus simulating the natural conditions of warmth and moisture +under which the lice thrive. The specimen was fed twice daily, while it +clung to the cloth upon which it rested. Under these conditions she +lived for one month. Copulation commenced five days after the female had +hatched and was repeated a number of times, sexual union lasting for +hours. The female laid one hundred and twenty-four eggs within +twenty-five days. + +The eggs hatched after eight days, under favorable conditions, such as +those under which the female was kept. They did not hatch in the cold. +Eggs kept near the person during the day and hung in clothing by the +bedside at night, during the winter, in a cold room, did not hatch until +the thirty-fifth day. When the nymphs emerge from the eggs, they feed at +once, if given a chance to do so. They are prone to scatter about the +person and abandon the fragment of cloth to which the adult clings. + +The adult stage is reached on the eleventh day, after three molts, about +four days apart. Adults enter into copulation about the fifth day and as +the eggs require eight days for development, the total cycle, under +favorable conditions, is about twenty-four days. Warburton's data differ +considerably from those commonly quoted and serve to emphasize the +necessity for detailed studies of some of the commonest of parasitic +insects. + +Body lice are voracious feeders, producing by their bites and the +irritating saliva which they inject, rosy elevations and papules which +become covered with a brownish crust. The intense itching provokes +scratching, and characteristic white scars (fig. 67) surrounded by +brownish pigment (fig. 68) are formed. The skin may become thickened and +take on a bronze tinge. This melanoderma is especially marked in the +region between the shoulders but it may become generalized, a prominent +characteristic of "vagabond's disease." According to Dubre and Beille, +this melanoderma is due to a toxic substance secreted by the lice, which +indirectly provokes the formation of pigment. + +[Illustration: 67. Pediculosis in man caused by the body louse. After +Morrow.] + +Control measures, in the case of the body louse, consist in boiling or +steaming the clothes or in some cases, sterilizing by dry heat. The +dermatitis may be relieved by the use of zinc-oxide ointment, to which +Pusey recommends that there be added, on account of their parasiticidal +properties, sulphur and balsam of Peru, equal parts, 15 to 30 grains to +the ounce. + +_Phthirius pubis_ (= _P. inguinalis_), the pubic louse, or so-called +"crab louse," differs greatly from the preceding in appearance. It is +characterized by its relatively short head which fits into a broad +depression in the thorax. The latter is broad and flat and merges into +the abdomen. The first pair of legs is slender and terminated by a +straight claw. The second and third pairs of legs are thicker and are +provided with powerful claws fitted for clinging to hairs. The females +(fig. 69) measure 1.5 to 2 mm. in length by 1.5 mm. in breadth. The male +averages a little over half as large. The eggs, or nits, are fixed at +the base of the hairs. Only a few, ten to fifteen are deposited by a +single female, and they hatch in about a week's time. The young lice +mature in two weeks. + +The pubic louse usually infests the hairs of the pubis and the perineal +region. It may pass to the arm pits or even to the beard or moustache. +Rarely, it occurs on the eyelids, and it has even been found, in a very +few instances, occurring in all stages, on the scalp. Infestation may be +contracted from beds or even from badly infested persons in a crowd. We +have seen several cases which undoubtedly were due to the use of public +water closets. It produces papular eruption and an intense pruritis. +When abundant, there occurs a grayish discoloration of the skin which +Duguet has shown is due to a poisonous saliva injected by the louse, as +is the melanoderma caused by the body louse. + +[Illustration: 68. Melanoderma caused by the body louse. From Portfolio +of Dermochromes, by permission of Rebman & Co., New York, Publishers.] + +The pubic louse may be exterminated by the measures recommended for the +head louse, or by the use of officinal mercurial ointment. + +[Illustration: 69. Phthirius pubis. Ventral aspect of female. (x12).] + + +HEMIPTERA + +Several species of HEMIPTERA-HETEROPTERA are habitual parasites of man, +and others occur as occasional or accidental parasites. Of all these, +the most important and widespread are the bed-bugs, belonging to the +genus _Cimex_ (= _Acanthia_). + + +THE BED-BUGS--The bed-bugs are characterized by a much flattened oval +body, with the short, broad head unconstricted behind, and fitting into +the strongly excavated anterior margin of the thorax. The compound eyes +are prominent, simple eyes lacking. Antennae four-jointed, the first +segment short, the second long and thick, and the third and fourth +slender. The tarsi are short and three segmented. + +It is often assumed in the literature of the subject that there is but a +single species of _Cimex_ attacking man, but several such species are to +be recognized. These are distinguishable by the characters given in +Chapter XII. We shall consider especially _Cimex lectularius_, the most +common and widespread species. + +_Cimex lectularius_ (= _Acanthia lectularia_, _Clinocoris +lectularius_), is one of the most cosmopolitan of human parasites but, +like the lice, it has been comparatively little studied until recent +years, when the possibility that it may be concerned with the +transmission of various diseases has awakened interest in the details of +its life-history and habits. + +[Illustration: 70. Cimex lectularius adult and eggs. Photograph by M. V. +S.] + +The adult insect (fig. 70) is 4-5 mm. long by 3 mm. broad, reddish brown +in color, with the beak and body appendages lighter in color. The short, +broad and somewhat rectangular head has no neck-like constriction but +fits into the broadly semilunar prothorax. The four segmented labium or +proboscis encloses the lancet-like maxillae and mandibles. The distal +of the four antennal segments is slightly club-shaped. The prothorax is +characteristic of the species, being deeply incised anteriorly and with +its thin lateral margins somewhat turned up. The mesothorax is +triangular, with the apex posteriorly, and bears the greatly atrophied +first pair of wings. There is no trace of the metathoracic pair. The +greatly flattened abdomen has eight visible segments, though in reality +the first is greatly reduced and has been disregarded by most writers. +The body is densely covered with short bristles and hairs, the former +being peculiarly saber-shaped structures sharply toothed at the apex and +along the convex side (fig. 159_b_). + +The peculiar disagreeable odor of the adult bed-bug is due to the +secretion of the stink glands which lie on the inner surface of the +mesosternum and open by a pair of orifices in front of the metacoxae, +near the middle line. In the nymphs, the thoracic glands are not +developed but in the abdomen there are to be found three unpaired dorsal +stink glands, which persist until the fifth molt, when they become +atrophied and replaced by the thoracic glands. The nymphal glands occupy +the median dorsal portion of the abdomen, opening by paired pores at the +anterior margin of the fourth, fifth and sixth segments. The secretion +is a clear, oily, volatile fluid, strongly acid in reaction. Similar +glands are to be found in most of the Hemiptera-Heteroptera and their +secretion is doubtless protective, through being disagreeable to the +birds. In the bed-bug, as Marlatt points out, "it is probably an +illustration of a very common phenomenon among animals, i.e., the +persistence of a characteristic which is no longer of any special value +to the possessor." In fact, its possession is a distinct disadvantage to +the bed-bug, as the odor frequently reveals the presence of the bugs, +before they are seen. + +The eggs of the bed-bug (fig. 70) are pearly white, oval in outline, +about a millimeter long, and possess a small operculum or cap at one +end, which is pushed off when the young hatches. They are laid +intermittently, for a long period, in cracks and crevices of beds and +furniture, under seams of mattresses, under loose wall paper, and +similar places of concealment of the adult bugs. Girault (1905) observed +a well-fed female deposit one hundred and eleven eggs during the +sixty-one days that she was kept in captivity. She had apparently +deposited some of her eggs before being captured. + +The eggs hatch in six to ten days, the newly emerged nymphs being about +1.5 mm. in length and of a pale yellowish white color. They grow slowly, +molting five times. At the last molt the mesathoracic wing pads appear, +characteristic of the adult. The total length of the nymphal stage +varies greatly, depending upon conditions of food supply, temperature +and possibly other factors. Marlatt (1907) found under most favorable +conditions a period averaging eight days between molting which, added to +an equal egg period, gave a total of about seven weeks from egg to adult +insect. Girault (1912) found the postembryonic period as low as +twenty-nine days and as high as seventy days under apparently similar +and normal conditions of food supply. Under optimum and normal +conditions of food supply, beginning August 27, the average nymphal life +was 69.9 days; average number of meals 8.75 and the molts 5. Under +conditions allowing about half the normal food supply the average +nymphal life was from 116.9 to 139 days. Nymphs starved from birth lived +up to 42 days. We have kept unfed nymphs, of the first stage, alive in a +bottle for 75 days. The interesting fact was brought out that under +these conditions of minimum food supply there were sometimes six molts +instead of the normal number. + +The adults are remarkable for their longevity, a factor which is of +importance in considering the spread of the insect and methods of +control. Dufour (1833) (not De Geer, as often stated) kept specimens for +a year, in a closed vial, without food. This ability, coupled with their +willingness to feed upon mice, bats, and other small mammals, and even +upon birds, accounts for the long periods that deserted houses and camps +may remain infested. There is no evidence that under such conditions +they are able to subsist on the starch of the wall paper, juices of +moistened wood, or the moisture in the accumulations of dust, as is +often stated. + +There are three or four generations a year, as Girault's breeding +experiments have conclusively shown. He found that the bed-bug does not +hibernate where the conditions are such as to allow it to breed and that +breeding is continuous unless interrupted by the lack of food or, during +the winter, by low temperature. + +Bed-bugs ordinarily crawl from their hiding places and attack the face +and neck or uncovered parts of the legs and arms of their victims. If +undisturbed, they will feed to repletion. We have found that the young +nymph would glut itself in about six minutes, though some individuals +fed continuously for nine minutes, while the adult required ten to +fifteen minutes for a full meal. When gorged, it quickly retreats to a +crack or crevice to digest its meal, a process which requires two or +three days. The effect of the bite depends very greatly on the +susceptibility of the individual attacked. Some persons are so little +affected that they may be wholly ignorant of the presence of a large +number of bugs. Usually the bite produces a small hard swelling, or +wheal, whitish in color. It may even be accompanied by an edema and a +disagreeable inflammation, and in such susceptible individuals the +restlessness and loss of sleep due to the presence of the insects may be +a matter of considerable importance. Stiles (1907) records the case of a +young man who underwent treatment for neurasthenia, the diagnosis being +agreed upon by several prominent physicians; all symptoms promptly +disappeared, however, immediately following a thorough fumigation of +his rooms, where nearly a pint of bed-bugs were collected. + +It is natural to suppose that an insect which throughout its whole life +is in such intimate relationship with man should play an important role +in the transmission of disease. Yet comparatively little is definitely +known regarding the importance of the bed-bug in this respect. It has +been shown that it is capable of transmitting the bubonic plague, and +South American trypanosomiasis. Nuttall succeeded in transmitting +European relapsing fever from mouse to mouse by its bite. It has been +claimed that Oriental sore, tuberculosis, and even syphilis may be so +carried. These phases of the subject will be considered later. + +The sources of infestation are many, and the invasion of a house is not +necessarily due to neglect, though the continued presence of the pests +is quite another matter. In apartments and closely placed houses they +are known to invade new quarters by migration. They are frequently to be +met with in boat and sleeper berths, and even the plush seats of day +coaches, whence a nucleus may be carried in baggage to residences. They +may be brought in the laundry or in clothes of servants. + +Usually they are a great scourge in frontier settlements and it is +generally believed that they live in nature under the bark of trees, in +lumber, and under similar conditions. This belief is founded upon the +common occurrence of bugs resembling the bed-bug, in such places. As a +matter of fact, they are no relation to bed-bugs but belong to +plant-feeding forms alone (fig. 19 _c_, _d_). + +It is also often stated that bed-bugs live in poultry houses, in +swallows nests, and on bats, and that it is from these sources that they +gain access to dwellings. These bugs are specifically distinct from the +true bed-bug, but any of them may, rarely, invade houses. Moreover, +chicken houses are sometimes thoroughly infested with the true _Cimex +lectularius_. + +Control measures consist in the use of iron bedsteads and the reduction +of hiding places for the bugs. If the infestation is slight they may be +exterminated by a vigilant and systematic hunt, and by squirting +gasoline or alcohol into cracks and crevices of the beds, and furniture. +Fumigation must be resorted to in more general infestations. + +The simplest and safest method of fumigation is by the use of flowers of +sulphur at the rate of two pounds to each one thousand cubic feet of +room space. The sulphur should be placed in a pan, a well made in the +top of the pile and a little alcohol poured in, to facilitate burning. +The whole should be placed in a larger pan and surrounded by water so as +to avoid all danger of fire. Windows should be tightly closed, beds, +closets and drawers opened, and bedding spread out over chairs in order +to expose them fully to the fumes. As metal is tarnished by the sulphur +fumes, ornaments, clocks, instruments, and the like should be removed. +When all is ready the sulphur should be fired, the room tightly closed +and left for twelve to twenty-four hours. Still more efficient in large +houses, or where many hiding places favor the bugs, is fumigation with +hydrocyanic acid gas. This is a deadly poison and must be used under +rigid precautions. Through the courtesy of Professor Herrick, who has +had much experience with this method, we give in the Appendix, the clear +and detailed directions taken from his bulletin on "Household Insects." + +Fumigation with formaldehyde gas, either from the liquid or "solid" +formalin, so efficient in the case of contagious diseases, is useless +against bed-bugs and most other insects. + + +OTHER BED-BUGS--_Cimex hemipterus_ (= _C. rotundatus_) is a tropical and +subtropical species, occurring in both the old and new world. Patton and +Cragg state that it is distributed throughout India, Burma, Assam, the +Malay Peninsula, Aden, the Island of Mauritius, Reunion, St. Vincent and +Porto Rico. "It is widely distributed in Africa, and is probably the +common species associated there with man." Brumpt also records it for +Cuba, the Antilles, Brazil, and Venezuela. + +This species, which is sometimes called the Indian bed-bug, differs from +_C. lectularius_ in being darker and in having a more elongate abdomen. +The head also is shorter and narrower, and the prothorax has rounded +borders. + +It has the same habits and practically the same life cycle as _Cimex +lectularius_. Mackie, in India, has found that it is capable of +transmitting the Asiatic type of recurrent fever. Roger suggested that +it was also capable of transmitting Kala-azar and Patton has described +in detail the developmental stages of _Leishmania_, the causative +organism of Kala-azar, in the stomach of this bug, but Brumpt declares +that the forms described are those of a common, non-pathogenic +flagellate to be found in the bug, and have nothing to do with the +human disease. Brumpt has shown experimentally that _Cimex hemipterus_ +may transmit _Trypanosoma cruzi_ in its excrement. + +_Cimex boueti_, occurring in French Guinea, is another species attacking +man. Its habits and general life history are the same as for the above +species. It is 3 to 4.5 mm. in length, has vestigial elytra, and much +elongated antennae and legs. The extended hind legs are about as long as +the body. + +_Cimex columbarius_, a widely distributed species normally living in +poultry houses and dove cotes, _C. inodorus_, infesting poultry in +Mexico, _C. hirundinis_, occurring in the nests of swallows in Europe +and _Oeciacus vicarius_ (fig. 19i) occurring in swallows' nests in this +country, are species which occasionally infest houses and attack man. + +[Illustration: 71. Conorhinus sanguisugus.] + +_Conorhinus sanguisugus_, the cone-nosed bed-bug. We have seen in our +consideration of poisonous insects, that various species of Reduviid +bugs readily attack man. Certain of these are nocturnal and are so +commonly found in houses that they have gained the name, of "big +bed-bugs." The most noted of these, in the United States, is _Conorhinus +sanguisugus_ (fig. 71), which is widely distributed in our Southern +States. + +Like its near relatives, _Conorhinus sanguisugus_ is carnivorous in +habit and feeds upon insects as well as upon mammalian and human blood. +It is reported as often occurring in poultry houses and as attacking +horses in barns. The life history has been worked out in considerable +detail by Marlatt, (1902), from whose account we extract the following. + +[Illustration: 72. Beak of Conorhinus sanguisugus. After Marlatt.] + +The eggs are white, changing to yellow and pink before hatching. The +young hatch within twenty days and there are four nymphal stages. In all +these stages the insect is active and predaceous, the mouth-parts (fig. +72) being powerfully developed. The eggs are normally deposited, and the +early stages are undoubtedly passed, out of doors, the food of the +immature forms being other insects. Immature specimens are rarely found +indoors. It winters both in the partly grown and adult stage, often +under the bark of trees or in any similar protection, and only in its +nocturnal spring and early summer flights does it attack men. Marlatt +states that this insect seems to be decidedly on the increase in the +region which it particularly infests,--the plains region from Texas +northward and westward. In California a closely related species of +similar habits is known locally as the "monitor bug." + +The effect of the bite of the giant bed-bug on man is often very severe, +a poisonous saliva apparently being injected into the wound. We have +discussed this phase of the subject more fully under the head of +poisonous insects. + +_Conorhinus megistus_ is a Brazilian species very commonly attacking +man, and of special interest since Chagas has shown that it is the +carrier of a trypanosomiasis of man. Its habits and life history have +been studied in detail by Neiva, (1910). + +This species is now pre-eminently a household insect, depositing its +eggs in cracks and crevices in houses, though this is a relatively +recent adaptation. The nymphs emerge in from twenty to forty days, +depending upon the temperature. There are five nymphal stages, and as in +the case of true bed-bugs, the duration of these is very greatly +influenced by the availability of food and by temperature. Neiva reckons +the entire life cycle, from egg to egg, as requiring a minimum of three +hundred and twenty-four days. + +The nymphs begin to suck blood in three to five days after hatching. +They usually feed at night and in the dark, attacking especially the +face of sleeping individuals. The bite occasions but little pain. The +immature insects live in cracks and crevices in houses and invade the +beds which are in contact with walls, but the adults are active flyers +and attack people sleeping in hammocks. The males as well as the females +are blood suckers. + +Like many blood-sucking forms, _Conorhinus megistus_ can endure for long +periods without food. Neiva received a female specimen which had been +for fifty-seven days alive in a tightly closed box. They rarely feed on +two consecutive days, even on small quantities of blood, and were never +seen to feed on three consecutive days. + +Methods of control consist in screening against the adult bugs, and the +elimination of crevices and such hiding places of the nymphs. Where the +infestation is considerable, fumigation with sulphur is advisable. + + +PARASITIC DIPTERA OR FLIES + +Of the DIPTERA or two-winged flies, many species occasionally attack +man. Of these, a few are outstanding pests, many of them may also serve +to disseminate disease, a phase of our subject which will be considered +later. We shall now consider the most important of the group from the +viewpoint of their direct attacks on man. + + +Psychodidae or Moth-Flies + +The PSYCHODIDAE or Moth-flies, include a few species which attack man, +and at least one species, _Phlebotomus papatasii_, is known to transmit +the so-called "three-day fever" of man. Another species is supposed to +be the vector of Peruvian verruga. + +The family is made up of small, sometimes very small, nematocerous +Diptera, which are densely covered with hairs, giving them a moth-like +appearance. The wings are relatively large, oval or lanceolate in shape, +and when at rest are held in a sloping manner over the abdomen, or are +held horizontally in such a way as to give the insect a triangular +outline. Not only is the moth-like appearance characteristic, but the +venation of the wings (fig. 163, d) is very peculiar and, according to +Comstock, presents an extremely generalized form. All of the +longitudinal veins separate near the base of the wing except veins R_2 +and R_3 and veins M_1 and M_2. Cross veins are wanting in most cases. + +Comparatively little is known regarding the life-history and habits of +the Psychodidae, but one genus, _Phlebotomus_, contains minute, +blood-sucking species, commonly known as sand-flies. The family is +divided into two subfamilies, the PSYCHODINAE and the PHLEBOTOMINAE. +The second of these, the PHLEBOTOMINAE, is of interest to us. + + +THE PHLEBOTOMINAE--The Phlebotominae differ from the Psychodinae +in that the radical sector branches well out into the wing rather than +at the base of the wing. They are usually less hairy than the +Psychodinae. The ovipositor is hidden and less strongly chitinized. +The species attacking man belong to the genus _Phlebotomus_, small forms +with relatively large, hairy wings which are held upright, and with +elongate proboscis. The mandibles and maxillae are serrated and fitted +for biting. + +According to Miss Summers (1913) there are twenty-nine known species of +the genus _Phlebotomus_, five European, eleven Asiatic, seven African +and six American. One species only, _Phlebotomus vexator_, has been +reported for the United States. This was described by Coquillett, +(1907), from species taken on Plummer's Island, Maryland. It measures +only 1.5 mm. in length. As it is very probable that this species is much +more widely distributed, and that other species of these minute flies +will be found to occur in our fauna, we quote Coquillett's description. + +_Phlebotomus vexator_, Coq.: Yellow, the mesonotum brown, hairs chiefly +brown; legs in certain lights appear brown, but are covered with a white +tomentum; wings hyaline, unmarked; the first vein (R_1) terminates +opposite one-fifth of the length of the first submarginal cell (cell +R_2); this cell is slightly over twice as long as its petiole; terminal, +horny portion of male claspers slender, bearing many long hairs; the +apex terminated by two curved spines which are more than one-half as +long as the preceding part, and just in front of these are two similar +spines, while near the middle of the length of this portion is a fifth +spine similar to the others. Length 1.5 mm. + +The life-history of the Phlebotomus flies has been best worked out for +the European _Phlebotomus papatasii_ and we shall briefly summarize the +account of Doerr and Russ (1913) based primarily on work on this +species. The European Phlebotomus flies appear at the beginning of the +warm season, a few weeks after the cessation of the heavy rains and +storms of springtime. They gradually become more abundant until they +reach their first maximum, which in Italy is near the end of July +(Grassi). They then become scarcer but reach a second maximum in +September. At the beginning of winter they vanish completely, +hibernating individuals not being found. + +After fertilization there is a period of eight to ten days before +oviposition. The eggs are then deposited, the majority in a single mass +covered by a slimy secretion from the sebaceous glands. The larvae emerge +in fourteen to twenty days. There is uncertainty as to the length of +larval life, specimens kept in captivity remaining fifty or more days +without transforming. Growth may be much more rapid in nature. The larvae +do not live in fluid media but in moist detritus in dark places. Marett +believes that they live chiefly on the excrement of pill-bugs (Oniscidae) +and lizards. Pupation always occurs during the night. The remnants of +the larval skin remain attached to the last two segments of the +quiescent pupa and serve to attach it to the stone on which it lives. +The pupal stage lasts eleven to sixteen days, the adult escaping at +night. + +Only the females suck blood. They attack not only man but all +warm-blooded animals and, according to recent workers, also cold-blooded +forms, such as frogs, lizards, and larvae. Indeed, Townsend (1914) +believes that there is an intimate relation between _Phlebotomus_ and +lizards, or other reptiles the world over. The Phlebotomus passes the +daylight hours within the darkened recesses of the loose stone walls and +piles of rock in order to escape wind and strong light. Lizards inhabit +the same places, and the flies, always ready to suck blood in the +absence of light and wind, have been found more prone to suck reptilian +than mammalian blood. + +On hot summer nights, when the wind is not stirring, the Phlebotomus +flies, or sand-flies, as they are popularly called, invade houses and +sleeping rooms in swarms and attack the inmates. As soon as light begins +to break the flies either escape to the breeding places, or cool, dark +places protected from the wind, or a part of them remain in the rooms, +hiding behind pictures, under garments, and in similar places. Wherever +the Phlebotomus flies occur they are an intolerable nuisance. On account +of their small size they can easily pass through the meshes of ordinary +screens and mosquito curtains. They attack silently and inflict a very +painful, stinging bite, followed by itching. The ankles, dorsum of the +feet, wrists, inner elbow, knee joint and similar places are favorite +places of attack, possibly on account of their more delicate skin. + +Special interest has been attracted to these little pests in recent +years, since it has been shown that they transmit the European +"pappatici fever" or "three day fever." More recently yet, it appears +that they are the carriers of the virus of the Peruvian "verruga." This +phase of the subject will be discussed later. + +Control measures have not been worked out. As Newstead says, "In +consideration of the facts which have so far been brought to light +regarding the economy of Phlebotomus, it is clearly evident that the +task of suppressing these insects is an almost insurmountable one. Had +we to deal with insects as large and as accessible as mosquitoes, the +adoption of prophylactic measures would be comparatively easy, but owing +to the extremely minute size and almost flea-like habits of the adult +insects, and the enormous area over which the breeding-places may occur, +we are faced with a problem which is most difficult of solution." For +these reasons, Newstead considers that the only really prophylactic +measures which can at present be taken, are those which are considered +as precautionary against the bites of the insects. + +Of repellents, he cites as one of the best a salve composed of the +following: + + Ol. Anisi 3 grs. + Ol. Eucalypti 3 grs. + Ol. Terebenth 3 grs. + Unq. Acid Borac. + +Of sprays he recommends as the least objectionable and at the same time +one of the most effective, formalin. "The dark portions and angles of +sleeping apartments should be sprayed with a one per cent. solution of +this substance every day during the season in which the flies are +prevalent. A fine spraying apparatus is necessary for its application +and an excessive amount must not be applied. It is considered an +excellent plan also to spray the mosquito curtains regularly every day +towards sunset; nets thus treated are claimed to repel the attacks of +these insects." This effectiveness of formalin is very surprising for, +as we have seen, it is almost wholly ineffective against bed-bugs, +mosquitoes, house flies and other insects, where it has been tried. + +A measure which promises to be very effective, where it can be adopted, +is the use of electric fans so placed as to produce a current of air in +the direction of the windows of sleeping apartments. On account of the +inability of the Phlebotomus flies to withstand even slight breezes, it +seems very probable that they would be unable to enter a room so +protected. + + +Culicidae or Mosquitoes + +From the medical viewpoint, probably the most interesting and important +of the blood-sucking insects are the mosquitoes. Certainly this is true +of temperate zones, such as those of the United States. The result is +that no other group of insects has aroused such widespread interest, or +has been subjected to more detailed study than have the mosquitoes, +since their role as carriers of disease was made known. There is an +enormous literature dealing with the group, but fortunately for the +general student, this has been well summarized by a number of workers. +The most important and helpful of the general works are those of Howard +(1901), Smith (1904), Blanchard (1905), Mitchell (1907), and especially +of Howard, Dyar, and Knab, whose magnificent monograph is still in +course of publication. + +Aside from their importance as carriers of disease, mosquitoes are +notorious as pests of man, and the earlier literature on the group is +largely devoted to references to their enormous numbers and their +blood-thirstiness in certain regions. They are to be found in all parts +of the world, from the equator to the Arctic and Antarctic regions. +Linnaeus, in the "Flora Lapponica," according to Howard, Dyar and Knab, +"dwells at some length upon the great abundance of mosquitoes in Lapland +and the torments they inflicted upon man and beast. He states that he +believes that nowhere else on earth are they found in such abundance and +he compares their numbers to the dust of the earth. Even in the open, +you cannot draw your breath without having your mouth and nostrils +filled with them; and ointments of tar and cream or of fish grease are +scarcely sufficient to protect even the case-hardened cuticle of the +Laplander from their bite. Even in their cabins, the natives cannot take +a mouthful of food or lie down to sleep unless they are fumigated almost +to suffocation." In some parts of the Northwestern and Southwestern +United States it is necessary to protect horses working in the fields by +the use of sheets or burlaps, against the ferocious attacks of these +insects. It is a surprising fact that even in the dry deserts of the +western United States they sometimes occur in enormous numbers. + +Until comparatively recent years, but few species of mosquitoes were +known and most of the statements regarding their life-history were based +upon the classic work of Reaumur (1738) on the biology of the rain +barrel mosquito, _Culex pipiens_. In 1896, Dr. Howard refers to +twenty-one species in the United States, now over fifty are known; +Giles, in 1900, gives a total of two hundred and forty-two for the world +fauna, now over seven hundred species are known. We have found eighteen +species at Ithaca, N. Y. + +All of the known species of mosquitoes are aquatic in the larval stage, +but in their life-histories and habits such great differences occur that +we now know that it is not possible to select any one species as typical +of the group. For our present purpose we shall first discuss the general +characteristics and structure of mosquitoes, and shall then give the +life-history of a common species, following this by a brief +consideration of some of the more striking departures from what have +been supposed to be the typical condition. + +The CULICIDAE are slender, nematocerous Diptera with narrow wings, +antennae plumose in the males, and usually with the proboscis much longer +than the head, slender, firm and adapted for piercing in the female. +The most characteristic feature is that the margins of the wings and, in +most cases, the wing veins possess a fringe of scale-like hairs. These +may also cover in part, or entirely, the head, thorax, abdomen and legs. +The females, only, suck blood. + +On account of the importance of the group in this country and the +desirability of the student being able to determine material in various +stages, we show in the accompanying figures the characters most used in +classification. + +The larvae (fig. 73) are elongate, with the head and thorax sharply +distinct. The larval antennae are prominent, consisting of a single +cylindrical and sometimes curved segment. The outer third is often +narrower and bears at its base a fan-shaped tuft of hairs, the +arrangement and abundance of which is of systematic importance. About +the mouth are the so-called rotary mouth brushes, dense masses of long +hairs borne by the labrum and having the function of sweeping food into +the mouth. The form and arrangement of thoracic, abdominal, and anal +tufts of hair vary in different species and present characteristics of +value. On either side of the eighth abdominal segment is a patch of +scales varying greatly in arrangement and number and of much value in +separating species. Respiration is by means of tracheae which open at the +apex of the so-called anal siphon, when it is present. In addition, +there are also one or two pairs of tracheal gills which vary much in +appearance in different species. On the ventral side of the anal siphon +is a double row of flattened, toothed spines whose number and shape are +likewise of some value in separating species. They constitute the comb +or pecten. + +[Illustration: 73. Culex larva showing details of external structure.] + +The pupa (fig. 139, b) unlike that of most insects, is active, though it +takes no food. The head and thorax are not distinctly separated, but the +slender flexible abdomen in sharply marked off. The antennae, +mouth-parts, legs, and wings of the future adult are now external, but +enclosed in chitinous cases. On the upper surface, near the base of the +wings are two trumpets, or breathing tubes, for the pupal spiracles are +towards the anterior end instead of at the caudal end, as in the larva. +At the tip of the abdomen is a pair of large chitinous swimming paddles. + +As illustrative of the life cycle of a mosquito we shall discuss the +development of a common house mosquito, _Culex pipiens_, often referred +to in the Northern United States as the rain barrel mosquito. Its life +cycle is often given as typical for the entire group, but, as we have +already emphasized, no one species can serve this purpose. + +The adults of _Culex pipiens_ hibernate throughout the winter in +cellars, buildings, hollow trees, or similar dark shelters. Early in the +spring they emerge and deposit their eggs in a raft-like mass. The +number of eggs in a single mass is in the neighborhood of two hundred, +recorded counts varying considerably. A single female may deposit +several masses during her life time. The duration of the egg stage is +dependent upon temperature. In the warm summer time the larvae may emerge +within a day. The larvae undergo four molts and under optimum conditions +may transform into pupae in about a week's time. Under the same favorable +conditions, the pupal stage may be completed in a day's time. The total +life cycle of _Culex pipiens_, under optimum conditions, may thus be +completed in a week to ten days. This period may be considerably +extended under less favorable conditions of temperature and food supply. + +_Culex pipiens_ breeds continuously throughout the summer, developing in +rain barrels, horse troughs, tin cans, or indeed in any standing water +about houses, which lasts for a week or more. The catch basins of sewers +furnish an abundant supply of the pests under some conditions. Such +places, the tin gutters on residences, and all possible breeding places +must be considered in attempts to exterminate this species. + +Other species of mosquitoes may exhibit radical departures from _Culex +pipiens_ in life-history and habits. To control them it is essential +that the biological details be thoroughly worked out for, as Howard, +Dyar, and Knab have emphasized, "much useless labor and expense can be +avoided by an accurate knowledge of the habits of the species." For a +critical discussion of the known facts the reader is referred to their +monograph. We shall confine ourselves to a few illustrations. + +The majority of mosquitoes in temperate climates hibernate in the egg +stage, hatching in the spring or even mild winter days in water from +melting snow. It is such single-brooded species which appear in +astounding numbers in the far North. Similarly, in dry regions the eggs +may stand thorough dessication, and yet hatch out with great promptness +when submerged by the rains. "Another provision to insure the species +against destruction in such a case, exists in the fact * * * that not +all the eggs hatch, a part of them lying over until again submerged by +subsequent rains." In temperate North America, a few species pass the +winter in the larval state. An interesting illustration of this is +afforded by _Wyeomia smithii_, whose larvae live in pitcher plants and +are to be found on the coldest winter days imbedded in the solid ice. +Late in the spring, the adults emerge and produce several broods during +the summer. + +In the United States, one of the most important facts which has been +brought out by the intensive studies of recent years is that certain +species are migratory and that they can travel long distances and become +an intolerable pest many miles from their breeding places. This was +forcibly emphasized in Dr. Smith's work in New Jersey, when he found +that migratory mosquitoes, developing in the salt marshes along the +coast, are the dominant species largely responsible for the fame of the +New Jersey mosquito. The species concerned are _Aedes sollicitans_, _A. +cantator_ and _A. taeniorhynchus_. Dr. Smith decided that the first of +these might migrate at least forty miles inland. It is obvious that +where such species are the dominant pest, local control measures are a +useless waste of time and money. Such migratory habits are rare, +however, and it is probable that the majority of mosquitoes do not fly +any great distance from their breeding places. + +While mosquitoes are thought of primarily as a pest of man, there are +many species which have never been known to feed upon human or mammalian +blood, no matter how favorable the opportunity. According to Howard, +Dyar, and Knab, this is true of _Culex territans_, one of the common +mosquitoes in the summer months in the Northern United States. There are +some species, probably many, in which the females, like the males, are +plant feeders. In experimental work, both sexes are often kept alive for +long periods by feeding them upon ripe banana, dried fig, raisins, and +the like, and in spite of sweeping assertions that mosquitoes must have +a meal of blood in order to stimulate the ovaries to development, some +of the common blood-sucking species, notably _Culex pipiens_, have been +bred repeatedly without opportunity to feed upon blood. + +The effect of the bite varies greatly with different species and depends +upon the susceptibility of the individual bitten. Some persons are +driven almost frantic by the attacks of the pests when their companions +seem almost unconscious of any inconvenience. Usually, irritation and +some degree of inflammation appear shortly following the bite. Not +infrequently a hardened wheal or even a nodule forms, and sometimes +scratching leads to secondary infection and serious results. + +The source of the poison is usually supposed to be the salivary glands +of the insect. As we have already pointed out, (p. 34), Macloskie +believed that one lobe of the gland, on each side, was specialized for +forming the poison, while a radically different view is that of +Schaudinn, who believed that the irritation is due to the expelled +contents of the oesophageal diverticula, which contain a gas and a +peculiar type of fungi or bacteria. In numerous attempts, Schaudinn was +unable to produce any irritation by applying the triturated salivary +glands to a wound, but obtained the typical result when he used the +isolated diverticula. + +The irritation of the bite may be relieved to some extent by using +ammonia water, a one per cent. alcoholic solution of menthol, or +preparations of cresol, or carbolic acid. Dr. Howard recommends rubbing +the bite gently with a piece of moist toilet soap. Castellani and +Chalmers recommend cleansing inflamed bites with one in forty carbolic +lotion, followed by dressing with boracic ointment. Of course, +scratching should be avoided as much as possible. + +Repellents of various kinds are used, for warding off the attacks of the +insects. We have often used a mixture of equal parts of oil of +pennyroyal and kerosene, applied to the hands and face. Oil of +citronella is much used and is less objectionable to some persons. A +recommended formula is, oil of citronella one ounce, spirits of camphor +one ounce, oil of cedar one-half ounce. A last resort would seem to be +the following mixture recommended by Howard, Dyar, and Knab for use by +hunters and fishermen in badly infested regions, against mosquitoes and +blackflies. + +Take 2-1/4 lbs. of mutton tallow and strain it. While still hot add 1/2 +lb. black tar (Canadian tar). Stir thoroughly and pour into the +receptacle in which it is to be contained. When nearly cool stir in +three ounces of oil of citronella and 1-1/4 oz. of pennyroyal. + +At night the surest protection is a good bed net. There are many types +of these in use, but in order to be serviceable and at the same time +comfortable it should be roomy and hung in such a way as to be stretched +tightly in every direction. We prefer one suspended from a broad, square +frame, supported by a right-angled standard which is fastened to the +head of the bed. It must be absolutely free from rents or holes and +tucked in securely under the mattress or it will serve merely as a +convenient cage to retain mosquitoes which gain an entrance. While such +nets are a convenience in any mosquito ridden community, they are +essential in regions where disease-carrying species abound. Screening of +doors, windows and porches, against the pests is so commonly practiced +in this country that its importance and convenience need hardly be +urged. + +Destruction of mosquitoes and prevention of breeding are of fundamental +importance. Such measures demand first, as we have seen, the correct +determination of the species which is to be dealt with, and a knowledge +of its life-history and habits. If it prove to be one of the migratory +forms, it is beyond mere local effort and becomes a problem demanding +careful organization and state control. An excellent illustration of the +importance and effectiveness of work along these lines is afforded by +that in New Jersey, begun by the late Dr. John B. Smith and being pushed +with vigor by his successor, Dr. Headlee. + +In any case, there is necessity for community action. Even near the +coast, where the migratory species are dominant, there are the local +species which demand attention and which cannot be reached by any +measures directed against the species of the salt marshes. The most +important of local measures consist in the destruction of breeding +places by filling or draining ponds and pools, clearing up of more +temporary breeding places, such as cans, pails, water barrels and the +like. Under conditions where complete drainage of swamps is +impracticable or undesirable, judicious dredging may result in a pool or +series of steep-sided pools deep enough to maintain a supply of fish, +which will keep down the mosquito larvae. Where water receptacles are +needed for storage of rain water, they should be protected by careful +screening or a film of kerosene over the top of the water, renewed every +two weeks or so, so as to prevent mosquitoes from depositing their eggs. +When kerosene is used, Water drawn from the bottom of the receptacle +will not be contaminated by it to any injurious extent. Where ponds +cannot be drained much good will be accomplished by spraying kerosene +oil on the surface of the water, or by the introduction of fish which +will feed on the larvae. + +Detailed consideration of the most efficient measures for controlling +mosquitoes is to be found in Dr. Howard's Bulletin No. 88 of the Bureau +of Entomology, "Preventive and remedial work against mosquitoes" or, in +more summarized form, in Farmers' Bulletin No. 444. One of these should +be obtained by any person interested in the problems of mosquito control +and public health. + +[Illustration: 74. Mouth parts of Simulium. After Grunberg.] + + +The Simuliidae, or Black Flies + +The SIMULIIDAE, or black flies, are small, dark, or black flies, with a +stout body and a hump-back appearance. The antennae are short but +eleven-segmented, the wings broad, without scales or hairs, and with the +anterior veins stout but the others very weak. The mouth-parts (fig. 74) +are fitted for biting. + +The larvae of the Simuliidae (fig. 75) are aquatic and, unlike those of +mosquitoes, require a well aerated, or swiftly running water. Here they +attach to stones, logs, or vegetation and feed upon various +micro-organisms. They pupate in silken cocoons open at the top. Detailed +life-histories have not been worked out for most of the species. We +shall consider as typical that of _Simulium pictipes_, an inoffensive +species widely distributed in the Eastern United States, which has been +studied especially at Ithaca, N.Y. (Johannsen, 1903). + +[Illustration: 75. Larva of Simulium, (x8). After Garman.] + +The eggs are deposited in a compact yellowish layer on the surface of +rock, on the brinks of falls and rapids where the water is flowing +swiftly. They are elongate ellipsoidal in shape, about .4 by .18 mm. As +myriads of females deposit in the same place the egg patches may be +conspicuous coatings of a foot or much more in diameter. When first laid +they are enveloped in a yellowish white slime, which becomes darker, +until finally it becomes black just before the emerging of the larvae. +The egg stage lasts a week. + +The larvae (fig. 75) are black, soft skinned, somewhat cylindrical in +shape, enlarged at both ends and attenuated in the middle. The posterior +half is much stouter than the anterior part and almost club-shaped. The +head bears two large fan-shaped organs which aid in procuring food. +Respiration is accomplished by means of three so-called blood gills +which are pushed out from the dorsal part of the rectum. The larvae occur +in enormous numbers, in moss-like patches. If removed from their natural +habitat and placed in quiet water they die within three or four hours. +Fastened to the rock by means of a disk-like sucker at the caudal end of +the body, they ordinarily assume an erect position. They move about on +the surface of the rocks, to a limited extent, with a looping gait +similar to that of a measuring worm, and a web is secreted which +prevents their being washed away by the swiftly flowing water. They feed +chiefly upon algae and diatoms. + +The complete larval stage during the summer months occupies about four +weeks, varying somewhat with the temperature and velocity of the water. +At the end of this period they spin from cephalic glands, boot-shaped +silken cocoons within which they pupate. The cocoon when spun is firmly +attached to the rock and also to adjacent cocoons. Clustered +continuously over a large area and sometimes one above another, they +form a compact, carpet-like covering on the rocks, the reddish-brown +color of which is easily distinguishable from the jet-black appearance +of the larvae. The pupal stage lasts about three weeks. The adult fly, +surrounded by a bubble of air, quickly rises to the surface of the water +and escapes. The adults (fig. 76) are apparently short lived and thus +the entire life cycle, from egg to egg is completed in approximately +eight weeks. + +[Illustration: 76. Simulium venustum, (x8). After Garman.] + +In the case of _Simulium pictipes_ at Ithaca, N. Y., the first brood of +adults emerges early in May and successive generations are produced +throughout the summer and early autumn. This species winters in the +larval condition. Most of the other species of _Simulium_ which have +been studied seem to be single brooded. + +While _Simulium pictipes_ does not attack man, there are a number of the +species which are blood-sucking and in some regions they are a veritable +scourge. In recent years the greatest interest in the group has been +aroused by Sambon's hypothesis that they transmit pellagra from man to +man. This has not been established, and, indeed, seems very doubtful, +but the importance of these insects as pests and the possibility that +they may carry disease make it urgent that detailed life-histories of +the hominoxious species be worked out. + +As pests a vivid account of their attacks is in Agassiz's "Lake +Superior" (p. 61), quoted by Forbes (1912). + +"Neither the love of the picturesque, however, nor the interests of +science, could tempt us into the woods, so terrible were the black +flies. This pest of flies which all the way hither had confined our +ramblings on shore pretty closely to the rocks and the beach, and had +been growing constantly worse, here reached its climax. Although +detained nearly two days, * * * we could only sit with folded hands, or +employ ourselves in arranging specimens, and such other operations as +could be pursued in camp, and under the protection of a 'smudge.' One, +whom scientific ardor tempted a little way up the river in a canoe, +after water plants, came back a frightful spectacle, with blood-red +rings round his eyes, his face bloody, and covered with punctures. The +next morning his head and neck were swollen as if from an attack of +erysipelas." + +There are even well authenticated accounts on record of death of humans +from the attacks of large swarms of these gnats. In some regions, and +especially in the Mississippi Valley in this country, certain species of +black flies have been the cause of enormous losses to farmers and +stockmen, through their attacks on poultry and domestic animals. C. V. +Riley states that in 1874 the loss occasioned in one county in Tennessee +was estimated at $500,000. + +The measures of prevention and protection against these insects have +been well summarized by Forbes (1912). They are of two kinds: "the use +of repellents intended to drive away the winged flies, and measures for +the local destruction of the aquatic larvae. The repellents used are +either smudges, or surface applications made to keep the flies from +biting. The black-fly will not endure a dense smoke, and the well-known +mosquito smudge seems to be ordinarily sufficient for the protection of +man. In the South, leather, cloth, and other materials which will make +the densest and most stifling smoke, are often preserved for this use in +the spring. Smudges are built in pastures for the protection of stock, +and are kept burning before the doors of barns and stables. As the +black-flies do not readily enter a dark room, light is excluded from +stables as much as possible during the gnat season. If teams must be +used in the open field while gnats are abroad, they may be protected +against the attacks of the gnats by applying cotton-seed oil or axle +grease to the surface, especially to the less hairy parts of the +animals, at least twice a day. A mixture of oil and tar and, indeed, +several other preventives, are of practical use in badly infested +regions; but no definite test or exact comparison has been made with any +them in a way to give a record of the precise results." + +"It is easy to drive the flies from houses or tents by burning pyrethrum +powder inside; this either kills the flies or stupifies them so that +they do not bite for some time thereafter." * * * "Oil of tar is +commonly applied to the exposed parts of the body for the purpose of +repelling the insects, and this preparation is supplied by the Hudson +Bay Company to its employees. Minnesota fishermen frequently grease +their faces and hands with a mixture of kerosene and mutton tallow for +the same purpose." We have found a mixture of equal parts of kerosene +and oil of pennyroyal efficient. + +Under most circumstances very little can be done to destroy this insect +in its early stage, but occasionally conditions are such that a +larvicide can be used effectively. Weed (1904), and Sanderson (1910) +both report excellent results from the use of phinotas oil, a +proprietary compound. The first-mentioned also found that in some places +the larvae could be removed by sweeping them loose in masses with stiff +stable brooms and then catching them downstream on wire netting +stretched in the water. + + +Chironomidae or Midges + +The flies of this family, commonly known as midges, resemble mosquitoes +in form and size but are usually more delicate, and the wing-veins, +though sometimes hairy, are not fringed with scales. The venation is +simpler than in the mosquitoes and the veins are usually less distinct. + +These midges, especially in spring or autumn, are often seen in immense +swarms arising like smoke over swamps and producing a humming noise +which can be heard for a considerable distance. At these seasons they +are frequently to be found upon the windows of dwellings, where they are +often mistaken for mosquitoes. + +The larvae are worm-like, but vary somewhat in form in the different +genera. Most of them are aquatic, but a few live in the earth, in +manure, decaying wood, under bark, or in the sap of trees, especially in +the sap which collects in wounds. + +[Illustration: 77. Culicoides guttipennis; (_a_) adult, (x15); (_b_) +head of same; (_c_) larva; (_d_) head; (_e_) pupa. After Pratt.] + +Of the many species of CHIRONOMIDAE, (over eight hundred known), the vast +majority are inoffensive. The sub-family Ceratopogoninae, however, forms +an exception, for some of the members of this group, known as sandflies, +or punkies, suck blood and are particularly troublesome in the +mountains, along streams, and at the seashore. Most of these have been +classed under the genus _Ceratopogon_, but the group has been broken up +into a number of genera and _Ceratopogon_, in the strict sense, is not +known to contain any species which sucks the blood of vertebrates. + +THE CERATOPOGONINAE--The Ceratopogoninae are among the smallest of the +Diptera, many of them being hardly a millimeter long and some not even +so large. They are Chironomidae in which the thorax is not prolonged over +the head. The antennae are filiform with fourteen (rarely thirteen) +segments in both sexes, those of the male being brush-like. The basal +segment is enlarged, the last segment never longer than the two +preceding combined, while the last five are sub-equal to, or longer than +the preceding segment. The legs are relatively stouter than in the other +Chironomidae. The following three genera of this subfamily are best known +as blood suckers in this country. + +Of the genus _Culicoides_ there are many species occurring in various +parts of the world. A number are known to bite man and animals and it is +probable that all are capable of inflicting injury. In some localities +they are called punkies, in others, sand-flies, a name sometimes also +applied to the species of _Simulium_ and _Phlebotomus_. Owing to their +very small size they are known by some tribes of Indians as No-see-ums. +The larvae are found in ponds, pools, water standing in hollow tree +stumps, and the like. Though probably living chiefly in fresh water, we +have found a species occurring in salt water. The larvae are small, +slender, legless, worm-like creatures (fig. 77_c_) with small brown head +and twelve body segments. The pupae (fig. 77_e_) are slender, more +swollen at the anterior end and terminating in a forked process. They +float nearly motionless in a vertical position, the respiratory tubes in +contact with the surface film. The adults are all small, rarely +exceeding 2-1/4 mm. in length. The wings are more or less covered with +erect setulae or hairs and in many species variously spotted and marked +with iridescent blotches. The antennae have fourteen segments, the palpi +usually five. The wing venation and mouth-parts are shown in figures 77 +and 78. Of the twenty or more species of this genus occurring in the +United States the following are known to bite: _C. cinctus_, _C. +guttipennis_, _C. sanguisuga_, _C. stellifer_, _C. variipennis_, _C. +unicolor_. + +[Illustration: 78. Culicoides guttipennis; mouth parts of adult. After +Pratt.] + +One of the most widely distributed and commonest species in the Eastern +States is _C. guttipennis_ (fig. 77a). It is black with brown legs, a +whitish ring before the apex of each femur and both ends of each tibia; +tarsi yellow, knobs of halteres yellow. Mesonotum opaque, brown, two +vittae in the middle, enlarging into a large spot on the posterior half, +also a curved row of three spots in front of each wing, and the narrow +lateral margins, light gray pruinose. Wings nearly wholly covered with +brown hairs, gray, with markings as shown in the figure. Length one mm. + +_Johannseniella_ Will. is a wide-spread genus related to the foregoing. +Its mouth-parts are well adapted for piercing and it is said to be a +persistent blood sucker, particularly in Greenland. This genus is +distinguished from _Culicoides_ by its bare wings, the venation (fig. +163, c), and the longer tarsal claws. There are over twenty North +American species. + +[Illustration: 79. Chrysops univittatus, (x4). After Osborn.] + +In the Southwestern United States, _Tersesthes torrens_ Towns. occurs, a +little gnat which annoys horses, and perhaps man also, by its bite. It +is related to _Culicoides_ but differs in the number of antennal +segments and in its wing venation (fig. 163, e). The fly measures but +two mm. in length and is blackish in color. The antennae of the female +have thirteen segments, the palpi but three, of which the second is +enlarged and swollen. + + +Tabanidae or Horse-Flies + +The TABANIDAE,--horse-flies, ear-flies, and deer-flies,--are well-known +pests of cattle and horses and are often extremely annoying to man. The +characteristics of the family and of the principal North American genera +are given in the keys of Chapter XII. There are over 2500 recorded +species. As in the mosquitoes, the females alone are blood suckers. The +males are flower feeders or live on plant juices. This is apparently +true also of the females of some of the genera. + +The eggs are deposited in masses on water plants or grasses and sedges +growing in marshy or wet ground. Those of a common species of _Tabanus_ +are illustrated in figure 80, _a_. They are placed in masses of several +hundred, light colored when first deposited but turning black. In a week +or so the cylindrical larvae, tapering at both ends (fig. 80, _b_), +escape to the water, or damp earth, and lead an active, carnivorous +life, feeding mainly on insect larvae, and worms. In the forms which have +been best studied the larval life is a long one, lasting for months or +even for more than a year. Until recently, little was known concerning +the life-histories of this group, but the studies of Hart (1895), and +Hine (1903 +) have added greatly to the knowledge concerning North +American forms. + +Many of the species attack man with avidity and are able to inflict +painful bites, which may smart for hours. In some instances the wound is +so considerable that blood will continue to flow after the fly has left. +We have seen several cases of secondary infection following such bites. + +[Illustration: 80. (_a_) Eggs of Tabanus. Photograph by J. T. Lloyd.] + +[Illustration: 80. (_b_) Larva of Tabanus. Photograph by M. V. S.] + +The horse-flies have been definitely convicted of transferring the +trypanosome of surra from diseased to healthy animals and there is good +evidence that they transfer anthrax. The possibility of their being +important agents in the conveyal of human diseases should not be +overlooked. Indeed, Leiper has recently determined that a species of +_Chrysops_ transfers the blood parasite _Filaria diurna_. + + +Leptidae or Snipe-Flies + +The family LEPTIDAE is made up of moderate or large sized flies, +predaceous in habit. They are sufficiently characterized in the keys of +Chapter XII. Four blood-sucking species belonging to three genera have +been reported. Of these _Symphoromyia pachyceras_ is a western species. +Dr. J. C. Bradley, from personal experience, reports it as a vicious +biter. + +[Illustration: 80. (_c_) Mouth parts of Tabanus. After Grunberg.] + + +Oestridae or Bot-flies + +To the family OESTRIDAE belong the bot and warble-flies so frequently +injurious to animals. The adults are large, or of medium size, heavy +bodied, rather hairy, and usually resemble bees in appearance. + +The larvae live parasitically in various parts of the body of mammals, +such as the stomach (horse bot-fly), the subcutaneous connective tissue +(warble-fly of cattle), or the nasal passage (sheep bot-fly or head +maggot). + +There are on record many cases of the occurrence of the larvae of +Oestridae as occasional parasites of man. A number of these have been +collected and reviewed in a thesis by Mme. Petrovskaia (1910). The +majority of them relate to the following species. + +_Gastrophilus haemorrhoidalis_, the red tailed bot-fly, is one of the +species whose larvae are most commonly found in the stomach of the horse. +Schoch (1877) cites the case of a woman who suffered from a severe case +of chronic catarrh of the stomach, and who vomited, and also passed from +the anus, larvae which apparently belonged to this species. Such cases +are exceedingly rare but instances of subcutaneous infestation are +fairly numerous. In the latter type these larvae are sometimes the cause +of the peculiar "creeping myasis." This is characterized at its +beginning by a very painful swelling which gradually migrates, producing +a narrow raised line four to twenty-five millimeters broad. When the +larva is mature, sometimes after several months, it becomes stationary +and a tumor is formed which opens and discharges the larva along with +pus and serum. + +_Gastrophilus equi_ is the most widespread and common of the horse +bot-flies. Portschinsky reports it as commonly causing subcutaneous +myasis of man in Russia. + +_Hypoderma bovis_ (= _Oestrus bovis_), and _Hypoderma lineata_ are the +so-called warble-flies of cattle. The latter species is the more common +in North America but Dr. C. G. Hewitt has recently shown that _H. bovis_ +also occurs. Though warbles are very common in cattle in this country, +the adult flies are very rarely seen. They are about half an inch in +length, very hairy, dark, and closely resemble common honey-bees. + +They deposit their eggs on the hairs of cattle and the animals in +licking themselves take in the young larvae. These pass out through the +walls of the oesophagus and migrate through the tissues of the animal, +to finally settle down in the subcutaneous tissue of the back. The +possibility of their entering directly through the skin, especially in +case of infestation of man, is not absolutely precluded, although it is +doubtful. + +For both species of _Hypoderma_ there are numerous cases on record of +their occurrence in man. Hamilton (1893) saw a boy, six years of age, +who had been suffering for some months from the glands on one side of +his neck being swollen and from a fetid ulceration around the back teeth +of the lower jaw of the same side. Three months' treatment was of no +avail and the end seemed near; one day a white object, which was seen to +move, was observed in the ulcer at the root of the tongue, and on being +extracted was recognized as a full grown larva of _Hypoderma_. It was of +usual tawny color, about half an inch long when contracted, about one +third that thickness, and quite lively. The case resulted fatally. The +boy had been on a dairy farm the previous fall, where probably the egg +(or larva) was in some way taken into his mouth, and the larva found +between the base of the tongue and the jaw suitable tissue in which to +develop. + +Topsent (1901) reports a case of "creeping myasis" caused by _H. +lineata_ in the skin of the neck and shoulders of a girl eight years of +age. The larva travelled a distance of nearly six and a half inches. The +little patient suffered excruciating pain in the place occupied by the +larva. + +_Hypoderma diana_ infests deer, and has been known to occur in man. + +_Oestris ovis_, the sheep bot-fly, or head maggot, is widely distributed +in all parts of the world. In mid-summer the flies deposit living +maggots in the nostrils of sheep. These larvae promptly pass up the nasal +passages into the frontal and maxillary sinuses, where they feed on the +mucous to be found there. In their migrations they cause great +irritation to their host, and when present in numbers may cause vertigo, +paroxysms, and even death. Portschinsky in an important monograph on +this species, has discussed in detail its relation to man. He shows that +it is not uncommon for the fly to attack man and that the minute living +larvae are deposited in the eyes, nostrils, lips, or mouth. A typical +case in which the larvae were deposited in the eye was described by a +German oculist Kayser, in 1905. A woman brought her six year old +daughter to him and said that the day before, about noontime, a flying +insect struck the eye of the child and that since then she had felt a +pain which increased towards evening. In the morning the pain ceased but +the eye was very red. She was examined at about noon, at which time she +was quiet and felt no pain. She was not sensitive to light, and the only +thing noticed was a slight congestion and accumulation of secretion in +the corner of the right eye. A careful examination of the eye disclosed +small, active, white larvae that crawled out from the folds of the +conjunctiva and then back and disappeared. Five of these larvae were +removed and although an uncomfortable feeling persisted for a while, the +eye became normal in about three weeks. + +Some of the other recorded cases have not resulted so favorably, for the +eyesight has been seriously affected or even lost. + +According to Edmund and Etienne Sergent (1907), myasis caused by the +larvae of _Oestris ovis_ is very common among the shepherds in Algeria. +The natives say that the fly deposits its larvae quickly, while on the +wing, without pause. The greatest pain is caused when these larvae +establish themselves in the nasal cavities. They then produce severe +frontal headaches, making sleep impossible. This is accompanied by +continuous secretion from the nasal cavities and itching pains in the +sinuses. If the larvae happen to get into the mouth, the throat becomes +inflamed, swallowing is painful, and sometimes vomiting results. The +diseased condition may last for from three to ten days or in the case of +nasal infection, longer, but recovery always follows. The natives remove +the larvae from the eye mechanically by means of a small rag. When the +nose is infested, tobacco fumigations are applied, and in case of throat +infestation gargles of pepper, onion, or garlic extracts are used. + +_Rhinoestrus nasalis_, the Russian gad-fly, parasitizes the +nasopharyngeal region of the horse. According to Portschinsky, it not +infrequently attacks man and then, in all the known cases deposits its +larvae in the eye, only. This is generally done while the person is +quiet, but not during sleep. The fly strikes without stopping and +deposits its larva instantaneously. Immediately after, the victim +experiences lancinating pains which without intermission increase in +violence. There is an intense conjunctivitis and if the larvae are not +removed promptly the envelopes of the eye are gradually destroyed and +the organ lost. + +[Illustration: 81. Larvae of Dermatobia cyaniventris. After Blanchard.] + +[Illustration: 82. Young larva of Dermatobia cyaniventris. After +Surcouf.] + +_Dermatobia cyaniventris_--This fly (fig. 83) is widely distributed +throughout tropical America, and in its larval stage is well known as a +parasite of man. The larvae (figs. 81 and 82) which are known as the "ver +macaque," "torcel," "ver moyocuil" or by several other local names, +enter the skin and give rise to a boil-like swelling, open at the top, +and comparable with the swelling produced by the warble fly larvae, in +cattle. They cause itching and occasional excruciating pain. When +mature, nearly an inch in length, they voluntarily leave their host, +drop to the ground and complete their development. The adult female is +about 12 mm. in length. The face is yellow, the frons black with a +grayish bloom; antennae yellow, the third segment four times as long as +the second, the arista pectinate. The thorax is bluish black with +grayish bloom; the abdomen depressed, brilliant metallescent blue with +violet tinge. The legs are yellowish, the squamae and wings brownish. + +The different types of larvae represented in figure 81 were formerly +supposed to belong to different species but Blanchard regards them as +merely various stages of the same species. It is only very recently that +the early stage and the method by which man becomes infested were made +known. + +[Illustration: 83. Dermatobia cyaniventris (x1-3/4). After Manson.] + +[Illustration: 84. Mosquito carrying eggs of Dermatobia cyaniventris. +After Surcouf.] + +About 1900, Blanchard observed the presence of packets of large-sized +eggs under the abdomen of certain mosquitoes from Central America; and +in 1910, Dr. Morales, of Costa Rica, declared that the Dermatobia +deposited its eggs directly under the abdomen of the mosquito and that +they were thus carried to vertebrates. Dr. Nunez Tovar observed the +mosquito carriers of the eggs and placing larvae from this source on +animals, produced typical tumors and reared the adult flies. It remained +for Surcouf (1913) to work out the full details. He found that the +Dermatobia deposits its eggs in packets covered by a very viscid +substance, on leaves. These become attached to mosquitoes of the species +_Janthinosoma lutzi_ (fig. 84) which walk over the leaves. The eggs +which adhere to the abdomen, remain attached and are thus transported. +The embryo develops, but the young larva (fig. 82) remains in the egg +until it has opportunity to drop upon a vertebrate fed upon by the +mosquito. + + +Muscidae + +The following MUSCIDAE, characterized elsewhere, deserve special mention +under our present grouping of parasitic species. Other important species +will be considered as facultative parasites. + +_Stomoxys calcitrans_, the stable-fly, or the biting house-fly, is often +confused with _Musca domestica_ and therefore is discussed especially in +our consideration of the latter species as an accidental carrier of +disease. Its possible relation to the spread of infantile paralysis is +also considered later. + +The _tsetse flies_, belonging to the genus _Glossina_, are African +species of blood-sucking Muscidae which have attracted much attention +because of their role in transmitting various trypanosome diseases of +man and animals. They are characterized in Chapter XII and are also +discussed in connection with the diseases which they convey. + +_Chrysomyia macellaria_, (= _Compsomyia_), the "screw worm"-fly is one +of the most important species of flies directly affecting man, in North +America. It is not normally parasitic, however, and hence will be +considered with other facultative parasites in Chapter IV. + +[Illustration: 85. Larva of Auchmeromyia luteola. After Graham-Smith.] + +[Illustration: 86. Auchmeromyia luteola (x4). After Graham-Smith.] + +_Auchmeromyia luteola_, the Congo floor maggot. This is a muscid of +grewsome habits, which has a wide distribution throughout Africa. The +fly (fig. 86) deposits its eggs on the ground of the huts of the +natives. The whitish larvae (fig. 85) on hatching are slightly flattened +ventrally, and each segment bears posteriorly three foot-pads +transversely arranged. At night the larvae find their way into the low +beds or couches of the natives and suck their blood. The adult flies do +not bite man and, as far as known, the larvae do not play any role in the +transmission of sleeping sickness or other diseases. + +This habit of blood-sucking by muscid larvae is usually referred to as +peculiar to _Auchmeromyia luteola_ but it should be noted that the larvae +of _Protocalliphora_ frequent the nests of birds and feed upon the +young. Mr. A. F. Coutant has studied especially the life-history and +habits of _P. azurea_, whose larvae he found attacking young crows at +Ithaca, N.Y. He was unable to induce the larvae to feed on man. + +[Illustration: 87. Cordylobia anthropophaga (x3). After Fulleborn.] + +[Illustration: 88. Larva of Cordylobia anthropophaga. After Blanchard.] + +_Cordylobia anthropophaga_, (_Ochromyia anthropophaga_), or Tumbu-fly +(fig. 87) is an African species whose larvae affect man much as do those +of _Dermatobia cyniventris_, of Central and South America. The larva +(fig. 88), which is known as "ver du Cayor" because it was first +observed in Cayor, in Senegambia, develops in the skin of man and of +various animals, such as dogs, cats, and monkeys. It is about 12 mm. in +length, and of the form of the larvae of other muscids. Upon the +intermediate segments are minute, brownish recurved spines which give to +the larva its characteristic appearance. The life-history is not +satisfactorily worked out, but Fuller (1914), after reviewing the +evidence believes that, as a rule, it deposits its young in the sleeping +places of man and animals, whether such be a bed, a board, the floor, or +the bare ground. In the case of babies, the maggots may be deposited on +the scalp. The minute maggots bore their way painlessly into the skin. +As many as forty parasites have been found in one individual and one +author has reported finding more than three hundred in a spaniel puppy. +Though their attacks are at times extremely painful, it is seldom that +any serious results follow. + + +THE SIPHONAPTERA OR FLEAS + +The SIPHONAPTERA, or fleas (fig. 89) are wingless insects, with highly +chitinized and laterally compressed bodies. The mouth-parts are formed +for piercing and sucking. Compound eyes are lacking but some species +possess ocelli. The metamorphosis is complete. + +This group of parasites, concerning which little was known until +recently, has assumed a very great importance since it was learned that +fleas are the carriers of bubonic plague. Now over four hundred species +are known. Of these, several species commonly attack man. The most +common hominoxious species are _Pulex irritans_, _Xenopsylla cheopis_, +_Ctenocephalus canis_, _Ctenocephalus felis_, _Ceratophyllus fasciatus_ +and _Dermatophilus penetrans_, but many others will feed readily on +human blood if occasion arises. + +[Illustration: 89. Xenopsylla cheopis, male (x25). After Jordan and +Rothschild.] + +We shall treat in this place of the general biology and habits of the +hominoxious forms and reserve for the systematic section the discussion +of the characteristics of the different genera. + +The most common fleas infesting houses in the Eastern United States are +the cosmopolitan dog and cat fleas, _Ctenocephalus canis_ (fig. 90) and +_C. felis_. Their life cycles will serve as typical. These two species +have until recently been considered as one, under the name _Pulex +serraticeps_. See figure 92. + +[Illustration: 90. Dog flea (x15). After Howard.] + +The eggs are oval, slightly translucent or pearly white, and measure +about .5 mm. in their long diameter. They are deposited loosely in the +hairs of the host and readily drop off as the animal moves around. +Howard found that these eggs hatch in one to two days. The larvae are +elongate, legless, white, worm-like creatures. They are exceedingly +active, and avoid the light in every way possible. They cast their first +skin in from three to seven days and their second in from three to four +days. They commenced spinning in from seven to fourteen days after +hatching and the imago appeared five days later. Thus in summer, at +Washington, the entire life cycle may be completed in about two weeks. +(cf. fig. 91, 92). + +Strickland's (1914) studies on the biology of the rat flea, +_Ceratophyllus fasciatus_, have so important a general bearing that we +shall cite them in considerable detail. + +[Illustration: 91. Larva of Xenopsylla cheopis. After Bacot and +Ridewood.] + +He found, to begin with, that there is a marked inherent range in the +rate of development. Thus, of a batch of seventy-three eggs, all laid in +the same day and kept together under the same conditions, one hatched in +ten days; four in eleven days; twenty-five in twelve days; thirty-one in +thirteen days; ten in fourteen days; one in fifteen days; and one in +sixteen days. Within these limits the duration of the egg period seems +to depend mainly on the degree of humidity. The incubation period is +never abnormally prolonged as in the case of lice, (Warburton) and +varying conditions of temperature and humidity have practically no +effect on the percentage of eggs which ultimately hatch. + +The same investigator found that the most favorable condition for the +larva is a low temperature, combined with a high degree of humidity; and +that the presence of rubbish in which the larva may bury itself is +essential to its successful development. When larvae are placed in a +bottle containing either wood-wool soiled by excrement, or with feathers +or filter paper covered with dried blood they will thrive readily and +pupate. They seem to have no choice between dried blood and powdered rat +feces for food, and also feed readily on flea excrement. They possess +the curious habit of always devouring their molted skins. + +[Illustration: 92. Head and pronotum of (_a_) dog flea; (_b_) of cat +flea; (_c_) of hen flea. After Rothschild. (_d_) Nycteridiphilus +(Ischnopsyllus) hexactenus. After Oudemans.] + +An important part of Strickland's experiments dealt with the question of +duration of the pupal stage under the influence of temperature and with +the longevity and habits of the adult. In October, he placed a batch of +freshly formed cocoons in a small dish that was kept near a white rat in +a deep glass jar in the laboratory. Two months later one small and +feeble flea had emerged, but no more until February, four months after +the beginning of the experiment. Eight cocoons were then dissected and +seven more found to contain the imago fully formed but in a resting +state. The remainder of the batch was then placed at 70^o F. for one +night, near a white rat. The next day all the cocoons were empty and the +fleas were found on the white rat. + +Thus, temperature greatly influences the duration of the pupal period, +which in _Ceratophyllus fasciatus_ averages seventeen days. Moreover, +when metamorphosis is complete a low temperature will cause the imago to +remain within the cocoon. + +Sexually mature and ovipositing fleas, he fed at intervals and kept +alive for two months, when the experiment was discontinued. In the +presence of rubbish in which they could bury themselves, unfed rat fleas +were kept alive for many months, whereas in the absence of any such +substratum they rarely lived a month. In the former case, it was found +that the length of life is influenced to some degree by the temperature +and humidity. In an experiment carried out at 70^o F. and 45 per cent +humidity, the fleas did not live for more than four months, while in an +experiment at 60^o F. and 70 per cent humidity they lived for at least +seventeen months. There was no indication that fleas kept under these +conditions sucked moisture from surrounding objects, and those kept in +bell jars, with an extract of flea-rubbish on filter paper, did not live +any longer than those which were not so supplied. + +Curiously enough, although the rat is the normal host of _Ceratophyllus +fasciatus_, it was found that when given the choice these fleas would +feed upon man in preference to rats. However, none of the fleas laid +eggs unless they fed on rat blood. + +The experiments of Strickland on copulation and oviposition in the rat +flea showed that fleas do not copulate until they are sexually mature +and that, at least in the case of _Ceratophyllus fasciatus_, the +reproductive organs are imperfectly developed for some time (more than a +week) after emerging from the pupa. When mature, copulation takes place +soon after the fleas have fed on their true host--the rat--but not if +they have fed on a facultative host only, such as man. Copulation is +always followed by oviposition within a very short time. + +The effect of the rat's blood on the female with regard to egg-laying, +Strickland concludes, is stimulating rather than nutritive, as fleas +that were without food for many months were observed to lay eggs +immediately after one feed. Similarly, the male requires the stimulus of +a meal of rat's blood before it displays any copulatory activity. + +Mitzmain (1910) has described in detail the act of biting on man, as +observed in the squirrel flea, _Ceratophyllus acutus_. "The flea when +permitted to walk freely on the arm selects a suitable hairy space where +it ceases abruptly in its locomotion, takes a firm hold with the tarsi, +projects its proboscis, and prepares to puncture the skin. A puncture is +drilled by the pricking epipharynx, the saw-tooth mandibles +supplementing the movement by lacerating the cavity formed. The two +organs of the rostrum work alternately, the middle piece boring, while +the two lateral elements execute a sawing movement. The mandibles, owing +to their basal attachments, are, as is expressed by the advisory +committee on plague investigations in India (_Journal of Hygiene_, vol. +6, No. 4, p. 499), 'capable of independent action, sliding up and down +but maintaining their relative positions and preserving the lumen of the +aspiratory channel.' The labium doubles back, the V-shaped groove of +this organ guiding the mandibles on either side." + +"The action of the proboscis is executed with a forward movement of the +head and a lateral and downward thrust of the entire body. As the +mouth-parts are sharply inserted, the abdomen rises simultaneously. The +hind and middle legs are elevated, resembling oars. The forelegs are +doubled under the thorax, the tibia and tarsi resting firmly on the +epidermis serve as a support for the body during the feeding. The +maxillary palpi are retracted beneath the head and thorax. The labium +continues to bend, at first acting as a sheath for the sawing mandibles, +and as these are more deeply inserted, it bends beneath the head with +the elasticity of a bow, forcing the mandibles into the wound until the +maxillae are embedded in the skin of the victim. When the proboscis is +fully inserted, the abdomen ceases for a time its lateral swinging." + +"The acute pain of biting is first felt when the mandibles have not +quite penetrated and subsequently during each distinct movement of the +abdomen. The swinging of the abdomen gradually ceases as it becomes +filled with blood. The sting of the biting gradually becomes duller and +less sensitive as feeding progresses. The movements of the elevated +abdomen grow noticeably feebler as the downward thrusts of the springy +bow-like labium becomes less frequent." + +"As the feeding process advances one can discern through the translucent +walls of the abdomen a constant flow of blood, caudally from the +pharynx, accompanied by a peristaltic movement. The end of the meal is +signified in an abrupt manner. The flea shakes its entire body, and +gradually withdraws its proboscis by lowering the abdomen and legs and +violently twisting the head." + +"When starved for several days the feeding of the rat fleas is conducted +in a rather vigorous manner. As soon as the proboscis is buried to the +full length the abdomen is raised and there ensues a gradual lateral +swaying motion, increasing the altitude of the raised end of the abdomen +until it assumes the perpendicular. The flea is observed at this point +to gain a better foothold by advancing the fore tarsi, and then, +gradually doubling back the abdomen, it turns with extreme agility, +nearly touching with its dorsal side the skin of the hand upon which it +is feeding. Meanwhile, the hungry parasite feeds ravenously." + +"It is interesting to note the peculiar nervous action which the rodent +fleas exhibit immediately when the feeding process is completed or when +disturbed during the biting. Even while the rostrum is inserted to the +fullest the parasite shakes its head spasmodically; in a twinkling the +mouth is withdrawn and then the flea hops away." + +A habit of fleas which we shall see is of significance in considering +their agency in the spread of bubonic plague, is that of ejecting blood +from the anus as they feed. + +Fleas are famous for their jumping powers, and in control measures it is +of importance to determine their ability along this line. It is often +stated that they can jump about four inches, or, according to the Indian +Plague Commission _Xenopsylla cheopis_ cannot hop farther than five +inches. Mitzmain (1910) conducted some careful experiments in which he +found that the human flea, _Pulex irritans_, was able to jump as far as +thirteen inches on a horizontal plane. The mean average of five +specimens permitted to jump at will was seven and three-tenths inches. +The same species was observed to jump perpendicularly to a height of at +least seven and three-fourths inches. Other species were not able to +equal this record. + +The effect of the bite of fleas on man varies considerably according to +the individual susceptibility. According to Patton and Cragg, this was +borne out in a curious manner by the experiments of Chick and Martin. +"In these, eight human hosts were tried; in seven, little or no +irritation was produced, while in one quite severe inflammation was set +up around each bite." Of two individuals, equally accustomed to the +insects, going into an infested room, one may be literally tormented by +them while the other will not notice them. Indeed it is not altogether +a question of susceptibility, for fleas seem to have a special +predilection for certain individuals. The typical itching wheals +produced by the bites are sometimes followed, especially after +scratching, by inflammatory papules. + +The itching can be relieved by the use of lotions of carbolic acid (2-3 +per cent), camphor, menthol lotion, or carbolated vaseline. If forced to +sleep in an infested room, protection from attacks can be in a large +measure gained by sprinkling pyrethrum, bubach, or California insect +powder between the sheets. The use of camphor, menthol, or oil of +eucalyptus, or oil of pennyroyal is also said to afford protection to a +certain extent. + +In the Eastern United States the occurrence of fleas as household pests +is usually due to infested cats and dogs which have the run of the +house. We have seen that the eggs are not attached to the host but drop +to the floor when they are laid. Verrill, cited by Osborn, states that +on one occasion he was able to collect fully a teaspoonful of eggs from +the dress of a lady in whose lap a half-grown kitten had been held for a +short time. Patton and Cragg record seeing the inside of a hat in which +a kitten had spent the night, so covered with flea eggs that it looked +"as if it had been sprinkled with sugar from a sifter." It is no wonder +that houses in which pets live become overrun with the fleas. + +One of the first control measures, then, consists in keeping such +animals out of the house or in rigorously keeping them free from fleas. +The latter can best be accomplished by the use of strong tar soap or +Armour's "Flesope," which may be obtained from most druggists. The use +of a three per cent solution of creolin, approximately four teaspoonfuls +to a quart of warm water, has also been recommended. While this is +satisfactory in the case of dogs, it is liable to sicken cats, who will +lick their fur in an effort to dry themselves. Howard recommends +thoroughly rubbing into the fur a quantity of pyrethrum powder. This +partially stupifies the fleas which should be promptly swept up and +burned. + +He also recommends providing a rug for the dog or cat to sleep on and +giving this rug a frequent shaking and brushing, afterwards sweeping up +and burning the dust thus removed. + +Since the larvae of fleas are very susceptible to exposure, the use of +bare floors, with few rugs, instead of carpets or matting, is to be +recommended. Thorough sweeping, so as to allow no accumulation of dust +in cracks and crevices will prove efficient. If a house is once +infested it may be necessary to thoroughly scrub the floors with hot +soapsuds, or to spray them with gasoline. If the latter method is +adopted, care must be taken to avoid the possibility of fire. + +To clear a house of fleas Skinner recommends the use of flake +naphthalene. In a badly infested house he took one room at a time, +scattering on the floor five pounds of flake naphthalene, and closed it +for twenty-four hours. It proved to be a perfect and effectual remedy +and very inexpensive, as the naphthalene could be swept up and +transferred to other rooms. Dr. Skinner adds, "so far as I am concerned, +the flea question is solved and if I have further trouble I know the +remedy. I intend to keep the dog and cat." + +The late Professor Slingerland very effectively used hydrocyanic acid +gas fumigation in exterminating fleas in houses. In one case, where +failure was reported, he found on investigation that the house had +become thoroughly reinfested from pet cats, which had been left +untreated. Fumigation with sulphur is likewise efficient. + +The fact that adult fleas are usually to be found on the floor, when not +on their hosts, was ingeniously taken advantage of by Professor S. H. +Gage in ridding an animal room at Cornell University of the pests. He +swathed the legs of a janitor with sticky fly-paper and had him walk +back and forth in the room. Large numbers of the fleas were collected in +this manner. + +In some parts of the southern United States hogs are commonly infested +and in turn infest sheds, barns and even houses. Mr. H. E. Vick informs +us that it is a common practice to turn sheep into barn-lots and sheds +in the spring of the year to collect in their wool, the fleas which +abound in these places after the hogs have been turned out. + +It is a common belief that adult fleas are attracted to fresh meat and +that advantage of this can be taken in trapping them. Various workers, +notably Mitzman (1910), have shown that there is no basis for such a +belief. + + +THE TRUE CHIGGERS--The chigoes, or true chiggers, are the most +completely parasitic of any of the fleas. Of the dozen or more known +species, one commonly attacks man. This is _Dermatophilus penetrans_, +more commonly known as _Sarcopsylla penetrans_ or _Pulex penetrans_. + +This species occurs in Mexico, the West Indies, Central and South +America. There are no authentic records of its occurrence in the United +States although, as Baker has pointed out, there is no reason why it +should not become established in Florida and Texas. It is usually +believed that Brazil was its original home. Sometime about the middle of +the nineteenth century it was introduced into West Africa and has spread +across that continent. + +The males and the immature females of _Dermatophilus penetrans_ (fig. +93) closely resemble those of other fleas. They are very active little +brown insects about 1-1.2 mm. in size, which live in the dust of native +huts and stables, and in dry, sandy soil. In such places they often +occur in enormous numbers and become a veritable plague. + +[Illustration: 93. Dermatophilus penetrans. Much enlarged. After +Karsten.] + +They attack not only man but various animals. According to Castellani +and Chalmers, "Perhaps the most noted feature is the way in which it +attacks pigs. On the Gold Coast it appeared to be largely kept in +existence by these animals. It is very easily captured in the free state +by taking a little pig with a pale abdomen, and placing it on its back +on the ground on which infected pigs are living. After watching a few +moments, a black speck will appear on the pig's abdomen, and quickly +another and another. These black specks are jiggers which can easily be +transferred to a test tube. On examination they will be found to be +males and females in about equal numbers." + +Both the males and females suck blood. That which characterizes this +species as distinguished from other fleas attacking man is that when the +impregnated female attacks she burrows into the skin and there swells +until in a few days she has the size and appearance of a small pea (fig. +94). Where they are abundant, hundreds of the pests may attack a single +individual (fig. 95). Here they lie with the apex of the abdomen +blocking the opening. According to Fulleborn (1908) they do not +penetrate beneath the epidermis. The eggs are not laid in the flesh of +the victim, as is sometimes stated, but are expelled through this +opening. The female then dies, withers and falls away or is expelled by +ulceration. According to Brumpt, she first quits the skin and then, +falling to the ground, deposits her eggs. The subsequent development in +so far as known, is like that of other fleas. + +[Illustration: 94. Dermatophilus penetrans, gravid female. After +Moniez.] + +[Illustration: 95. Chiggers in the sole of foot of man. Manson's +Tropical Diseases. Permission of Cassell and Co.] + +The chigoe usually enters between the toes, the skin about the roots of +the nails, or the soles of the feet, although it may attack other parts +of the body. Mense records the occurrence in folds of the epidermis, as +in the neighborhood of the anus. They give rise to irritation and unless +promptly and aseptically removed there often occurs pus formation and +the development of a more or less serious abscess. Gangrene and even +tetanus may ensue. + +Treatment consists in the careful removal of the insect, an operation +more easily accomplished a day or two after its entrance, than at first, +when it is unswollen. The ulcerated point should then be treated with +weak carbolic acid, or tincture of iodine, or dusted thoroughly with an +antiseptic powder. + +[Illustration: 96. Echidnophaga gallinacea.] + +[Illustration: 97. Echidnophaga gallinacea infesting head of chicken. +After Enderlein.] + +Castellani and Chalmers recommend as prophylactic measures, keeping the +house clean and keeping pigs, poultry, and cattle away therefrom. "High +boots should be used, and especial care should be taken not to go to a +ground floor bathroom with bare feet. The feet, especially the toes, and +under the nails, should be carefully examined every morning to see if +any black dots can be discovered, when the jigger should be at once +removed, and in this way suppuration will be prevented. It is +advisable, also, to sprinkle the floors with carbolic lotion, Jeyes' +fluid, or with pyrethrum powder, or with a strong infusion of native +tobacco, as recommended by Law and Castellani." + +_Echidnophaga gallinacea_ (fig. 96) is a widely distributed Hectopsyllid +attacking poultry (fig. 97). It occurs in the Southern and Southwestern +United States and has been occasionally reported as attacking man, +especially children. It is less highly specialized than _Dermatophilus +penetrans_, and does not ordinarily cause serious trouble in man. + + + + +CHAPTER IV + +ACCIDENTAL OR FACULTATIVE PARASITES + + +In addition to the many species of Arthropods which are normally +parasitic on man and animals, there is a considerable number of those +which may be classed as _accidental_ or _facultative_ parasites. + +Accidental or facultative parasites are species which are normally +free-living, but which are able to exist as parasites when accidentally +introduced into the body of man or other animal. A wide range of forms +is included under this grouping. + + +ACARINA + +A considerable number of mites have been reported as accidental or even +normal, endoparasites of man, but the authentic cases are comparatively +few. + +In considering such reports it is well to keep in mind von Siebold's +warning that in view of the universal distribution of mites one should +be on his guard. In vessels in which animal and other organic fluids and +moist substances gradually dry out, mites are very abundantly found. If +such vessels are used without very careful preliminary cleaning, for the +reception of evacuations of the sick, or for the reception of parts +removed from the body, such things may be readily contaminated by mites, +which have no other relation whatever to them. + +Nevertheless, there is no doubt but that certain mites, normally +free-living, have occurred as accidental parasites of man. Of these the +most commonly met with is _Tyroglyphus siro_, the cheese-mite. + +_Tyroglyphus siro_ is a small mite of a whitish color. The male measures +about 500u long by 250u wide, the female slightly larger. They live in +cheese of almost any kind, especially such as is a little decayed. "The +individuals gather together in winter in groups or heaps in the hollows +and chinks of the cheese and there remain motionless. As soon as the +temperature rises a little, they gnaw away at the cheese and reduce it +to a powder. The powder is composed of excrement having the appearance +of little grayish microscopic balls; eggs, old and new, cracked and +empty; larvae, nymphs, and perfect mites, cast skins and fragments of +cheese, to which must be added numerous spores of microscopic +fungi."--Murray. + +_Tyroglyphus siro_, and related species, have been found many times in +human feces, under conditions which preclude the explanation that the +contamination occurred outside of the body. They have been supposed to +be the cause of dysentery, or diarrhoea, and it is probable that the +_Acarus dysenteriae_ of Linnaeus, and Latreille, was this species. +However, there is little evidence that the mites cause any noteworthy +symptoms, even when taken into the body in large numbers. + +_Histiogaster spermaticus_ (fig. 152) is a Tyroglyphid mite which was +reported by Trouessart (1902) as having been found in a cyst in the +groin, adherent to the testis. When the cyst was punctured, it yielded +about two ounces of opalescent fluid containing spermatozoa and numerous +mites in all stages of development. The evidence indicated that a +fecundated female mite had been introduced into the urethra by means of +an unclean catheter. Though Trouessart reported the case as that of a +Sarcoptid, Banks places the genus _Histiogaster_ with the Tyroglyphidae. +He states that our species feeds on the oyster-shell bark louse, +possibly only after the latter is dead, and that in England a species +feeds within decaying reeds. + +_Nephrophages sanguinarius_ is a peculiarly-shaped, angular mite which +was found by Miyake and Scriba (1893) for eight successive days in the +urine of a Japanese suffering from fibrinuria. Males, .117 mm. long by +.079 mm. wide, females .36 mm. by. 12 mm., and eggs were found both in +the spontaneously emitted urine and in that drawn by means of a +catheter. All the mites found were dead. The describers regarded this +mite as a true endoparasite, but it is more probable that it should be +classed as an accidental parasite. + + +MYRIAPODA + +There are on record a number of cases of myriapods occurring as +accidental parasites of man. The subject has been treated in detail by +Blanchard (1898 and 1902), who discussed forty cases. Since then at +least eight additions have been made to the list. + +Neveau-Lamaire (1908) lists thirteen species implicated, representing +eight different genera. Of the _Chilognatha_ there are three, _Julus +terrestris_, _J. londinensis_ and _Polydesmus complanatus_. The +remainder are _Chilopoda_, namely, _Lithobius forficatus_, _L. +malenops_, _Geophilus carpophagus_, _G. electricus_, _G. similis_, _G. +cephalicus_, _Scutigera coleoptrata_, _Himantarium gervaisi_, +_Chaetechelyne vesuviana_ and _Stigmatogaster subterraneus_. + +The majority of the cases relate to infestation of the nasal fossae, or +the frontal sinus, but intestinal infestation also occurs and there is +one recorded case of the presence of a species in _Julus_ (fig. 13) in +the auditory canal of a child. + +In the nose, the myriapods have been known to live for months and +according to some records, even for years. The symptoms caused by their +presence are inflammation, with or without increased flow of mucus, +itching, more or less intense headache, and at times general symptoms +such as vertigo, delirium, convulsions, and the like. These symptoms +disappear suddenly when the parasites are expelled. + +In the intestine of man, myriapods give rise to obscure symptoms +suggestive of infestation by parasitic worms. In a case reported by +Verdun and Bruyant (1912), a child twenty months of age had been +affected for fifteen days by digestive disturbances characterized by +loss of appetite, nausea and vomiting. The latter had been particularly +pronounced for three days, when there was discovered in the midst of the +material expelled a living myriapod of the species _Chaetechelyne +vesuviana_. Anthelminthics had been administered without result. In some +of the other cases, the administration of such drugs had resulted in the +expulsion of the parasite through the anus. + +One of the extreme cases on record is that reported by Shipley (1914). +Specimens of _Geophilus gorizensis_ (= _G. subterraneus_) "were vomited +and passed by a woman of 68 years of age. Some of the centipedes emerged +through the patient's nose, and it must be mentioned that she was also +suffering from a round worm. One of her doctors was of the opinion that +the centipedes were certainly breeding inside the lady's intestines, and +as many as seven or eight, sometimes more, were daily leaving the +alimentary canal." + +"According to her attendant's statements those centipedes had left the +body in some hundreds during a period of twelve or eighteen months. +Their presence produced vomiting and some haematemesis, and treatment +with thymol, male-fern and turpentine had no effect in removing the +creatures." + +The clinical details, as supplied by Dr. Theodore Thompson were as +follows: + +"Examined by me July, 1912, her tongue was dry and glazed. There was +bleeding taking place from the nose and I saw a living centipede she had +just extracted from her nostril. Her heart, lungs and abdomen appeared +normal. She was not very wasted, and did not think she had lost much +flesh, nor was there any marked degree of anemia." + +Shipley gives the following reasons for believing it impossible that +these centipedes could have multiplied in the patient's intestine. "The +breeding habits of the genus _Geophilus_ are peculiar, and ill adapted +for reproducing in such a habitat. The male builds a small web or nest, +in which he places his sperm, and the female fertilizes herself from +this nest or web, and when the eggs are fertilized they are again laid +in a nest or web in which they incubate and in two or three weeks hatch +out. The young _Geophilus_ differ but very little from the adult, except +in size. It is just possible, but improbable, that a clutch of eggs had +been swallowed by the host when eating some vegetables or fruit, but +against this is the fact that the _Geophilus_ does not lay its eggs upon +vegetables or fruit, but upon dry wood or earth. The egg-shell is very +tough and if the eggs had been swallowed the egg-shells could certainly +have been detected if the dejecta were examined. The specimens of the +centipede showed very little signs of being digested, and it is almost +impossible to reconcile the story of the patient with what one knows of +the habits of the centipedes." + +In none of the observed cases have there been any clear indications as +to the manner of infestation. It is possible that the myriapods have +been taken up in uncooked fruit or vegetables. + + +LEPIDOPTEROUS LARVAE + +SCHOLECIASIS--Hope (1837) brought together six records of infestation of +man by lepidopterous larvae and proposed to apply the name scholeciasis +to this type of parasitism. The clearest case was that of a young boy +who had repeatedly eaten raw cabbage and who vomited larvae of the +cabbage butterfly, _Pieris brassicae_. Such cases are extremely rare, and +there are few reliable data relative to the subject. In this connection +it may be noted that Spuler (1906) has described a moth whose larvae live +as ectoparasites of the sloth. + + +COLEOPTERA + +CANTHARIASIS--By this term Hope designated instances of accidental +parasitism by the larvae or adults of beetles. Reports of such cases +are usually scouted by parasitologists but there seems no good basis for +wholly rejecting them. Cobbold refers to a half dozen cases of +accidental parasitism by the larvae of _Blaps mortisaga_. In one of +these cases upwards of 1200 larvae and several perfect insects were +said to have been passed _per annum_. French (1905) reports the case of +a man who for a considerable period voided adult living beetles of the +species _Nitidula bipustulata_. Most of the other cases on record relate +to the larvae of _Dermestidae_ (larder beetles _et al._) or +_Tenebrionidae_ (meal infesting species). Infestation probably occurs +through eating raw or imperfectly cooked foods containing eggs or minute +larvae of these insects. + +[Illustration: 98. Larva of Piophila casei. Caudal aspect of larva. +Posterior stigmata.] + +Brumpt cites a curious case of accidental parasitism by a coleopterous +larva belonging to the genus _Necrobia_. This larva was extracted from a +small tumor, several millimeters long, on the surface of the conjunctiva +of the eye. The larvae of this genus ordinarily live in decomposing flesh +and cadavers. + + +DIPTEROUS LARVAE + +[Illustration: 99. Piophila casei. After Graham-Smith.] + +MYASIS--By this term (spelled also myiasis, and myiosis), is meant +parasitism by dipterous larvae. Such parasitism may be normal, as in the +cases already described under the heading parasitic Diptera, or it may +be facultative, due to free-living larvae being accidentally introduced +into wounds or the body-cavities of man. Of this latter type, there is a +multitude of cases on record, relating to comparatively few species. The +literature of the subject, like that relating to facultative parasitism +in general, is unsatisfactory, for most of the determinations of species +have been very loose. Indeed, so little has been known regarding the +characteristics of the larvae concerned that in many instances they could +not be exactly determined. Fortunately, several workers have undertaken +comparative studies along this line. The most comprehensive publication +is that of Banks (1912), entitled "The structure of certain dipterous +larvae, with particular reference to those in human food." + +Without attempting an exhaustive list, we shall discuss here the more +important species of Diptera whose larvae are known to cause myasis, +either external or internal. The following key will serve to determine +those most likely to be encountered. The writers would be glad to +examine specimens not readily identifiable, if accompanied by exact data +relative to occurrence. + +_a._ Body more or less flattened, depressed; broadest in the middle, + each segment with dorsal, lateral, and ventral fleshy processes, of + which the laterals, at least, are more or less spiniferous (fig. + 101). _Fannia_ (= _Homalomyia_). + + In _F. canicularis_ the dorsal processes are nearly as long as the + laterals; in _F. scalaris_ the dorsal processes are short spinose + tubercles. + +_aa._ Body cylindrical, or slender conical tapering toward the head; + without fleshy lateral processes (fig. 105). + +_b._ With the posterior stigmata at the end of shorter or longer + tubercles, or if not placed upon tubercles, then not in pit; usually + without a "marginal button" and without a chitinous ring surrounding + the three slits; the slits narrowly or broadly oval, not bent (fig. + 171 i). _Acalyptrate muscidae_ and some species of + _Anthomyiidae_. To this group belong the cheese skipper (_Piophila + casei_, figs. 98, 99), the pomace-fly (_Drosophila ampelophila_), + the apple maggot (_Rhagoletis pomonella_), the cherry fruit fly + (_Rhagoletis cingulata_), the small dung fly (_Sepsis violacea_, + fig. 170), the beet leaf-miner (_Pegomyia vicina_, fig. 171 i), the + cabbage, bean and onion maggots (_Phorbia_ spp.) et. al. + +_bb._ Posterior stigmata of various forms, if the slits are narrowly + oval (fig. 171) then they are surrounded by a chitin ring which may + be open ventro-mesally. + +_c._ Integument leathery and usually strongly spinulose; larvae + hypodermatic or endoparasitic. Bot flies (fig. 171, f, g, + k).--_Oestridae_ + +_cc._ Integument not leathery and (except in _Protocalliphora_) spinulae + restricted to transverse patches near the incisures of the segments. + +_d._ The stigmal plates in a pit; the lip-like margin of the pit with a + number of fleshy tubercles; chitin ring of the stigma not complete; + open ventro-mesally, button absent (fig. 171 e). Flesh + flies.--_Sarcophaga_ + +_dd._ Stigmata not in a pit. + +_e._ The chitin ring open ventra-mesally; button absent (fig. 171 c). + Screw-worm fly. _Chrysomyia_ + +_ee._ The chitin ring closed. + +_f._ Slits of the posterior stigmata straight; marginal "button" present + (fig. 171 b); two distinct mouth hooks, fleshy tubercles around the + anal area. _Phormia_ (fig. 171 f), _Lucilia_ and _Calliphora_ (fig. + 172, a, b), _Protocalliphora_ (fig. 171, j), _Cynomyia_ (fig. 171, + a). Blow flies, bluebottle flies. _Calliphorinae_ + +_ff._ Slits of the posterior stigmata sinuous or bent. Subfamily + Muscinae. + +_g._ Slits of the posterior stigmata bent; usually two mouth hooks. + _Muscina stabulans_ (fig. 171, l), _Muscina similis_, _Myiospila + meditabunda_ (fig. 172, i), and some of the higher _Anthomyiidae_. + +_gg._ Slits of the posterior stigmata sinuous; mouth hooks usually + consolidated into one. The house-fly (_Musca domestica_ fig. 171, + d), the stable fly (_Stomoxys calcitrans_), the horn fly (_Lyperosia + irritans_), _Pyrellia_, _Pseudopyrellia_, _Morellia_, _Mesembrina_. + _Polietes_, et. al. (fig. 172 in part). + +_Eristalis_--The larvae of _Eristalis_ are the so-called rat-tailed +maggots, which develop in foul water. In a few instances these larvae +have been known to pass through the human alimentary canal uninjured. +Hall and Muir (1913) report the case of a boy five years of age, who had +been ailing for ten weeks and who was under treatment for indigestion +and chronic constipation. For some time he had vomited everything he +ate. On administration of a vermifuge he voided one of the rat-tailed +maggots of _Eristalis_. He admitted having drunk water from a ditch full +of all manner of rotting matter. It was doubtless through this that he +became infested. It is worth noting that the above described symptoms +may have been due to other organisms or substances in the filthy water. + +_Piophila casei_, the cheese-fly (fig. 99), deposits its eggs not only +in old cheeses, but on ham, bacon, and other fats. The larvae (fig. 98) +are the well-known cheese skippers, which sometimes occur in great +abundance on certain kinds of cheese. Indeed, some people have a +comfortable theory that such infested cheese is especially good. Such +being the case, it is small wonder that this species has been repeatedly +reported as causing intestinal myasis. Thebault (1901) describes the +case of a girl who, shortly after consuming a large piece of badly +infested cheese, became ill and experienced severe pains in the region +of the navel. Later these extended through the entire alimentary canal, +the excrement was mixed with blood and she suffered from vertigo and +severe headaches. During the four following days the girl felt no +change, although the excretion of the blood gradually diminished and +stopped. On the fourth day she voided two half-digested larvae and, +later, seven or eight, of which two were alive and moving. + +That these symptoms may be directly attributed to the larvae, or +"skippers," has been abundantly shown by experimental evidence. +Portschinsky cites the case of a dog fed on cheese containing the larvae. +The animal suffered much pain and its excrement contained blood. On +_post mortem_ it was found that the small intestine throughout almost +its entire length was marked by bloody bruises. The papillae on these +places were destroyed, although the walls were not entirely perforated. +In the appendix were found two or three dead larvae. Alessandri (1910) +has likewise shown that the larvae cause intestinal lesions. + +According to Graham-Smith, Austen (1912) has recorded a case of myasis +of the nose, attended with a profuse watery discharge of several weeks +duration and pain, due to the larvae of _Piophila casei_. + + +ANTHYOMYIIDAE--The characteristic larvae of two species of _Fannia_ (= +_Homalomyia_ or _Anthomyia_, in part) (fig. 101) are the most commonly +reported of dipterous larvae causing intestinal myasis. Hewitt (1912) has +presented a valuable study of the bionomics and of the larvae of these +flies, a type of what is needed for all the species concerned in myasis. +We have seen two cases of their having been passed in stools, without +having caused any special symptoms. In other instances their presence in +the alimentary canal has given rise to symptoms vaguely described as +those of tapeworm infestation, or helminthiasis. More specifically, they +have been described as causing vertigo, severe headache, nausea and +vomiting, severe abdominal pains, and in some instances, bloody +diarrhoea. + +[Illustration: 100. Fannia canicularis (x4). After Graham-Smith.] + +One of the most striking cases is that reported by Blankmeyer (1914), of +a woman whose illness began fourteen years previously with nausea and +vomiting. After several months of illness she began passing larvae and +was compelled to resort to enemas. Three years previous to the report, +she noticed frequent shooting pains in the rectal region and at times +abdominal tenderness was marked. There was much mucus in the stools and +she "experienced the sensation of larvae crawling in the intestine." +Occipital headaches were marked, with remissions, and constipation +became chronic. The appetite was variable, there was a bad taste in the +mouth, tongue furred and ridged, and red at the edges. Her complexion +was sallow, and general nervousness was marked. As treatment, there were +given doses of magnesium sulphate before breakfast and at 4 P. M., with +five grain doses of salol four times a day. The customary parasiticides +yielded no marked benefit. At the time of the report the patient passed +from four to fifty larvae per day, and was showing some signs of +improvement. The nausea had disappeared, her nervousness was less +evident, and there was a slight gain in weight. + +The case was complicated by various other disorders, but the symptoms +given above seem to be in large part attributable to the myasis. There +is nothing in the case to justify the assumption that larvae were +continuously present, for years. It seems more reasonable to suppose +that something in the habits of the patient favored repeated +infestation. Nevertheless, a study of the various cases of intestinal +myasis caused by these and other species of dipterous larvae seems to +indicate that the normal life cycle may be considerably prolonged under +the unusual conditions. + +The best authenticated cases of myasis of the urinary passage have been +due to larvae of _Fannia_. Chevril (1909) collected and described twenty +cases, of which seven seemed beyond doubt. One of these was that of a +woman of fifty-five who suffered from albuminuria, and urinated with +much difficulty, and finally passed thirty to forty larvae of _Fannia +canicularis_. + +It is probable that infestation usually occurs through eating partially +decayed fruit or vegetables on which the flies have deposited their +eggs. Wellman points out that the flies may deposit their eggs in or +about the anus of persons using outside privies and Hewitt believes that +this latter method of infection is probably the common one in the case +of infants belonging to careless mothers. "Such infants are sometimes +left about in an exposed and not very clean condition, in consequence of +which flies are readily attracted to them and deposit their eggs." + +[Illustration: 101. Larva of Fannia scalaris.] + + +MUSCINAE--The larvae of the common house-fly, _Musca domestica_, are +occasionally recorded as having been passed with the feces or vomit of +man. While such cases may occur, it is probable that in most instances +similar appearing larvae of other insects have been mistakenly +identified. + +_Muscina stabulans_ is regarded by Portschinsky (1913) as responsible +for many of the cases of intestinal myasis attributed to other species. +He records the case of a peasant who suffered from pains in the lower +part of the breast and intestines, and whose stools were mixed with +blood. From November until March he had felt particularly ill, being +troubled with nausea and vomiting in addition to the pain in his +intestines. In March, his physician prescribed injections of a +concentrated solution of tannin, which resulted in the expulsion of +fifty living larvae of _Muscina stabulans_. Thereafter the patient felt +much better, although he suffered from intestinal catarrh in a less +severe form. + +[Illustration: 102 Muscina stabulans (x4). After Graham-Smith.] + + +CALLIPHORINAE--Closely related to the Sarcophagidae are the +_Calliphorinae_, to which group belong many of the so-called "blue +bottle" flies. Their larvae feed upon dead animals, and upon fresh and +cooked meat. Those of _Protocalliphora_, already mentioned, are +ectoparasitic on living nestling birds. Larva of _Lucilia_, we have +taken from tumors on living turtles. To this sub-family belongs also +_Auchmeromyia luteola_, the Congo floor maggot. Some of these, and at +least the last mentioned, are confirmed, rather than faculative +parasites. Various species of Calliphorinae are occasionally met with as +facultative parasites of man. + +[Illustration: 103. Lucilia caesar, (x3). After Howard.] + +_Chrysomyia macellaria_, the screw worm fly (fig. 107), is the fly which +is responsible for the most serious cases of human myasis in the United +States. It is widely distributed in the United States but is especially +abundant in the south. While the larvae breed in decaying matter in +general, they so commonly breed in the living flesh of animals that they +merit rank as true parasites. The females are attracted to open wounds +of all kinds on cattle and other animals and quickly deposit large +numbers of eggs. Animals which have been recently castrated, dehorned, +or branded, injured by barbed wire, or even by the attacks of ticks are +promptly attacked in the regions where the fly abounds. Even the navel +of young calves or discharges from the vulva of cows may attract the +insect. + +[Illustration: 104. Calliphora erythrocephala, (x6). After +Graham-Smith.] + +Not infrequently the fly attacks man, being attracted by an offensive +breath, a chronic catarrh, or a purulent discharge from the ears. Most +common are the cases where the eggs are deposited in the nostrils. The +larvae, which are hatched in a day or two, are provided with strong +spines and proceed to bore into the tissues of the nose, even down into +or through the bone, into the frontal sinus, the pharynx, larynx, and +neighboring parts. + +Osborn (1896) quotes a number of detailed accounts of the attacks of the +_Chrysomyia_ on man. A vivid picture of the symptomology of rhinal +myasis caused by the larvae of this fly is given by Castellani and +Chalmers: "Some couple of days after a person suffering from a chronic +catarrh, foul breath, or ozaena, has slept in the open or has been +attacked by a fly when riding or driving,--_i.e._, when the hands are +engaged--signs of severe catarrh appear, accompanied with inordinate +sneezing and severe pain over the root of the nose or the frontal bone. +Quickly the nose becomes swollen, and later the face also may swell, +while examination of the nose may show the presence of the larvae. Left +untreated, the patient rapidly becomes worse, and pus and blood are +discharged from the nose, from which an offensive odor issues. Cough +appears as well as fever, and often some delirium. If the patient lives +long enough, the septum of the nose may fall in, the soft and hard +palates may be pierced, the wall of the pharynx may be destroyed. By +this time, however, the course of the disease will have become quite +evident by the larvae dropping out of the nose, and if the patient +continues to live all the larvae may come away naturally." + +For treatment of rhinal myasis these writers recommend douching the nose +with chloroform water or a solution of chloroform in sweet milk (10-20 +per cent), followed by douches of mild antiseptics. Surgical treatment +may be necessary. + +[Illustration: 105. Larva of a flesh fly (Sarcophaga). Caudal aspect. +Anterior stigmata. Pharyngeal skeleton.] + + +SARCOPHAGIDAE--The larvae (fig. 105) of flies of this family usually +feed upon meats, but have been found in cheese, oleomargerine, pickled +herring, dead and living insects, cow dung and human feces. Certain +species are parasitic in insects. Higgins (1890) reported an instance of +"hundreds" of larvae of _Sarcophaga_ being vomited by a child eighteen +months of age. There was no doubt as to their origin for they were +voided while the physician was in the room. There are many other reports +of their occurrence in the alimentary canal. We have recorded elsewhere +(Riley, 1906) a case in which some ten or twelve larvae of +_Sarcophaga_ were found feeding on the diseased tissues of a malignant +tumor. The tumor, a melanotic sarcoma, was about the size of a small +walnut, and located in the small of the back of an elderly lady. +Although they had irritated and caused a slight haemorrhage, neither the +patient nor others of the family knew of their presence. Any discomfort +which they had caused had been attributed to the sarcomatous growth. The +infestation occurred in mid-summer. It is probable that the adult was +attracted by the odor of the discharges and deposited the living maggots +upon the diseased tissues. + +[Illustration: 106. A flesh fly (Sarcophaga), (x4). After Graham-Smith.] + +According to Kuchenmeister, _Sarcophaga carnaria_ (fig. 106), attracted +by the odor, deposits its eggs and larvae in the vagina of girls and +women when they lie naked in hot summer days upon dirty clothes, or when +they have a discharge from the vagina. In malignant inflammations of the +eyes the larvae even nestle under the eyelids and in Egypt, for example, +produce a very serious addition to the effects of small-pox upon the +cornea, as according to Pruner, in such cases perforation of the cornea +usually takes place. + +[Illustration: 107. Chrysomyia macellaria, (x3).] + +_Wohlfartia magnifica_ is another Sarcophagid which commonly infests man +in the regions where it is abundant. It is found in all Europe but is +especially common in Russia, where Portschinsky has devoted much +attention to its ravages. It deposits living larvae in wounds, the nasal +fossae, the ears and the eyes, causing injuries even more revolting than +those described for _Chrysomyia_. + + + + +CHAPTER V + +ARTHROPODS AS SIMPLE CARRIERS OF DISEASE + + +The fact that certain arthropods are poisonous, or may affect the health +of man as direct parasites has always received attention in the medical +literature. We come now to the more modern aspect of our subject,--the +consideration of insects and other arthropods as transmitters and +disseminators of disease. + +The simplest way in which arthropods may function in this capacity is as +_simple carriers_ of pathogenic organisms. It is conceivable that any +insect which has access to, and comes in contact with such organisms and +then passes to the food, or drink, or to the body of man, may in a +wholly accidental and incidental manner convey infection. That this +occurs is abundantly proved by the work of recent years. We shall +consider as typical the case against the house-fly, which has attracted +so much attention, both popular and scientific. The excellent general +treatises of Hewitt (1910), Howard (1911), and Graham-Smith (1913), and +the flood of bulletins and popular literature render it unnecessary to +consider the topic in any great detail. + + +THE HOUSE-FLY AS A CARRIER OF DISEASE + +Up to the past decade the house-fly has usually been regarded as a mere +pest. Repeatedly, however, it had been suggested that it might +disseminate disease. We have seen that as far back as the sixteenth +century, Mercurialis suggested that it was the agent in the spread of +bubonic plague, and in 1658, Kircher reiterated this view. In 1871, +Leidy expressed the opinion that flies were probably a means of +communicating contagious diseases to a greater degree than was generally +suspected. From what he had observed regarding gangrene in hospitals, he +thought flies should be carefully excluded from wounds. In the same +year, the editor of the _London Lancet_, referring to the belief that +they play a useful role in purifying the air said, "Far from looking +upon them as dipterous angels dancing attendance on Hygeia, regard them +rather in the light of winged sponges spreading hither and thither to +carry out the foul behests of Contagion." + +These suggestions attracted little attention from medical men, for it is +only within very recent years that the charges have been supported by +direct evidence. Before considering this evidence, it is necessary that +we define what is meant by "house-fly" and that we then consider the +life-history of the insect. + +There are many flies which are occasionally to be found in houses, but +according to various counts, from 95 per cent to 99 per cent of these in +warm weather in the Eastern United States belong to the one species +_Musca domestica_ (fig. 108). This is the dominant house-fly the world +over and is the one which merits the name. It has been well +characterized by Schiner (1864), whose description has been freely +translated by Hewitt, as follows: + +"Frons of male occupying a fourth part of the breadth of the head. +Frontal stripe of female narrow in front, so broad behind that it +entirely fills up the width of the frons. The dorsal region of the +thorax dusty grey in color with four equally broad longitudinal stripes. +Scutellum gray with black sides. The light regions of the abdomen +yellowish, transparent, the darkest parts at least at the base of the +ventral side yellow. The last segment and a dorsal line blackish brown. +Seen from behind and against the light, the whole abdomen shimmering +yellow, and only on each side of the dorsal line on each segment a dull +transverse band. The lower part of the face silky yellow, shot with +blackish brown. Median stripe velvety black. Antennae brown. Palpi black. +Legs blackish brown. Wings tinged with pale gray with yellowish base. +The female has a broad velvety back, often reddishly shimmering frontal +stripe, which is not broader at the anterior end than at the bases of +the antennae, but become so very much broader above that the light +dustiness of the sides is entirely obliterated. The abdomen gradually +becoming darker. The shimmering areas on the separate segments generally +brownish. All the other parts are the same as in the male." + +The other species of flies found in houses in the Eastern United States +which are frequently mistaken for the house or typhoid fly may readily +be distinguished by the characters of the following key: + +_a._ Apical cell (R_5) of the wide wing open, i.e., the bounding veins + parallel or divergent (fig. 100). Their larvae are flattened, the + intermediate body segments each fringed with fleshy, more or less + spinose, processes. _Fannia_ + + b. Male with the sides of the second and third abdominal segments + translucent yellowish. The larva with three pairs of nearly equal + spiniferous appendages on each segment, arranged in a + longitudinal series and in addition two pairs of series of smaller + processes (fig. 100) _F. canicularis_ + + bb. Male with blackish abdomen, middle tibia with a tubercle beyond + the middle. The larva with spiniferous appendages of which the + dorsal and ventral series are short, the lateral series long and + feathered (fig. 101) _F. scalaris_ + +aa. Apical cell (R) of the wing more or less narrowed in the margin; i. + e., the bounding veins more or less converging (fig. 108). + + b. The mouth-parts produced and pointed, fitted for piercing. + + c. Palpi much shorter than the proboscis; a brownish gray fly, its + thorax with three rather broad whitish stripes; on each border + of the middle stripe and on the mesal borders of the lateral + stripes is a blackish brown line. Abdomen yellowish brown; on + the second, third and fourth segments are three brown spots + which may be faint or even absent. The larvae live in dung. The + stable-fly (fig. 110) _Stomoxys calcitrans_ + + cc. Palpi nearly as long as the proboscis. Smaller species than the + house-fly. The horn-fly (fig. 167) _Haematobia irritans_ + + bb. Mouth-parts blunt, fitted for lapping. + + c. Thorax, particularly on the sides and near the base of the wings + with soft golden yellow hairs among the bristles. This fly is + often found in the house in very early spring or even in the + winter. The cluster-fly, _Pollenia rudis_ + + cc. Thorax without golden yellow hairs among the bristles. + + d. The last segment of the vein M with an abrupt angle. (fig. + 108). The larvae live in manure, etc. House-fly, _Musca + domestica_ + + dd. The last segment of vein M with a broad, gentle curve (fig. + 102). + + e. Eyes microscopically hairy; each abdominal segment with two + spots. Larvae in dung. _Myiospila meditabunda_ + + ee. Eyes bare; abdomen gray and brown marbled. _Muscina_ + + f. With black legs and palpi. _M. assimilis_ + + ff. With legs more or less yellowish; palpi yellow. Larvae in + decaying vegetable substances, dung, etc. _M. stabulans_ + +It is almost universally believed that the adults of _Musca domestica_ +hibernate, remaining dormant throughout the winter in attics, around +chimneys, and in sheltered but cold situations. This belief has been +challenged by Skinner (1913), who maintains that all the adult flies die +off during the fall and early winter and that the species is carried +over in the pupal stage, and in no other way. The cluster-fly, _Pollenia +rudis_, undoubtedly does hibernate in attics and similar situations and +is often mistaken for the house-fly. In so far as concerns _Musca +domestica_, the important question as to hibernation in the adult stage +is an open one. Many observations by one of the writers (Johannsen) tend +to confirm Dr. Skinner's conclusion, in so far as it applies to +conditions in the latitude of New York State. Opposed, is the fact that +various experimenters, notably Hewitt (1910) and Jepson (1909) wholly +failed to carry pupae through the winter. + +[Illustration: 108. The house or typhoid fly (Musca domestica (x4)). +After Howard.] + +The house-fly breeds by preference in horse manure. Indeed, Dr. Howard, +whose extensive studies of the species especially qualify him for +expressing an opinion on the subject, has estimated that under ordinary +city and town conditions, more than ninety per cent of the flies present +in houses have come from horse stables or their vicinity. They are not +limited to such localities, by any means, for it has been found that +they would develop in almost any fermenting organic substance. Thus, +they have been bred from pig, chicken, and cow manure, dirty waste +paper, decaying vegetation, decaying meat, slaughter-house refuse, +sawdust-sweepings, and many other sources. A fact which makes them +especially dangerous as disease-carriers is that they breed readily in +human excrement. + +The eggs are pure white, elongate ovoid, somewhat broader at the +anterior end. They measure about one millimeter (1-25 inch) in length. +They are deposited in small, irregular clusters, one hundred and twenty +to one hundred and fifty from a single fly. A female may deposit as many +as four batches in her life time. The eggs hatch in from eight to +twenty-four hours. + +The newly hatched larva, or maggot (fig. 108), measures about two +millimeters (1-12 inch) in length. It is pointed at the head end and +blunt at the opposite end, where the spiracular openings are borne. It +grows rapidly, molts three times and reaches maturity in from six to +seven days, under favorable conditions. + +The pupal stage, like that of related flies, is passed in the old larval +skin which, instead of being molted, becomes contracted and heavily +chitinized, forming the so-called _puparium_ (fig. 108). The pupal stage +may be completed in from three to six days. + +Thus during the warm summer months a generation of flies may be produced +in ten to twelve days. Hewitt at Manchester, England, found the minimum +to be eight days but states that larvae bred in the open air in horse +manure which had an average daily temperature of 22.5^o C., occupied +fourteen to twenty days in their development, according to the air +temperature. + +After emergence, a period of time must elapse before the fly is capable +of depositing eggs. This period has been tuned the _preoviposition_ +period. Unfortunately we have few exact data regarding this period. +Hewitt found that the flies became sexually mature in ten to fourteen +days after their emergence from the pupal state and four days after +copulation they began to deposit their eggs; in other words the +preoviposition stage was fourteen days or longer. Griffith (1908) found +this period to be ten days. Dr. Howard believes that the time "must +surely be shorter, and perhaps much shorter, under midsummer conditions, +and in the freedom of the open air." He emphasizes that the point is of +great practical importance, since it is during this period that the +trapping and other methods of destroying the adult flies, will prove +most useful. + +Howard estimates that there may be nine generations of flies a year +under outdoor conditions in places comparable in climate to Washington. +The number may be considerably increased in warmer climates. + +The rate at which flies may increase under favorable conditions is +astounding. Various writers have given estimates of the numbers of flies +which may develop as the progeny of a single individual, providing all +the eggs and all the individual flies survived. Thus, Howard estimates +that from a single female, depositing one hundred and twenty eggs on +April 15th, there may be by September 10th, 5,598,720,000,000 adults. +Fortunately, living forms do not produce in any such mathematical manner +and the chief value of the figures is to illustrate the enormous +struggle for existence which is constantly taking place in nature. + +Flies may travel for a considerable distance to reach food and shelter, +though normally they pass to dwellings and other sources of food supply +in the immediate neighborhood of their breeding places. Copeman, Howlett +and Merriman (1911) marked flies by shaking them in a bag containing +colored chalk. Such flies were repeatedly recovered at distances of +eight to one thousand yards and even at a distance of seventeen hundred +yards, nearly a mile. + +Hindle and Merriman (1914) continued these experiments on a large scale +at Cambridge, England. They "do not think it likely that, as a rule, +flies travel more than a quarter of a mile in thickly-housed areas." In +one case a single fly was recovered at a distance of 770 yards but a +part of this distance was across open fen-land. The surprising fact was +brought out that flies tend to travel either _against_ or across the +wind. The actual direction followed may be determined either directly by +the action of the wind (positive anemotropism), or indirectly owing to +the flies being attracted by any odor that it may convey from a source +of food. They conclude that it is likely that the chief conditions +favoring the disposal of flies are fine weather and a warm temperature. +The nature of the locality is another considerable factor. Hodge (1913) +has shown that when aided by the wind they may fly to much greater +distances over the water. He reports that at Cleveland, Ohio, the cribs +of the water works, situated a mile and a quarter, five miles, and six +miles out in Lake Erie are invaded by a regular plague of flies when the +wind blows from the city. Investigation showed that there was absolutely +nothing of any kind in which flies could breed on the crib. + +The omnivorous habits of the house-fly are matters of everyday +observation. From our view point, it is sufficient to emphasize that +from feeding on excrement, on sputum, on open sores, or on putrifying +matter, the flies may pass to the food or milk upon the table or to +healthy mucous membranes, or uncontaminated wounds. There is nothing in +its appearance to tell whether the fly that comes blithely to sup with +you is merely unclean, or whether it has just finished feeding upon +dejecta teeming with typhoid bacilli. + +[Illustration: 100. Pulvillus of foot of house-fly, showing glandular +hairs.] + +The method of feeding of the house-fly has an important bearing on the +question of its ability to transmit pathogenic organisms. Graham-Smith +(1910) has shown that when feeding, flies frequently moisten soluble +substances with "vomit" which is regurgitated from the crop. This is, of +course, loaded with bacteria from previous food. When not sucked up +again these drops of liquid dry, and produce round marks with an opaque +center and rim and an intervening less opaque area. Fly-specks, then, +consist of both vomit spots and feces. Graham-Smith shows a photograph +of a cupboard window where, on an area six inches square, there were +counted eleven hundred and two vomit marks and nine fecal deposits. + +From a bacteriologist's viewpoint a discussion of the possibility of a +fly's carrying bacteria would seem superfluous. Any exposed object, +animate or inanimate, is contaminated by bacteria and will transfer them +if brought into contact with suitable culture media, whether such +substance be food, or drink, open wounds, or the sterile culture media +of the laboratory. A needle point may convey enough germs to produce +disease. Much more readily may the house-fly with its covering of hairs +and its sponge-like pulvilli (fig. 109) pick up and transfer bits of +filth and other contaminated material. + +For popular instruction this inevitable transfer of germs by the +house-fly is strikingly demonstrated by the oft copied illustration of +the tracks of a fly on a sterile culture plate. Two plates of gelatine +or, better, agar medium are prepared. Over one of these a fly (with +wings clipped) is allowed to walk, the other is kept as a check. Both +are put aside at room temperature, to be examined after twenty-four to +forty-eight hours. At the end of that time, the check plate is as clear +as ever, the one which the fly has walked is dotted with colonies of +bacteria and fungi. The value in the experiment consists in emphasizing +that by this method we merely render visible what is constantly +occurring in nature. + +A comparable experiment which we use in our elementary laboratory work +is to take three samples of _clean_ (preferably, sterile) fresh milk in +sterile bottles. One of them is plugged with a pledget of cotton, into +the second is dropped a fly from the laboratory and into the third is +dropped a fly which has been caught feeding upon garbage or other filth. +After a minute or two the flies are removed and the vials plugged as was +number one. The three are then set aside at room temperature. When +examined after twenty-four hours the milk in the first vial is either +still sweet or has a "clean" sour odor; that of the remaining two is +very different, for it has a putrid odor, which is usually more +pronounced in the case of sample number three. + +Several workers have carried out experiments to determine the number of +bacteria carried by flies under natural conditions. One of the most +extended and best known of these is the series by Esten and Mason +(1908). These workers caught flies from various sources in a sterilized +net, placed them in a sterile bottle and poured over them a known +quantity of sterilized water, in which they were shaken so as to wash +the bacteria from their bodies. They found the number of bacteria on a +single fly to range from 550 to 6,600,000. Early in the fly season the +numbers of bacteria on flies are comparatively small, while later the +numbers are comparatively very large. The place where flies live also +determines largely the numbers that they carry. The lowest number, 550, +was from a fly caught in the bacteriological laboratory, the highest +number, 6,600,000 was the average from eighteen swill-barrel flies. +Torrey (1912) made examination of "wild" flies from a tenement house +district of New York City. He found "that the surface contamination of +these 'wild' flies may vary from 570 to 4,400,000 bacteria per insect, +and the intestinal bacterial content from 16,000 to 28,000,000." + +Less well known in this country is the work of Cox, Lewis, and Glynn +(1912). They examined over four hundred and fifty naturally infected +house-flies in Liverpool during September and early October. Instead of +washing the flies they were allowed to swim on the surface of sterile +water for five, fifteen, or thirty minutes, thus giving natural +conditions, where infection occurs from vomit and dejecta of the flies, +as well as from their bodies. They found, as might be expected, that +flies from either insanitary or congested areas of the city contain far +more bacteria than those from the more sanitary, less congested, or +suburban areas. The number of aerobic bacteria from the former varied +from 800,000 to 500,000,000 per fly and from the latter from 21,000 to +100,000. The number of intestinal forms conveyed by flies from +insanitary or congested areas was from 10,000 to 333,000,000 as compared +with from 100 to 10,000 carried by flies from the more sanitary areas. + +Pathogenic bacteria and those allied to the food poisoning group were +only obtained from the congested or moderately congested areas and not +from the suburban areas, where the chances of infestation were less. + +The interesting fact was brought out that flies caught in milk shops +apparently carry and obtain more bacteria than those from other shops +with exposed food in a similar neighborhood. The writers explained this +as probably due to the fact that milk when accessible, especially during +the summer months, is suitable culture medium for bacteria, and the +flies first inoculate the milk and later reinoculate themselves, and +then more of the milk, so establishing a vicious circle. + +They conclude that in cities where food is plentiful flies rarely +migrate from the locality in which they are bred, and consequently the +number of bacteria which they carry depends upon the general standard +of cleanliness in that locality. Flies caught in a street of modern, +fairly high class, workmen's dwellings forming a sanitary oasis in the +midst of a slum area, carried far less bacteria than those caught in the +adjacent neighborhood. + +Thus, as the amount of dirt carried by flies in any particular locality, +measured in the terms of bacteria, bears a definite relation to the +habits of the people and to the state of the streets, it demonstrates +the necessity of efficient municipal and domestic cleanliness, if the +food of the inhabitants is to escape pollution, not only with harmless +but also with occasional pathogenic bacteria. + +The above cited work is of a general nature, but, especially in recent +years, many attempts have been made to determine more specifically the +ability of flies to transmit pathogenic organisms. The critical reviews +of Nuttall and Jepson (1909), Howard (1911), and Graham-Smith (1913) +should be consulted by the student of the subject. We can only cite here +a few of the more striking experiments. + +Celli (1888) fed flies on pure cultures of _Bacillus typhosus_ and +declared that he was able to recover these organisms from the intestinal +contents and excrement. + +Firth and Horrocks (1902), cited by Nuttall and Jepson, "kept _Musca +domestica_ (also bluebottles) in a large box measuring 4 x 3 x 3 feet, +with one side made of glass. They were fed on material contaminated with +cultures of _B. typhosus_. Agar plates, litmus, glucose broth and a +sheet of clean paper were at the same time exposed in the box. After a +few days the plates and broth were removed and incubated with a positive +result." Graham-Smith (1910) "carried out experiments with large numbers +of flies kept in gauze cages and fed for eight hours on emulsions of _B. +typhosus_ in syrup. After that time the infested syrup was removed and +the flies were fed on plain syrup. _B. typhosus_ was isolated up to 48 +hours (but not later) from emulsions of their feces and from plates over +which they walked." + +Several other workers, notably Hamilton (1903), Ficker (1903), +Bertarelli (1910) Faichnie (1909), and Cochrane (1912), have isolated +_B. typhosus_ from "wild" flies, naturally infected. The papers of +Faichnie and of Cochrane we have not seen, but they are quoted in +_extenso_ by Graham-Smith (1913). + +On the whole, the evidence is conclusive that typhoid germs not only may +be accidentally carried on the bodies of house-flies but may pass +through their bodies and be scattered in a viable condition in the feces +of the fly for at least two days after feeding. Similar, results have +been reached in experiments with cholera, tuberculosis and yaws, the +last-mentioned being a spirochaete disease. Darling (1913) has shown that +murrina, a trypanosome disease of horses and mules in the Canal zone is +transmitted by house-flies which feed upon excoriated patches of +diseased animals and then pass to cuts and galls of healthy animals. + +Since it is clear that flies are abundantly able to disseminate viable +pathogenic bacteria, it is important to consider whether they have +access to such organisms in nature. A consideration of the method of +spread of typhoid will serve to illustrate the way in which flies may +play an important role. + +Typhoid fever is a specific disease caused by _Bacillus typhosus_, and +by it alone. The causative organism is to be found in the excrement and +urine of patients suffering from the disease. More than that, it is +often present in the dejecta for days, weeks, or even months and years, +after the individual has recovered from the disease. Individuals so +infested are known as "typhoid carriers" and they, together with those +suffering from mild cases, or "walking typhoid," are a constant menace +to the health of the community in which they are found. + +Human excrement is greedily visited by flies, both for feeding and for +ovipositing. The discharges of typhoid patients, or of chronic +"carriers," when passed in the open, in box privies, or camp latrines, +or the like, serve to contaminate myriads of the insects which may then +spread the germ to human food and drink. Other intestinal diseases may +be similarly spread. There is abundant epidaemiological evidence that +infantile diarrhoea, dysentery, and cholera may be so spread. + +Stiles and Keister (1913) have shown that spores of _Lamblia +intestinalis_, a flagellate protozoan living in the human intestine, may +be carried by house-flies. Though this species is not normally +pathogenic, one or more species of _Entamoeba_ are the cause of a type +of a highly fatal tropical dysentery. Concerning it, and another +protozoan parasite of man, they say, "If flies can carry _Lamblia_ +spores measuring 10 to 7u, and bacteria that are much smaller, and +particles of lime that are much larger, there is no ground to assume +that flies may not carry _Entamoeba_ and _Trichomonas_ spores." + +Tuberculosis is one of the diseases which it is quite conceivable may be +carried occasionally. The sputum of tubercular patients is very +attractive to flies, and various workers, notably Graham-Smith, have +found that _Musca domestica_ may distribute the bacillus for several +days after feeding on infected material. + +A type of purulent opthalmia which is very prevalent in Egypt is often +said to be carried by flies. Nuttall and Jepson (1909) consider that the +evidence regarding the spread of this disease by flies is conclusive and +that the possibility of gonorrhoeal secretions being likewise conveyed +cannot be denied. + +Many studies have been published, showing a marked agreement between the +occurrence of typhoid and other intestinal diseases and the prevalence +of house-flies. The most clear-cut of these are the studies of the Army +Commission appointed to investigate the cause of epidemics of enteric +fever in the volunteer camps in the Southern United States during the +Spanish-American War. Though their findings as presented by Vaughan +(1909), have been quoted very many times, they are so germane to our +discussion that they will bear repetition: + +"Flies swarmed over infected fecal matter in the pits and fed upon the +food prepared for the soldiers in the mess tents. In some instances +where lime had recently been sprinkled over the contents of the pits, +flies with their feet whitened with lime were seen walking over the +food." Under such conditions it is no wonder that "These pests had +inflicted greater loss upon American soldiers than the arms of Spain." + +Similar conditions prevailed in South Africa during the Boer War. Seamon +believes that very much of the success of the Japanese in their fight +against Russia was due to the rigid precautions taken to prevent the +spread of disease by these insects and other means. + +Veeder has pointed out that the characteristics of a typical fly-borne +epidemic of typhoid are that it occurs in little neighborhood epidemics, +extending by short leaps from house to house, without regard to water +supply or anything else in common. It tends to follow the direction of +prevailing winds (cf. the conclusions of Hindle and Merriman). It occurs +during warm weather. Of course, when the epidemic is once well under +way, other factors enter into its spread. + +In general, flies may be said to be the chief agency in the spread of +typhoid in villages and camps. In cities with modern sewer systems they +are less important, though even under the best of such conditions, they +are important factors. Howard has emphasized that in such cities there +are still many uncared-for box privies and that, in addition, the +deposition of feces overnight in uncared-for waste lots and alleys is +common. + +Not only unicellular organisms, such as bacteria and protozoa, but also +the eggs, embryos and larvae of parasitic worms have been found to be +transported by house-flies. Ransom (1911) has found that _Habronema +muscae_, a nematode worm often found in adult flies, is the immature +stage of a parasite occurring in the stomach of the horse. The eggs or +embryos passing out with the feces of the horse, are taken up by fly +larvae and carried over to the imago stage. + +Grassi (1883), Stiles (1889), Calandruccio (1906), and especially Nicoll +(1911), have been the chief investigators of the ability of house-flies +to carry the ova and embryos of human intestinal parasites. Graham-Smith +(1913) summarizes the work along this line as follows: + +"It is evident from the investigations that have been quoted that +house-flies and other species are greatly attracted to the ova of +parasitic worms contained in feces and other materials, and make great +efforts to ingest them. Unless the ova are too large they often succeed, +and the eggs are deposited uninjured in their feces, in some cases up to +the third day at least. The eggs may also be carried on their legs or +bodies. Under suitable conditions, food and fluids may be contaminated +with the eggs of various parasitic worms by flies, and in one case +infection of the human subject has been observed. Feces containing +tape-worm segments may continue to be a source of infection for as long +as a fortnight. Up to the present, however, there is no evidence to show +what part flies play in the dissemination of parasitic worms under +natural conditions." + +Enough has been said to show that the house-fly must be dealt with as a +direct menace to public health. Control measures are not merely matters +of convenience but are of vital importance. + +Under present conditions the speedy elimination of the house-fly is +impossible and the first thing to be considered is methods of protecting +food and drink from contamination. The first of these methods is the +thorough screening of doors and windows to prevent the entrance of +flies. In the case of kitchen doors, the flies, attracted by odors, are +likely to swarm onto the screen and improve the first opportunity for +gaining an entrance. This difficulty can be largely avoided by +screening-in the back porch and placing the screen door at one end +rather than directly before the door. + +The use of sticky fly paper to catch the pests that gain entrance to the +house is preferable to the various poisons often used. Of the latter, +formalin (40 per cent formaldehyde) in the proportion of two +tablespoonfuls to a pint of water is very efficient, if all other +liquids are removed or covered, so that the flies must depend on the +formalin for drink. The mixture is said to be made more attractive by +the addition of sugar or milk, though we have found the plain solution +wholly satisfactory, under proper conditions. It should be emphasized +that this formalin mixture is not perfectly harmless, as so often +stated. There are on record cases of severe and even fatal poisoning +from the accidental drinking of solutions. + +When flies are very abundant in a room they can be most readily gotten +rid of by fumigation with sulphur, or by the use of pure pyrethrum +powder either burned or puffed into the air. Herrick (1913) recommends +the following method: "At night all the doors and windows of the kitchen +should be closed; fresh powder should be sprinkled over the stove, on +the window ledges, tables, and in the air. In the morning flies will be +found lying around dead or stupified. They may then be swept up and +burned." This method has proved very efficaceous in some of the large +dining halls in Ithaca. + +The writers have had little success in fumigating with the vapors of +carbolic acid, or carbolic acid and gum camphor, although these methods +will aid in driving flies from a darkened room. + +All of these methods are but makeshifts. As Howard has so well put it, +"the truest and simplest way of attacking the fly problem is to prevent +them from breeding, by the treatment or abolition of all places in which +they can breed. To permit them to breed undisturbed and in countless +numbers, and to devote all our energy to the problem of keeping them out +of our dwellings, or to destroy them after they have once entered in +spite of all obstacles, seems the wrong way to go about it." + +We have already seen that _Musca domestica_ breeds in almost any +fermenting organic material. While it prefers horse manure, it breeds +also in human feces, cow dung and that of other animals, and in refuse +of many kinds. To efficiently combat the insect, these breeding places +must be removed or must be treated in some such way as to render them +unsuitable for the development of the larvae. Under some conditions +individual work may prove effective, but to be truly efficient there +must be extensive and thorough cooperative efforts. + +Manure, garbage, and the like should be stored in tight receptacles and +carted away at least once a week. The manure may be carted to the fields +and spread. Even in spread manure the larvae may continue their +development. Howard points out that "it often happens that after a lawn +has been heavily manured in early summer the occupants of the house will +be pestered with flies for a time, but finding no available breeding +place these disappear sooner or later. Another generation will not breed +in the spread manure." + +Hutchinson (1914) has emphasized that the larvae of houseflies have +deeply engrained the habit of migrating in the prepupal stage and has +shown that this offers an important point of attack in attempts to +control the pest. He has suggested that maggot traps might be developed +into an efficient weapon in the warfare against the house-fly. Certain +it is that the habit greatly simplifies the problem of treating the +manure for the purpose of killing the larvae. + +There have been many attempts to find some cheap chemical which would +destroy fly larvae in horse manure without injuring the bacteria or +reducing the fertilizing values of the manure. The literature abounds in +recommendations of kerosene, lime, chloride of lime, iron sulphate, and +other substances, but none of them has met the situation. The whole +question has been gone into thoroughly by Cook, Hutchinson and Scales +(1914), who tested practically all of the substances which have been +recommended. They find that by far the most effective, economical, and +practical of the substances is borax in the commercial form in which it +is available throughout the country. + +"Borax increases the water-soluble nitrogen, ammonia and alkalinity of +manure and apparently does not permanently injure the bacterial flora. +The application of manure treated with borax at the rate of 0.62 pound +per eight bushels (10 cubic feet) to soil does not injure the plants +thus far tested, although its cumulative effect, if any, has not been +determined." + +As their results clearly show that the substances so often recommended +are inferior to borax, we shall quote in detail their directions for +treating manure so as to kill fly eggs and maggots. + +"Apply 0.62 pound borax or 0.75 pound calcined colemanite to every 10 +cubic feet (8 bushels) of manure immediately on its removal from the +barn. Apply the borax particularly around the outer edges of the pile +with a flour sifter or any fine sieve, and sprinkle two or three gallons +of water over the borax-treated manure. + +"The reason for applying the borax to the fresh manure immediately after +its removal from the stable is that the flies lay their eggs on the +fresh manure, and borax, when it comes in contact with the eggs, +prevents their hatching. As the maggots congregate at the outer edge of +the pile, most of the borax should be applied there. The treatment +should be repeated with each addition of fresh manure, but when the +manure is kept in closed boxes, less frequent applications will be +sufficient. When the calcined colemanite is available, it may be used at +the rate of 0.75 pound per 10 cubic feet of manure, and is a cheaper +means of killing the maggots. In addition to the application of borax to +horse manure to kill fly larvae, it may be applied in the same proportion +to other manures, as well as to refuse and garbage. Borax may also be +applied to the floors and crevices in barns, stables, markets, etc., as +well as to street sweepings, and water should be added as in the +treatment of horse manure. After estimating the amount of material to be +treated and weighing the necessary amount of borax, a measure may be +used which will hold the proper amount, thus avoiding the subsequent +weighings. + +"While it can be safely stated that no injurious action will follow the +application of manure treated with borax at the rate of 0.62 pound for +eight bushels, or even larger amounts in the case of some plants, +nevertheless the borax-treated manure has not been studied in connection +with the growth of all crops, nor has its cumulative effect been +determined. It is therefore recommended that not more than 15 tons per +acre of the borax-treated manure should be applied to the field. As +truckmen use considerably more than this amount, it is suggested that +all cars containing borax-treated manure be so marked, and that +public-health officials stipulate in their directions for this treatment +that not over 0.62 pound for eight bushels of manure be used, as it has +been shown that larger amounts of borax will injure most plants. It is +also recommended that all public-health officials and others, in +recommending the borax treatment for killing fly eggs and maggots in +manure, warn the public against the injurious effects of large amounts +of borax on the growth of plants." + +"The amount of manure from a horse varies with the straw or other +bedding used, but 12 or 15 bushels per week represent the approximate +amount obtained. As borax costs from five to six cents per pound in +100-pound lots in Washington, it will make the cost of the borax +practically one cent per horse, per day. And if calcined colemanite is +purchased in large shipments the cost should be considerably less." + +Hodge (1910) has approached the problem of fly extermination from +another viewpoint. He believes that it is practical to trap flies out of +doors during the preoviposition period, when they are sexually immature, +and to destroy such numbers of them that the comparatively few which +survive will not be able to lay eggs in sufficient numbers to make the +next generation a nuisance. To the end of capturing them in enormous +numbers he has devised traps to be fitted over garbage cans, into stable +windows, and connected with the kitchen window screens. Under some +conditions this method of attack has proved very satisfactory. + +One of the most important measures for preventing the spread of disease +by flies is the abolition of the common box privy. In villages and rural +districts this is today almost the only type to be found. It is the +chief factor in the spread of typhoid and other intestinal diseases, as +well as intestinal parasites. Open and exposed to myriads of flies which +not only breed there but which feed upon the excrement, they furnish +ideal conditions for spreading contamination. Even where efforts are +made to cover the contents with dust, or ashes, or lime, flies may +continue to breed unchecked. Stiles and Gardner have shown that +house-flies buried in a screened stand-pipe forty-eight inches under +sterile sand came to the surface. Other flies of undetermined species +struggled up through seventy-two inches of sand. + +So great is the menace of the ordinary box privy that a number of +inexpensive and simple sanitary privies have been designed for use where +there are not modern sewer systems. Stiles and Lumsden (1911) have given +minute directions for the construction of one of the best types, and +their bulletin should be obtained by those interested. + +Another precaution which is of fundamental importance in preventing the +spread of typhoid, is that of disinfecting all discharges from patients +suffering with the disease. For this purpose, quick-lime is the cheapest +and is wholly satisfactory. In chamber vessels it should be used in a +quantity equal to that of the discharge to be treated. It should be +allowed to act for two hours. Air-slaked lime is of no value whatever. +Chloride of lime, carbolic acid, or formalin may be used, but are more +expensive. Other intestinal diseases demand similar precautions. + + +STOMOXYS CALCITRANS, THE STABLE-FLY--It is a popular belief that +house-flies bite more viciously just before a rain. As a matter of +fact, the true house-flies never bite, for their mouth-parts are not +fitted for piercing. The basis of the misconception is the fact that a +true biting fly, _Stomoxys calcitrans_ (fig. 110), closely resembling +the house-fly, is frequently found in houses and may be driven in in +greater numbers by muggy weather. From its usual habitat this fly is +known as the "stable-fly" or, sometimes as the "biting house-fly." + +_Stomoxys calcitrans_ may be separated from the house-fly by the use of +the key on p. 145. It may be more fully characterized as follows: + +The eyes of the male are separated by a distance equal to one-fourth of +the diameter of the head, in the female by one-third. The frontal stripe +is black, the cheeks and margins of the orbits silvery-white. The +antennae are black, the arista feathered on the upper side only. The +proboscis is black, slender, fitted for piercing and projects forward in +front of the head. The thorax is grayish, marked by four conspicuous, +more or less complete black longitudinal stripes; the scutellum is +paler; the macrochaetae are black. The abdomen is gray, dorsally with +three brown spots on the second and third segments and a median spot on +the fourth. These spots are more pronounced in the female. The legs are +black, the pulvilli distinct. The wings are hyaline, the vein M_{1+2} +less sharply curved than in the house-fly, the apical cell being thus +more widely open (cf. fig. 110). Length 7 mm. + +[Illustration: 110. Stomoxys calcitrans; adult, larva, puparium and +details, (x5). After Howard.] + +This fly is widely distributed, being found the world over. It was +probably introduced into the United States, but has spread to all parts +of the country. Bishopp (1913) regards it as of much more importance as +a pest of domestic animals in the grain belt than elsewhere in the +United States. The life-history and habits of this species have assumed +a new significance since it has been suggested that it may transmit the +human diseases, infantile paralysis and pellagra. In this country, the +most detailed study of the fly is that of Bishopp (1913) whose data +regarding the life cycle are as follows: + +The eggs like those of the house-fly, are about one mm. in length. Under +a magnifying glass they show a distinct furrow along one side. When +placed on any moist substance they hatch in from one to three days after +being deposited. + +The larva or maggots (fig. 110) have the typical shape and actions of +most maggots of the Muscid group. They can be distinguished from those +of the house-fly as the stigma-plates are smaller, much further apart, +with the slits less sinuous. Development takes place fairly rapidly when +the proper food conditions are available and the growth is completed +within eleven to thirty or more days. + +The pupa (fig. 110), like that of related flies, undergoes its +development within the contracted and hardened last larval skin, or +puparium. This is elongate oval, slightly thicker towards the head end, +and one-sixth to one-fourth of an inch in length. The pupal stage +requires six to twenty days, or in cool weather considerably longer. + +The life-cycle of the stable-fly is therefore considerably longer than +that of _Musca domestica_. Bishopp found that complete development might +be undergone in nineteen days, but that the average period was somewhat +longer, ranging from twenty-one to twenty-five days, where conditions +are very favorable. The longest period which he observed was forty-three +days, though his finding of full grown larvae and pupae in straw during +the latter part of March, in Northern Texas, showed that development may +require about three months, as he considered that these stages almost +certainly developed from eggs deposited the previous December. + +The favorite breeding place, where available, seems to be straw or +manure mixed with straw. It also breeds in great numbers in +horse-manure, in company with _Musca domestica_. + +Newstead considers that in England the stable-fly hibernates in the +pupal stage. Bishopp finds that in the southern part of the United +States there is no true hibernation, as the adults have been found to +emerge at various times during the winter. He believes that in the +northern United States the winter is normally passed in the larval and +pupal stages, and that the adults which have been observed in heated +stables in the dead of winter were bred out in refuse within the warm +barns and were not hibernating adults. + +Graham-Smith (1913) states that although the stable-fly frequents stable +manure, it is probably not an important agent in distributing the +organisms of intestinal diseases. Bishopp makes the important +observation that "it has never been found breeding in human excrement +and does not frequent malodorous places, which are so attractive to the +house-fly. Hence it is much less likely to carry typhoid and other germs +which may be found in such places." + +Questions of the possible agency of _Stomoxys calcitrans_ in the +transmission of infantile paralysis and of pellagra, we shall consider +later. + + +OTHER ARTHROPODS WHICH MAY SERVE AS SIMPLE CARRIERS OF PATHOGENIC +ORGANISMS--It should be again emphasized that any insect which has +access to, and comes in contact with, pathogenic organisms and then +passes to the food, or drink, or the body of man, may serve as a simple +carrier of disease. In addition to the more obvious illustrations, an +interesting one is the previously cited case of the transfer of +_Dermatobia cyaniventris_ by a mosquito (fig. 81-84). Darling (1913) has +shown that in the tropics, the omnipresent ants may be important factors +in the spread of disease. + + + + +CHAPTER VI + +ARTHROPODS AS DIRECT INOCULATORS OF DISEASE GERMS + + +We have seen that any insect which, like the house-fly, has access to +disease germs and then comes into contact with the food or drink of man, +may serve to disseminate disease. Moreover, it has been clearly +established that a contaminated insect, alighting upon wounded or +abraded surfaces, may infect them. These are instances of mere +accidental, mechanical transfer of pathogenic organisms. + +Closely related are the instances of direct inoculation of disease germs +by insects and other arthropods. In this type, a blood-sucking species +not only takes up the germs but, passing to a healthy individual, it +inserts its contaminated mouth-parts and thus directly inoculates its +victim. In other words, the disease is transferred just as blood +poisoning may be induced by the prick of a contaminated needle, or as +the laboratory worker may inoculate an experimental animal. + +Formerly, it was supposed that this method of the transfer of disease by +arthropods was a very common one and many instances are cited in the +earlier literature of the subject. It is, however, difficult to draw a +sharp line between such cases and those in which, on the one hand, the +arthropod serves as a mere passive carrier or, on the other hand, serves +as an essential host of the pathogenic organism. More critical study of +the subject has led to the belief that the importance of the role of +arthropods as direct inoculators has been much overestimated. + +The principal reason for regarding this phase of the subject as +relatively unimportant, is derived from a study of the habits of the +blood-sucking species. It is found that, in general, they are +intermittent feeders, visiting their hosts at intervals and then +abstaining from feeding for a more or less extended period, while +digesting their meal. In the meantime, most species of bacteria or of +protozoan parasites with which they might have contaminated their +mouth-parts, would have perished, through inability to withstand drying. + +In spite of this, it must be recognized that this method of transfer +does occur and must be reckoned with in any consideration of the +relations of insects to disease. We shall first cite some general +illustrations and shall then discuss the role of fleas in the spreading +of bubonic plague, an illustration which cannot be regarded as typical, +since it involves more than mere passive carriage. + + +SOME ILLUSTRATIONS OF DIRECT INOCULATION OF DISEASE GERMS BY ARTHROPODS + +In discussing poisonous arthropods, we have already emphasized that +species which are of themselves innocuous to man, may occasionally +introduce bacteria by their bite or sting and thus cause more or less +severe secondary symptoms. That such cases should occur, is no more than +is to be expected. The mouth-parts or the sting of the insect are not +sterile and the chances of their carrying pyogenic organisms are always +present. + +More strictly falling in the category of transmission of disease germs +by direct inoculation are the instances where the insect, or related +form, feeds upon a diseased animal and passes promptly to a healthy +individual which it infects. Of such a nature are the following: + +Various species of biting flies are factors in the dissemination of +anthrax, an infectious and usually fatal disease of animals and, +occasionally, of man. That the bacteria with which the blood of diseased +animals teem shortly before death might be transmitted by such insects +has long been contended, but the evidence in support of the view has +been unsatisfactory. Recently, Mitzmain (1914) has reported a series of +experiments which show conclusively that the disease may be so conveyed +by a horse-fly, _Tabanus striatus_, and by the stable-fly, _Stomoxys +calcitrans_. + +Mitzmain's experiments were tried with an artificially infected guinea +pig, which died of the disease upon the third day. The flies were +applied two and one-half hours, to a few minutes, before the death of +the animal. With both species the infection was successfully transferred +to healthy guinea pigs by the direct method, in which the flies were +interrupted while feeding on the sick animal. The evidence at hand does +not warrant the conclusion that insect transmission is the rule in the +case of this disease. + +The nagana, or tsetse-fly disease of cattle is the most virulent disease +of domestic animals in certain parts of Africa. It is caused by a +protozoan blood parasite, _Trypanosoma brucei_, which is conveyed to +healthy animals by the bite of _Glossina morsitans_ and possibly other +species of tsetse-flies. The flies remain infective for forty-eight +hours after feeding on a diseased animal. The insect also serves as an +essential host of the parasite. + +Surra, a similar trypanosomiasis affecting especially horses and mules, +occurs in southern Asia, Malaysia, and the Philippines where the +tsetse-flies are not to be found. It is thought to be spread by various +species of blood-sucking flies belonging to the genera _Stomoxys_, +_Haematobia_, and _Tabanus_. Mitzmain (1913) demonstrated that in the +Philippines it is conveyed mechanically by _Tabanus striatus_. + +The sleeping sickness of man, in Africa, has also been supposed to be +directly inoculated by one, or several, species of tsetse-flies. It is +now known that the fly may convey the disease for a short time after +feeding, but that there is then a latent period of from fourteen to +twenty-one days, after which it again becomes infectious. This indicates +that in the meantime the parasite has been undergoing some phase of its +life-cycle and that the fly serves as an intermediate host. We shall +therefore consider it more fully under that grouping. + +These are a few of the cases of direct inoculation which may be cited as +of the simpler type. We shall next consider the role of the flea in the +dissemination of the bubonic plague, an illustration complicated by the +fact that the bacillus multiples within the insect and may be indirectly +inoculated. + + +THE ROLE OF FLEAS IN THE TRANSMISSION OF THE PLAGUE + +The plague is a specific infectious disease caused by _Bacillus pestis_. +It occurs in several forms, of which the bubonic and the pneumonic are +the most common. According to Wyman, 80 per cent of the human cases are +of the bubonic type. It is a disease which, under the name of oriental +plague, the pest, or the black death, has ravaged almost from time +immemorial the countries of Africa, Asia, and Europe. The record of its +ravages are almost beyond belief. In 542 A. D. it caused in one day ten +thousand deaths in Constantinople. In the 14th century it was introduced +from the East and prevailed throughout Armenia, Asia Minor, Egypt and +Northern Africa and Europe. Hecker estimates that one-fourth of the +population of Europe, or twenty-five million persons, died in the +epidemic of that century. From then until the 17th century it was almost +constantly present in Europe, the great plague of London, in 1665 +killing 68,596 out of a population of 460,000. Such an epidemic would +mean for New York City a proportionate loss of over 600,000 in a single +year. It is little wonder that in the face of such an appalling disaster +suspicion and credulity were rife and the wildest demoralization ensued. + +During the 14th century the Jews were regarded as responsible for the +disease, through poisoning wells, and were subjected to the most +incredible persecution and torture. In Milan the visitation of 1630 was +credited to the so-called anointers,--men who were supposed to spread +the plague by anointing the walls with magic ointment--and the most +horrible tortures that human ingenuity could devise were imposed on +scores of victims, regardless of rank or of public service (fig. 112, +a). Manzoni's great historical novel, "The Betrothed" has well pictured +conditions in Italy during this period. + +[Illustration: 111. A contemporaneous engraving of the pest hospital in +Vienna in 1679. After Peters.] + +In modern times the plague is confined primarily to warm climates, a +condition which has been brought about largely through general +improvement in sanitary conditions. + +At present, the hotbed of the disease is India, where there were +1,040,429 deaths in 1904 and where in a period of fifteen years, ending +with January 1912, there were over 15,000,000 deaths. The reported +deaths in that country for 1913 totaled 198,875. + +During the winter of 1910-11 there occurred in Manchuria and North China +a virulent epidemic of the pneumonic plague which caused the death of +nearly 50,000 people. The question as to its origin and means of spread +will be especially referred to later. + +[Illustration: 112 a. A medieval method of combating the plague. The +persecution of the anointers in Milan in 1630. From a copy of "Il +processi originale degli untori" in the library of Cornell University.] + +Until recent years, the plague had not been known to occur in the New +World but there were outbreaks in Brazil and Hawaii in 1899, and in 1900 +there occurred the first cases in San Francisco. In California there +were 125 cases in the period 1900-04; three cases in the next three +years and then from May 1907 to March 1908, during the height of the +outbreak, 170 cases. Since that time there have been only sporadic +cases, the last case reported being in May 1914. Still more recent were +the outbreaks in the Philippine Islands, Porto Rico, and Cuba. + +On June 24, 1914, there was recognized a case of human plague in New +Orleans. The Federal Health Service immediately took charge, and +measures for the eradication of the disease were vigorously enforced. Up +to October 10, 1914 there had been reported 30 cases of the disease in +man, and 181 cases of plague in rats. + +[Illustration: 112 b. The modern method of combating the plague. A day's +catch of rats in the fight against plague in San Francisco. Courtesy of +Review of Reviews.] + +The present-day methods of combating bubonic plague are well illustrated +by the fight in San Francisco. Had it not been for the strenuous and +radical anti-plague campaign directed by the United States Marine +Hospital Service we might have had in our own country an illustration of +what the disease can accomplish. On what newly acquired knowledge was +this fight based? + +The basis was laid in 1894, when the plague bacillus was first +discovered. All through the centuries, before and during the Christian +era, down to 1894, the subject was enveloped in darkness and there had +been a helpless, almost hopeless struggle in ignorance on the part of +physicians, sanitarians, and public health officials against the ravages +of this dread disease. Now its cause, method of propagation and means to +prevent its spread are matters of scientific certainty. + +After the discovery of the causative organism, one of the first advances +was the establishment of the identity of human plague and that of +rodents. It had often been noted that epidemics of the human disease +were preceded by great epizootics among rats and mice. So well +established was this fact that with the Chinese, unusual mortality among +these rodents was regarded as foretelling a visitation of the human +disease. That there was more than an accidental connection between the +two was obvious when Yersin, the discoverer of _Bacillus pestis_, +announced that during an epidemic the rats found dead in the houses and +in the streets almost always contain the bacillus in great abundance in +their organs, and that many of them exhibit veritable buboes. + +Once it was established that the diseases were identical, the attention +of the investigators was directed to a study of the relations between +that of rats and of humans, and evidence accumulated to show that the +bubonic plague was primarily a disease of rodents and that in some +manner it was conveyed from them to man. + +There yet remained unexplained the method of transfer from rat to man. +As long ago as the 16th century, Mercuralis suggested that house-flies +were guilty of disseminating the plague but modern investigation, while +blaming the fly for much in the way of spreading disease, show that it +is an insignificant factor in this case. + +Search for blood-sucking insects which would feed on both rodents and +man, and which might therefore be implicated, indicated that the fleas +most nearly met the conditions. At first it was urged that rat fleas +would not feed upon man and that the fleas ordinarily attacking man +would not feed upon rats. More critical study of the habits of fleas +soon showed that these objections were not well-founded. Especially +important was the evidence that soon after the death of their host, rat +fleas deserted its body and might then become a pest in houses where +they had not been noticed before. + +Attention was directed to the fact that while feeding, fleas are in the +habit of squirting blood from the anus and that in the case of those +which had fed upon rats and mice dying of the plague, virulent plague +bacilli were to be found in such blood. Liston (1905) even found, and +subsequent investigations confirmed, that the plague bacilli multiply in +the stomach of the insect and that thus the blood ejected was richer in +the organisms than was that of the diseased animal. It was found that a +film of this infected blood spread out under the body of the flea and +that thus the bacilli might be inoculated by the bite of the insect and +by scratching. + +Very recently, Bacot and Martin (1914) have paid especial attention to +the question of the mechanism of the transmission of the plague bacilli +by fleas. They believe that plague infested fleas regurgitate blood +through the mouth, and that under conditions precluding the possibility +of infection by dejecta, the disease may be thus transmitted. The +evidence does not seem sufficient to establish that this is the chief +method of transmission. + +Conclusive experimental proof that fleas transmit the disease is further +available from a number of sources. The most extensive series of +experiments is that of the English Plague Commission in India, which +reported in 1906 that: + +On thirty occasions a healthy rat contracted plague in sequence of +living in the neighborhood of a plague infected rat under circumstances +which prevented the healthy rat coming in contact with either the body +or excreta of the diseased animal. + +In twenty-one experiments out of thirty-eight, healthy rats living in +flea-proof cages contracted plague when exposed to rat fleas +(_Xenopsylla cheopis_), collected from rats dead or dying of septicaemic +plague. + +Close contact of plague-infected with healthy animals, if fleas are +excluded, does not give rise to an epizootic among the latter. As the +huts were never cleaned out, close contact included contact with feces +and urine of infected animals, and contact with, and eating of food +contaminated with feces and urine of infected animals, as well as pus +from open plague ulcers. Close contact of young, even when suckled by +plague-infected mothers, did not give the disease to the former. + +If fleas are present, then the epizootic, once started, spreads from +animal to animal, the rate of progress being in direct proportion to the +number of fleas. + +Aerial infection was excluded. Thus guinea-pigs suspended in a cage two +feet above the ground did not contract the disease, while in the same +hut those animals allowed to run about and those placed two inches above +the floor became infected. It had previously been found that a rat flea +could not hop farther than about five inches. + +Guinea pigs and monkeys were placed in plague houses in pairs, both +protected from soil contact infection and both equally exposed to aerial +infection, but one surrounded with a layer of tangle-foot paper and the +other surrounded with a layer of sand. The following observations were +made: + +(_a_) Many fleas were caught in the tangle-foot, a certain proportion of +which were found on dissection to contain in their stomachs abundant +bacilli microscopically identical with plague bacilli. Out of +eighty-five human fleas dissected only one contained these bacilli, +while out of seventy-seven rat fleas twenty-three were found thus +infected. + +(_b_) The animals surrounded with tangle-foot in no instance developed +plague, while several (24 per cent) of the non-protected animals died of +the disease. + +Thus, the experimental evidence that fleas transmit the plague from rat +to rat, from rats to guinea pigs, and from rats to monkeys is +indisputable. There is lacking direct experimental proof of its transfer +from rodents to man but the whole chain of indirect evidence is so +complete that there can be no doubt that such a transfer does occur so +commonly that in the case of bubonic plague it must be regarded as the +normal method. + +Rats are not the only animals naturally attacked by the plague but as +already suggested, it occurs in various other rodents. In California the +disease has spread from rats to ground squirrels (_Otospermophilus +beecheyi_), a condition readily arising from the frequency of +association of rats with the squirrels in the neighborhood of towns, and +from the fact that the two species of fleas found on them are also found +on rats. While the danger of the disease being conveyed from squirrels +to man is comparatively slight, the menace in the situation is that the +squirrels may become a more or less permanent reservoir of the disease +and infect rats, which may come into more frequent contact with man. + +The tarbagan (_Arctomys bobac_), is a rodent found in North Manchuria, +which is much prized for its fur. It is claimed that this animal is +extremely susceptible to the plague and there is evidence to indicate +that it was the primary source of the great outbreak of pneumonic plague +which occurred in Manchuria and North China during the winter of +1910-11. + +Of fleas, any species which attacks both rodents and man may be an agent +in the transmission of the plague. We have seen that in India the +species most commonly implicated is the rat flea, _Xenopsylla cheopis_, +(= _Loemopsylla_ or _Pulex cheopis_) (fig. 89). This species has also +been found commonly on rats in San Francisco. The cat flea, +_Ctenocephalus felis_, the dog flea, _Ctenocephalus canis_, the human +flea, _Pulex irritans_, the rat fleas, _Ceratophyllus fasciatus_ and +_Ctenopsyllus musculi_ have all been shown to meet the conditions. + +But, however clear the evidence that fleas are the most important agent +in the transfer of plague, it is a mistake fraught with danger to assume +that they are the only factor in the spread of the disease. The +causative organism is a bacillus and is not dependent upon any insect +for the completion of its development. + +Therefore, any blood-sucking insect which feeds upon a plague infected +man or animal and then passes to a healthy individual, conceivably might +transfer the bacilli. Verjbitski (1908) has shown experimentally that +bed-bugs may thus convey the disease. Hertzog found the bacilli in a +head-louse, _Pediculus humanus_, taken from a child which had died from +the plague, and McCoy found them in a louse taken from a plague-infected +squirrel. On account of their stationary habits, the latter insects +could be of little significance in spreading the disease. + +Contaminated food may also be a source of danger. While this source, +formerly supposed to be the principal one, is now regarded as +unimportant, there is abundant experimental evidence to show that it +cannot be disregarded. It is believed that infection in this way can +occur only when there is some lesion in the alimentary canal. + +Still more important is the proof that in pneumonic plague the patient +is directly infective and that the disease is spread from man to man +without any intermediary. Especially conclusive is the evidence obtained +by Drs. Strong and Teague during the Manchurian epidemic of 1910-11. +They found that during coughing, in pneumonic plague cases, even when +sputum visible to the naked eye is not expelled, plague bacilli in large +numbers may become widely disseminated into the surrounding air. By +exposing sterile plates before patients who coughed a single time, very +numerous colonies of the bacillus were obtained. + +But the great advance which has been made rests on the discovery that +bubonic plague is in the vast majority of cases transmitted by the flea. +The pneumonic type forms a very small percentage of the human cases and +even with it, the evidence indicates that the original infection is +derived from a rodent through the intermediary of the insect. + +So modern prophylactic measures are directed primarily against the rat +and fleas. Ships coming from infected ports are no longer disinfected +for the purpose of killing the plague germs, but are fumigated to +destroy the rats and the fleas which they might harbor. When anchored at +infected ports, ships must observe strenuous precautions to prevent the +ingress of rats. Cargo must be inspected just before being brought on +board, in order to insure its freedom from rats. Even lines and hawsers +must be protected by large metal discs or funnels, for rats readily run +along a rope to reach the ship. Once infested, the ship must be +thoroughly fumigated, not only to avoid carrying the disease to other +ports but to obviate an outbreak on board. + +When an epidemic begins, rats must be destroyed by trapping and +poisoning. Various so-called biological poisons have not proved +practicable. Sources of food supply should be cut off by thorough +cleaning up, by use of rat-proof garbage cans and similar measures. Hand +in hand with these, must go the destruction of breeding places, and the +rat-proofing of dwellings, stables, markets, warehouses, docks and +sewers. All these measures are expensive, and a few years ago would have +been thought wholly impossible to put into practice but now they are +being enforced on a large scale in every fight against the disease. + +Rats and other rodents are regularly caught in the danger zone and +examined for evidence of infection, for the sequence of the epizootic +and of the human disease is now understood. In London, rats are +regularly trapped and poisoned in the vicinity of the principal docks, +to guard against the introduction of infected animals in shipping. +During the past six years infected rats have been found yearly, thirteen +having been found in 1912. In Seattle, Washington, seven infected rats +were found along the water front in October, 1913, and infected ground +squirrels are still being found in connection with the anti-plague +measures in California. + +The procedure during an outbreak of the human plague was well +illustrated by the fight in San Francisco. The city was districted, and +captured rats, after being dipped in some fluid to destroy the fleas, +were carefully tagged to indicate their source, and were sent to the +laboratory for examination. If an infected rat was found, the officers +in charge of the work in the district involved were immediately notified +by telephone, and the infected building was subjected to a thorough +fumigation. In addition, special attention was given to all the +territory in the four contiguous blocks. + +By measures such as these, this dread scourge of the human race is being +brought under control. Incidentally, the enormous losses due to the +direct ravages of rats are being obviated and this alone would justify +the expenditure many times over of the money and labor involved in the +anti-rat measures. + + + + +CHAPTER VII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC ORGANISMS + + +We now have to consider the cases in which the arthropod acts as the +essential host of a pathogenic organism. In other words, cases in which +the organism, instead of being passively carried or merely accidentally +inoculated by the bite of its carrier, or vector, is taken up and +undergoes an essential part of its development within the arthropod. + +[Illustration: 113. Dipylidium caninum. The double pored tapeworm of the +dog.] + +In some cases, the sexual cycle of the parasite is undergone in the +arthropod, which then serves as the _definitive_ or _primary host_. In +other cases, it is the asexual stage of the parasite which is undergone, +and the arthropod then acts as the _intermediate host_. This distinction +is often overlooked and all the cases incorrectly referred to as those +in which the insect or other arthropod acts as intermediate host. + +We have already emphasized that this is the most important way in which +insects may transmit disease, for without them the particular organisms +concerned could never complete their development. Exterminate the +arthropod host and the life cycle of the parasite is broken, the disease +is exterminated. + +As the phenomenon of alternation of generations, as exhibited by many of +the parasitic protozoa, is a complicated one and usually new to the +student, we shall first take up some of the grosser cases illustrated by +certain parasitic worms. There is the additional reason that these were +the first cases known of arthropod transmission of pathogenic organisms. + + +INSECTS AS INTERMEDIATE HOSTS OF TAPEWORMS + +A number of tapeworms are known to undergo their sexual stage in an +insect or other arthropod. Of these at least two are occasional +parasites of man. + +_Dipylidium caninum_ (figs. 113 and 114), more generally known as +_Taenia cucumerina_ or _T. elliptica_, is the commonest intestinal +parasite of pet dogs and cats. It is occasionally found as a human +parasite, 70 per cent of the cases reported being in young children. + +In 1869, Melnikoff found in a dog louse, _Trichodectes canis_, some +peculiar bodies which Leuckart identified as the larval form of this +tapeworm. The worm is, however, much more common in dogs and cats than +is the skin parasite, and hence it appears that the _Trichodectes_ could +not be the only intermediate host. In 1888, Grassi found that it could +also develop in the cat and dog fleas, _Ctenocephalus felis_ and _C. +canis_, and in the human flea, _Pulex irritans_. + +[Illustration: 114. Dipylidium caninum. Rostrum evaginated and +invaginated. After Blanchard.] + +[Illustration: 115. Dipylidium caninum. Immature cysticercoid. After +Grassi and Rovelli.] + +The eggs, scattered among the hairs of the dog or cat, are ingested by +the insect host and in its body cavity they develop into pyriform +bodies, about 300u in length, almost entirely destitute of a bladder, +but in the immature stage provided with a caudal appendage (fig. 115). +Within the pear-shaped body (fig. 116) are the invaginated head and +suckers of the future tapeworm. This larval form is known as a +cysticercoid, in contradistinction to the bladder-like cysticercus of +many other cestodes. It is often referred to in literature as +_Cryptocystis trichodectis_ Villot. + +As many as fifty of the cysticercoids have been found in the body cavity +of a single flea. When the dog takes up an infested flea or louse, by +biting itself, or when the cat licks them up, the larvae quickly develop +into tapeworms, reaching sexual maturity in about twenty days in the +intestine of their host. Puppies and kittens are quickly infested when +suckling a flea-infested mother, the developing worms having been found +in the intestines of puppies not more than five or six days old. + +[Illustration: 116. Dipylidium caninum. Cysticercoid. After Villet.] + +Infestation of human beings occurs only through accidental ingestion of +an infested flea. It is natural that such cases should occur largely in +children, where they may come about in some such way as illustrated in +the accompanying figures 117 and 118. + +_Hymenolepis diminuta_, very commonly living in the intestine of mice +and rats, is also known to occur in man. Its cysticercoid develops in +the body cavity of a surprising range of meal-infesting insects. Grassi +and Rovelli (abstract in Ransom, 1904) found it in the larvae and adult +of a moth, _Asopia farinalis_, in the earwig, _Anisolabis annulipes_, +the Tenebrionid beetles _Akis spinosa_ and _Scaurus striatus_. Grassi +considers that the lepidopter is the normal intermediate host. The +insect takes up the eggs scattered by rats and mice. It has been +experimentally demonstrated that man may develop the tapeworm by +swallowing infested insects. Natural infection probably occurs by +ingesting such insects with cereals, or imperfectly cooked foods. + +[Illustration: 117. One way in which Dipylidium infection in children +may occur. After Blanchard.] + +_Hymenolepis lanceolata_, a parasite of geese and ducks, has been +reported once for man. The supposed cysticercoid occurs in various small +crustaceans of the family Cyclopidae. + +[Illustration: 118. The probable method by which Dipylidium infection +usually occurs.] + +Several other cestode parasites of domestic animals are believed to +develop their intermediate stage in certain arthropods. Among these may +be mentioned: + +_Choanotaenia infundibulformis_, of chickens, developing in the house-fly +(Grassi and Rovelli); + +_Davainea cesticillus_, of chickens, in some lepidopter or coleopter +(Grassi and Rovelli); + +_Hymenolepis anatina_, _H. gracilis_, _H. sinuosa_, _H. coronula_ and +_Fimbriaria fasciolaris_, all occurring in ducks, have been reported as +developing in small aquatic crustaceans. In these cases, cysticercoids +have been found which, on account of superficial characters, have been +regarded as belonging to the several species, but direct experimental +evidence is scant. + + +ARTHROPODS AS INTERMEDIATE HOSTS OF NEMATODE WORMS + + +FILARIASIS AND MOSQUITOES--A number of species of Nematode worms +belonging to the genus _Filaria_, infest man and other vertebrates and +in the larval condition are to be found in the blood. Such infestation +is known as _filariasis_. The sexually mature worms are to be found in +the blood, the lymphatics, the mesentery and subcutaneous connective +tissue. In the cases best studied it has been found that the larval +forms are taken up by mosquitoes and undergo a transformation before +they can attain maturity in man. + +The larvae circulating in the blood are conveniently designated as +microfilariae. In this stage they are harmless and only one species, +_Filaria bancrofti_, appears to be of any great pathological +significance at any stage. + +_Filaria bancrofti_ in its adult state, lives in the lymphatics of man. +Though often causing no injury it has been clearly established that they +and their eggs may cause various disorders due to stoppage of the +lymphatic trunks (fig. 119). Manson lists among other effects, abscess, +varicose groin glands, lymph scrotum, chyluria, and elephantiasis. + +The geographical distribution of this parasite is usually given as +coextensive with that of elephantiasis, but it is by no means certain +that it is the only cause of this disease and so actual findings of the +parasites are necessary. Manson reports that it is "an indigenous +parasite in almost every country throughout the tropical and subtropical +world, as far north as Spain in Europe and Charlestown in the United +States, and as far south as Brisbane in Australia." In some sections, +fully 50 per cent of the natives are infested. Labredo (1910) found +17.82 per cent infestation in Havana. + +[Illustration: 119. Elephantiasis in Man. From "New Sydenham Society's +Atlas."] + +The larval forms of _Filaria bancrofti_ were first discovered in 1863, +by Demarquay, in a case of chylous dropsy. They were subsequently noted +under similar conditions, by several workers, and by Wucherer in the +urine of twenty-eight cases of tropical chyluria, but in 1872 Lewis +found that the blood of man was the normal habitat, and gave them the +name _Filaria sanguinis hominis_. The adult worm was found in 1876 by +Bancroft, and in 1877, Cobbold gave it the name _Filaria bancrofti_. It +has since been found repeatedly in various parts of the lymphatic +system, and its life-history has been the subject of detailed studies by +Manson (1884), Bancroft (1899), Low (1900), Grassi and Noe (1900), Noe +(1901) and Fulleborn (1910). + +The larvae as they exist in the circulating blood, exhibit a very active +wriggling movement, without material progression. They may exist in +enormous numbers, as many as five or six hundred swarming in a single +drop of blood. This is the more surprising when we consider that they +measure about 300u x 8u, that is, their width is equal to the diameter +of the red blood corpuscle of their host and their length over +thirty-seven times as great. + +Their organs are very immature and the structure obscure. When they have +quieted down somewhat in a preparation it may be seen that at the head +end there is a six-lipped and very delicate prepuce, enclosing a short +"fang" which may be suddenly exserted and retracted. Completely +enclosing the larva is a delicate sheath, which is considerably longer +than the worm itself. To enter into further details of anatomy is beyond +the scope of this discussion and readers interested are referred to the +work of Manson and of Fulleborn. + +One of the most surprising features of the habits of these larvae is the +periodicity which they exhibit in their occurrence in the peripheral +blood. If a preparation be made during the day time there may be no +evidence whatever of filarial infestation, whereas a preparation from +the same patient taken late in the evening or during the night may be +literally swarming with the parasites. Manson quotes Mackenzie as having +brought out the further interesting fact that should a "filarial subject +be made to sleep during the day and remain awake at night, the +periodicity is reversed; that is to say, the parasites come into the +blood during the day and disappear from it during the night." There have +been numerous attempts to explain this peculiar phenomenon of +periodicity but in spite of objections which have been raised, the most +plausible remains that of Manson, who believes that it is an adaptation +correlated with the life-habits of the liberating agent of the parasite, +the mosquito. + +The next stages in the development of _Filaria nocturna_ occur in +mosquitoes, a fact suggested almost simultaneously by Bancroft and +Manson in 1877, and first demonstrated by the latter very soon +thereafter. The experiments were first carried out with _Culex +quinquefasciatus_ (= _fatigans_) as a host, but it is now known that a +number of species of mosquitoes, both anopheline and culicine, may serve +equally well. + +When the blood of an infested individual is sucked up and reaches the +stomach of such a mosquito, the larvae, by very active movements, escape +from their sheaths and within a very few hours actively migrate to the +body cavity of their new host and settle down primarily in the thoracic +muscles. There in the course of sixteen to twenty days they undergo a +metamorphosis of which the more conspicuous features are the formation +of a mouth, an alimentary canal and a trilobed tail. At the same time +there is an enormous increase in size, the larvae which measured .3 mm. +in the blood becoming 1.5 mm. in length. This developmental period may +be somewhat shortened in some cases and on the other hand may be +considerably extended. The controlling factor seems to be the one of +temperature. + +The transformed larvae then reenter the body cavity and finally the +majority of them reach the interior of the labium (fig. 120). A few +enter the legs and antennae, and the abdomen, but these are wanderers +which, it is possible, may likewise ultimately reach the labium, where +they await the opportunity to enter their human host. + +It was formerly supposed that when the infested mosquito punctured the +skin of man, the mature larvae were injected into the circulation. The +manner in which this occurred was not obvious, for when the insect feeds +it inserts only the stylets, the labium itself remaining on the surface +of the skin. Fulleborn has cleared up the question by showing that at +this time the filariae escape and, like the hookworm, actively bore into +the skin of their new host. + +[Illustration: 120. Filaria in the muscles and labium of Culex. After +Blanchard.] + +Once entered, they migrate to the lymphatics and there quickly become +sexually mature. The full grown females measure 85-90 mm. in length by +.24-.28 mm. in diameter, while the males are less than half this size, +being about 40 mm. by .1 mm. Fecundation occurs and the females will be +found filled with eggs in various stages of development, for they are +normally viviparous. + +_Filaria philippinensis_ is reported by Ashburn and Craig (1907) as a +common blood filaria in the Philippine Islands. As they describe it, it +differs from _Filaria bancrofti_ primarily in that it does not exhibit +periodicity. Its development has been found to occur in _Culex +quinquefasciatus_, where it undergoes metamorphosis in about fourteen or +fifteen days. There is doubt as to the species being distinct from +_bancrofti_. + +Several other species occur in man and are thought to be transferred by +various insects, among which have been mentioned Tabanidae and +tsetse-flies, but there is no experimental proof in support of such +conjectures. + +_Filaria immitis_ is a dangerous parasite of the dog, the adult worm +living in the heart and veins of this animal. It is one of the species +which has been clearly shown to undergo its development in the mosquito, +particularly in _Anopheles maculipennis_ and _Aedes calopus_ (= +Stegomyia). The larval form occurs in the peripheral blood, especially +at night. When taken up by mosquitoes they differ from _Filaria +bancrofti_ in that they undergo their development in the Malpighian +tubules rather than in the thoracic muscles. In about twelve days they +have completed their growth in the tubules, pierce the distal end, and +pass to the labium. This species occurs primarily in China and Japan, +but is also found in Europe and in the United States. It is an +especially favorable species for studying the transformations in the +mosquito. + +[Illustration: 121. Dracunculus medinensis; female; mouth; embryo. After +Bastian and Leuckart.] + +_Filariae_ are also commonly found in birds, and in this country this is +the most available source of laboratory material. We have found them +locally (Ithaca, N. Y.) in the blood of over sixty per cent of all the +crows examined, at any season of the year, and have also found them in +English sparrows. + +In the crows, they often occur in enormous numbers, as many as two +thousand having been found in a single drop of the blood of the most +heavily infested specimen examined. For study, a small drop of blood +should be mounted on a clean slide and the coverglass rung with vaseline +or oil to prevent evaporation. In this way they can be kept for hours. + +Permanent preparations may be made by spreading out the blood in a film +on a perfectly clean slide and staining. This is easiest done by +touching the fresh drop of blood with the end of a second slide which is +then held at an angle of about 45^o to the first slide and drawn over it +without pressure. Allow the smear to dry in the air and stain in the +usual way with haematoxylin. + + +OTHER NEMATODE PARASITES OF MAN AND ANIMALS DEVELOPING IN ARTHROPODS + +_Dracunculus medinensis_ (fig. 121), the so-called guinea-worm, is a +nematode parasite of man which is widely distributed in tropical Africa, +Asia, certain parts of Brazil and is occasionally imported into North +America. + +The female worm is excessively long and slender, measuring nearly three +feet in length and not more than one-fifteenth of an inch in diameter. +It is found in the subcutaneous connective tissue and when mature +usually migrates to some part of the leg. Here it pierces the skin and +there is formed a small superficial ulcer through which the larvae reach +the exterior after bursting the body of the mother. + +[Illustration: 122. Cyclops, the intermediate host of Dracunculus.] + +Fedtschenko (1879) found that when these larvae reach the water they +penetrate the carapace of the little crustacean, _Cyclops_ (fig. 122). +Here they molt several times and undergo a metamorphosis. Fedtschenko, +in Turkestan, found that these stages required about five weeks, while +Manson who confirmed these general results, found that eight or nine +weeks were required in the cooler climate of England. + +Infection of the vertebrate host probably occurs through swallowing +infested cyclops in drinking water. Fedtschenko was unable to +demonstrate this experimentally and objection has been raised against +the theory, but Leiper (1907), and Strassen (1907) succeeded in +infesting monkeys by feeding them on cyclops containing the larvae. + +_Habronema muscae_ is a worm which has long been known in its larval +stage, as a parasite of the house-fly. Carter found them in 33 per cent +of the house-flies examined in Bombay during July, 1860, and since that +time they have been shown to be very widely distributed. Italian workers +reported them in 12 per cent to 30 per cent of the flies examined. +Hewitt reported finding it rarely in England. In this country it was +first reported by Leidy who found it in about 20 per cent of the flies +examined at Philadelphia, Pa. Since then it has been reported by several +American workers. We have found it at Ithaca, N. Y., but have not made +sufficient examinations to justify stating percentage. Ransom (1913) +reports it in thirty-nine out of one hundred and thirty-seven flies, or +28 per cent. + +[Illustration: 123. An Echinorhynchid, showing the spinose retractile +proboscis.] + +[Illustration: 124. June beetle (Lachnosterna).] + +[Illustration: Larva] + +Until very recently the life-history of this parasite was unknown but +the thorough work of Ransom (1911, 1913) has shown clearly that the +adult stage occurs in the stomach of horses. The embryos, produced by +the parent worms in the stomach of the horse, pass out with the feces +and enter the bodies of fly larvae which are developing in the manure. In +these they reach their final stage of larval development at about the +time the adult flies emerge from the pupal stage. In the adult fly they +are commonly found in the head, frequently in the proboscis, but they +occur also in the thorax and abdomen. Infested flies are accidentally +swallowed by horses and the parasite completes its development to +maturity in the stomach of its definitive host. + +_Gigantorhynchus hirudinaceus_ (= _Echinorhynchus gigas_) is a common +parasite of the pig and has been reported as occurring in man. The adult +female is 20-35 cm. long and 4-9 mm. in diameter. It lacks an alimentary +canal and is provided with a strongly spined protractile rostrum, by +means of which it attaches to the intestinal mucosa of its host. + +The eggs are scattered with the feces of the host and are taken up by +certain beetle larvae. In Europe the usual intermediate hosts are the +larvae of the cockchafer, _Melolontha vulgaris_, or of the flower beetle, +_Cetonia aurata_. Stiles has shown that in the United States the +intermediate host is the larva of the June bug, _Lachnosterna_ (fig. +124). It is probable that several of the native species serve in this +capacity. + +A number of other nematode parasites of birds and mammals have been +reported as developing in arthropods but here, as in the case of the +cestodes, experimental proof is scant. The cases above cited are the +better established and will serve as illustrations. + + + + +CHAPTER VIII + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA + + +MOSQUITOES AND MALARIA + +Under the name of malaria is included a group of morbid symptoms +formerly supposed to be due to a miasm or bad air, but now known to be +caused by protozoan parasites of the genus _Plasmodium_, which attack +the red blood corpuscles. It occurs in paroxysms, each marked by a +chill, followed by high fever and sweating. The fever is either +intermittent or remittent. + +There are three principal types of the disease, due to different species +of the parasite. They are: + +1. The benign-tertian, caused by _Plasmodium vivax_, which undergoes its +schizogony or asexual cycle in the blood in forty-eight hours or even +less. This type of the disease,--characterized by fever every two days, +is the most wide-spread and common. + +2. The quartan fever is due to the presence of _Plasmodium malariae_, +which has an asexual cycle of seventy-two hours, and therefore the fever +recurs every three days. This type is more prevalent in temperate and +sub-tropical regions, but appears to be rare everywhere. + +3. The sub-tertian "aestivo-autumnal," or "pernicious" fever is caused by +_Plasmodium falciparum_. Schizogony usually occurs in the internal +organs, particularly in the spleen, instead of in the peripheral +circulation, as is the case of the tertian and quartan forms. The fever +produced is of an irregular type and the period of schizogony has not +been definitely determined. It is claimed by some that the variations +are due to different species of malignant parasites. + +It is one of the most wide-spread of human diseases, occurring in almost +all parts of the world, except in the polar regions and in waterless +deserts. It is most prevalent in marshy regions. + +So commonplace is malaria that it causes little of the dread inspired by +most of the epidemic diseases, and yet, as Ross says, it is perhaps the +most important of human diseases. Figures regarding its ravages are +astounding. Celli estimated that in Italy it caused an average annual +mortality of fifteen thousand, representing about two million cases. In +India alone, according to Ross (1910) "it has been officially estimated +to cause a mean annual death-rate of five per thousand; that is, to kill +every year, on the average, one million one hundred and thirty +thousand." In the United States it is widespread and though being +restricted as the country develops, it still causes enormous losses. +During the year 1911, "in Alabama alone there were seventy thousand +cases and seven hundred and seventy deaths." The weakening effects of +the disease, the invasion of other diseases due to the attacks of +malaria, are among the very serious results, but they cannot be +estimated. + +Not only is there direct effect on man, but the disease has been one of +the greatest factors in retarding the development of certain regions. +Everywhere pioneers have had to face it, and the most fertile regions +have, in many instances been those most fully dominated by it. Herrick +(1903) has presented an interesting study of its effects on the +development of the southern United States and has shown that some parts, +which are among the most fertile in the world, are rendered practically +uninhabitable by the ravages of malaria. Howard (1909) estimates that +the annual money loss from the disease in the United States is not less +than $100,000,000. + +It was formerly supposed that the disease was due to a miasm, to a +noxious effluvia, or infectious matter rising in the air from swamps. In +other words its cause was, as the name indicated "mal aria," and the +deep seated fear of night air is based largely on the belief that this +miasm was given off at night. Its production was thought to be favored +by stirring of the soil, dredging operations and the like. + +The idea of some intimate connection between malaria and mosquitoes is +not a new one. According to Manson, Lancisi noted that in some parts of +Italy the peasants for centuries have believed that malaria is produced +by the bite of mosquitoes. Celli states that one not rarely hears from +such peasants the statement that "In such a place, there is much fever, +because it is full of mosquitoes." Koch points out that in German East +Africa the natives call malaria and the mosquito by the same name, +_Mbu_. The opinion was not lacking support from medical men. Celli +quotes passages from the writings of the Italian physician, Lancisi, +which indicate that he favored the view in 1717. + +Dr. Josiah Nott is almost universally credited with having supported the +theory, in 1848, but as we have already pointed out his work has been +misinterpreted. The statements of Beauperthuy, (1853) were more +explicit. + +The clearest early presentation of the circumstantial evidence in favor +of the theory of mosquito transmission was that of A. F. A. King, an +American physician, in 1883. He presented a series of epidemiological +data and showed "how they may be explicable by the supposition that the +mosquito is the real source of the disease, rather than the inhalation +or cutaneous absorption of a marsh vapor." We may well give the space to +summarizing his argument here for it has been so remarkably +substantiated by subsequent work: + +1. Malaria, like mosquitoes, affects by preference low and moist +localities, such as swamps, fens, jungles, marshes, etc. + +2. Malaria is hardly ever developed at a lower temperature than 60^o +Fahr., and such a temperature is necessary for the development of the +mosquito. + +3. Mosquitoes, like malaria, may both accumulate in and be obstructed by +forests lying in the course of winds blowing from malarious localities. + +4. By atmospheric currents malaria and mosquitoes are alike capable of +being transported for considerable distances. + +5. Malaria may be developed in previously healthy places by turning up +the soil, as in making excavations for the foundation of houses, tracks +for railroads, and beds for canals, because these operations afford +breeding places for mosquitoes. + +6. In proportion as countries, previously malarious, are cleared up and +thickly settled, periodical fevers disappear, because swamps and pools +are drained so that the mosquito cannot readily find a place suitable to +deposit her eggs. + +7. Malaria is most dangerous when the sun is down and the danger of +exposure after sunset is greatly increased by the person exposed +sleeping in the night air. Both facts are readily explicable by the +mosquito malaria theory. + +8. In malarial districts the use of fire, both indoors and to those who +sleep out, affords a comparative security against malaria, because of +the destruction of mosquitoes. + +9. It is claimed that the air of cities in some way renders the poison +innocuous, for, though a malarial disease may be raging outside, it does +not penetrate far into the interior. We may easily conceive that +mosquitoes, while invading cities during their nocturnal pilgrimages +will be so far arrested by walls and houses, as well as attracted by +lights in the suburbs, that many of them will in this way be prevented +from penetrating "far into the interior." + +10. Malarial diseases and likewise mosquitoes are most prevalent toward +the latter part of summer and in the autumn. + +11. Various writers have maintained that malaria is arrested by canvas +curtains, gauze veils and mosquito nets and have recommended the rise of +mosquito curtains, "through which malaria can seldom or never pass." It +can hardly be conceived that these intercept marsh-air but they +certainly do protect from mosquitoes. + +12. Malaria spares no age, but it affects infants much less frequently +than adults, because young infants are usually carefully housed and +protected from mosquito inoculation. + +Correlated with the miasmatic theory was the belief that some animal or +vegetable organism which lived in marshes, produced malaria, and +frequent searches were made for it. Salisbury (1862) thought this +causative organism to be an alga, of the genus _Palmella_; others +attributed it to certain fungi or bacteria. + +In 1880, the French physician, Laveran, working in Algeria, discovered +an amoeboid organism in the blood of malarial patients and definitely +established the parasitic nature of this disease. Pigmented granules had +been noted by Meckel as long ago as 1847, in the spleen and blood of a +patient who had died of malaria, and his observations had been +repeatedly verified, but the granules had been regarded as degeneration +products, and the fact that they occurred in the body of a foreign +organism had been overlooked. + +Soon after the discovery of the parasites in the blood, Gerhardt (1884) +succeeded in transferring the disease to healthy individuals by +inoculation of malarious blood, and thus proved that it is a true +infection. This was verified by numerous experimenters and it was found +that inoculation with a very minute quantity of the diseased blood would +not only produce malaria but the particular type of disease. + +Laveran traced out the life cycle of the malarial parasite as it occurs +in man. The details as we now know them and as they are illustrated by +the accompanying figure 125, are as follows: + +The infecting organism or _sporozoite_, is introduced into the +circulation, penetrates a red blood corpuscle, and forms the amoeboid +_schizont_. This lives at the expense of the corpuscle and as it +develops there are deposited in its body scattered black or reddish +black particles. These are generally called melanin granules, but are +much better referred to as haemozoin, as they are not related to +melanin. The haemozoin is the most conspicuous part of the parasite, a +feature of advantage in diagnosing from unstained preparations. + +[Illustration: 125. Life cycle of the malaria parasite. Adapted from +Leuckart's chart, by Miss Anna Stryke.] + +As the schizont matures, its nucleus breaks up into a number of daughter +nuclei, each with a rounded mass of protoplasm about it, and finally the +corpuscles are broken down and these rounded bodies are liberated in the +plasma as _merozoites_. These merozoites infect new corpuscles and thus +the asexual cycle is continued. The malarial paroxysm is coincident with +sporulation. + +As early as Laveran's time it was known that under conditions not yet +determined there are to be found in the blood of malarious patients +another phase of the parasite, differing in form according to the type +of the disease. In the pernicious type these appear as large, +crescent-shaped organisms which have commonly been called "crescents." +We now know that these are sexual forms. + +When the parasite became known there immediately arose speculations as +to the way in which it was transferred from man to man. It was thought +by some that in nature it occurred as a free-living amoeba, and that +it gained access to man through being taken up with impure water. +However, numerous attempts to infect healthy persons by having them +drink or inhale marsh water, or by injecting it into their circulation +resulted in failure, and influenced by Leuckart's and Melnikoff's work +on _Dipylidium_, that of Fedtschenko on _Dracunculus_, and more +especially by that of Manson on _Filaria_, search was made for some +insect which might transfer the parasite. + +Laveran had early suggested that the role of carrier might be played by +the mosquito, but Manson first clearly formulated the hypothesis, and it +was largely due to his suggestions that Ross in India, undertook to +solve the problem. With no knowledge of the form or of the appearance in +this stage, or of the species of mosquito concerned, Ross spent almost +two and a half years of the most arduous work in the search and finally +in August, 1897, seventeen years after the discovery of the parasite in +man, he obtained his first definite clue. In dissecting a +"dappled-winged mosquito," "every cell was searched and to my intense +disappointment nothing whatever was found, until I came to the insect's +stomach. Here, however, just as I was about to abandon the examination, +I saw a very delicate circular cell, apparently lying amongst the +ordinary cells of the organ and scarcely distinguishable from them. On +looking further, another and another similar object presented itself. I +now focused the lens carefully on one of these, and found that it +contained a few minute granules of some black substance, exactly like +the pigment of the parasite of malaria. I counted altogether twelve of +these cells in the insect." + +Further search showed that "the contents of the mature pigment cells did +not consist of clear fluid but of a multitude of delicate, thread-like +bodies which on the rupture of the parent cell, were poured into the +body cavity of the insect. They were evidently spores." + +With these facts established, confirmation and extension of Ross's +results quickly followed, from many different sources. We cannot trace +this work in detail but will only point out that much of the credit is +due to the Italian workers, Grassi, Bignami, and Bastianelli, and to +Koch and Daniels. + +It had already been found that when fresh blood was mounted and properly +protected against evaporation, a peculiar change occurred in these +crescents after about half an hour's time. From certain of them there +were pushed out long whip-like processes which moved with a very active, +lashing movement. The parasite at this stage is known as the +"flagellated body." Others, differing somewhat in details of structure, +become rounded but do not give off "flagella." + +The American worker, MacCallum (1897), in studying bird malaria as found +in crows, first recognized the true nature of these bodies. He regarded +them as sexual forms and believed that the so-called flagella played the +part of spermatozoa. Thus, the "flagellated body" is in reality a +_microgametoblast_, producing _microgametes_, or the male sexual +element, while the others constitute the _macrogametes_, or female +elements. + +It was found that when blood containing these sexual forms was sucked up +by an Anopheline mosquito and taken into its stomach, a microgamete +penetrated and fertilized a macrogamete in a way analogous to what takes +place in the fertilization of the egg in higher forms. The resultant, +mobile organism is known as the _migratory ookinete_. In this stage the +parasite bores through the epithelial lining of the "stomach" +(mid-intestine) of the mosquito and becomes encysted under the muscle +layers. Here the _oocyst_, as it is now known, matures and breaks up +into the body cavity and finally its products come to lie in the +salivary glands of the mosquito. Ten to twelve days are required for +these changes, after which the mosquito is infective, capable of +introducing the parasite with its saliva, when feeding upon a healthy +person. + +Thus the malarial parasite is known to have a double cycle, an +alternation of generations, of which the asexual stage is undergone in +man, the sexual in certain species of mosquitoes. The mosquito is +therefore the definitive host rather than the _intermediate_, as usually +stated. + +The complicated cycle may be made clearer by the diagram of Miss Stryke +(1912) which, by means of a double-headed mosquito (fig. 126) endeavors +to show how infection takes place through the biting of the human +victim, (at A), in whom asexual multiplication then takes place, and how +the sexual stages, taken up at B in the diagram, are passed in the body +of the mosquito. + +[Illustration: 126. Life cycle of the malarial parasite. After Miss Anna +Stryke.] + +The experimental proof that mosquitoes of the Anopheline group are +necessary agents in the transmission of malaria was afforded in 1900 +when two English physicians, Drs. Sambon and Low lived for the three +most malarial months in the midst of the Roman Campagna, a region +famous for centuries as a hot-bed of malaria. The two experimenters +moved about freely throughout the day, exposed themselves to rains and +all kinds of weather, drank marsh water, slept exposed to the marsh air, +and, in short, did everything which was supposed to cause malaria, +except that they protected themselves thoroughly from mosquito bites, +retiring at sunset to a mosquito-proof hut. Though they took no quinine +and all of their neighbors suffered from malaria, they were absolutely +free from the disease. + +To complete the proof, mosquitoes which had fed in Rome on malarious +patients were sent to England and allowed to bite two volunteers, one of +them Dr. Manson's own son, who had not been otherwise exposed to the +disease. Both of these gentlemen contracted typical cases of malaria and +the parasites were to be found in abundance in their blood. + +[Illustration: 127. Eggs of Anopheles. After Howard.] + +Since that time there have been many practical demonstrations of the +fact that malaria is transmitted exclusively by the bite of mosquitoes +and that the destruction of the mosquitoes means the elimination of the +disease. + +We have said that the malarial parasite is able to undergo its +development only in certain species of mosquitoes belonging to the +Anopheline group. It is by no means certain that all of this group even, +are capable of acting as the definitive host of the parasites, and much +careful experiment work is still needed along this line. In the United +States, several species have been found to be implicated, _Anopheles +quadrimaculatus_ and _Anopheles crucians_ being the most common. The +characteristics of these species and the distinctions between them and +other mosquitoes will be discussed in Chapter XII. + +In antimalarial work it is desirable to distinguish the anopheline +mosquitoes from the culicine species in all stages. The following +tabulation presents the more striking distinctions between the groups as +represented in the United States. + +_Anopheles_ _Culex, Aedes, etc_. + +_Eggs_: Laid singly in small Deposited in clumps in the +numbers upon the surface of the form of a raft (Culex group) or +water. Eggs lie upon their sides deposited singly in the water or +and float by means of lateral on the ground in places which +expansions (fig. 127). may later be submerged. + +_Larva_: When at rest floats in When at rest (with few exceptions) +a horizontal position beneath the floats suspended in an +surface film. No respiratory oblique or vertical position, or +tube but instead a flattened more rarely nearly horizontal, +area on the eighth abdominal with the respiratory tube in +segment into which the two contact with the surface film +spiracles open (fig. 128). (fig. 128). + +_Adults_: Palpi in both sexes Palpi short in the female, in +nearly or quite as long as the the male usually elongate. +proboscis. Proboscis projecting Proboscis projects forward at an +forward nearly on line with the angle with the axis of the body. +axis of the body. When at rest When at rest on a vertical wall +on a vertical wall the body is the body is usually held parallel +usually held at an angle with the or the tip of the abdomen inclined +vertical (fig. 128). Wings frequently towards the wall (fig. 128). +spotted (fig. 130). Wings usually not spotted. + +[Illustration: 128. (_a_) Normal position of the larvae of Culex and +Anopheles in the water. Culex, left; Anopheles, middle; Culex pupa, +right hand figure.] + +These malarial-bearing species are essentially domesticated mosquitoes. +They develop in any accumulation of water which stands for a week or +more. Ponds, puddles, rain barrels, horse troughs, cess-pools, cans, +even the foot-prints of animals in marshy ground may afford them +breeding places. + +[Illustration: 128. (_b_) Normal position of Culex and Anopheles on the +wall.] + +It is clear from what has been said regarding the life cycle of the +malarial parasite that the mosquito is harmless if not itself diseased. +Hence malarial-bearing species may abound in the neighborhood where +there is no malaria, the disease being absent simply because the +mosquitoes are uninfected. Such a locality is potentially malarious and +needs only the introduction of a malarial patient who is exposed to the +mosquitoes. It is found that such patients may harbor the parasites in +their blood long after they are apparently well and thus may serve as a +menace, just as do the so-called typhoid carriers. In some malarious +regions as high as 80-90 per cent of the natives are such +malaria-carriers and must be reckoned with in antimalaria measures. + +Based upon our present day knowledge of the life cycle of the malarial +parasite the fight against the disease becomes primarily a problem in +economic entomology,--it is a question of insect control, in its +broadest interpretation. + +[Illustration: 129. Larva of Anopheles. After Howard.] + +The lines of defence and offence against the disease as outlined by +Boyce (1909) are: + + 1. Measures to avoid the reservoir (man): + + Segregation. + + Screening of patients. + + 2. Measures to avoid Anopheles: + + Choice of suitable locality, when possible. + + Screening of houses and porches. + + Sleeping under mosquito nets. + + 3. Measures to exterminate the Anopheles: + + Use of natural enemies. + + Use of culicides, oiling ponds, etc. + + Drainage and scavenging to destroy breeding places. + + Enforcement of penalties for harboring larvae or keeping stagnant + water. + + Educational methods. + + 4. Systematic treatment with quinine to exterminate the parasites. + + +MOSQUITOES AND YELLOW FEVER + +Yellow fever was until recently one of the most dreaded of epidemic +diseases. It is an acute, specific and infectious disease, +non-contagious in character but occurring in epidemics, or endemics, +within a peculiarly limited geographical area. It is highly fatal, but +those who recover are generally immune from subsequent attacks. + +It is generally regarded as an American disease, having been found by +Cortez, in Mexico, and being confined principally to the American +continents and islands. It also occurs in Africa and attempts have been +made to show that it was originally an African disease but there is not +sufficient evidence to establish this view. + +There have been many noted outbreaks in the United States. Boston +suffered from it in 1691 and again in 1693; New York in 1668 and as late +as 1856; Baltimore in 1819. In 1793 occurred the great epidemic in +Philadelphia, with a death rate of one in ten of the population. In the +past century it was present almost every year in some locality of our +Southern States, New Orleans being the greatest sufferer. In the latter +city there were 7848 deaths from the disease in 1853, 4854 in 1858, and +4046 in 1878. The last notable outbreak was in 1905. Reed and Carroll +(1901) estimated that during the period from 1793 to 1900 there had not +been less than 500,000 cases in the United States. + +[Illustration: 130. Anopheles quadrimaculatus, male and female, +(x3-1/2). After Howard.] + +As in the case of the plague, the most stringent methods of control +proved ineffective and helplessness, almost hopelessness marked the +great epidemics. A vivid picture of conditions is that given by Mathew +Cary, 1793 (quoted by Kelly, 1906) in "A Short Account of the Malignant +Fever Lately Prevalent in Philadelphia." + +[Illustration: 131. Anopheles punctipennis. Female, (x4). After Howard.] + +"The consternation of the people of Philadelphia at this period was +carried beyond all bounds. Dismay and affright were visible in the +countenance of almost every person. Of those who remained, many shut +themselves in their houses and were afraid to walk the streets. * * * +The corpses of the most respectable citizens, even those who did not die +of the epidemic, were carried to the grave on the shafts of a chair +(chaise), the horse driven by a negro, unattended by friends or +relative, and without any sort of ceremony. People hastily shifted their +course at the sight of a hearse coming toward them. Many never walked on +the footpath, but went into the middle of the streets to avoid being +infected by passing by houses wherein people had died. Acquaintances and +friends avoided each other in the streets and only signified their +regard by a cold nod. The old custom of shaking hands fell into such +disuse that many shrunk back with affright at even the offer of the +hand. A person with a crape, or any appearance of mourning was shunned +like a viper. And many valued themselves highly on the skill and address +with which they got to the windward of every person they met. Indeed, it +is not probable that London, at the last stage of the plague, exhibited +stronger marks of terror than were to be seen in Philadelphia from the +24th or 25th of August until pretty late in September." + +[Illustration: 132. Anopheles crucians. Female (x4). After Howard.] + +Such was the condition in Philadelphia in 1793 and, as far as methods of +control of the disease were concerned, there was practically no advance +during the last century. The dominant theory was that yellow fever was +spread by _fomites_, that is, exposed bedding, clothing, baggage, and +the like. As late as 1898 a bulletin of the United States Marine +Hospital Service stated: + +"While yellow fever is a communicable disease, it is not contagious in +the ordinary acceptance of the term, but is spread by the infection of +places and articles of bedding, clothing, and furniture." + +Based upon this theory, houses, baggage, freight, even mail, were +disinfected, and the most rigid quarantine regulations were enforced. +The hardships to which people of the stricken regions were subjected and +the financial losses are incalculable. And withal, the only efficient +check upon the disease seemed to be the heavy frosts. It was found that +for some reason, the epidemic abated with cold weather,--a measure +beyond human control. + +[Illustration: 133. Culex sollicitans. Female (x4). After Howard.] + +It is not strange that among the multitude of theories advanced to +explain the cause and method of dissemination of the disease there +should be suggestions that yellow fever was transmitted by the mosquito. +We have seen that Beauperthuy (1855) clearly urged this theory. + +More detailed, and of the greatest influence in the final solution of +the problem were the arguments of Dr. Carlos Finlay, of Havana. In 1881, +in a paper presented before the "Real Academia de Ciencias Medicas, +Fisicas y Naturales de la Habana," he said: + +"I feel convinced that any theory which attributes the origin and the +propagation of yellow fever to atmospheric influences, to miasmatic or +meteorological conditions, to filth, or to the neglect of general +hygienic precautions, must be considered as utterly indefensible." + +He postulated the existence of a material transportable substance +causing yellow fever,--"something tangible which requires to be conveyed +from the sick to the healthy before the disease can be propagated" and +after discussing the peculiarities of the spread of the disease and the +influence of meteorological conditions, he decides that the carriers of +the disease must be sought among insects. He continues: + +"On the other hand, the fact of yellow fever being characterized both +clinically and (according to recent findings) histologically, by lesions +of the blood vessels and by alterations of the physical and chemical +conditions of the blood, suggested that the insect which should convey +the infectious particles from the patient to the healthy should be +looked for among those which drive their sting into blood vessels in +order to suck human blood. Finally, by reason of other considerations +which need not be stated here, I came to think that the mosquito might +be the transmitter of yellow fever." + +"Assimilating the disease to small-pox and to vaccination, it occurred +to me that in order to inoculate yellow fever it would be necessary to +pick out the inoculable material from within the blood vessels of a +yellow fever patient and to carry it likewise into the interior of a +blood vessel of a person who was to be inoculated. All of which +conditions the mosquito satisfies most admirably through its bite." + +In the course of his study of the problem, Finlay made detailed studies +of the life history and habits of the common mosquitoes at Havana, and +arrived at the conclusion that the carrier of the yellow fever was the +_Culex mosquito_ or _Aedes calopus_, as it is now known. With this +species he undertook direct experimental tests, and believed that he +succeeded in transmitting the disease by the bite of infected mosquitoes +in three cases. Unfortunately, possibility of other exposure was not +absolutely excluded, and the experiments attracted little attention. + +Throughout the next twenty years Finlay continued his work on yellow +fever, modifying his original theory somewhat as time went on. Among his +later suggestions was that in the light of Smith's work on Texas fever, +his theory must be "somewhat modified so as to include the important +circumstance that the faculty of transmitting the yellow fever germ need +not be limited to the parent insect, directly contaminated by stinging a +yellow fever patient (or perhaps by contact with or feeding from his +discharges), but may be likewise inherited by the next generation of +mosquitoes issued from the contaminated parent." He believed that the +bite of a single mosquito produced a light attack of the disease and was +thus effective in immunizing the patient. Throughout the period, many +apparently successful attempts to transmit the disease by mosquitoes +were made. In the light of present day knowledge we must regard these as +defective not only because possibility of other infection was not +absolutely excluded but because no account was taken of the incubation +period within the body of the mosquito. + +In 1900, while the American army was stationed in Cuba there occurred an +epidemic of yellow fever and an army medical board was appointed for +"the purpose of pursuing scientific investigations with reference to the +acute infectious diseases prevalent on the island." This was headed by +Walter Reed and associated with him were James Carroll, Jesse W. Lazear +and Aristides Agramonte, the latter a Cuban immune. For a detailed +summary of this work the lay reader cannot do better than read Dr. +Kelly's fascinating biography "Walter Reed and Yellow Fever." + +Arriving at the army barracks near Havana the Commission first took up +the study of _Bacillus icteroides_, the organism which Sanarelli, an +Italian physician, had declared the causative agent in yellow fever. +They were unable to isolate this bacillus either from the blood during +life or from the blood and organs of cadavers and therefore turned their +attention to Finlay's theory of the propagation of yellow fever by means +of the mosquito. In this work they had the unselfish and enthusiastic +support of Dr. Finlay himself, who not only consulted with them and +placed his publications at their disposal, but furnished eggs from which +their experimental mosquitoes were obtained. Inoculations of eleven +non-immunes through the bite of infected mosquitoes were made, and of +these, two gave positive results. The first of the two was Dr. Carroll +who allowed himself to be bitten by a mosquito which had been caused to +feed upon four cases of yellow fever, two of them severe and two mild. +The first patient had been bitten twelve days before. + +Three days after being bitten, Dr. Carroll came down with a typical case +of yellow fever. So severe was the attack that for three days his life +hung in the balance. During his convalescence an incident occurred which +showed how the theory of mosquito transmission of the disease was +generally regarded. We quote from Dr. Kelly: "One of his nurses who came +from Tennessee had had considerable experience with yellow fever, having +indeed, lost her husband and several children from it. One day early in +his illness Dr. Carroll mentioned to her that he had contracted the +disease through the bite of a mosquito, and noticed that she looked +surprised. Some time later, when well enough to look over the daily +records of his condition, he found this entry: 'Says he got his illness +through the bite of a mosquito,--delirious'." + +The second case was that of an American who was bitten by four +mosquitoes, two of which had bitten severe (fatal) cases of yellow fever +twelve days previously, one of which had bitten a severe case (second +day) sixteen days before and one which had bitten a severe case eight +days before. Five days later, the subject developed a well pronounced +but mild case of the disease. + +In the meantime, another member of the Commission, Dr. Lazear, was +accidentally bitten by a mosquito while collecting blood from yellow +fever patients. Five days later he contracted a typical case which +resulted fatally. + +So clear was the evidence from these preliminary experiments that the +commission felt warranted in announcing, October 27, 1900, that, "The +mosquito serves as the intermediate host for the parasite of yellow +fever, and it is highly probable that the disease is only propagated +through the bite of this insect." + +In order to extend the experimental evidence under conditions which +could leave no possibility of infection from other sources, a special +experimental sanitary station, named in honor of the deceased member of +the Commission, was established in an open field near the town of +Quemados, Cuba. Here there were constructed two small buildings known +respectively as the "infected clothing building" and the "infected +mosquito building." + +The infected clothing building, 14 x 20 feet in size, was purposely so +constructed as to exclude anything like efficient ventilation, but was +thoroughly screened to prevent the entrance of mosquitoes. Into this +building were brought sheets, pillow-slips, blankets, etc., contaminated +by contact with cases of yellow fever and their discharges,--many of +them purposely soiled with a liberal quantity of black vomit, urine, and +fecal matter from patients sick with yellow fever. Nothing could better +serve as the fomites which were supposed to convey the dread disease. + +Three non-immunes unpacked these articles, giving each a thorough +handling and shaking in order to disseminate through the air of the room +the specific agent of the disease. They were then used in making up the +beds which the volunteers occupied each night for a period of twenty +days. The experiment was repeated three times, volunteers even sleeping +in the soiled garments of yellow fever victims but in not a single case +was there the slightest symptom of disease. The theory of the spread of +yellow fever by fomites was completely demolished. + +The infected mosquito building, equal in size to its companion, was the +antithesis as far as other features were concerned. It was so +constructed as to give the best possible ventilation, and bedding which +was brought into it was thoroughly sterilized. Like the infected +clothing building it was carefully screened, but in this case it was in +order to keep mosquitoes in it as well as to prevent entrance of others. +Through the middle of the room ran a mosquito-proof screen. + +On December 5, 1900, a non-immune volunteer who had been in the +quarantine camp for fifteen days and had had no other possible exposure, +allowed himself to be bitten by five mosquitoes which had fed on yellow +fever patients fifteen or more days previously. The results were fully +confirmatory of the earlier experiments of the Commission--at the end of +three days, nine and a half hours, the subject came down with a well +marked case of yellow fever. + +In all, ten cases of experimental yellow fever, caused by the bite of +infected mosquitoes were developed in Camp Lazear. Throughout the period +of the disease, other non-immunes slept in the little building, +separated from the patient only by the mosquito-proof screen, but in no +circumstances did they suffer any ill effects. + +It was found that a yellow fever patient was capable of infecting +mosquitoes only during the first three or four days after coming down +with the disease. Moreover, after the mosquito has bitten such a +patient, a period of at least twelve days must elapse before the insect +is capable of transmitting the disease. + +Once the organism has undergone its twelve day development, the mosquito +may remain infective for weeks. In experiments of the Commission, two of +the mosquitoes transmitted the disease to a volunteer fifty-seven days +after their contamination. No other volunteers presenting themselves, +one of these mosquitoes died the sixty-ninth and one the seventy-first +day after their original contamination, without it being determined +whether they were still capable of transmitting the disease. + +So carefully carried out was this work and so conclusive were the +results that Dr. Reed was justified in writing: + +"Six months ago, when we landed on this island, absolutely nothing was +known concerning the propagation and spread of yellow fever--it was all +an unfathomable mystery--but today the curtain has been drawn--its mode +of propagation is established and we know that a case minus mosquitoes +is no more dangerous than one of chills and fever." + +The conclusions of the Commission were fully substantiated by numerous +workers, notably Dr. Guiteras of the Havana Board of Health, who had +taken a lively interest in the work and whose results were made known in +1901, and by the Brazilian and French Commission at Sao Paulo, Brazil, +in 1903. + +Throughout the work of the Army Commission and down to the present time +many fruitless efforts have been made to discover the specific organism +of yellow fever. It was clearly established that the claims of Sanarelli +for _Bacillus icteroides_ were without foundation. It was found, too, +that whatever the infective agent might be it was capable of passing +through a Berkefeld filter and thus belongs to the puzzling group of +"filterable viruses." It was further found that the virus was destroyed +by heating up to 55^o C for ten minutes. It is generally believed that +the organism is a Protozoan. + +The question of the hereditary transmission of the yellow fever organism +within the mosquito was left unsettled by the Army Commission, though, +as we have seen, it was raised by Finlay. Marchoux and Simond, of the +French Commission devoted much attention to this phase of the problem +and basing their conclusions on one apparently positive case, they +decided that the disease could be transmitted through the egg of an +infected _Aedes calopus_ to the second generation and thence to man. The +conclusion, which is of very great importance in the control of yellow +fever, has not been verified by other workers. + +Once clearly established that yellow fever was transmitted solely by +mosquitoes, the question of the characteristics, habits, and +geographical distribution of the insect carrier became of vital +importance. + +_Aedes calopus_, more commonly known as _Stegomyia fasciata_ or +_Stegomyia calopus_ (fig. 134) is a moderate sized, rather strikingly +marked mosquito. The general color is dark-brown or reddish-brown, but +the thorax has a conspicuous broad, silvery-white curved line on each +side, with two parallel median silvery lines. Between the latter there +is a slender, broken line. The whole gives a lyre-shaped pattern to the +thorax. The abdomen is dark with silvery-white basal bands and silvery +white spots on each side of the abdominal segments. Legs black with +rings of pure white at the base of the segments. + +Size of the female 3.3 to 5 mm.; male 3 to 4.5 mm. + +[Illustration: 134. The yellow fever mosquito (Aedes calopus). (x7). +After Howard.] + +It is preeminently a domesticated species, being found almost +exclusively about the habitation of man. "Its long association with man +is shown by many of its habits. It approaches stealthily from behind. It +retreats upon the slightest alarm. The ankles and, when one is sitting +at a table or desk, the underside of the hands and wrists are favorable +points of attack. It attacks silently, whereas other mosquitoes have a +piping or humming note. The warning sound has doubtless been suppressed +in the evolutionary process of its adaptation to man. It is extremely +wary. It hides whenever it can, concealing itself in garments, working +into the pockets, and under the lapels of coats, and crawling up under +the clothes to bite the legs. In houses, it will hide in dark corners, +under picture moldings and behind the heads of old-fashioned bedsteads. +It will enter closets and hide in the folds of garments."--Howard. + +It was claimed by the French Commission, and subsequently often stated +in discussions of the relation of the mosquito to yellow fever that the +mature _Aedes calopus_ will bite only at night. If this were true it +would be of the greatest importance in measures to avoid the disease. +Unfortunately, the claim was illy founded and numerous workers have +clearly established that the exact converse is more nearly true, this +mosquito being pre-eminently a day species, feeding most actively in +early morning, about sunrise, and late in the afternoon. On cloudy days +it attacks at any time during the day. Thus there is peril in the +doctrine that infected regions may be visited with perfect safety during +the daytime and that measures to avoid the mosquito attack need be taken +only at night. + +[Illustration: 135_a_. Aedes calopus. Pupa. After Howard.] + +Dr. Finlay maintained that the adult, even when starved, would not bite +when the temperature was below 23^o C, but subsequent studies have shown +that this statement needs modification. The French Commission, working +at Rio de Janeiro, found that _Aedes calopus_ would bite regularly at +temperatures between 22^o and 25^o and that the optimum temperature was +between 27^o and 30^o C, but their experiments led them to believe that it +would bite in nature at a temperature as low as 17^o C. + +The yellow fever mosquito breeds in cisterns, water barrels, pitchers +and in the various water receptacles about the house. In our own +Southern States it very commonly breeds in the above-ground cisterns +which are in general use. Often the larvae (fig. 135b) are found in +flower vases, or even in the little cups of water which are placed under +the legs of tables to prevent their being overrun by ants. They have +been repeatedly found breeding in the holy water font in churches. In +short, they breed in any collection of water in close proximity to the +dwellings or gathering places of man. + +The life cycle under favorable conditions is completed in from twelve to +fifteen days. These figures are of course very dependent upon the +temperature. The Army Commission in Cuba found that the cycle might be +completed in as brief a period as nine and a half days. Under less +favorable conditions it may be greatly lengthened. + +The adults are long lived. We have seen that during the experimental +work in Cuba specimens were kept in captivity for sixty-nine and +seventy-one days, respectively, and that they were proved to retain +their infectivity for at least fifty-seven days. Dr. Guiteras +subsequently kept an infected adult for one hundred and fifty-four days. + +Low temperatures have a very great effect not only on development, but +on the activity and even life of the adults. Long before the method of +transmission of yellow fever was discovered it was well known that the +epidemics were brought to a close by heavy frosts, and it is now known +that this is due to the killing of the mosquitoes which alone could +spread the disease. + +[Illustration: 135_b_. Aedes calopus; larva. (x7). After Howard.] + +_Aedes calopus_ has a very wide distribution since, as Howard says, +being a domestic mosquito, having a fairly long life in the adult stage, +and having the custom of hiding itself in the most ingenious ways, it is +particularly subject to carriage for long distances on board vessels, in +railway trains, even packed in baggage. In general, its permanent +distribution is from 40 degrees north latitude to 40 degrees south +latitude (Brumpt), in a belt extending around the world. In the United +States it breeds in most of our Southern States. + +Thus, as in the case of malaria, there are many places where the insect +carrier is abundant but where yellow fever does not occur. Such, for +instance, are Hawaii, Australia and Asia. An outbreak may occur at any +time that a patient suffering from the disease is allowed to enter and +become a source of infection for the mosquitoes. In this connection +various writers have called attention to the menace from the Panama +Canal. When it is completed, it will allow of direct passage from +regions where yellow fever is endemic and this will greatly increase the +possibility of its introduction into these places where it is now +unknown. The result, with a wholly non-immune population, would be +appalling. + +On the other hand, there are places wholly outside of the normal range +of _Aedes calopus_ where the disease has raged. Such are New York, +Boston, and even Philadelphia, which have suffered notable epidemics. +These outbreaks have been due to the introduction of infected mosquitoes +during the heat of summer, when they have not only conveyed the disease +but have found conditions favorable for their multiplication. Or, +uninfected mosquitoes have been thus accidentally brought in and +developed in large numbers, needing then only the accidental +introduction of cases of the disease to start an epidemic. + +Methods of control of various diseases have been revolutionized by the +discovery that they were insect-borne, but in no other case has the +change been as radical or the results as spectacular as in the case of +yellow fever. The "shot-gun quarantine," the sufferings and horrors, the +hopelessness of fighting absolutely blindly have given way to an +efficient, clear-cut method of control, based upon the knowledge that +the disease is carried from man to man solely by the mosquito, _Aedes +calopus_. The lines of defense and offense are essentially as follows: + +In the first place, when a case of yellow fever occurs, stringent +precautions must be adopted to prevent the infection of mosquitoes and +the escape of any already infected. This means that the patient must be +removed to a mosquito-proof room, or ward beyond reach of the insects, +and that the infected room must be thoroughly fumigated at once, to kill +the mosquitoes hiding within it. All cracks and openings should be +closed with strips of paper and fumigation with burning sulphur or +pyrethrum carefully carried out. + +It should be remembered that if the first case noted is that of a +resident rather than imported, it means that the mosquito carriers +became infected more than two weeks before the case was diagnosed, for +as we have seen, the germ must undergo a twelve-day period of +development within its insect host. Therefore a careful search must be +made for mild cases which, though unrecognized, may serve as foci for +the spread of the disease. + +In face of a threatened epidemic one of the most essential measures is +to educate the citizens and to gain their complete cooperation in the +fight along modern lines. This may be done through the schools, the +pulpit, places of amusement, newspapers and even bulletin boards. + +Emphasis should be placed on the necessity of both non-immunes and +immunes using mosquito curtains, and in all possible ways avoiding +exposure to the mosquitoes. + +Then the backbone of the fight must be the anti-mosquito measures. In +general, these involve screening and fumigating against adults, and +control of water supply, oiling, and drainage against the larvae. The +region involved must be districted and a thorough survey undertaken to +locate breeding places, which must, if possible, be eradicated. If they +are necessary for water supplies, such as casks, or cisterns, they +should be carefully screened to prevent access of egg-laying adults. + +The practical results of anti-mosquito measures in the fight against +yellow fever are well illustrated by the classic examples of the work in +Havana, immediately following the discoveries of the Army Commission and +by the stamping out of the New Orleans epidemic in 1905. + +The opportunities for an immediate practical application of the theories +of the Army Commission in Havana were ideal. The city had always been a +hotbed of yellow fever and was the principal source from which the +disease was introduced year after year into our Southern States. It was +under martial law and with a military governor who was himself a +physician and thoroughly in sympathy with the views of the Commission, +the rigid enforcement of the necessary regulations was possible. The +story of the first campaign has been often told, but nowhere more +clearly than in Dr. Reed's own account, published in the _Journal of +Hygiene_ for 1902. + +Closer home was the demonstration of the efficacy of these measures in +controlling the yellow fever outbreak in New Orleans in 1905. During the +spring and early summer of the year the disease had, unperceived, gained +a firm foothold in that city and when, in early July the local Board of +Health took cognizance of its existence, it was estimated that there had +been in the neighborhood of one hundred cases. + +Conditions were not as favorable as they had been under martial law in +Havana for carrying on a rigid fight along anti-mosquito lines. The +densely populated city was unprepared, the public had to be educated, +and an efficient organization built up. The local authorities actively +began a general fight against the mosquito but in spite of their best +efforts the disease continued to spread. It was recognized that more +rigid organization was needed and on August 12th the United States +Public Health and Marine Hospital Service was put in absolute charge of +the fight. Up to this time there had been one hundred and forty-two +deaths from a total of nine hundred and thirteen cases and all of the +conditions seemed to threaten an outbreak to exceed the memorable one of +1878 when, as we have seen there were four thousand and forty-six +deaths. + +With the hearty cooperation of the citizens,--physicians and laymen +alike,--the fight was waged and long before frost or any near approach +thereto the disease was stamped out,--a thing unheard of in previous +epidemics. The total loss of life was four hundred and sixty--about 11 +per cent as great as that from the comparable epidemic of 1878. If the +disease had been promptly recognized and combated with the energy which +marked the fight later in the summer, the outbreak would have made +little headway and the great proportion of these lives would have been +saved. + + + + +CHAPTER IX + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA (Continued) + + +INSECTS AND TRYPANOSOMIASES + +By trypanosomiasis is meant a condition of animal parasitism, common to +man and the lower animals, in which trypanosomes, peculiar flagellate +protozoa, infest the blood. Depending upon the species, they may be +harmless, producing no appreciable ill-effect, or pathogenic, giving +rise to conditions of disease. A number of these are known to be +transferred by insects. + +In order that we may consider more fully the developmental stage of +these parasites within their insect host, it is necessary that we +describe briefly the structure of the blood-inhabiting stage. + +[Illustration: 136. Trypanosoma brucei. After Bruce.] + +The trypanosomes are elongated, usually pointed, flagellated protozoa +(fig. 136) in which the single flagellum, bent under the body, forms the +outer limit of a delicate undulating membrane. It arises near one end of +the organism from a minute centrosome-like body which is known as the +blepheroplast, and at the opposite end extends for a greater or less +distance as a free flagellum. Enclosing, or close beside the +blepheroplast is the small kinetonucleus. The principal nucleus, round +or oval in form, is situated near the center of the body. Asexual +reproductions occurs in this stage, by longitudinal fission, the nucleus +and the blepheroplast dividing independently of one another. From the +blepheroplast of one of the daughter cells a new flagellum is formed. + +Among the pathogenic species are to be found the causative organisms of +some of the most serious diseases of domestic animals and even of man. +It is probable that these pathogenic species secrete a specific poison. +The majority of them are tropical in distribution. + +Though we are concerned especially with the species which infest man, we +shall first consider two of the trypanosomes of lower animals, known +long before any of those of man had been found. + + +FLEAS AND LICE AS CARRIERS OF TRYPANOSOMA LEWISI.--_Trypanosoma lewisi_, +the first mammalian trypanosome known, is to be found in the blood of +wild rats. Like its host, it appears to be cosmopolitan in distribution, +having been reported from several localities in the United States, +Brazil, Argentine, England, Germany, France, Italy, Russia, Asia and +Africa. + +This species is usually regarded as non-pathogenic, but in experimental +work, especially with white rats, heavy infestations often result +fatally, and naturally infested specimens sometimes show evidence of +injury. Rats which have been infested exhibit at least temporary +immunity against new infection. + +_Trypanosoma lewisi_ is transmitted from rat to rat by fleas and by +lice. Rabinowitsch and Kempner (1899) first found that healthy rats +which were kept with infested rats, showed trypanosomes in their blood +after about two weeks. They found the trypanosomes in the alimentary +canal of fleas which had fed on the diseased rats. On teasing such fleas +in physiological salt solution and inoculating them into fresh rats they +were able to produce the infection. Finally, they showed that the fleas +which had fed upon infested rats were able to carry the parasites to +healthy rats. Corresponding experiments with lice were not successful. +Prowazek (1905) found in the rat louse (_Haematopinus spinulosus_) +organisms which he regarded as developmental stages of the _Trypanosoma +lewisi_. He believed that the sexual cycle was undergone in this insect. + +Nuttall (1908) readily transmitted the trypanosomes through the agency +of fleas, (_Ceratophyllus fasciatus_ and _Ctenopthalmus agyrtes_). He +believes that these insects are probably the chief transmitters of the +parasite. He was also able to transmit it from diseased to healthy rats +through the agency of the rat louse. He was unable to trace any +developmental stages in the louse and inclined to the opinion that +Prowazek was deceived by the presence of extraneous flagellates such as +are known to exist in a number of blood-sucking arthropods. + +Nuttall concludes that since three distinct kinds of blood-sucking +insects are capable of transmitting _Trypanosoma lewisi_ it appears +doubtful that this flagellate is a parasite of the invertebrate "host" +in the sense claimed by Prowazek and other investigators. + + +TSETSE-FLIES AND NAGANA--One of the greatest factors in retarding the +development of certain regions of Africa has been the presence of a +small fly, little larger than the common house-fly. This is the +tsetse-fly, _Glossina morsitans_ (fig. 165) renowned on account of the +supposed virulence of its bite for cattle, horses and other domestic +mammals. + +The technical characteristics of the tsetse-flies, or Glossinas, and +their several species, will be found in a later chapter. We need +emphasize only that they are blood-sucking Muscidae and that, unlike the +mosquitoes, the sexes resemble each other closely in structure of the +mouth-parts, and in feeding habits. + +In 1894, Colonel David Bruce discovered that the fly was not in itself +poisonous but that the deadly effect of its bite was due to the fact +that it transmitted a highly pathogenic blood parasite, _Trypanosoma +brucei_. This trypanosome Bruce had discovered in the blood of South +African cattle suffering from a highly fatal disease known as "nagana". +On inoculating the blood of infected cattle into horses and dogs he +produced the disease and found the blood teeming with the causative +organism. In the course of his work he established beyond question that +the "nagana" and the tsetse-fly disease were identical. + +Tsetse-flies of the species _Glossina morsitans_, which fed upon +diseased animals, were found capable of giving rise to the disease in +healthy animals up to forty-eight hours after feeding. Wild tsetse-flies +taken from an infected region to a region where they did not normally +occur were able to transmit the disease to healthy animals. It was found +that many of the wild animals in the tsetse-fly regions harbored +_Trypanosoma brucei_ in their blood, though they showed no evidence of +disease. As in the case of natives of malarial districts, these animals +acted as reservoirs of the parasite. Non-immune animals subjected to the +attacks of the insect carrier, quickly succumbed to the disease. + +A question of prime importance is as to whether the insect serves as an +essential host of the pathogenic protozoan or whether it is a mere +mechanical carrier. Bruce inclined to the latter view. He was unable to +find living trypanosomes in the intestines or excrements of the fly or +to produce the disease on the many occasions when he injected the +excrement into healthy animals. Moreover, he had found that the +experimental flies were infective only during the first forty-eight +hours and that if wild flies were taken from the infected region, "kept +without food for three days and then fed on a healthy dog, they never +gave rise to the disease." + +Koch had early described what he regarded as sexual forms from the +intestine of the fly but it remained for Kleine (1909) to experimentally +demonstrate that a part of the life cycle of the parasite was undergone +in the fly. Working with _Glossina palpalis_, he found that for a period +of ten days or longer after feeding on an animal suffering from nagana +it was non-infective, but that then it became infective and was able to +transmit the disease for weeks thereafter. He discovered and described +developmental stages of the parasite within the intestine of the insect. +In other words, the tsetse-fly (in nature, _Glossina morsitans_), serves +as an essential host, within which an important part of the life cycle +of the parasite is undergone. These conclusions were quickly verified by +Bruce and numerous other workers and are no longer open to question. +Klein and Taute are even inclined to think that mechanical transmission +plays practically no role in nature, unless the fly is interrupted while +feeding and passes immediately to a new animal. + + +TSETSE-FLIES AND SLEEPING SICKNESS OF MAN--About the beginning of the +present century a hitherto little known disease of man began to attract +great attention on account of its ravages in Uganda and the region of +Victoria Nyanza in South Africa. It was slow, insiduous and absolutely +fatal, characterized in its later stages by dullness, apathy, and +finally absolute lethargy all day long, symptoms which gave it the name +of "sleeping sickness." + +It was soon found that the disease was not a new one but that it had +been known for over a hundred years on the west coast of Africa. Its +introduction into Central and East Africa and its rapid spread have been +attributed primarily to the development of the country, the formation of +new trade routes and the free mingling of native tribes formerly +isolated. It is estimated that in the first ten years of the present +century there were approximately two hundred thousand deaths from the +disease in the Uganda protectorate. In the British province Bugosa, on +the Victoria Nyanza there were thirty thousand deaths in the period from +1902-1905. + +While the disease is peculiarly African there are a number of instances +of its accidental introduction into temperate regions. Slaves suffering +from it were occasionally brought to America in the early part of the +last century and cases have sometimes been imported into England. In +none of the cases did the disease gain a foothold or spread at all to +other individuals. + +In 1902 Dutton described a trypanosome, _T. gambiense_, which he and +Forde had found the year before in the blood of a patient suffering from +a peculiar type of fever in Gambia. In 1902-1903 Castellani found the +same parasite in the cerebro-spinal fluid of sleeping-sickness patients +and definitely reported it as the causative organism of the disease. His +work soon found abundant confirmation, and it was discovered that the +sleeping sickness was but the ultimate phase of the fever discovered by +Dutton and Forde. + +When Castellani made known his discovery of the trypanosome of sleeping +sickness, Brumpt, in France, and Sambon, in England, independently +advanced the theory that the disease was transmitted by the tsetse-fly, +_Glossina palpalis_. This theory was based upon the geographical +distribution and epidemiology of the disease. Since then it has been +abundantly verified by experimental evidence. + +Fortunately for the elucidation of problems relating to the methods of +transfer of sleeping sickness, _Trypanosoma gambiense_ is pathogenic for +many species of animals. In monkeys it produces symptoms very similar to +those caused in man. Bruce early showed that _Glossina palpalis_ "fed on +healthy monkeys eight, twelve, twenty-four and forty-eight hours after +having fed on a native suffering from trypanosomiasis, invariably +transmitted the disease. After three days the flies failed to transmit +it." In his summary in Osler's Modern Medicine, he continues "But this +is not the only proof that these flies can carry the infective agent. On +the lake shore there was a large native population among whom we had +found about one-third to be harboring trypanosomes in their blood. The +tsetse-flies caught on this lake shore, brought to the laboratory in +cages, and placed straightway on healthy monkeys, gave them the disease +in every instance, and furnished a startling proof of the danger of +loitering along the lake shore among those infected flies." + +As in the case of nagana, Bruce and most of the earlier investigators +supposed the transmission of the sleeping sickness trypanosome by +_Glossina palpalis_ to be purely mechanical. The work of Kleine (1909) +clearly showed that for _Trypanosoma gambiense_ as well as for +_Trypanosoma brucei_ the fly served as an essential host. Indeed, Kleine +and many subsequent investigators are inclined to think that there is +practically no mechanical transmission of trypanosomes from animal to +animal by _Glossina_ in nature, and that the many successful experiments +of the earlier investigators were due to the fact that they used wild +flies which already harbored the transformed parasite rather than +directly inoculated it from the blood of the diseased experimental +animals. While the criticism is applicable to some of the work, this +extreme view is not fully justified by the evidence at hand. + +Kleine states (1912) that _Glossina palpalis_ can no longer be regarded +as the sole transmitter of sleeping sickness. Taute (1911) had shown +that under experimental conditions _Glossina morsitans_ was capable of +transferring the disease and Kleine calls attention to the fact that in +German East Africa, in the district of the Rovuma River, at least a +dozen cases of the disease have occurred recently, though only _Glossina +morsitans_ exists in the district. It appears, however, that these cases +are due to a different parasite, _Trypanosoma rhodesiense_. This +species, found especially in north-east Rhodesia and in Nyassaland, is +transferred by _Glossina morsitans_. + +Other workers maintain that the disease may be transmitted by various +blood-sucking flies, or even bugs and lice which attack man. Fulleborn +and Mayer (1907) have shown by conclusive experiments that _Aedes +(Stegomyia) calopus_ may transmit it from one animal to another if the +two bites immediately succeed each other. + +It is not possible that insects other than the tsetse-flies (and only +certain species of these), play an important role in the transmission of +the disease, else it would be much more wide-spread. Sambon (1908) +pointed out that the hypothesis that is spread by _Aedes calopus_ is +opposed by the fact that the disease never spread in the Antilles, +though frequently imported there by West African slaves. The same +observation would apply also to conditions in our own Southern States in +the early part of the past century. + +Since _Glossina palpalis_ acts as an essential host of the parasite and +the chief, if not the only, transmitter, the fight against sleeping +sickness, like that against malaria and yellow fever, becomes primarily +a problem in economic entomology. The minutest detail of the +life-history, biology, and habits of the fly, and of its parasites and +other natural enemies becomes of importance in attempts to eradicate the +disease. Here we can consider only the general features of the subject. + +_Glossina palpalis_ lives in limited areas, where the forest and +undergrowth is dense, along the lake shore or river banks. According to +Hodges, the natural range from shore is under thirty yards, though the +distance to which the flies may follow man greatly exceed this. + +It is a day feeder, a fact which may be taken advantage of in avoiding +exposure to its attacks. The young are brought forth alive and +full-grown, one every nine or ten days. Without feeding, they enter the +ground and under favorable conditions, complete their development in a +month or more. + +[Illustration: 137. Sleeping sickness concentration camp in German East +Africa. Report of German Commission.] + +Methods of control of the disease must look to the prevention of +infection of the flies, and to their avoidance and destruction. Along +the first line, much was hoped from temporary segregation of the sick in +regions where the fly was not found. On the assumption that the flies +acted as carriers only during the first two or three days, it was +supposed that even the "fly belts" would become safe within a few days +after the sick were removed. The problem was found to be a much more +difficult one when it was learned that after a given brief period the +fly again became infective and remained so for an indeterminate period. +Nevertheless, isolation of the sick is one of the most important +measures in preventing the spread of the disease into new districts. +Much, too, is being accomplished by moving native villages from the fly +belts. (c.f. fig. 137.) + +All measures to avoid the flies should be adopted. This means locating +and avoiding the fly belts as far as possible, careful screening of +houses, and protection of the body against bites. + +Clearing the jungle along the water courses for some yards beyond the +natural range of the fly has proved a very important measure. Castellani +recommends that the area be one hundred yards and around a village three +hundred yards at least. + +Detailed studies of the parasites and the natural enemies of the +tsetse-fly are being undertaken and may ultimately yield valuable +results. + + +SOUTH AMERICAN TRYPANOSOMIASIS--The tsetse-flies are distinctively +African in distribution and until recently there were no trypanosomes +known to infest man in America. In 1909 Dr. Chagas, of Rio de Janeiro +described a new species, _Trypanosoma cruzi_, pathogenic to man. + +_Trypanosoma cruzi_ is the causative organism of a disease common in +some regions of Brazil, where it is known as "opilacao." It is +especially to be met with in children and is characterized by extreme +anemia, wasting, and stunted development associated with fever, and +enlargement of the thyroid glands. The disease is transmitted by the +bites of several species of assassin-bugs, or Reduviidae, notably by +_Conorhinus megistus_. The evolution of the parasite within the bug has +been studied especially by Chagas and by Brumpt. From the latter's text +we take the following summary. + +The adult trypanosomes, ingested by a _Conorhinus megistus_, of any +stage, first change into Crithidia-like forms and then those which +remain in the stomach become ovoid and non-motile. Brumpt found these +forms in immense numbers, in a _Conorhinus_ which had been infested +fourteen months before. The forms which pass into the intestine quickly +assume the _Crithidia_ form and continue to develop rapidly under this +form. Some weeks later they evolve into the trypanosome forms, +pathogenic for man. They then pass out with the excrement of the bug and +infect the vertebrate host as soon as they come in contact with any +mucous layer (buccal, ocular or rectal). More rarely they enter through +the epidermis. + +Brumpt showed that the development could take place in three species; +bed-bugs (_Cimex lectularius_, _C. hemipterus_) and in the tick +_Ornithodoros moubata_. The evolution proceeds in the first two species +of bed-bugs as rapidly as in Conorhinus, or even more rapidly, but they +remain infective for a much shorter time and hence Brumpt considers that +they play a much less important role in the spread of the disease. + +_Conorhinus megistus_, like related forms in our Southern States, very +commonly frequents houses and attacks man with avidity. Chagas states +that the bites are painless and do not leave any traces. They are +usually inflicted on the lips, or the cheeks and thus the buccal mucosa +of a sleeper may be soiled by the dejections of the insect and the bite +serving as a port of entry of the virus, remain unnoticed. + +The possibility of some of our North American Reduviidae playing a +similar role in the transmission of disease should not be overlooked. + + +LEISHMANIOSES AND INSECTS--Closely related to the trypanosomes is a +group of intracellular parasites which have recently been grouped by +Ross under the genus _Leishmania_. Five species are known to affect man. +Three of these produce local skin infestations, but two of them, +_Leishmania donovani_ and _L. infantum_, produce serious and often fatal +systemic diseases. + +The first of these, that produced by _L. donovani_, is an exceedingly +virulent disease common in certain regions of India and China. It is +commonly known as "Kala-azar," or "dum-dum" fever, and more technically +as tropical leishmaniasis. Patton (1907) believes that the parasite is +transmitted by the bed-bug _Cimex hemipterus_, and has described a +developmental cycle similar to that which can be found in artificial +cultures. On the other hand, Donovan was unable to confirm Patton's work +and believes that the true intermediate host is a Reduviid bug, +_Conorhinus rubrofasciatus_. + +_Leishmania infantum_ is the cause of the so-called infantile splenic +leishmaniasis, occurring in northern Africa, Spain, Portugal, Italy, and +possibly other parts of Europe. The parasite occurs habitually in the +dog and is only accidentally transferred to children. Alvares and da +Silva, in Portugal (according to Brumpt, 1913) have found that the +excrement of a flea from a diseased dog contains flagellates, and they +suggest that the infection may be transmitted by the accidental +inoculation of this excrement by means of the proboscis of the flea, as +has been thought to occur in the case of the plague. To this Brumpt +objects that they and other workers who thought to trace the development +of _Leishmania infantum_ were apparently misled by the presence of a +harmless _Herpetomonas_ which infests dog fleas in all countries, even +where the leishmaniasis is unknown. + +Basile (1910 and 1911) however, carried on numerous experiments +indicating that the disease was transferred from children to dogs and +from dog to dog by the dog flea, and was able to find in the tissues of +the insects forms perfectly identical with those found in children and +in dogs suffering from leishmaniasis. He also found that _Pulex +irritans_ was capable of acting as the carrier. + +Of the cutaneous type of leishmaniasis, the best known is the so-called +"Oriental sore," an ulcerative disease of the skin which is epidemic in +many tropical and subtropical regions. The causative organism is +_Leishmania tropica_, which occurs in the diseased tissues as bodies +very similar to those found in the spleen in cases of Kala-azar. The +disease is readily inoculable and there is no doubt that it may be +transferred from the open sores to abraded surfaces of a healthy +individual by house-flies. It is also believed by a number of +investigators that it may be transferred and directly inoculated by +various blood-sucking insects. + + +TICKS AND DISEASES OF MAN AND ANIMALS + +We have seen that the way to the discoveries of the relations of +arthropods to disease was pointed out by the work of Leuckart and +Melnikoff on the life cycle of _Dipylidium_, and of Fedtschenko and +Manson on that of _Filaria_. They dealt with grosser forms, belonging to +well-recognized parasitic groups. + +This was long before the role of any insect as a carrier of pathogenic +micro-organisms had been established, and before the Protozoa were +generally regarded as of importance in the causation of disease. The +next important step was taken in 1889 when Smith and Kilbourne +conclusively showed that the so-called Texas fever of cattle, in the +United States, is due to an intracorpuscular blood parasite transmitted +exclusively by a tick. This discovery, antedating by eight years the +work on the relation of the mosquito to malaria, had a very great +influence on subsequent studies along these lines. + +While much of the recent work has dealt with the true insects, or +hexapods, it is now known that several of the most serious diseases of +animals, and at least two important diseases of man are tick borne. +These belong to the types known collectively as _babesioses_ (or +"_piroplasmoses_"), and _spirochaetoses_. + +The term _babesiosis_ is applied to a disease of man or animals which is +caused by minute protozoan parasites of the genus _Babesia_, living in +the red blood corpuscles. These parasites have usually been given the +generic name _Piroplasma_ and hence the type of disease which they cause +is often referred to as "_piroplasmosis_." The best known illustration +is the disease known in this country as Texas fever of cattle. + + +CATTLE TICKS AND TEXAS FEVER--The cattle disease, which in the United +States is known as Texas fever, is a widely distributed, exceedingly +acute disease. In Australia it is known as _redwater fever_ and in +Europe as haemoglobinuria, due to the fact that the urine of the diseased +animals is discolored by the breaking down of the red blood corpuscles +infested by the parasite. + +In their historical discussion, Smith and Kilbourne, point out that as +far back as 1796 it was noted that Southern cattle, in a state of +apparent health, might spread a fatal disease among Northern herds. As +observations accumulated, it was learned that this infection was carried +only during the warm season of the year and in the depth of winter +Southern cattle were harmless. Moreover, Southern cattle after remaining +for a short time in the North lost their power to transmit the disease, +and the same was true of cattle which had been driven for a considerable +distance. + +Very significant was the fact that the infection was not communicated +directly from the Southern to Northern cattle but that the ground over +which the former passed was infected by them, and that the infection was +transmitted thence to susceptible cattle _after a period of not less +than thirty days had elapsed_. + +Of course a disease as striking as this, and which caused such enormous +losses of cattle in the region invaded was fruitful in theories +concerning its causation. The most widespread was the belief that +pastures were infected by the saliva, urine, or manure of Southern +cattle. There were not wanting keen observers who suggested that the +disease was caused by ticks, but little weight was given to their view. + +Various workers had described bacteria which they had isolated from the +organs of the diseased animals, but their findings could not be +verified. In 1889, Smith and Kilbourne discovered a minute, pear-shaped +organism (fig. 138) in the red blood corpuscles of a cow which had +succumbed to Texas fever. On account of their shape they were given the +generic name _Pyrososma_ and because they were usually found two in a +corpuscle, the specific name, _bigeminum_. It is now generally accepted +that the parasite is the same which Babes had observed the year before +in Roumanian cattle suffering from haemoglobinuria, and should be known +as _Babesia bovis_ (Babes). + +[Illustration: 138. Babesia bovis in blood corpuscles. After Calli.] + +[Illustration: 139. The cattle tick (Boophilus annulatus). (_a_) Female; +(_b_) male. After Comstock.] + +By a series of perfectly conclusive experiments carried on near +Washington, D.C., Smith and Kilbourne showed that this organism was +carried from Southern cattle to non-immune animals by the so-called +Southern cattle tick, _Boophilus annulatus_ (= _Margaropus annulatus_) +(fig. 139). + +Of fourteen head of native cattle placed in a field with tick-infested +Northern cattle all but two contracted the disease. This experiment was +repeated with similar results. Four head of native cattle kept in a plot +with three North Carolina cattle which had been carefully freed from +ticks remained healthy. A second experiment the same year gave similar +results. + +Still more conclusive was the experiment showing that fields which had +not been entered by Southern cattle but which had been infected by +mature ticks taken from such animals would produce Texas fever in native +cattle. On September 13, 1889, several thousand ticks collected from +cattle in North Carolina three and four days before, were scattered in a +small field near Washington. Three out of four native animals placed in +this field contracted the disease. The fourth animal was not examined as +to its blood but it showed no external symptoms of the disease. + +[Illustration: 140. Hyalomma aegypticum. After Nuttall and Warburton.] + +In these earlier experiments it was believed that the cattle tick acted +as a carrier of the disease between the Southern cattle and the _soil_ +of the Northern pastures. "It was believed that the tick obtained the +parasite from the blood of its host and in its dissolution on the +pasture a certain resistant spore form was set free which produced the +disease when taken in with the food." The feeding of one animal for some +time with grass from the most abundantly infected field, without any +appearance of the disease, made this hypothesis untenable. + +In the experimental work in 1890 the astonishing fact was brought out +that the disease was conveyed neither by infected ticks disintegrating +nor by their directly transferring the parasite, but that it was +conveyed by the young hatched from eggs of infected ticks. In other +words, the disease was hereditarily transferred to ticks of the second +generation and they alone were capable of conveying it. + +Thus was explained the fact that Texas fever did not appear immediately +along the route of Southern cattle being driven to Northern markets but +that after a certain definite period it manifested itself. It was +conveyed by the progeny of ticks which had dropped from the Southern +cattle and deposited their eggs on the ground. + +These results have been fully confirmed by workers in different parts of +the world,--notably by Koch, in Africa, and by Pound, in Australia. + +The disease is apparently transmitted by _Boophilus annulatus_ alone, in +the United States, but it, or an almost identical disease, is conveyed +by _Ixodes hexagonus_ in Norway, _Ixodes ricinus_ in Finland and France +and by the three species, _Boophilus decoloratus_, _Hyalomma aegypticum_ +(fig. 140 and 141), and _Haemaphysalis punctata_ in Africa. + +[Illustration: 141. Hyalomma aegypticum. Capitulum of female; (_a_) +dorsal, (_b_) ventral aspect.] + +In spite of the detailed study which it has received, the life cycle of +_Babesia bovis_ has not been satisfactorily worked out. The asexual +reproduction in the blood of the vertebrate host has been described but +the cycle in the tick is practically unknown. + +More successful attempts have been made to work out the life cycle of a +related species, _Babesia canis_, which causes malignant jaundice in +dogs in Africa and parts of Southern Europe. In this instance, also, the +disease is transmitted by heredity to the ticks of the second +generation. Yet the larval, or "seed ticks," from an infected female are +not capable of conveying the disease, but only the nymphs and adults. +Still more complicated is the condition in the case of _Babesia ovis_ of +sheep, which Motas has shown can be conveyed solely by the adult, +sexually mature ticks of the second generation. + +In _Babesia canis_, Christopher (1907) observed developmental stages in +the tick. He found in the stomach of adult ticks, large motile +club-shaped bodies which he considered as ookinetes. These bodies pass +to the ovaries of the tick and enter the eggs where they become globular +in form and probably represent an oocyst. This breaks up into a number +of sporoblasts which enter the tissues of the developing tick and give +rise to numerous sporozoites, which collect in the salivary glands and +thence are transferred to the vertebrate host. A number of other species +of _Babesia_ are known to infest vertebrates and in all the cases where +the method has been worked out it has been found that the conveyal was +by ticks. We shall not consider the cases more fully here, as we are +concerned especially with the method of transfer of human diseases. + + +TICKS AND ROCKY MOUNTAIN SPOTTED FEVER OF MAN--Ever since 1873 there has +been known in Montana and Idaho a peculiar febrile disease of man, which +has gained the name of "Rocky Mountain spotted fever." Its onset is +marked by chills and fever which rapidly become acute. In about four to +seven days there appears a characteristic eruption on the wrists, ankles +or back, which quickly covers the body. + +McClintic (1912) states that the disease has now been reported from +practically all of the Rocky Mountain States, including Arizona, +California, Colorado, Idaho, Montana, Nevada, Oregon, Utah, Washington, +and Wyoming. "Although the disease is far more prevalent in Montana and +Idaho than in any of the other States, its spread has assumed such +proportions in the last decade as to call for the gravest consideration +on the part of both the state and national health authorities. In fact, +the disease has so spread from state to state that it has undoubtedly +become a very serious interstate problem demanding the institution of +measures for its control and suppression." + +A peculiar feature of the Rocky Mountain spotted fever is a marked +variation in its severity in different localities. In Montana, and +especially in the famous Bitter Root Valley, from 33 per cent to 75 per +cent of the cases result fatally. On the other hand, the fatality does +not exceed four per cent in Idaho. + +In 1902, Wilson and Chowning reported the causative organism of spotted +fever to be a blood parasite akin to the _Babesia_ of Texas fever, and +made the suggestion that the disease was tick-borne. The careful studies +of Stiles (1905) failed to confirm the supposed discovery of the +organism, and the disease is now generally classed as due to an +invisible virus. On the other hand, the accumulated evidence has fully +substantiated the hypothesis that it is tick-borne. + +According to Ricketts (1907) the experimental evidence in support of +this hypothesis was first afforded by Dr. L. P. McCalla and Dr. H. A. +Brereton, in 1905. These investigators transmitted the disease from man +to man in two experiments. "The tick was obtained 'from the chest of a +man very ill with spotted fever' and 'applied to the arm of a man who +had been in the hospital for two months and a half, and had lost both +feet from gangrene due to freezing.' On the eighth day the patient +became very ill and passed through a mild course of spotted fever, +leaving a characteristic eruption. The experiment was repeated by +placing the tick on a woman's leg and she likewise was infected with +spotted fever." + +The most detailed studies were those of the late Dr. H. T. Ricketts, and +it was he who clearly established the tick hypothesis. In the summer of +1906 he found that guinea pigs and monkeys are very susceptible to +spotted fever and can readily be infected by inoculation of blood from +patients suffering from the disease. This opened the way to experimental +work on tick transmission. A female tick was fed upon an infected guinea +pig for two days, removed and isolated for two days and then placed upon +a healthy guinea pig. After an incubation period of three and a half +days the experimental animal contracted a well-marked case of the +disease. + +A similar result was obtained at the same time by King, and later in the +season Ricketts proved that the male tick was also capable of +transmitting the disease. He found that there was a very intimate +relation of the virus to the tick and that the transmission must be +regarded as biological throughout. Ticks remained infective as long as +they lived and would feed for a period of several months. If they +acquired the disease in the larval or nymphal stage they retained it +during molting and were infective in the subsequent stages. In a few +cases the larvae from an infected female were infective. + +The evidence indicated that the tick suffers from a relatively harmless, +generalized infection and the virus proliferates in its body. The +disease probably is transferred through the salivary secretion of the +tick since inoculation experiments show that the salivary glands of the +infected adult contain the virus. + +It is probable that in nature the reservoir of the virus of spotted +fever is some one or more of the native small animals. Infected ticks +have been found in nature, and as various wild animals are susceptible +to the disease, it is obvious that it may exist among them unnoticed. +Wilson and Chowning suggested that the ground squirrel plays the +principal role. + +Unfortunately, much confusion exists regarding the correct name of the +tick which normally conveys the disease. In the medical literature it is +usually referred to as _Dermacentor occidentalis_, but students of the +group now agree that it is specifically distinct. Banks has designated +it as _Dermacentor venustus_ and this name is used in the publications +of the Bureau of Entomology. On the other hand, Stiles maintains that +the common tick of the Bitter Root Valley, and the form which has been +collected by the authors who have worked on Rocky Mountain spotted fever +in that region, is separable from _D. venustus_, and he has described it +under the name of _Dermacentor andersoni_. + +Mayer (1911) has shown experimentally that spotted fever may be +transmitted by several different species of ticks, notably _Dermacentor +marginatus_, _Dermacentor variabilis_ and _Amblyomma americanum_. This +being the case, the question of the exact systematic status of the +species experimented upon in the Bitter Root Valley becomes less +important, for since _Dermacentor occidentalis_, _Dermacentor venustus_ +and _Dermacentor andersoni_ all readily attack man, it is probable that +either species would readily disseminate the disease if it should spread +into their range. + +Hunter and Bishop (1911) have emphasized the fact that in the eastern +and southern United States there occur several species which attack man, +and any one of which might transmit the disease from animal to animal +and from animal to man. The following species, they state, would +probably be of principal importance in the Southern and Eastern States: +the lone star tick (_Amblyomma americanum_); the American dog tick +(_Dermacentor variabilis_); and the gulf-coast tick (_Amblyomma +maculatum_). In the extreme southern portions of Texas, _Amblyomma +cajennense_, is a common pest of man. + +Since the evidence all indicates that Rocky Mountain spotted fever is +transmitted solely by the tick, and that some of the wild animals serve +as reservoirs of the virus, it is obvious that personal prophylaxis +consists in avoiding the ticks as fully as possible, and in quickly +removing those which do attack. General measures along the line of tick +eradication must be carried out if the disease is to be controlled. That +such measures are feasible has been shown by the work which has been +done in controlling the tick-borne Texas fever of cattle, and by such +work as has already been done against the spotted fever tick, which +occurs on both wild and domestic animals. Detailed consideration of +these measures is to be found in the publications of the Public Health +and Marine Hospital Service, and the Bureau of Entomology. Hunter and +Bishopp give the following summarized recommendations for control or +eradication measures in the Bitter Root Valley. + +(1) A campaign of education, whereby all the residents of the valley +will be made thoroughly familiar with the feasibility of the plan of +eradication, and with what it will mean in the development of the +valley. + +(2) The obtaining of legislation to make it possible to dip or oil all +live stock in the Bitter Root Valley. + +(3) The obtaining of an accurate census of the horses, cattle, sheep, +mules, and dogs in the valley. + +(4) The construction of ten or more dipping vats. + +(5) The providing of materials to be used in the dipping mixture. + +(6) The organization of a corps of workers to carry on the operations. + +(7) The systematic dipping of the horses, cattle, sheep, and dogs of the +valley on a definite weekly schedule from approximately March 10 to June +9. + +(8) The treatment by hand of the animals in localities remote from vats, +on the same schedule. + +They estimate that after three seasons' operations a very small annual +expenditure would provide against reinfestation of the valley by the +incoming of cattle from other places. + +Supplementary measures consist in the killing of wild mammals which may +harbor the tick; systematic burning of the brush and debris on the +mountain side; and in clearing, since the tick is seldom found on land +under cultivation. + + + + +CHAPTER X + +ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA [_Continued_] + + +ARTHROPODS AND SPIROCHAETOSES OF MAN AND ANIMALS + +The term spirochaetoses is applied to diseases of man or animals which +are due to protistan parasites belonging to the group of slender, spiral +organisms known as spirochaetes. + +There has been much discussion concerning the relationship Of the +spirochaetes. Formerly, they were regarded as bacteria closely related to +the forms grouped in the genus _Spirillum_. The results of the detailed +study which the spirochaetes have received in recent years, have led most +of the workers to consider them as belonging to the protozoa. The merits +of the discussion we are not concerned with here, but rather with the +fact that a number of diseases caused by spirochaetes are +arthropod-borne. The better known of these we shall discuss. + + +AFRICAN RELAPSING FEVER OF MAN--It has long been known to the natives of +Africa and to travelers in that country, that the bite of a certain +tick, _Ornithodoros moubata_, may be followed by severe or even fatal +fever of the relapsing type. Until recent years, it was supposed that +the effect was due to some special virulence of the tick, just as nagana +of cattle was attributed to the direct effect of the bite of the +tsetse-fly. The disease is commonly known as "tick-fever" or by the +various native names of the tick. + +In 1904, Ross and Milne, in Uganda, and Dutton and Todd on the Congo, +discovered that the cause of the disease is a spirochaete which is +transmitted by the tick. This organism has been designated by Novy and +Knapp as _Spirochaeta duttoni_. + +_Ornithodoros moubata_ (fig. 142), the carrier of African relapsing +fever, or "tick-fever," is widely distributed in tropical Africa, and +occurs in great numbers in the huts of natives, in the dust, cracks and +crevices of the dirt floors, or the walls. It feeds voraciously on man +as well as upon birds and mammals. Like others of the _Argasidae_, it +resembles the bed-bug in its habit of feeding primarily at night. Dutton +and Todd observed that the larval stage is undergone in the egg and that +the first free stage is that of the octopod nymph. + +[Illustration: 142. Ornithodoros moubata. (_a_) Anterior part of venter; +(_b_) second stage nymph; (_c_) capitulum; (_d_) dorsal and (_e_) +ventral aspect of female; (_f_) ventral aspect of nymph; (_g_) capitulum +of nymph. After Nuttall and Warburton.] + +The evidence that the fever is transmitted by this tick is conclusive. +Koch found that from five per cent to fifteen per cent, and in some +places, fifty per cent of the ticks captured, harbored the spirochaete. +The disease is readily transmitted to monkeys, rats, mice and other +animals and the earlier experiments along these lines have been +confirmed by many workers. + +Not only are the ticks which have fed on infected individuals capable of +conveying the disease to healthy animals but they transmit the causative +organism to their progeny. Thus Mollers (1907), working in Berlin, +repeatedly infected monkeys through the bites of nymphs which had been +bred in the laboratory from infected ticks. Still more astonishing was +his discovery that ticks of the third generation were infective. In +other words, if the progeny of infected ticks were fed throughout life +on healthy animals, and on maturity deposited eggs, the nymphs which +hatched from these eggs would still be capable of carrying the +infection. + +The developmental cycle of the spirochaete within the tick has not been +fully worked out, though the general conclusions of Leishman (1910) have +been supported by the recent works of Balfour (1911 and 1912), and +Hindle (1912), on the life cycle of spirochaetes affecting fowls. + +_Spirochaeta duttoni_ ingested by _Ornithodoros moubata_ apparently +disappear within a few days, but Leishman believed that in reality they +break up into minute granules which are to be found in the alimentary +canal, the salivary glands and the Malpighian tubes of the tick. These +granules, or "coccoid bodies," as Hindle calls them, are supposed to be +the form in which the spirochaetes infect the new host. We shall see +later that Marchoux and Couvy (1913) dissent wholly from this +interpretation. + +According to Leishman, and Hindle, the coccoid bodies are not injected +into the vertebrate host with the saliva of the tick, as are the +sporozoites of malaria with that of the mosquito. Instead, they pass out +with the excrement and secondarily gain access to the wound inflicted by +the tick. + +Nuttall (1912) calls attention to the fact that the geographical +distribution of _Ornithodoros moubata_ is far wider than our present +records show for the distribution of the relapsing fever in man and that +there is every reason to fear the extension of the disease. Huts where +the ticks occur should be avoided and it should be remembered that in +infected localities there is special danger in sleeping on the ground. + + +EUROPEAN RELAPSING FEVER--There is widely distributed in Europe a type +of relapsing fever which is caused by _Spirochaeta recurrentis_. It has +long been supposed that this disease is spread by the bed-bug and there +is some experimental evidence to show that it may be conveyed by these +insects. + +In 1897, Tictin found that he could infect monkeys by inoculating the +contents of bed-bugs which had fed upon a patient within forty-eight +hours. Nuttall, in 1907, in one experiment succeeded in transmitting +_Spirochaeta recurrentis_ from mouse to mouse by bites of bed-bugs. The +bugs, thirty-five in number, were transferred at short intervals from +one mouse to another, not being allowed to take a full meal on the +first, or infected mouse. + +On the other hand, there is much clinical evidence to show that the +European relapsing fever like various other types of the disease is +transmitted from man to man by head and body lice (_Pediculus humanus_ +and _Pediculus corporis_). + +Interesting supplementary evidence is that of Bayon's observations +(1912), in Moscow. "Having visited the big municipal night hospitals at +Moscow I soon noticed that they were kept with such scrupulous +cleanliness, disinfected so lavishly, the beds of iron, the floor +cemented, that it was not possible for bed-bugs to thrive to any extent +on the premises. The people sleeping there were allowed, however, to +sleep in their own clothes. The introduction of these model homes had +not had any effect on the incidence of relapsing fever, for the places +were still hot-beds of the fever during winter. On the other hand, +though I changed my rooms several times, I found bugs in every +successive lodging, and I was told in Moscow, this can hardly be +avoided. Yet no foreigner, or Russian of the better class, ever catches +relapsing fever. To this may be added the fact that when I asked for +clothes-lice and promised to pay a kopec for two, the attendants from +the night hostel brought me next morning a small ounce bottle crammed +with _Pediculus capitis_ (= _P. humanus_), and _Pediculus vestimentorum_ +(= _P. corporis_) collected off the sleepers. If relapsing fever were +transmitted by bed-bugs, it would be much more disseminated than it is +at present in Moscow." + +Direct experimental evidence of the agency of lice in transmitting +relapsing fever is especially clear in the case of a type of the disease +prevalent in parts of North Africa. We shall consider this evidence +later. + + +OTHER TYPES OF RELAPSING FEVER OF MAN--In addition to the three types of +human relapsing fever already referred to, several others have been +distinguished and have been attributed to distinct species of +spirochaetes. The various spirochaetoses of man are: + +African, caused by _S. duttoni_; European, caused by _S. recurrentis_; +North African, caused by _S. berbera_; East African, caused by _S. +rossi_; East Indian, caused by _S. carteri_; North American, caused by +_S. novyi_; South American, caused by _S. duttoni_ (?). + +Nuttall (1912) in his valuable resume of the subject, has emphasized +that "in view of the morphological similarity of the supposedly +different species of spirochaetes and their individual variations in +virulence, we may well doubt if any of the 'species' are valid. As I +pointed out four years ago, the various specific names given to the +spirochaetes causing relapsing fever in man may be used merely for +convenience _to distinguish strains or races_ of different origin. They +cannot be regarded as valid names, in the sense of scientific +nomenclature, for virulence and immunity reactions are not adequate +tests of specificity." + + +NORTH AFRICAN RELAPSING FEVER OF MAN--The type of human relapsing fever +to be met with in Algeria, Tunis, and Tripoli, is due to a _Spirochaeta_ +to which does not differ morphologically from _Spirochaeta duttoni_, but +which has been separated on biological grounds as _Spirochaeta berberi_. + +Experimenting with this type of disease in Algeria, Sergent and Foly +(1910), twice succeeded in transmitting it from man to monkeys by +inoculation of crushed body lice and in two cases obtained infection of +human subjects who had received infected lice under their clothing and +who slept under coverings harboring many of the lice which had fed upon +a patient. Their results were negative with _Argas persicus_, _Cimex +lectularius_, _Musca domestica_, _Haematopinus spinulosus_ and +_Ceratophyllus fasciatus_. They found body lice associated with every +case of relapsing fever which they found in Algeria. + +Nicolle, Blaizot, and Conseil (1912) showed that the louse did not +transmit the parasite by its bite. Two or three hours after it has fed +on a patient, the spirochaetes begin to break up and finally they +disappear, so that after a day, repeated examinations fail to reveal +them. They persist, nevertheless, in some unknown form, for if the +observations are continued they reappear in eight to twelve days. These +new forms are virulent, for a monkey was infected by inoculating a +single crushed louse which had fed on infected blood fifteen days +before. + +Natural infection is indirect. Those attacked by the insect scratch, and +in this act they excoriate the skin, crush the lice and contaminate +their fingers. The least abrasion of the skin serves for the entrance of +the spirochaetes. Even the contact of the soiled fingers on the various +mucosa, such as the conjunctive of the eye, is sufficient. + +As in the case of _Spirochaeta duttoni_, the organism is transmitted +hereditarily in the arthropod vector. The progeny of lice which have fed +on infected blood may themselves be infective. + + +SPIROCHAETOSIS OF FOWLS--One of the best known of the spirochaetes +transmitted by arthropods is _Spirochaeta gallinarum_, the cause of a +very fatal disease of domestic fowls in widely separated regions of the +world. According to Nuttall, it occurs in Southeastern Europe, Asia, +Africa, South America and Australia. + +In 1903, Marchoux and Salimbeni, working in Brazil, made the first +detailed study of the disease, and showed that the causative organism is +transmitted from fowl to fowl by the tick _Argas persicus_. They found +that the ticks remained infective for at least five months. Specimens +which had fed upon diseased birds in Brazil were sent to Nuttall and he +promptly confirmed the experiments. Since that date many investigators, +notably Balfour and Hindle, have contributed to the elucidating of the +life-cycle of the parasite. Since it has been worked out more fully than +has that of any of the human spirochaetes, we present Hindle's diagram +(fig. 143) and quote the brief summary from his preliminary paper +(1911_b_). + +"Commencing with the ordinary parasite in the blood of the fowl, the +spirochaete grows until it reaches a certain length (16-19u) and then +divides by transverse division. This process is repeated, and is +probably the only method of multiplication of the parasite within the +blood. When the spirochaetes disappear from the circulation, some of them +break up into the coccoid bodies which, however, do not usually develop +in the fowl. When the spirochaetes are ingested by _Argas persicus_, some +of them pass through the gut wall into the coelomic fluid. From this +medium they bore their way into the cells of the various organs of the +tick and there break up into a number of coccoid bodies. These +intracellular forms multiply by ordinary fission in the cells of the +Malpighian tubules and gonads. Some of the coccoid bodies are formed in +the lumen of the gut and Malpighian tubules. The result is that some of +the coccoid bodies may be present in the Malpighian secretion and +excrement of an infected tick and when mixed with the coxal fluid may +gain entry into another fowl by the open wound caused by the tick's +bite. They then elongate and redevelop into ordinary spirochaetes in the +blood of the fowl, and the cycle may be repeated." + +[Illustration: 143. Spirochaeta gallinarum. After Hindle.] + +Hindle's account is clear cut and circumstantial, and is quite in line +with the work of Balfour, and of Leishman. Radically different is the +interpretation of Marchoux and Couvy (1913). These investigators +maintain that the granules localized in the Malpighian tubules in the +larvae and, in the adult, also in the ovules and the genital ducts of the +male and female, are not derived from spirochaetes but that they exist +normally in many acariens. They interpret the supposed disassociation +of the spirochaete into granules as simply the first phase, not of a +process of multiplication, but of a degeneration ending in the death of +the parasite. The fragmented chromatin has lost its affinity for stains, +remaining always paler than that of the normal spirochaetes. On the other +hand, the granules of Leishman stain energetically with all the basic +stains. + +Further, according to Marchoux and Couvy, infection takes place without +the emission of the coxal fluid and indeed, soiling of the host by the +coxal fluid diluting the excrement is exceptional. All of the organs of +the Argasid are invaded by the parasites, but they pass from the +coelom into the acini of the salivary glands and collect in its +efferent canal. The saliva serves as the vehicle of infection. + +Thus, the question of the life cycle of _Spirochaeta gallinarum_, and of +spirochaetes in general, is an open one. + +It should be noted that _Argas persicus_, the carrier of _Spirochaeta +gallinarum_, is a common pest of poultry in the southwestern United +States. Though the disease has not been reported from this country, +conditions are such that if accidentally introduced, it might do great +damage. + + +OTHER SPIROCHAETE DISEASES OF ANIMALS--About a score of other blood +inhabiting spirochaetes have been reported as occurring in mammals, but +little is known concerning their life-histories. One of the most +important is _Spirochaeta theileri_ which produces a spirochaetosis of +cattle in the Transvaal. Theiler has determined that it is transmitted +by an Ixodid tick, _Margaropus decoloratus_. + + +TYPHUS FEVER AND PEDICULIDAE + +Typhus is an acute, and continued fever, formerly epidemically prevalent +in camps, hospitals, jails, and similar places where persons were +crowded together under insanitary conditions. It is accompanied by a +characteristic rash, which gives the disease the common name of +"spotted" or "lenticular" fever. The causative organism is unknown. + +Typhus fever has not generally been supposed to occur in the United +States, but there have been a few outbreaks and sporadic cases +recognized. According to Anderson and Goldberger (1912_a_), it has been +a subject of speculation among health authorities why, in spite of the +arrival of occasional cases in this country and of many persons from +endemic foci of the disease, typhus fever apparently does not gain a +foothold in the United States. These same workers showed that the +so-called Brill's disease, studied especially in New York City, is +identical with the typhus fever of Mexico and of Europe. + +The conditions under which the disease occurs and under which it spreads +most rapidly are such as to suggest that it is carried by some parasitic +insect. On epidemiological grounds the insects most open to suspicion +are the lice, bed-bugs and fleas. + +In 1909, Nicolle, Comte and Conseil, succeeded in transmitting typhus +fever from infected to healthy monkeys by means of the body louse +(_Pediculus corporis_). Independently of this work, Anderson and +Goldberger had undertaken work along this line in Mexico, and in 1910 +reported two attempts to transmit the disease to monkeys by means of +body lice. The first experiment resulted negatively, but the second +resulted in a slight rise in temperature, and in view of later results +it seems that this was due to infection with typhus. + +Shortly after, Ricketts and Wilder (1910) succeeded in transmitting the +disease to the monkey by the bite of body lice in two experiments, the +lice in one instance deriving their infection from a man and in another +from the monkey. Another monkey was infected by typhus through the +introduction of the feces and abdominal contents of infested lice into +small incisions. Experiments with fleas and bed-bugs resulted +negatively. + +Subsequently, Goldberger and Anderson (1912_b_) indicated that the head +louse (_Pediculus humanus_) as well, may become infected with typhus. In +an attempt to transmit typhus fever (Mexican virus) from man to monkey +by subcutaneous injection of a saline suspension of crushed head lice, +the monkeys developed a typical febrile reaction with subsequent +resistance to an inoculation of virulent typhus (Mexican) blood. In one +of the three experiments to transmit the disease from man to monkey by +means of the bite of the head louse, the animal bitten by the presumably +infected head lice proved resistant to two successive immunity tests +with virulent typhus blood. + +In 1910, Ricketts and Wilder reported an experiment undertaken with a +view to determining whether the young of infected lice were themselves +infected. Young lice were reared to maturity on the bodies of typhus +patients, so that if the eggs were susceptible to infection at any stage +of their development, they would have every opportunity of being +infected within the ovary. The eggs of these infected lice were +obtained, they were incubated, and the young lice of the second +generation were placed on a normal rhesus monkey. The experimenters were +unable to keep the monkey under very close observation during the +following three or four weeks, but from the fact that he proved +resistant to a subsequent immunity test they concluded that he probably +owed this immunity to infection by these lice of the second generation. + +Anderson and Goldberger (1912_b_) object that due consideration was not +given to the possibility of a variable susceptibility of the monkey to +typhus. Their similar experiment was "frankly negative." + +Prophylaxis against typhus fever is, therefore, primarily a question of +vermin extermination. A brief article by Dr. Goldberger (1914) so +clearly shows the practical application of his work and that of the +other investigators of the subject, that we abstract from it the +following account: + +"In general terms it may be stated that association with a case of +typhus fever in the absence of the transmitting insect is no more +dangerous than is association with a case of yellow fever in the absence +of the yellow fever mosquito. Danger threatens only when the insect +appears on the scene." + +"We may say, therefore, that to prevent infection of the individual it +is necessary for him only to avoid being bitten by the louse. In theory +this may readily be done, for we know that the body louse infests and +attaches itself almost entirely to the body linen, and that boiling +kills this insect and its eggs. Individual prophylaxis is based +essentially, therefore, on the avoidance of contact with individuals +likely to harbor lice. Practically, however, this is not always as easy +as it may seem, especially under the conditions of such intimate +association as is imposed by urban life. Particularly is this the case +in places such as some of the large Mexican cities, where a large +proportion of the population harbors this vermin. Under such +circumstances it is well to avoid crowds or crowded places, such as +public markets, crowded streets, or public assemblies at which the +'peon' gathers." + +"Community prophylaxis efficiently and intelligently carried out is, +from a certain point of view, probably easier and more effective in +protecting the individual than is the individual's own effort to guard +himself. Typhus emphasizes, perhaps better than any other disease, the +fact that fundamentally, sanitation and health are economic problems. In +proportion as the economic condition of the masses has improved--that +is, in proportion as they could afford to keep clean--the notorious +filth disease has decreased or disappeared. In localities where it still +prevails, its further reduction or complete eradication waits on a +further improvement in, or extension of, the improved economic status of +those afflicted. Economic evolution is very slow process, and, while +doing what we can to hasten it, we must take such precautions as +existing conditions permit, looking to a reduction in or complete +eradication of the disease." + +"When possible, public bath houses and public wash houses, where the +poor may bathe and do their washings at a minimum or without cost, +should be provided. Similar provision should be made in military and +construction camps. Troops in the field should be given the opportunity +as frequently as possible to wash and _scald_ or _boil_ their body +linen." + +"Lodging houses, cheap boarding houses, night shelters, hospitals, jails +and prisons, are important factors in the spread and frequently +constitute foci of the disease. They should receive rigid sanitary +supervision, including the enforcement of measures to free all inmates +of such institutions of lice on admission." + +"So far as individual foci of the disease are concerned these should be +dealt with by segregating and keeping under observation all exposed +individuals for 14 days--the period of incubation--from the last +exposure, by disinfecting (boiling or steaming) the suspected bedding, +body linen, and clothes, for the destruction of any possible vermin that +they may harbor, and by fumigating (with sulphur) the quarters that they +may have occupied." + +"It will be noted that nothing has been said as to the disposition of +the patient. So far as the patient is concerned, he should be removed to +'clean' surroundings, making sure that he does not take with him any +vermin. This may be done by bathing, treating the hair with an +insecticide (coal oil, tincture of larkspur), and a complete change of +body linen. Aside from this, the patient may be treated or cared for in +a general hospital ward or in a private house, provided the sanitary +officer is satisfied that the new surroundings to which the patient has +been removed are 'clean,' that is, free from vermin. Indeed, it is +reasonably safe to permit a 'clean' patient to remain in his own home if +this is 'clean,' for, as has already been emphasized, there can be no +spread in the absence of lice. This is a common experience in native +families of the better class and of Europeans in Mexico City." + +"Similarly the sulphur fumigation above prescribed may be dispensed with +as unnecessary in this class of cases." + + + + +CHAPTER XI + +SOME POSSIBLE, BUT IMPERFECTLY ESTABLISHED CASES OF ARTHROPOD +TRANSMISSION OF DISEASE + + +INFANTILE PARALYSIS OR ACUTE ANTERIOR POLIOMYELITIS + +The disease usually known in this country as infantile paralysis or, +more technically, as acute anterior poliomyelitis, is one which has +aroused much attention in recent years. + +The causative organism of infantile paralysis is unknown, but it has +been demonstrated that it belongs to the group of filterable viruses. It +gives rise to a general infection, producing characteristic lesions in +the central nervous system. The result of the injury to the motor nerves +is a more or less complete paralysis of the corresponding muscle. This +usually manifests itself in the legs and arms. The fatal cases are +usually the result of paralysis of the muscles of respiration. Of the +non-fatal cases about 60 per cent remain permanently crippled in varying +degrees. + +Though long known, it was not until about 1890 that it was emphasized +that the disease occurs in epidemic form. At this time Medin reported +his observations on an epidemic of forty-three cases which occurred in +and around Stockholm in 1887. Since then, according to Frost (1911), +epidemics have been observed with increasing frequency in various parts +of the world. The largest recorded epidemics have been those in Vermont, +1894, 126 cases; Norway and Sweden, 1905, about 1,500 cases; New York +City, 1907, about 2,500 cases. Since 1907 many epidemics have been +reported in the United States, and especially in the Northern States +east of the Dakotas. In 1912 there were over 300 cases of the disease in +Buffalo, N. Y., with a mortality of somewhat over 11 per cent. + +In view of the sudden prominence and the alarming spread of infantile +paralysis, there have been many attempts to determine the cause, and the +manner in which the disease spreads and develops in epidemic form. In +the course of these studies, the question of possible transmission by +insects was naturally suggested. + +C. W. Howard and Clark (1912) presented the results of studies in this +phase of the subject. They dealt especially with the house-fly, bedbug, +head, and body lice, and mosquitoes. It was found that the house-fly +(_Musca domestica_) can carry the virus of poliomyelitis in an active +state for several days upon the surface of the body and for several +hours within the gastro-intestinal tract. Mosquitoes and lice were found +not to take up or maintain the virus. On the other hand, the bedbug +(_Cimex lectularius_) was found to take the virus from the infected +monkeys and to maintain it in a living state within the body for a +period of seven days. This was demonstrated by grinding up in salt +solution, insects which had fed on poliomyelitic animals and injecting +the filtrate into a healthy monkey. The experimenters doubted that the +bedbug is a carrier of the virus in nature. + +Earlier in the same year, Brues and Sheppard published the results of an +intensive epidemiological study of the outbreak of 1911, in +Massachusetts. Special attention had been paid to the possibility of +insect transfer and the following conclusion was reached: + +"Field work during the past summer together with a consideration of the +epidemiology of the disease so far as known, points strongly toward +biting flies as possible carriers of the virus. It seems probable that +the common stable-fly (_Stomoxys calcitrans_ L.) may be responsible to a +certain extent for the spread of acute epidemic poliomyelitis, possibly +aided by other biting flies, such as _Tabanus lineola_. No facts which +disprove such a hypothesis have as yet been adduced, and experiments +based upon it are now in progress." + +As stated by Brues (1913), especial suspicion fell upon the stable-fly +because: + +1. The blood-sucking habits of the adult fly suit it for the transfer of +virus present in the blood. + +2. The seasonal abundance of the fly is very closely correlated with the +incidence of the disease, rising rapidly during the summer and reaching +a maximum in July and August, then slowly declining in September and +October. + +3. The geographical distribution of the fly is, so far as can be +ascertained, wider, or at least co-extensive with that of poliomyelitis. + +4. _Stomoxys_ is distinctly more abundant under rural conditions, than +in cities and thickly populated areas. + +5. While the disease spreads over districts quickly and in a rather +erratic way, it often appears to follow along lines of travel, and it is +known that _Stomoxys_ flies will often follow horses for long distances +along highways. + +6. In a surprisingly large number of cases, it appeared probable that +the children affected had been in the habit of frequenting places where +_Stomoxys_ is particularly abundant, i.e., about stables, barnyards, +etc. + +The experiments referred to were carried on during the summer of 1912 +and in September Dr. Rosenau announced that the disease was transferred +by the bite of the stable-fly. + +A monkey infected by inoculation was exposed to the bites of upwards of +a thousand of the _Stomoxys_ flies daily, by stretching it at full +length and rolling it in a piece of chicken wire, and then placing it on +the floor of the cage in which the flies were confined. The flies fed +freely from the first, as well as later, after paralysis had set in. +Alternating with the inoculated monkey, healthy monkeys were similarly +introduced into the cage at intervals. New monkeys were inoculated to +keep a supply of such infected animals and additional healthy ones were +exposed to the flies, which fed willingly and in considerable numbers on +each occasion. "Thus the flies were given every opportunity to obtain +infection from the monkeys, since the animals were bitten during +practically every stage of the disease from the time of the inoculation +of the virus till their death following the appearance of paralysis. By +the same arrangement the healthy monkeys were likely to be bitten by +flies that had previously fed during the various stages of the disease +on the infected monkeys. The flies had meanwhile enjoyed the opportunity +of incubating the virus for periods varying from the day or two which +usually elapses between consecutive feedings, to the two or three-week +period for which at least some (although a very small percentage) of the +flies lived in the cage." + +"In all, twelve apparently healthy monkeys of a small Japan species were +exposed to the flies in the manner described for the infected monkeys. +Some were placed in the cage only once or twice and others a number of +times after varying intervals. These exposures usually lasted for about +half an hour, but were sometimes more protracted. No results were +apparent until two or three weeks after the experiment was well under +way, and then in rather rapid succession six of the animals developed +symptoms of poliomyelitis. In three, the disease appeared in a virulent +form, resulting in death, while the other three experienced transient +tremblings, diarrhoea, partial paralysis and recovery."--Brues, 1913. + +Very soon after the announcement of the results of experiments by +Rosenau and Brues, they were apparently conclusively confirmed by +Anderson and Frost (1912), who repeated the experiments, at Washington. +They announced that through the bites of the _Stomoxys_ flies that had +previously fed on infected monkeys, they had succeeded in experimentally +infecting three healthy monkeys. + +The results of these experiments gained much publicity and in spite of +the conservative manner in which they had been announced, it was widely +proclaimed that infantile paralysis was conveyed in nature by the +stable-fly and by it alone. + +Serious doubt was cast on this theory by the results of further +experiments by Anderson and Frost, reported in May of 1913. Contrary to +the expectations justified by their first experience, the results of all +the later, and more extended, experiments were wholly negative. Not once +were these investigators again able to transmit the infection of +poliomyelitis through _Stomoxys_. They concluded that it was extremely +doubtful that the insect was an important factor in the natural +transmission of the disease, not only because of their series of +negative results, "but also because recent experiments have afforded +additional evidence of the direct transmissibility or contagiousness of +poliomyelitis, and because epidemiological studies appear to us to +indicate that the disease is more likely transmitted largely through +passive human virus carriers." + +Soon after this, Kling and Levaditi (1913) published their detailed +studies on acute anterior poliomyelitis. They considered that the +experiments of Flexner and Clark (and Howard and Clark), who fed +house-flies on emulsion of infected spinal cord, were under conditions +so different from what could occur in nature that one could not draw +precise conclusions from them regarding the epidemiology of the disease. +They cited the experiments of Josefson (1912), as being under more +reasonable conditions. He sought to determine whether the inoculation of +monkeys with flies caught in the wards of the Hospital for Contagious +Diseases at Stockholm, where they had been in contact with cases of +poliomyelitis, would produce the disease. The results were completely +negative. + +Kling and Lavaditi made four attempts of this kind. The flies were +collected in places where poliomyelitics had dwelt, three, four and +twenty-four after the beginning of the disease in the family and one, +three, and fifteen days after the patient had left the house. These +insects were for the greater part living and had certainly been in +contact with the infected person. In addition, flies were used which had +been caught in the wards of the Hospital for Contagious Diseases at +Soderkoping, when numbers of poliomyelitics were confined there. +Finally, to make the conditions as favorable as possible, the emulsions +prepared from these flies were injected without previous filtering, +since filtration often causes a weakening of the virus. In spite of +these precautions, all their results were negative, none of the +inoculated animals having contracted poliomyelitis. They also +experimented with bedbugs which had fed upon infected patients at +various stages of the disease, but the results in these cases also were +wholly negative. + +Kling and Levaditi considered at length the possibility of transmission +of the disease by _Stomoxys_. As a result of their epidemiological +studies, they found that infantile paralysis continued to spread in +epidemic form in the dead of winter, when these flies were very rare and +torpid, or were even completely absent. Numerous cases developed in the +northern part of Sweden late in October and November, long after snow +had fallen. On account of the rarity of the Stomoxys flies during the +period of their investigations they were unable to conduct satisfactory +experiments. In one instance, during a severe epidemic, they found a +number of the flies in a stable near a house inhabited by an infected +family, though none was found in the house itself. These flies were used +in preparing an emulsion which, after filtering, was injected into the +peritoneal cavity of a monkey. The result was wholly negative. + +As for the earlier experiments, Kling and Levaditi believe if the flies +were responsible for the transmission of the disease in the cases +reported by Rosenau and Brues, and the first experiments of Anderson and +Frost, it was because the virus of infantile paralysis is eliminated +with the nasal secretions of paralyzed monkeys and the flies, becoming +contaminated, had merely acted as accidental carriers. + +Still further evidence against the hypothesis of the transmission of +acute anterior poliomyelitis by _Stomoxys calcitrans_ was brought +forward by Sawyer and Herms (1913). Special precautions were used to +prevent the transference of saliva or other possibly infectious material +from the surface of one monkey to that of another, and to avoid the +possibility of complicating the experiments by introducing other +pathogenic organisms from wild flies, only laboratory-bred flies were +used. In a series of seven carefully performed experiments, in which the +conditions were varied, Sawyer and Herms were unable to transmit +poliomyelitis from monkey to monkey through the agency of _Stomoxys_, or +to obtain any indication that the fly is the usual agent for spreading +the disease in nature. + +The evidence at hand to date indicates that acute anterior +poliomyelitis, or infantile paralysis, is transmitted by contact with +infected persons. Under certain conditions insects may be agents in +spreading the disease, but their role is a subordinate one. + + +Pellagra + +PELLAGRA is an endemic and epidemic disease characterized by a peculiar +eruption or erythema of the skin (figs 144 and 145), digestive +disturbances and nervous trouble. Insanity is a common result, rather +than a precursor of the disease. The manifestations of pellagra are +periodic and its duration indeterminate. + +[Illustration: 144. Pellagrous eruption on the face. After Watson.] + +The disease is one the very name of which was almost unknown in the +United States until within the past decade. It has usually been regarded +as tropical, though it occurs commonly in Italy and in various parts of +Europe. Now it is known that it not only occurs quite generally in the +United States but that it is spreading. Lavinder (1911) says that "There +are certainly many thousand cases of the disease in this country, and +the present situation must be looked upon with grave concern." + +It is not within the scope of this book to undertake a general +discussion of pellagra. The subject is of such importance to every +medical man that we cannot do better than refer to Lavinder's valuable +precis. We can only touch briefly upon the entomological phases of the +problems presented. + +The most commonly accepted theories regarding the etiology of the +disease have attributed it to the use of Indian corn as an article of +diet. This supposed relationship was explained either on the basis of, +(a) insufficiency of nutriment and inappropriateness of corn as a prime +article of food; (b) toxicity of corn or, (c) parasitism of certain +organisms--fungi or bacteria--ingested with either sound or deteriorated +corn. + +In 1905, Sambon proposed the theory of the protozoal origin of pellagra +and in 1910 he marshalled an imposing array of objections to the theory +that there existed any relationship between corn and the disease. He +presented clear evidence that pellagra existed in Europe before the +introduction of Indian corn from America, as an article of diet, and +that its spread was not _pari passu_ with that of the use of corn. Cases +were found in which the patients had apparently never used corn, though +that is obviously difficult to establish. He showed that preventive +measures based on the theory had been a failure. Finally, he believed +that the recurrence of symptoms of the disease for successive springs, +in patients who abstained absolutely from the use of corn, militated +against the theory. + +[Illustration: 145. Pellagrous eruption on the hand. After Watson.] + +On the other hand, Sambon believed that the periodicity of the symptoms, +peculiarities of distribution and seasonal incidence, and analogies of +the symptoms to those of other parasitic diseases indicated that +pellagra was of protozoal origin, and that it was insect-borne. + +The insect carriers, he believed to be one or more species of Simuliidae, +or black-flies. In support of this he stated that _Simulium_ appears to +effect the same topographical conditions as pellagra, that in its imago +stage it seems to present the same seasonal incidence, that it has a +wide geographical distribution which seems to cover that of pellagra, +and that species of the genus are known to cause severe epizootics. +Concluding from his studies in Italy, that pellagra was limited almost +wholly to agricultural laborers, he pointed out that the Simulium flies +are found only in rural districts, and as a rule do not enter towns, +villages, or houses. + +[Illustration: 146. A favorite breeding place of Simulium. Ithaca, N. +Y.] + +When Sambon's detailed report was published in 1910, his theory was +seized upon everywhere by workers who were anxious to test it and who, +in most cases, were favorably disposed towards it because of the +wonderful progress which had been made in the understanding of other +insect-borne diseases. In this country, the entomological aspects of the +subject have been dealt with especially by Forbes (1912), and by King +and Jennings, under the direction of W. D. Hunter, of the Bureau of +Entomology, and in cooperation with the Thompson-McFadden Pellagra +Commission of the Department of Tropical Medicine of the New York +Post-Graduate Medical School. An important series of experiments with +monkeys has been undertaken by S. J. Hunter, of Kansas, but +unfortunately we have as yet no satisfactory evidence that these animals +are susceptible to the disease--a fact which renders the whole problem +difficult. + +The accumulated evidence is increasingly opposed to Sambon's hypothesis +of the transmission of pellagra by _Simulium_. This has been so clearly +manifested in the work of the Thompson-McFadden Commission that we quote +here from the report by Jennings (1914): + +"Our studies in 1912 convinced us that there was little evidence to +support the incrimination of any species of _Simulium_ in South Carolina +in the transmission of pellagra. Reviewing the group as a whole, we find +that its species are essentially "wild" and lack those habits of +intimate association with man which would be expected in the vector of +such a disease as pellagra. Although these flies are excessively +abundant in some parts of their range and are moderately so in +Spartanburg County, man is merely an incidental host, and no disposition +whatever to seek him out or to invade his domicile seems to be +manifested. Critically considered, it is nearer the fact that usually +man is attacked only when he invades their habitat." + +"As our knowledge of pellagra accumulates, it is more and more evident +that its origin is in some way closely associated with the domicile. The +possibility that an insect whose association with man and his immediate +environment is, at the best, casual and desultory, can be active in the +causation of the disease becomes increasingly remote." + +"Our knowledge of the biting habits of _Simulium_ is not complete, but +it is evident, as regards American species at least, that these are +sometimes not constant for the same species in different localities. +Certain species will bite man freely when opportunity offers, while +others have never been known to attack him. To assume that the proximity +of a _Simulium_-breeding stream necessarily implies that persons in its +vicinity must be attacked and bitten is highly fallacious. In +Spartanburg County attacks by _Simulium_ seems to be confined to the +immediate vicinity of the breeding-places. Our records and observations, +exceedingly few in number, refer almost exclusively to such locations. +Statements regarding such attacks, secured with much care and +discrimination from a large number of persons, including many +pellagrins, indicate conclusively that these insects are seldom a pest +of man in this county. A certain number of the persons questioned were +familiar with the gnats in other localities, but the majority were +seemingly ignorant of the existence of such flies with biting habits. +This is especially striking, in view of the fact that the average +distance of streams from the homes of the pellagra cases studied was +about 200 yards, many being at a distance of less than 200 yards, and +that 78 per cent of these streams were found to be infested by larval +_Simulium_. Such ignorance in a large number of persons cannot be +overlooked and indicates strongly that our belief in the negligible +character of local attacks by _Simulium_ is well founded." + +"In localities infested by 'sand-flies,' mosquitoes, etc., these pests +are always well known and the ignorance described above is very +significant." + +"Such positive reports as we received nearly always referred to bites +received in the open, along streams, etc., and observations made of +their attack were of those on field laborers in similar situations. +Males engaged in agricultural pursuits are almost exempt from pellagra +in Spartanburg County. During the season of 1913, in some two or three +instances, observations were made of the biting of _Simulium_ and some +additional and entirely creditable reports were received. These +observations and reports were under conditions identical with those +referred to in the reports of 1912 and confirm the conclusions based on +the observations of that year. I would repeat with emphasis that it is +inconceivable that a fly of the appearance and habits of the prevalent +species of _Simulium_ could be present in such a region, especially +about the haunts of man and attack him with sufficient frequency and +regularity to satisfactorily account for so active and prevalent a +disease as pellagra without being a well-known and recognized pest." + +"In connection with the conditions in the Piedmont region of South +Carolina, it may be well to cite the results of a study of those in the +arid region of western Texas." + +"In May, 1913, in company with Capt. J. F. Siler of the +Thompson-McFadden Pellagra Commission, I visited the region of which +Midland in Midland County is the center. This region is very dry and +totally devoid of running water for a long distance in every direction. +The only natural source of water-supply, a few water holes and ponds, +were visited and found to be of such a nature that the survival of +_Simulium_, far less its propagation in them, is absolutely impossible. +The nearest stream affording possibilities as a source of _Simulium_ is +60 miles away, while the average distance of such possibility is not +less than 100 miles." + +"Artificial sources of water-supply were also investigated carefully and +were found to offer no opportunity for the breeding of _Simulium_." + +"At Midland the histories of five cases of pellagra were obtained, which +gave clear evidence that this place or its immediate vicinity was the +point of origin. Persons of long residence in the country were +questioned as to the occurrence of such flies as _Simulium_ and returned +negative answers. These included a retired cattle owner, who is a man of +education and a keen observer, an expert veterinarian stationed in the +country who has the cattle of the country under constant observation, +and a practical cattle man, manager of a ranch and of wide experience. +The latter had had experience with 'Buffalo gnats' in other localities +(in the East) and is well acquainted with them. His close personal +supervision of the cattle under his charge, makes it practically certain +that he would have discovered these gnats had they been present in the +country." + +"At the time the study was made, _Simulium_ was breeding and active in +the adult state in the vicinity of Dallas, Texas, in the eastern part of +the state. We have here a region in which cases of pellagra have +originated, yet in which _Simulium_ does not and cannot breed." + +Other possible insect vectors of pellagra have been studied in great +detail and the available evidence indicates that if _any_ insect plays a +role in the spread of the disease, _Stomoxys calcitrans_ most nearly +fills the conditions. This conclusion was announced by Jennings and King +in 1912, and has been supported by their subsequent work. + +Yet, after all the studies of the past decade, the old belief that +pellagra is essentially of dietary origin is gaining ground. Goldberger, +Waring and Willets (1914) of the United States Public Health Service +summarize their conclusions in the statement, (1) that it is dependent +on some yet undetermined fault in a diet in which the animal or +leguminous protein component is disproportionately large and (2) that no +pellagra develops in those who _consume_ a mixed, well-balanced, and +varied diet, such, for example, as that furnished by the Government to +the enlisted men of the Army, Navy, and Marine Corps. + + +Leprosy + +LEPROSY is a specific, infectious disease due to _Bacillus leprae_, and +characterized by the formation of tubercular nodules, ulcerations, and +disturbances of sensation. In spite of the long time that the disease +has been known and the dread with which it is regarded, little is known +concerning the method of transfer of the causative organism or the means +by which it gains access to the human body. + +It is known that the bacilli are to be found in the tubercles, the scurf +of the skin, nasal secretions, the sputum and, in fact in practically +all the discharges of the leper. Under such conditions it is quite +conceivable that they may be transferred in some instances from diseased +to healthy individuals through the agency of insects and other +arthropods. Many attempts have been made to demonstrate this method of +spread of the disease, but with little success. + +Of the suggested insect carriers none seem to meet the conditions better +than mosquitoes, and there are many suggestions in literature that these +insects play an important role in the transmission of leprosy. The +literature has been reviewed and important experimental evidence +presented by Currie (1910). He found that mosquitoes feeding, under +natural conditions, upon cases of nodular leprosy so rarely, if ever, +imbibe the lepra bacillus that they cannot be regarded as one of the +ordinary means of transference of this bacillus from lepers to the skin +of healthy persons. He believes that the reason that mosquitoes that +have fed on lepers do not contain the lepra bacillus is that when these +insects feed they insert their proboscis directly into a blood vessel +and thus obtain bacilli-free blood, unmixed with lymph. + +The same worker undertook to determine whether flies are able to +transmit leprosy. He experimented with five species found in +Honolulu,--_Musca domestica_, _Sarcophaga pallinervis_, _Sarcophaga +barbata_, _Volucella obesa_ and an undetermined species of _Lucilia_. +The experiments with _Musca domestica_ were the most detailed. From +these experiments he concluded, first, that all of the above-named +flies, when given an opportunity to feed upon leprous fluids, will +contain the bacilli in their intestinal tracts and feces for several +days after such feeding. Second, that considering the habits of these +flies, and especially those of _Musca domestica_, it is certain that, +given an exposed leprous ulcer, these insects will frequently convey +immense numbers of lepra bacilli, directly or indirectly, to the skins, +nasal mucosa, and digestive tracts of healthy persons. Additional +evidence along this line has recently been brought forward by Honeij and +Parker (1914), who incriminate both _Musca domestica_ and _Stomoxys +calcitrans_. Whether or not such insect-borne bacilli are capable of +infecting persons whose skin and mucosa are thus contaminated, Currie +was unwilling to maintain, but he concludes that until we have more +accurate knowledge on this point, we are justified in regarding these +insects with grave suspicion of being one of the means of disseminating +leprous infection. + +Various students of the subject have suggested that bed-bugs may be the +carriers of leprosy and have determined the presence of acid-fast +bacilli in the intestines of bed-bugs which had fed on leprous patients. +Opposed to this, the careful experiments of Thompson (1913) and of +Skelton and Parkham (1913) have been wholly negative. + +Borrel has recently suggested that _Demodex_, may play a role in +spreading the infection in families. Many other insects and acariens +have been suggested as possible vectors, but the experimental data are +few and in no wise conclusive. The most that can be said is that it is +quite possible that under favorable conditions the infection might be +spread by any of the several blood-sucking forms or by house-flies. + + +Verruga peruviana + +VERRUGA PERUVIANA is defined by Castellani and Chalmers as "a chronic, +endemic, specific, general disorder of unknown origin, not contagious, +but apparently inoculable, and characterized by an irregular fever +associated with rheumatoid pains, anemia, followed by granulomatous +swellings in the skin, mucous membranes, and organs of the body." It has +been generally believed by medical men interested that the comparatively +benign eruptive verruga is identical with the so-called Oroya, or +Carrion's fever, a malignant type. This view is not supported by the +work of Strong, Tyzzer and Brues, (1913). + +The disease is confined to South America and to definitely limited areas +of those countries in which it does occur. It is especially prevalent in +some parts of Peru. + +The causative organism and the method of transfer of verruga are +unknown. Castellani and Chalmers pointed out in 1910 that the study of +the distribution of the disease in Peru would impress one with the +similarity to the distribution of the Rocky Mountain fever and would +lead to the conclusion that the aetiological cause must in some way be +associated with some blood-sucking animal, perhaps an arachnid, and that +this is supported by the fact that the persons most prone to the +infection are those who work in the fields. + +More recently, Townsend (1913), in a series of papers, has maintained +that verruga and Carrion's disease are identical, and that they are +transmitted to man by the bites of the Psychodid fly, _Phlebotomus +verrucarum_. He succeeded in producing the eruptive type of the disease +in experimental animals by injecting a physiological salt trituration of +wild Phlebotomus flies. A cebus monkey was exposed from October so to +November 6, by chaining him to a tree in the verruga zone, next to a +stone wall from which the flies emerged in large numbers every night. +Miliar eruption began to appear on the orbits November 13 and by +November 21, there were a number of typical eruptions, with exudation on +various parts of the body exactly like miliar eruptive sores commonly +seen on legs of human cases. + +An assistant in the verruga work, George E. Nicholson, contracted the +eruptive type of the disease, apparently as a result of being bitten by +the Phlebotomus flies. He had slept in a verruga zone, under a tight +net. During the night he evidently put his hands in contact with the +net, for in the morning there were fifty-five unmistakable Phlebotomus +bites on the backs of his hands and wrists. + +Townsend believes that in nature, lizards constitute the reservoir of +the disease and that it is from them that the Phlebotomus flies receive +the infection. + + +Cancer + +There are not wanting suggestions that this dread disease is carried, or +even caused, by arthropods. Borrel (1909) stated that he had found mites +of the genus _Demodex_ in carcinoma of the face and of the mammae. He +believed that they acted as carriers of the virus. + +Saul (1910) and Dahl (1910) go much further, since they attribute the +production of the malignant growth to the presence of mites which Saul +had found in cancers. These Dahl described as belonging to a new +species, which he designated _Tarsonemus hominis_. These findings have +since been confirmed by several workers. Nevertheless, the presence of +the mite is so rare that it cannot be regarded as an important factor in +the causation of the disease. The theory that cancer is caused by an +external parasite is given little credence by investigators in this +field. + + +IN CONCLUSION, it should be noted that the medical and entomological +literature of the past few years abounds in suggestions, and in +unsupported direct statements that various other diseases are +insect-borne. Knab (1912) has well said "Since the discovery that +certain blood-sucking insects are the secondary hosts of pathogenic +parasites, nearly every insect that sucks blood, whether habitually or +occasionally, has been suspected or considered a possible transmitter of +disease. No thought seems to have been given to the conditions and the +characteristics of the individual species of blood-sucking insects, +which make disease transmission possible." + +He points out that "in order to be a potential transmitter of human +blood-parasites, an insect must be closely associated with man and +normally have opportunity to suck his blood repeatedly. It is not +sufficient that occasional specimens bite man, as, for example, is the +case with forest mosquitoes. Although a person may be bitten by a large +number of such mosquitoes, the chances that any of these mosquitoes +survive to develop the parasites in question, (assuming such development +to be possible), and then find opportunity to bite and infect another +person, are altogether too remote. Applying this criterion, not only the +majority of mosquitoes but many other blood-sucking insects, such as +Tabanidae and Simuliidae, may be confidently eliminated. Moreover, +these insects are mostly in evidence only during a brief season, so that +we have an additional difficulty of a very long interval during which +there could be no propagation of the disease in question." He makes an +exception of tick-borne diseases, where the parasites are directly +transmitted from the tick host to its offspring and where, for this +reason, the insect remains a potential transmitter for a very long +period. He also cites the trypanosome diseases as possible exceptions, +since the causative organisms apparently thrive in a number of different +vertebrate hosts and may be transmitted from cattle, or wild animals, to +man. + +Knab's article should serve a valuable end in checking irresponsible +theorizing on the subject of insect transmission of disease. +Nevertheless, the principles which he laid down cannot be applied to the +cases of accidental carriage of bacterial diseases, or to those of +direct inoculation of pyogenic organisms, or of blood parasites such as +the bacillus of anthrax, or of bubonic plague. Accumulated evidence has +justified the conclusion that certain trypanosomes pathogenic to man are +harbored by wild mammals, and so form an exception. Townsend believes +that lizards constitute the natural reservoir of verruga; and it seems +probable that field mice harbor the organism of tsutsugamushi disease. +Such instances are likely to accumulate as our knowledge of the relation +of arthropods to disease broadens. + + + + +CHAPTER XII + +HOMINOXIOUS ARTHROPODS + + +The following synoptic tables are presented in the hope that they may be +of service in giving the reader a perspective of the relationships of +the Arthropoda in general and enabling him to identify the more +important species which have been found noxious to man. Though +applicable chiefly to the arthropods found in the United States, exotic +genera and species which are concerned in the transmission of disease +are also included. For this reason the keys to the genera of the Muscids +of the world are given. As will be seen, the tables embrace a number of +groups of species which are not injurious. This was found necessary in +order that the student might not be lead to an erroneous determination +which would result were he to attempt to identify a species which +heretofore had not been considered noxious, by means of a key containing +only the noxious forms. The names printed in BOLD FACED TYPE indicate +the hominoxious arthropods which have been most commonly mentioned in +literature. + + +CRUSTACEA + +Arthropods having two pairs of antennae which are sometimes modified for +grasping, and usually with more than five pairs of legs. With but few +exceptions they are aquatic creatures. Representatives are: Crabs, +lobsters, shrimps, crayfish, water-fleas, and woodlice. To this class +belongs the CYCLOPS (fig. 122) a genus of minute aquatic crustaceans of +which at least one species harbors _Dracunculus medinensis_, the Guinea +worm (fig. 121). + + +MYRIAPODA + +Elongate, usually vermiform, wingless, terrestrial creatures having one +pair of antennae, legs attached to each of the many intermediate body +segments. This group is divided into two sections, now usually given +class rank: the DIPLOPODA or millipedes (fig. 13), commonly known as +thousand legs, characterized by having two pairs of legs attached to +each intermediate body segment, and the CHILOPODA or centipedes (fig. +14) having only one pair of legs to each body segment. + + +ARACHNIDA + +In this class the antennae are apparently wanting, wings are never +present, and the adults are usually provided with four pairs of legs. +Scorpions, harvest-men, spiders, mites, etc. + + +HEXAPODA (Insects) + +True insects have a single pair of antennae, which is rarely vestigial, +and usually one or two pairs of wings in the adult stage. Familiar +examples are cockroaches, crickets, grasshoppers, bugs, dragon-flies, +butterflies, moths, mosquitoes, flies, beetles, ants, bees and wasps. + + +ORDERS OF THE ARACHNIDA + +a. Abdomen distinctly segmented. A group of orders including scorpions, + (fig. 11), whip-scorpions (fig. 10), pseudo-scorpions, solpugids + (fig. 12) harvest-men (daddy-long-legs or harvestmen), etc. + ARTHROGASTRA + +aa. Abdomen unsegmented, though sometimes with numerous annulations + SPHAEROGASTRA + + b. A constriction between cephalothorax and abdomen (fig. 7). True + Spiders ARANEIDA + + bb. No deep constriction between these parts. + + c. Legs usually well developed, body more or less depressed (fig. + 49). Mites ACARINA + + cc. Legs stumpy or absent, body more or less elongate or vermiform, + or if shorter, the species is aquatic or semi-aquatic in habit. + + d. Four pairs of short legs; species inhabiting moss or water. + Water-bears. TARDIGRADA + + dd. Two pairs of clasping organs near the mouth, instead of legs, + in the adult; worm-like creatures parasitic within the nasal + passages, lungs, etc. of mammals and reptiles (fig. 148). + Tongue worms. LINGUATULINA + +[Illustration: 148. Linguatula. (_a_) larva; (enlarged). (_b_) adult; +(natural size).] + + +ACARINA[E] + +a. Abdomen annulate, elongate; very minute forms, often with but four + legs (fig. 62). DEMODICOIDEA + + b. With but four legs of five segments each. Living on plants, often + forming galls. ERIOPHYIDAE + + bb. With eight legs, of three segments each. Living in the skin of + mammals. DEMODICIDAE + + To this family belongs the genus DEMODEX found in the sebaceous + glands and hair follicles of various mammals, including man. _D. + phylloides_ Csokor has been found in Canada on swine, causing + white tubercles on the skin. _D. bovis_ Stiles has been reported + from the United States on cattle, upon the skin of which they + form swellings. D. FOLLICULORUM Simon is the species found on + man. See page 78. + +aa. Abdomen not annulate nor prolonged behind; eight legs in the adult + stage. + + b. With a distinct spiracle upon a stigmal plate on each side of the + body (usually ventral) above the third or fourth coxae or a little + behind (fig. 50); palpi free; skin often coriaceous or leathery; + tarsi often with a sucker. + + c. Hypostome large (fig. 50), furnished below with many recurved + teeth; venter with furrows, skin leathery; large forms, usually + parasitic. IXODOIDEA + + d. Without scutum but covered by a more or less uniform leathery + integument; festoons absent; coxae unarmed, tarsi without + ventral spurs; pulvilli absent or vestigial in the adults; + palpi cylindrical; sexual dimorphism slight. ARGASIDAE + + e. Body flattened, oval or rounded, with a distinct flattened + margin differing in structure from the general integument; + this margin gives the body a sharp edge which is not + entirely obliterated even when the tick is full fed. + Capitulum (in adults and nymphs) entirely invisible + dorsally, distant in the adult by about its own length from + the anterior border. Eyes absent. ARGUS Latr. + + f. Body oblong; margin with quadrangular cells; anterior tibiae + and metatarsi each about three times as long as broad. On + poultry, southwest United States. A. PERSICUS MINIATUS + + _A. brevipes_ Banks, a species with proportionately shorter + legs has been recorded from Arizona. + + ff. With another combination of characters. About six other + species of _Argas_ from various parts of the world, + parasitic on birds and mammals. + + ee. Body flattened when unfed, but usually becoming very convex + on distention; anterior end more or less pointed and + hoodlike; margin thick and not clearly defined, similar in + structure to the rest of the integument and generally + disappearing on distention; capitulum subterminal, its + anterior portions often visible dorsally in the adult; eyes + present in some species. + + f. Integument pitted, without rounded tubercles; body provided + with many short stiff bristles; eyes absent. On horses, + cattle and man (fig. 48). OTIOBIUS Banks. + + O. MEGNINI, a widely distributed species, is the type of + this genus. + + ff. Integument with rounded tubercles or granules; body + without stiff bristles. ORNITHODOROS Koch. + + g. Two pairs of eyes; tarsi IV with a prominent subterminal + spur above; leg I strongly roughened. On cattle and man. + O. CORIACEUS + + gg. No eyes; no such spur on the hind tarsi. + + h. Tarsi I without humps above. _O. talaje._ + + hh. Tarsi I with humps above. + + i. Tarsi IV without distinct humps above. On hogs, + cattle and man. O. TURICATA + + ii. Tarsi IV with humps nearly equidistant (fig. 142). + Africa. O. MOUBATA + +[Illustration: 149. Haemaphysalis wellingtoni. Note short palpi. After +Nuttall and Warburton.] + + dd. With scutum or shield (fig. 50); festoons usually present; + coxae usually armed with spurs, tarsi generally with one or two + ventral spurs; pulvilli present in the adults; sexual + dimorphism pronounced. IXODIDAE + + e. With anal grooves surrounding anus in front; inornate; + without eyes; no posterior marginal festoons; venter of the + male with non-salient plates. Numerous species, 14 from the + United States, among them I. RICINUS (fig. 49 and 50), + SCAPULARIS, COOKEI, _hexagonus_, _bicornis_. IXODES Latr. + (including Ceratixodes). + + ee. With anal groove contouring anus behind, or groove faint or + obsolete. + + f. With short palpi (fig. 149). + + g. Without eyes, inornate, with posterior marginal festoons; + male without ventral plates. Numerous species. _H. + chordeilis_ and _leporis-palustris_ from the United + States. _Haemaphysalis_ Koch. + +[Illustration: 150. Stigmal plate of Dermacentor andersoni; (_a_) of +male, (_b_) of female. After Stiles. (_c_) Dermacentor variabilis, male; +(_d_) Glyciphagus obesus; (_e_) Otodectes cynotis; (_f_) Tyroglyphus +lintneri; (_g_) Tarsonemus pallidus; (_h_) anal plate and mandible of +Liponyssus; (_c_) to (_h_) after Banks.] + + gg. With eyes. + + h. Anal groove distinct; posterior marginal festoons + present. + + i. Base of the capitulum (fig. 150c) rectangular + dorsally; usually ornate. DERMACENTOR Koch. + + j. Adults with four longitudinal rows of large + denticles on each half of hypostome; stigmal plate + nearly circular, without dorso-lateral + prolongation, goblets very large, attaining 43u to + 115u in diameter; not over 40 per plate, each + plate surrounded by an elevated row of regularly + arranged supporting cells; white rust wanting; + base of capitulum distinctly broader than long, + its postero-lateral angles prolonged slightly, if + at all; coxae T with short spurs; trochanter I with + small dorso-terminal blade. Texas, Arizona, etc. + _D. nitens_ + +[Illustration: 151. Rhipicephalus bursa, male. After Nuttall and +Warburton.] + + jj. Adults with three longitudinal rows of large + denticles on each half of hypostome; goblet cells + always more than 40 per plate; whitish rust + usually present. + + k. Dorso-lateral prolongation of stigmal plate small + or absent; plates of the adults distinctly + longer than broad; goblet cells large, usually + 30u to 85u in diameter, appearing as very coarse + punctations on untreated specimens, but on + specimens treated with caustic potash they + appear very distinct in outline; base of + capitulum distinctly (usually about twice) + broader than long, the postero-lateral angles + distinctly produced caudad; spurs of coxae I + long, lateral spur slightly longer than median; + trochanter I with dorso-terminal spur. _D. + albipictus_, (= _variegatus_), _salmoni_, + _nigrolineatus_. + + kk. Dorso-lateral prolongation of stigmal plate + distinct. + + l. Body of plate distinctly longer than broad; + goblet cells of medium size, usually 17.5u to + 35u or 40u in diameter, appearing as medium + sized punctuations on untreated specimens, but + on the specimens treated with caustic potash + they appear very distinct in outline, which is + not circular; base of capitulum usually less + than twice as broad as long, the + postero-lateral angles always distinctly + prolonged caudad. + + m. Trochanter I with distinct dorso-subterminal + retrograde sharp, digitate spur; + postero-lateral angles of capitulum + pronouncedly prolonged caudal, 112u to 160u + long; goblet cells attain 13u to 40u in + diameter; type locality California. D. + OCCIDENTALIS + + mm. Trochanter I with dorso-terminal blade; + postero-lateral angles of capitulum with + rather short prolongations. + + n. Stigmal plate small, goblet cells not + exceeding 45 in the male or 100 in the + female; scutum with little rust, coxa I + with short spurs, the inner distinctly + shorter than the outer. _D. + parumapertus-marginatus_ + + nn. Stigmal plate larger; goblet cells over 70 + in the male and over 100 in the female; + coxa I with longer spurs, inner slightly + shorter than the outer; scutum with + considerable rust. D. VENUSTUS[F] + + ll. Goblet cells small, rarely exceeding 17.6u, + occasionally reaching 19u in diameter; on + untreated specimens they appear as very fine + granulations, and on specimens treated with + caustic potash they may be difficult to see, + but their large number can be determined from + the prominent stems of the goblets; surface of + outline of the goblets distinctly circular; + base of the capitulum usually less than twice + as broad as long, the postero-lateral angle + distinctly prolonged caudad; spurs of coxae I + long. _D. reticulatus_ and _electus_ (= + _variabilis_?) + + ii. Base of the capitulum (fig. 151) usually hexagonal + (except in the male of _puchellus_); and usually + inornate. + + j. No ventral plate or shield in either sex (fig. + 153). R. BICORNIS from the United States. + RHIPICENTOR Nuttall + + jj. Males with a pair of adanal shields, and usually a + pair of accessory adanal shields. Numerous + species, among them _R. sanguineus_ (fig. 154) and + _texanus_, the latter from the United States. + _Rhipicephalus_ Koch + + hh. Anal grooves faint or obsolete; no marginal festoons. + + i. Short palpi; highly chitinized; unfed adults of large + size; coxae conical; male with a median plate + prolonged in two long spines projecting caudad; + segments of leg pair IV greatly swollen (fig. 155, + 156). _M. winthemi_ _Margaropus_ Karsch + +[Illustration: 152. Monieziella (Histiogaster) emtomophaga-spermatica, +ventral aspect, male and female. After Trouessart.] + + ii. Very short palpi, ridged dorsally and laterally; + slightly chitinized; unfed adults of smaller size; + coxae I bifid; male with adanal and accessory adanal + shields (fig. 139). B. ANNULATUS. BOOPHILUS Curtis + + ff. Palpi longer than broad (fig. 157). + + g. Male with pair of adanal shields, and two posterior + abdominal protrusions capped by chitinized points; + festoons present or absent. Several species, among them + H. AEGYPTICUM (fig. 140) from the old world. HYALOMMA + Koch + + gg. Male without adanal shields but small ventral plaques + are occasionally present close to the festoons. Many + species, a few from the United States (fig. 157). + AMBLYOMMA Koch + + + h. Coxa I with but one spine; metatarsi (except I) with + two thickened spurs at tips. _A. maculatum_ + + hh. Coxa I with two spines; metatarsi without stout spurs + at tips, only slender hairs. + + i. Projections of coxa I blunt and short. Large species + on the gopher tortoise in Florida. _A. tuberculatum_ + + ii. Projections of coxa I longer, and at least one of + them sharp pointed; second segment of palpus twice + as long as the third; coxa IV of the male with a + long spine. + + j. Porose areas nearly circular; shield of both sexes + pale yellowish, with some silvery streaks and + marks, and some reddish spots; shield of female as + broad as long. A. CAJENNENSE (= MIXTUM). + + jj. Porose areas elongate, shield brown, in the female + with an apical silvery mark, in the male with two + small and two or four other silvery spots; shield + of the female longer than broad (fig 158 e). A. + AMERICANUM. + +[Illustration: 153. Rhipicentor bicornis, ventral aspect, male. After +Nuttall and Warburton.] + + cc. Hypostome small, without teeth, venter without furrows; body + often with coriaceous shields, posterior margin of the body + never crenulate (i.e. without festoons); no eyes. GAMASOIDEA. + + d. Parasitic on vertebrates; mandibles fitted for piercing; body + sometimes constricted. DERMANYSSIDAE. + + e. Anal plate present. DERMANYSSINAE. + + f. Body short; legs stout, hind pair reaching much beyond the + tip of the body. On bats. _Pteroptus_ Dufour. + + ff. Body long; hind legs not reaching beyond the tip of the + body. + + g. Peritreme on the dorsum, very short; body distinctly + constricted. _Ptilonyssus_ Berl. + + gg. Peritreme on the venter, longer; body not distinctly + constricted. + + h. Mandibles in both sexes chelate. Parasitic on bats, + mice and birds (fig. 150, h). LIPONYSSUS Kol. + + The species L. (= LEIOGNATHUS) SYLVIARUM frequents the + nests of warblers. An instance is on record of + these mites attacking man, causing a pruritis. + + hh. Mandibles in the male chelate (fig. 158 j), in the + female long, styliform. Parasitic on birds. + DERMANYSSUS Dug. Two species of importance may be + noted, _D. hirundinus_ and D. GALLINAE. The latter + (fig. 51) is a serious pest of poultry, sometimes + attacking man, causing itching and soreness. + + ee. Anal plate absent. In lungs and air passages of some + mammals. HALARACHNINAE. + + dd. Free or attached to insects, rarely on vertebrates. + + e. First pair of legs inserted within the same body opening as + the oral tube; genital apertures surrounded by the sternum. + On insects. UROPODIDAE. + +[Illustration: 154. Rhicephalus sanguineus, male. After Nuttall and +Warburton.] + + ee. First pair of legs inserted at one side of the mouth + opening; male genital aperture usually on the anterior + margin of the sternal plate. GAMASIDAE. + + This family contains a number of genera, some of which are + found upon mammals, though the majority affect only other + arthropods. One species, LAELAPS STABULARIS, frequents the + bedding in stables, and in one instance at least, has + occasioned irritation and itching, in man. + + bb. No distinct spiracle in the stigmal plate on each side of the + body. + + c. Body usually coriaceous, with few hairs, with a specialized seta + arising from a pore near each posterior corner of the + cephalothorax; no eyes; mouth parts and palpi very small; + ventral openings of the abdomen large; tarsi without sucker. Not + parasitic. ORIBATOIDEA. + + cc. Body softer; without such specialized seta. + + d. Aquatic species. HYDRACHNOIDEA. + + dd. Not aquatic. + + e. Palpi small, three segmented, adhering for some distance to + the lip; ventral suckers at genital opening or near anal + opening usually present; no eyes; tarsi often end in + suckers; beneath the skin on the venter are seen rod-like + epimera that support the legs; body often entire. Adults + frequently parasitic. SARCOPTOIDEA. + + f. With tracheae; no ventral suckers; legs ending in claws; + body divided into cephalothorax and abdomen; the female + with a clavate hair between legs I and II. Usually not + parasitic on birds and mammals. TARSONEMIDAE + + g. Hind legs of female ending in claw and sucker as in the + other pairs. PEDICULOIDINAE + + To this sub-family belongs the genus PEDICULOIDES + + P. ventricosus is described on page 69. + +[Illustration: 155. Margaropus winthemi, male. After Nuttall and +Warburton.] + +[Illustration: 156. Margaropus winthemi, capitulum and scutum. +After Nuttall and Warburton.] + + gg. Hind legs of the female end in long hairs. TARSONEMINAE + + TARSONEMUS INTECTUS Karpelles, normally found upon grain, + is said to attack man in Hungary and Russia. Other + species of the genus affect various plants (c.f. fig. + 150, g). + + ff. Without tracheae; no such clavate hair. + + g. Genital suckers usually present; integument usually + without fine parallel lines. + + h. Legs short, without clavate hair on tarsi I and II. On + insects. CANESTRINIDAE. + + hh. Legs longer, with a clavate hair on tarsi I and II. + Not normally parasitic except on bees. TYROGLYPHIDAE + + i. Dorsal integument more or less granulate; claws very + weak, almost invisible; some hairs of the body + plainly feathered; ventral apertures large. + GLYCIPHAGUS Her. + + This genus occurs in the United States. In Europe the + mites have been found feeding on all sorts of + substances. They are known as sugar mites and + cause the disease known as grocer's itch. G. + DOMESTICUS and G. PRUNORUM are old world species + (fig. 150, d). + + ii. Dorsal integument not granulate; claws distinct; no + prominent feathered hairs; ventral aperture small. + + j. Mandibles not chelate; elongate, and toothed below; + body without long hairs; palpi enlarged at tip and + provided with two divergent bristles. Species feed + on decaying substances. _Histiostoma_ Kram. + + jj. Mandibles chelate; palpi not enlarged at the tip, + nor with two bristles. + + k. No clavate hair on the base of tarsi I and II; no + suture between cephalothorax and abdomen. Live + on bees or in their nests. _Trichotarsus_ Can. + + kk. A clavate or thickened hair at the base of tarsi + I and II. + + l. The bristle on the penultimate segment of the + legs arises from near the middle; no suture + between the cephalothorax and abdomen. The + species, some of which occur in the United + States, feed on dried fruit, etc. + _Carpoglyphus_ Robin. + + ll. The bristle on the penultimate segment of the + legs arise from near the tip; a suture between + cephalothorax and abdomen. + + m. Cephalothorax with four distinct and long + bristles in a transverse row; tarsi I and II + about twice as long as the preceding segment + (fig. 150 f). TYROGLYPHUS Latr. + + n. Some bristles on tarsi I and II near + middle, distinctly spine-like; the sense + hair about its length from the base of the + segment. Several species in the United + States belong to this group. + + nn. No spine-like bristles near the middle of + the tarsi; sense hair not its length from + the base of the segment. + + o. Of the terminal abdominal bristles, only + two are about as long as the abdomen; + leg I of the male greatly thickened and + with a spine at apex of the femur below. + T. FARINAE. + + oo. Of the terminal abdominal bristles at + least six or more are very long, nearly + as long as the body. + + p. Bristles of the body distinctly plumose + or pectinate; tarsi very long. T. + LONGIOR. + + pp. Bristles of the body not pectinate. + + q. In mills, stored foods, grains, etc. + Third and fourth joints of hind legs + scarcely twice as long as broad; + abdominal bristles not unusually + long; legs I and II of the male not + unusually stout. T. AMERICANUS. + + qq. With other characters and habits. + _T. lintneri_ (fig. 150 f) the + mushroom mite, and several other + species. + + mm. Cephalothorax with but two long distinct + bristles (besides the frontal pair), but + sometimes a very minute intermediate pair; + tarsi I and II unusually short and not twice + as long as the preceding segment. + + n. Tarsi with some stout spines. RHIZOGLYPHUS + Clap. + + The species of this genus are vegetable + feeders. Several occur in the United + States. R. PARASITICUS and R. + SPINITARSUS have been recorded from the + old world, attacking human beings who + handle affected plants. + + nn. Tarsi with only fine hairs. MONIEZIELLA + Berl. + + The species of this genus, as far as known, + are predaceous or feed on recently + killed animal matter. Several species + occur in the United States. M. (= + HISTIOGASTER) ENTOMOPHAGA (fig. 152) + from the old world has been recorded as + injurious to man. + + gg. Genital suckers absent; integument with fine parallel + lines. Parasitic on birds and mammals. + + h. Possessing a specially developed apparatus for clinging + to hairs of mammals. LISTROPHORIDAE. + + hh. Without such apparatus. + + i. Living on the plumage of birds. ANALGESIDAE. + + ii. In the living tissues of birds and mammals. + + j. Vulva longitudinal. In the skin and cellular + tissues of birds. CYTOLEICHIDAE. + + This family contains two species, both occurring in + the United States on the common fowl. + _Laminosioptes cysticola_ occurs on the skin and + also bores into the subcutaneous tissue where it + gives rise to a calcareous cyst. _Cytoleichus + nudus_ is most commonly found in the air + passages and air cells. + + jj. Vulva transverse. In the skin of mammals and + birds. SARCOPTIDAE + + k. Anal opening on the dorsum. + + l. Third pair of legs in the male without apical + suckers. On cats and rabbits. NOTOEDRES + Rail. + + The itch mite of the cat, N. CATI (fig. 61) has + been recorded on man. + + ll. Third leg in the male with suckers. On bats + _Prosopodectes_ Can. + + kk. Anal opening below. + + l. Pedicel of the suckers jointed; mandibles + styliform and serrate near the tip. PSOROPTES + Gerv. P. COMMUNIS OVIS is the cause of sheep + scab. + + ll. Pedicel of the suckers not jointed; mandibles + chelate. + + m. No suckers on the legs of the females; + parasitic on birds, including chickens. _C. + mutans_ is itch mite of chickens. + _Cnemidocoptes_ Furst. + + mm. Suckers at least on legs I and II; parasitic + on mammals. + + n. Legs very short; in the male the hind pairs + equal in size; body usually short. + SARCOPTES Latr. + + S. SCABIEI is the itch mite of man (fig. + 56). + +[Illustration: 157. Amblyomma, female. After Nuttall and Warburton.] + + nn. Legs more slender; in the male the third + pair is much larger than the fourth; body + more elongate. + + o. Female with suckers on the fourth pair of + legs. Species do not burrow in the skin, + but produce a scab similar to sheep + scab. They occur in the ox, horse, sheep + and goat. CHORIOPTES Gerv. C. SYMBIOTES + BOVIS of the ox has been recorded a few + times on man. + + oo. Female without suckers to the fourth + legs. + + p. Hind part of the male abdomen with two + lobes. On a few wild animals. + _Caparinia_ Can. + + pp. Hind part of the male abdomen without + lobes. Live in ears of dogs and cats. + _Otodectes_ Canestr. + + _O. cynotis_ Hering (fig. 150 e) has + been taken in the United States. + + ee. Palpi usually of four or five segments, free; rarely with + ventral suckers near genital or anal openings; eyes often + present; tarsi never end in suckers; body usually divided + into cephalothorax and abdomen; rod-like epimera rarely + visible; adults rarely parasitic. + + f. Last segment of the palpi never forms a thumb to the + preceding segment; palpi simple, or rarely formed to hold + prey; body with but few hairs. EUPODOIDEA. + + g. Palpi often geniculate, or else fitted for grasping prey; + mandibles large and snout like; cephalothorax with four + long bristles above, two in front, two behind; last + segment of leg I longer than the preceding segment, + often twice as long. BDELLIDAE. + + gg. Palpi never geniculate (fig. 158a), nor fitted for + grasping prey: beak small; cephalothorax with bristles + in different arrangement; last segment of leg I shorter + or but little longer than the preceding joint; eyes when + present near posterior border. EUPODIDAE + + Moniez has described a species from Belgium (TYDEUS + MOLESTUS) which attacks man. It is rose colored; + eyeless; its legs are scarcely as long as its body, + the hind femur is not thickened; the mandibles are + small and the anal opening is on the venter. The + female attains a length of about 0.3 mm. + + ff. Last segment of the palpus forms a thumb to the preceding, + which ends in a claw (with few exceptions); body often + with many hairs (fig. 158 k). TROMBIDOIDEA. + + g. Legs I and II with processes bearing spines; skin with + several shields; coxae contiguous. CAECULIDAE. + + gg. Legs I and II without such processes; few if any + shields. + + h. Palpi much thickened on the base, moving laterally, + last joint often with two pectinate bristles; no eyes; + legs I ending in several long hairs; adult sometimes + parasitic. CHEYLETIDAE + + CHEYLETUS ERUDITUS, which frequents old books, has once + been found in pus discharged from the ear of man. + + hh. Palpi less thickened, moving vertically; eyes usually + present; leg I not ending in long hairs. + + i. Coxae contiguous, radiate; legs slender, bristly; body + with few hairs; no dorsal groove; tarsi not swollen. + ERYTHRAEIDAE. + + ii. Coxae more or less in two groups; legs less bristly. + +[Illustration: 158. (_a_) Tydeus, beak and leg from below; (_b_) +Cheyletus pyriformis, beak and palpus; (_c_) beak and claw of +Pediculoides; (_d_) leg of Sarcoptes; (_e_) scutum of female of +Amblyomma americana; (_f_) leg I and tip of mandible of Histiostoma +americana; (_g_) Histiogaster malus, mandible and venter; (_h_) +Aleurobius farinae, palpus and leg I of male; (_i_) Otodectes cynotis, +tip of abdomen of male, (_j_) beak and anal plate of Dermanyssus +gallinae female; (_k_) palpus of Allothrombium. (_a_) to (_j_) after +Banks.] + + j. Body with fewer, longer hairs; often spinning + threads; no dorsal groove; tarsi never swollen; + mandibles styliform (for piercing). TETRANYCHIDAE + + The genus TETRANYCHUS may be distinguished from the + other genera occurring in the United States by + the following characters: No scale-like + projections on the front of the cephalothorax; + legs I as long or longer than the body; palp + ends in a distinct thumb; the body is about 1.5 + times as long as broad. T. MOLESTISSIMUS Weyenb. + from South America, and T. TELARIUS from Europe + and America ordinarily infesting plants, are + said also to molest man. + + jj. Body with many fine hairs or short spines; not + spinning threads; often with dorsal groove; tarsi + often swollen. + + k. Mandibles styliform for piercing. RHYCHOLOPHIDAE. + + kk. Mandibles chelate, for biting. TROMBIDIDAE + + The genus TROMBIDIUM has recently been subdivided + by Berlese into a number of smaller ones, of + which some five or six occur in the United + States. The mature mite is not parasitic but + the larvae which are very numerous in certain + localities will cause intense itching, + soreness, and even more serious complications. + They burrow beneath the skin and produce + inflamed spots. They have received the popular + name of "RED BUG." The names LEPTUS AMERICANUS + and L. IRRITANS have been applied to them, + although they are now known to be immature + stages. (Fig. 44.) + + +HEXAPODA (Insecta) + +The Thysanura (springtails and bristletails), the Neuropteroids +(may-flies, stone-flies, dragon-flies, caddis-flies, etc.), Mallophaga +(bird lice), Physopoda (thrips), Orthoptera (grasshoppers, crickets, +roaches), are of no special interest from our viewpoint. The remaining +orders are briefly characterized below. + + +SIPHUNCULATA (page 275) + +Mouth parts suctorial; beak fleshy, not jointed; insect wingless; +parasitic upon mammals. Metamorphosis incomplete. Lice. + + +HEMIPTERA (page 275) + +Mouth parts suctorial; beak or the sheath of the beak jointed; in the +mature state usually with four wings. In external appearance the +immature insect resembles the adult except that the immature form (i.e. +nymph) never has wings, the successive instars during the process of +growth, therefore, are quite similar; and the metamorphosis is thus +incomplete. To this order belong the true bugs, the plant lice, leaf +hoppers, frog hoppers, cicadas, etc. + + +LEPIDOPTERA + +The adult insect has the body covered with scales and (with the rare +exception of the females of a few species) with four wings also covered +with scales. Proboscis, when present, coiled, not segmented, adapted for +sucking. Metamorphosis complete, i.e. the young which hatches from the +egg is quite unlike the adult, and after undergoing several molts +transforms into a quiescent pupa which is frequently enclosed in a +cocoon from which the adult later emerges. The larvae are known as +caterpillars. Butterflies and moths. + + +DIPTERA (page 285) + +The adult insect is provided with two, usually transparent, wings, the +second pair of wings of other insects being replaced by a pair of +halteres or balancers. In a few rare species the wings, or halteres, or +both, are wanting. The mouth parts, which are not segmented, are adapted +for sucking. The tarsi are five-segmented. Metamorphosis complete. The +larvae, which are never provided with jointed legs, are variously known +as maggots, or grubs, or wrigglers. Flies, midges, mosquitoes. + + +SIPHONAPTERA (page 316) + +Mouth parts adapted for sucking; body naked or with bristles and spines; +prothorax well developed; body compressed; tarsi with five segments; +wings absent. Metamorphosis complete. The larva is a wormlike creature. +Fleas. + + +COLEOPTERA + +Adult with four wings (rarely wanting), the first pair horny or +leathery, veinless, forming wing covers which meet in a line along the +middle of the back. Mouth parts of both immature stages and adults +adapted for biting and chewing. Metamorphosis complete. The larvae of +many species are known as grubs. Beetles. + + +HYMENOPTERA + +Adult insect with four, usually transparent, wings, wanting in some +species. Mouth parts adapted for biting and sucking; palpi small; tarsi +four or five-segmented. Metamorphosis complete. Parasitic four-winged +flies, ants, bees, and wasps. + + +SIPHUNCULATA AND HEMIPTERA + +a. Legs with claws fitted for clinging to hairs; wings wanting; + spiracles of the abdomen on the dorsal surface. (= ANOPLURA = + PARASITICA) SIPHUNCULATA. + + b. Legs not modified into clinging hooks; tibia and tarsus very long + and slender; tibia without thumb-like process; antennae + five-segmented. HAEMATOMYZIDAE Endr. + + _Haematomyzus elephantis_ on the elephant. + + bb. Legs modified into clinging hooks; tibia and tarsus usually short + and stout; tibia with a thumb-like process; head not anteriorly + prolonged, tube-like. + + c. Body depressed; a pair of stigmata on the mesothorax, and + abdominal segments three to eight; antennae three to + five-segmented. + + d. Eyes large, projecting, distinctly pigmented; pharynx short and + broad; fulturae (inner skeleton of head) very strong and broad, + with broad arms; proboscis short, scarcely attaining the + thorax. PEDICULIDAE + + e. Antennae three-segmented. A few species occurring upon old + world monkeys. _Pedicinis_ Gerv. + + ee. Antennae five-segmented. + + f. All legs stout; thumb-like process of the tibia very long + and slender, beset with strong spines, fore legs stouter + than the others; abdomen elongate, segments without + lateral processes; the divided telson with a conical + process posteriorly upon the ventral side. PEDICULUS L. + + g. Upon man. + + h. Each abdominal segment dorsally with from one to three + more or less regular transverse rows of small setae; + antenna about as long as the width of the head. Head + louse (fig. 65). P. HUMANUS. + + hh. "No transverse rows of abdominal setae; antenna longer + than the width of the head; species larger." Piaget. + Body louse of man. P. CORPORIS. + + gg. Upon apes and other mammals. _P. pusitatus_ (?). + + ff. Fore legs delicate, with very long and slender claws; + other legs very stout with short and stout claws; + thumb-like process of the tibia short and stout; abdomen + very short and broad; segment one to five closely crowded, + thus the stigmata of segments three to five apparently + lying in one segment; segments five to eight with lateral + processes; telson without lateral conical appendages (fig. + 69). Crab louse of man. PHTHIRUS PUBIS. + + dd. Eyes indistinct or wanting; pharynx long and slender, fulturae + very slender and closely applied to the pharynx; proboscis + very long. Several genera found upon various mammals. + HAEMATOPINIDAE. + + cc. Body swollen; meso- and metathorax, and abdominal segments two + to eight each with a pair of stigmata; eyes wanting; antennae + four or five-segmented; body covered with stout spines. Three + genera found upon marine mammals. ECHINOPHTHIRIIDAE + +aa. Legs fitted for walking or jumping; spiracles of abdomen usually + ventral; beak segmented. + + b. Apex of head usually directed anteriorly; beak arising from its + apex; sides of the face remote from the front coxae; first pair of + wings when present thickened at base, with thinner margins. + HETEROPTERA + +[Illustration: 159. Taxonomic details of Hemiptera-Heteroptera. (_a_) +Dorsal aspect; (_b_) seta from bedbug; (_c_) wing of Heteropteron; (_d_) +leg; (_e_) wing of Sinea.] + + c. Front tarsi of one segment, spade-form (palaeformes); beak short, + at most two-segmented; intermediate legs long, slender; + posterior pair adapted for swimming. CORIXIDAE + + cc. Front tarsi rarely one-segmented, never spade-form; beak free, + at least three-segmented. + + d. Pulvilli wanting. + + e. Hemelytra usually with a distinct clavus (fig. 159), clavus + always ends behind the apex of the scutellum, forming the + commissure. (Species having the wings much reduced or + wanting should be sought for in both sections.) + + f. Antennae very short; meso- and metasternum composite; eyes + always present. + + g. Ocelli present; beak four-segmented. OCHTERIDAE and + NERTHRIDAE. + + gg. Ocelli wanting; antennae more or less hidden in a groove. + + h. Anterior coxae inserted at or near anterior margin of + the prosternum; front legs raptorial; beak + three-segmented BELOSTOMIDAE (with swimming legs), + NEPIDAE, NAUCORIDAE. + + i. Metasternum without a median longitudinal keel; + antennae always four-segmented. + + j. Beak short, robust, conical; the hairy fleck on the + corium elongate, large, lying in the middle + between the inner angle of the membrane and the + outer vein parallel to the membrane margin; + membrane margin S-shaped. + + k. The thick fore femur with a relatively deep + longitudinal furrow to receive the tibia. + Several American species (fig. 19f.). BELOSTOMA + (= Lethocerus Mayer) + + kk. The less thickened fore femur without such a + furrow. B. GRISEUS. BENACUS Stal. + + jj. Beak slender, cylindrical; the hairy spot on the + corium rounded lying next to the inner angle of + the membrane. + + k. Membrane large, furrow of the embolium broadened. + _Z. aurantiacum_, _fluminea_, etc. _Zaitha_ + + kk. Membrane very short; furrow of embolium not + broadened. Western genus. _Pedinocoris_ + + ii. Metasternum with a long median longitudinal keel. + Southwestern forms. _Abedus ovatus_ and _Deniostoma + dilatato_ + + hh. Anterior coxae inserted at the posterior margin of the + prosternum; legs natatorial. Back swimmers (fig. 19 b.). + NOTONECTIDAE + + i. Apices of the hemelytra entire; the three pairs of legs + similar in shape; beak three-segmented; abdomen not + keeled or hairy. _Plea_ Leach + + ii. Apices of hemelytra notched; legs dissimilar; beak + four-segmented; abdomen keeled and hairy. + + j. Hemelytra usually much longer than the abdomen; + fourth segment of the antenna longer than the third + segment; hind tarsi with claws. _Bueno_ Kirk. + + jj. Hemelytra but little longer than the abdomen; fourth + segment of the antenna shorter than the third + segment; hind tarsi without claws (fig. 19b). + NOTONECTA L. + + ff. Antennae longer than the head; or if shorter, then the eyes + and ocelli absent. + + g. Eyes, ocelli, and scutellum wanting; beak + three-segmented; head short; hemelytra always short; + membrane wanting. Insects parasitic on bats. POLYCTENIDAE + + gg. Eyes present. + + h. First two antennal segments very short, last two long, + pilose, third thickened at the base; ocelli present, + veins of the hemelytra forming cells. DIPSOCORIDAE (= + CERATOCOMBIDAE) including SCHIZOPTERIDAE. + + hh. Third segment of the antenna not thickened at the + base, second as long or longer than the third, rarely + shorter. + + i. Posterior coxae hinged (cardinate), if rarely + rotating, the cuneus is severed, the membrane is one + or two-celled, and the meso- and metasternum are + composite. + + j. Ocelli absent, clypeus dilated toward the apex; + hemelytra always short, membrane wanting. Species + parasitic. Bed bugs, etc. CIMICIDAE + + k. Beak short, reaching to about the anterior coxae; + scutellum acuminate at the apex; lateral margin + of the elytra but little reflexed, apical margin + more or less rounded; intermediate and posterior + coxae very remote. + + l. Body covered with short hairs, only the sides + of the pronotum and the hemelytra fringed with + longer hairs; antennae with the third and + fourth segments very much more slender than + the first and second; pronotum with the + anterior margin very _deeply sinuate_. CIMEX + L. + + m. Sides of the pronotum widely dilated, broader + than the breadth of one eye, and densely + fringed with backward curved hairs; apical + margin of the hemelytra nearly straight, + rounded toward the interior or exterior + angles. + + n. Body covered with very short hairs; second + segment of the antenna shorter than the + third; sides of the pronotum feebly + reflexed, fringed with shorter hairs than + the breadth of one eye; hemelytra with the + commissural (inner) margin rounded and + shorter than the scutellum, apical margin + rounded towards the interior angle. The + common bed bug (fig. 19h). C. LECTULARIUS + Linn + + nn. Body covered with longer hairs; second and + third segments of the antenna of equal + length; side of the pronotum narrowly, but + distinctly, reflexed, fringed with longer + hairs than the breadth of one eye; + hemelytra with the commissural margin + straight and longer than the scutellum, + apical margin rounded towards the exterior + angle. Species found on bats in various + parts of the United States. _C. + pillosellus_ Hov. + + mm. Sides of the pronotum neither dilated, nor + reflexed, fringed with less dense and nearly + straight hairs; hemelytra with the apical + margin distinctly rounded. Parasitic on man, + birds and bats. Occurs in the old world, + Brazil and the West Indies. C. HEMIPTERUS + Fabr. (= rotundatus) + + ll. Body clothed with rather longer silky hairs; + third and fourth segments of the antenna + somewhat more slender than the first and + second; anterior margin of the pronotum _very + slightly sinuate_ or nearly straight in the + middle, produced at the lateral angles. This + is the species which in American collections + is known as _C. hirundinis_, the latter being + an old world form. It is found in swallows + nests. O. VICARIUS. OECIACUS Stal + + kk. Beak long, reaching to the posterior coxae; + scutellum rounded at the apex; lateral margins + of the elytra strongly reflexed, apical margin + slightly sinuate toward the middle; intermediate + and posterior coxae sub-contiguous. This species + infests poultry in southwest United States and + in Mexico. H. INODORUS. HAEMATOSIPHON Champ. + +[Illustration: 160. Pselliopsis (Milyas) cinctus (x2). After C. V. +Riley.] + + jj. Ocelli present, if rarely absent in the female, + then the tarsus has two segments; or if with three + tarsal segments, the wing membrane with one or two + cells. + + k. Beak four-segmented, or with two-segmented tarsi. + ISOMETOPIDAE, MICROPHYSIDAE, and some + CAPSIDAE. + + kk. Beak three-segmented. + + l. Hemelytra with embolium; head horizontal, more + or less conical; membrane with one to four + veins, rarely wanting. ANTHOCORIDAE + + Several species of this family affecting man + have been noted, ANTHOCORIS KINGI and + CONGOLENSE, from Africa and LYCTOCORIS + CAMPESTRIS from various parts of the world. + LYCTOCORIS FITCHII Reuter (fig. 19 j), later + considered by Reuter as a variety of L. + CAMPESTRIS, occurs in the United States. + + ll. Hemelytra without embolium. Superfamily + ACANTHIOIDEA (= SALDAE Fieber and LEPTOPODAE + Fieber) + + ii. Posterior coxae rotating. + + j. Claws preapical; aquatic forms. GERRIDAE and + VELIADAE + + jj. Claws apical. + + k. Prosternum without stridulatory sulcus (notch for + beak). + + l. Tarsus with three segments; membrane with two + or three longitudinal cells from which veins + radiate; rarely with free longitudinal veins + (Arachnocoris) or veins nearly obsolete + (Arbela); clavus and corium coriaceous; ocelli + rarely absent. NABIDAE + + REDUVIOLUS (= CORISCUS) SUBCOLEOPTRATUS (fig. 19 + g), a species belonging to this family, + occurring in the United States, has been + accused of biting man. This insect is flat, + of a jet black color, bordered with yellow + on the sides of the abdomen, and with + yellowish legs. It is predaceous, feeding on + other insects. + + ll. With other combinations of characters. + HYDROMETRIDAE, HENICOCEPHALIDAE, + NAEOGEIDAE, MESOVELIADAE, JOPPEICIDAE + + kk. Prosternum with stridulatory sulcus (notch for + beak); with three segments, short, strong. + + l. Antennae filiform or sometimes more slender + apically, geniculate; wing membrane with two + or three large basal cells; scutellum small or + moderate/ REDUVIIDAE + + For a key to the genera and species see next + page. + + ll. Last antennal segment clavate or fusiform; win + membrane with the veins often forked and + anastomosing; scutellum large; tarsi each with + two segments; fore legs strong. (= PHYMATIDAE) + MACROCEPHALIDAE + + ee. Clavus noticeably narrowed towards the apex, never extending + beyond the scutellum, the two not meeting to form a + commissure; head horizontal, much prolonged between the + antennae, on each side with an antennal tubercle, sometimes + acute; ocelli absent; meso- and metasternum simple; tarsi + each with two segments; body flattened (fig. 19c). ARADIDAE, + including DYSODIIDAE. + + dd. Pulvilli present (absent in one Australian family + THAUMATOCORIDAE in which case there is a membranous + appendage at the tip of the tibia). CAPSIDAE (= + MIRIDAE),[G] _Eotrechus_ (in family GERRIDAE), + NAEOGAIDAE, TINGITIDAE, PIESMIDAE, MYODOCHIDAE, + CORIZIDAE, COREIDAE, ALYDIDAE, PENTATOMIDAE, + SCUTELLERIDAE, etc. + + bb. Apex of head directed ventrally, beak arising from the hinder part + of the lower side of the head; sides of face contiguous to the + front coxae; first pair of wings, when present, of uniform + thickness. Cicadas, scale insects, plant lice (Aphids), + spittle-insects, leaf hoppers, etc. HOMOPTERA + + +REDUVIIDAE OF THE UNITED STATES + +(Adapted from a key given by Fracker). + +a. Ocelli none; wings and hemelytra always present in the adults; no + discoidal areole in the corium near the apex of the clavus. + _Orthometrops decorata_, _Oncerotrachelus acuminatus_, etc., + Pennsylvania and south. _Sarcinae_ + +aa. Ocelli present in the winged individuals; anterior coxae not as long + as the femora. + + b. Hemelytra without a quadrangular or discoidal areole in the corium + near the apex of the clavus. + + c. Ocelli not farther cephalad than the caudal margins of the eyes; + segment two of the antenna single. + + d. Thorax usually constricted caudad of the middle; anterior coxae + externally flat or concave. PIRATINAE + + e. Middle tibiae without spongy fossa, head long, no lateral + tubercle on neck. _S. stria_, Carolina, Ill., Cal. + _Sirthenia_ Spinola + + ee. Middle tibiae with spongy fossa; fore tibiae convex above; + neck with a small tubercle on each side. + + f. Apical portion of anterior tibiae angularly dilated beneath, + the spongy fossa being preceded by a small prominence. + MELANOLESTES Stal + + g. Black, with piceous legs and antennae. N. E. States (fig. + 19a) M. PICIPES + + gg. Sides, and sometimes the whole dorsal surface of the + abdomen red. Ill., and southward. M. ABDOMINALIS + + ff. Tibiae not dilated as in "f"; spongy fossa elongate; + metapleural sulci close to the margin. R. BIGUTTATUS (fig. + 22). South RASAHUS A. and S. + + dd. Thorax constricted in the middle or cephalad of the middle; + anterior tarsi each three-segmented. + + e. Apex of the scutellum narrow, without spines or with a single + spine REDUVIINAE + + f. Antennae inserted in the lateral or dorso-lateral margins of + the head; antenniferous tubercles slightly projecting from + the sides of the head; head produced strongly cephalad; + ocelli at least as far apart as the eyes. + + g. Antennae inserted very near the apex of the head; segments + one and three of the beak short, segment two nearly four + times as long as segment one. R. PROLIXUS. W. I. + RHODNIUS Stal + + gg. Antennae inserted remote from the vertex of the head. + + h. Body slightly hairy; pronotum distinctly constricted; + angles distinct; anterior lobe four-tuberculate, with + the middle tubercles large and conical. _M. + phyllosoma_, large species from the southwest. + _Meccus_ Stal + + hh. Body smooth, margin of the pronotum sinuous, scarcely + constricted; anterior lobe lined with little + tubercles. CONORHINUS Lap. + + i. Surface of the pronotum and prosternum more or less + granular. + + j. Eyes small, head black; body very narrow, a fifth + as wide as long; beak reaches the middle of the + prosternum. California. C. PROTRACTUS + + jj. Eyes large, head fuscous; body at least a fourth + as wide as long. Southern species. _C. + rubrofasciatus_ + + ii. Pronotum and prosternum destitute of granules. + + j. Border of abdomen entirely black except for a + narrow yellowish spot at the apex of one segment. + Texas. _C. gerstaeckeri_ + + jj. Border of abdomen otherwise marked. + + k. Beak slender, joints one and two slightly pilose, + two more than twice as long as one; tubercles at + the apical angles of the pronotum slightly + acute, conical. Md. to Ill. and south. The + masked bed bug hunter (fig. 71). C. SANGUISUGUS + + kk. Beak entirely pilose, joint two a third longer + than joint one; joint one much longer than + three; tubercles at the apical angles of + pronotum slightly elevated, obtuse. Ga., Ill., + Tex., Cal. _C. variegatus_ + + ff. Antenna inserted on top of the head between margins, close + to the eyes; antenniferous tubercles not projecting from + the side of the head. + + g. Anterior lobe of the pronotum with a bispinous or + bituberculate disc; femora unarmed. _S. arizonica_, _S. + bicolor_. Southwestern species. _Spiniger_ Burm. + + gg. Disc of pronotum unarmed; apex of scutellum produced + into a spine; ocelli close to the eyes; eyes large and + close together. REDUVIUS Lamarck + + h. Color piceous. Widely distributed in the United States. + (Fig. 20). R. PERSONATUS + + hh. More or less testaceous in color. Southwestern states. + R. SENILIS + + ee. Apex of scutellum broad, with two or three spines. + ECTRICHODIINAE + + f. First segment of the antenna about as long as the head. _E. + cruciata_ Pa. and south; _E. cinctiventris_, Tex. and Mex. + _Ectrichodia_ L. et S. + + ff. First segment of the antennae short. _P. aeneo-nitens_. + South. _Pothea_ A. et S. + + cc. Ocelli cephalad of the hind margins of the eyes; first segment + of the antennae stout, second segment divided into many smaller + segments. South and west. _Homalocoris maculicollis_, and + _Hammatocerus purcis_. HAMMATOCERINAE + + bb. Hemelytra with a quadrangular or discoidal areole in the corium + near the apex of the clavus (fig. 159e). + + c. Anal areole of the membrane not extending as far proximad as the + costal areole; basal segment of the antenna thickened, porrect; + the other segments slender, folding back beneath the head and + the first segment STENOPODINAE + + d. Head armed with a ramous or furcate spine below each side, + caudad of the eyes. + + e. First segment of the antenna thickened, apex produced in a + spine beyond the insertion of the second segment. Species + from Va., Ill. and south. _Pnirontis_ Stal. + + ee. First segment of the antenna not produced beyond the + insertion of the second segment. _Pygolampis_, N. E. states + and south; _Gnathobleda_, S. W. and Mex. + + dd. Head unarmed below or armed with a simple spine; rarely with a + subfurcate spine at the side of the base. Carolina, Missouri + and south. _Stenopoda_, _Schumannia_, _Diaditus_, _Narvesus_, + _Oncocephalus_ + + cc. Anal areole of membrane extending farther proximad than the + costal areole. + + d. Ocelli farther apart than the eyes. _A. crassipes_, widely + distributed in the United States; other species occur in the + southwest. _Apiomerus_ Hahn. + + dd. Ocelli not so far apart as the eyes. ZELINAE + + e. Sides of mesosternum without a tubercle or fold in front. + + f. Fore femur as long as or longer than the hind femur; first + segment of the beak much shorter than the second. _Z. + audax_, in the north eastern states; other species south + and west. _Zelus_ Fabr. + + ff. Fore femur shorter than the hind femur, rarely of equal + length, in this case the first segment of the beak as long + or longer than the second. + + g. First segment of the beak shorter than the second; fore + femur a little shorter than the hind femur; the first + segment of the beak distinctly longer than the head + before the eyes. _P. cinctus_ a widely distributed + species (fig. 160). _P. punctipes_, _P. spinicollis_, + Cal., Mex. (= _Milyas_) _Pselliopus_ Berg. + + gg. First segment of the beak as long or longer than the + second. + + h. Pronotum armed with spines on the disc. + + i. Juga distinctly prominent at the apex and often acute + or subacute; fore femur distinctly thickened; + hemelytra usually not reaching the apex of the + abdomen. _Fitchia aptera_, N. Y., south and west; + _F. spinosula_, South; _Rocconata annulicornis_, + Texas, etc. + + ii. Juga when prominent, obtuse at apex; eyes full width + of the head; fore femur not thickened; pronotum with + four spines on posterior lobe. _R. taurus_, Pa., + south and west. _Repipta_ Stal. + + hh. Pronotum unarmed on the disc. + + i. Spines on each apical angle of the penultimate + abdominal segment. _A. cinereus_, Pa., and south. + _Atrachelus_ A. et S. + + ii. Apical angle of the penultimate abdominal segment + unarmed. _Fitchia_ (in part); _Castolus ferox_, + Arizona. + + ee. Sides of the mesosternum with a tubercle or fold in front at + the hind angles of the prosternum; first segment of the beak + longer than the part of the head cephalad of the eyes. + + f. Fore femur thickened, densely granulated; hind femur + unarmed. + +[Illustration: 161. Taxonomic details of Diptera. (_a_) Ventral aspect +of abdomen of Cynomyia; (_b_) antenna of Tabanus; (_c_) ventral aspect +of abdomen of Chortophila; (_d_) ventral aspect of abdomen of Stomoxys; +(_e_) claw of Aedes (Culex) sylvestris, male; (_f_) claw of Hippoboscid; +(_g_) foot of dipterous insect showing empodium developed pulvilliform; +(_h_) hind tarsal segment of Simulium vittatum, female; (_i_) foot of +dipterous insect showing bristle-like empodium.] + + g. Fore tibiae each with three long spines on the ventral + side. _S. diadema_ (fig. 159e), a widely distributed + species; and several southwestern species. _Sinea_ A. et + S. + + gg. Fore tibiae unarmed. _A. multispinosa_, widely + distributed; _A. tabida_, Cal. _Acholla_ Stal. + + ff. Fore femur unarmed, rarely a little thickened, a little + granulated. + + g. Pronotum produced caudad over the scutellum, with a high + mesal tuberculate ridge (fig. 19e). A. CRISTATUS. N. Y. + to Cal. and south. ARILUS Hahn. + + gg. Caudal lobe of the pronotum six sided, neither elevated + nor produced caudad. _H. americanus_, Southwest; also + several W. I. and Mexican genera. HARPACTOR Lap. + + +DIPTERA (Mosquitoes, Midges, Flies) + +a. Integument leathery, abdominal segments indistinct; wings often + wanting; parasitic forms. PUPIPARA + + b. Head folding back on the dorsum of the thorax; wingless flies + parasitic on bats. Genus _Nycteribia_. NYCTERIBIIDAE + + bb. Head not folding back upon the dorsum of the thorax; flies either + winged or wingless; parasitic on birds and on bats and other + mammals. + + c. Antennae reduced, wings when present, with distinct parallel veins + and outer crossveins; claws simple; palpi leaf-like, projecting + in front of the head. Flies chiefly found on bats. Several + genera occur in North America. STREBLIDAE + +[Illustration: 162. Hippobosca equina, x4. After Osborn.] + + cc. Antennae more elongate, segments more or less distinctly + separated; head sunk into an emargination of the thorax; wings + when present with the veins crowded toward the anterior margin; + palpi not leaf-like. HIPPOBOSCIDAE + + d. Wings absent or reduced and not adapted for flight. + + e. Wings and halteres (balancers) absent. _M. ovinus_, the sheep + tick. _Melophagus_ Latr. + + ee. Wing reduced (or cast off), halteres present. + + f. Claw bidentate; ocelli present. On deer after the wings are + cast off. _L. depressa_. _Lipoptena_ Nitsch + + ff. Claw tridentate (fig. 161 f). On _Macropis_. _B. + femorata._ _Brachypteromyia_ Will. + + dd. Wings present and adapted for flight. + + e. Claws bidentate. + + f. Ocelli present; head flat; wings frequently cast off. On + birds before casting of the wing. _Lipoptena_ Nitsch. + + ff. Ocelli absent; head round; wings present. The horse tick + H. EQUINA may attack man (fig. 162). HIPPOBOSCA L. + + ee. Claws tridentate (fig. 161 f.). + + f. Anal cell closed at apical margin by the anal crossvein. + + g. Ocelli absent. _Stilbometopa_ Coq. + + gg. Ocelli present. + + h. R_{4+5} does not form an angle at the crossvein. On + birds. There is a record of one species of this genus + attacking man. ORNITHOMYIA Latr. + + hh. R_{4+5} makes an angle at the crossvein. _O. + confluens_. _Ornithoica_ Rdi. + + ff. Anal cell not closed by an anal crossvein. _Lynchia_, + _Pseudolfersia_, and _Olfersia_ are chiefly bird + parasites. The first mentioned genus is said to be the + intermediate host of _Haemoproteus columbae_. + +aa. Abdominal segments chitinous; not parasitic in the adult stage. + + b. Antennae with six or more segments and empodium not developed + pulvilliform; palpi often with four segments. + + c. Ocelli present. BLEPHAROCERIDAE, RHYPHIDAE, BIBIONIDAE, + MYCETOPHILIDAE, besides some isolated genera of other families. + + cc. Ocelli absent. + + d. Dorsum of the thorax with a V-shaped suture; wings usually with + numerous veins; legs often very long and slender. Crane flies. + TIPULIDAE + + dd. Dorsum of the thorax without a V-shaped suture. + + e. Not more than four longitudinal veins ending in the wing + margin; wing usually hairy: antennae slender; coxae not + long; tibiae: without spurs, legs long and slender. Small, + delicate flies often called Gall gnats. CECIDOMYIIDAE. + + ee. More than four longitudinal veins ending in the wing margin. + + f. The costal vein is not produced beyond the tip of the wing; + radius with not more than three branches. + + g. Antennae short, composed of ten or eleven closely united + segments; legs stout; body stout; abdomen oval; anterior + veins stout, posterior ones weak (fig. 163 b); eyes of + the male contiguous over the antennae. Black flies, + buffalo flies, turkey gnats. Many North American + species, several of them notorious for their blood + sucking propensities. SIMULIIDAE + + h. Second joint of the hind tarsus with basal scale-like + process and dorsal excision (fig. 161 h); radial + sector not forked; no small cell at the base of the + wing. _S. forbesi_, _jenningsi_, _johannseni_, + _meridionale_, _piscicidium_, VENUSTUM, VITTATUM, etc. + Widely distributed species (= EUSIMULIUM) SIMULIUM + Latr. + + hh. No basal scale-like process on the second joint of the + hind tarsus; radial sector usually forked (fig. 163 + b). + + i. Face broad, small basal cell of the wing present. _P. + fulvum_, HIRTIPES, _mutatum_, PECUARUM, _pleurale_. + PROSIMULIUM Roub. + + ii. Face linear; small basal cell of the wing absent. + One species, _P. furcatum_, from California. + _Parasimulium_ Malloch + + gg. Flies of a different structure. + + h. Antennae composed of apparently two segments and a + terminal arista formed of a number of closely united + segments. Rare flies with aquatic larvae. + ORPHNEPHILIDAE + + hh. Antennae of six to fifteen segments, those of the male + usually plumose; legs frequently slender and wings + narrow. CHIRONOMIDAE + + i. Media forked (except in the European genus + _Brachypogon_); thorax without longitudinal fissure + and not produced over the head (except in four + exotic genera); antennae usually fourteen-jointed in + both sexes; fore tibia with a simple comb of setulae, + hind tibia with two unequal combs, middle tibia + without comb. CERATOPOGONINAE + + j. Thorax produced cap-like over the head, wing narrow + and very long. _Jenkinsia_, _Macroptilum_ and + _Calyptopogon_, eastern hemisphere; + _Paryphoconus_, Brazil. + + jj. Thorax not produced over the head. + + k. Eyes pubescent, empodium well developed, or if + short then R_{2+3} distinct and crossvein-like + or the branches of R coalescent; r-m crossvein + present; fore femora not thickened; wing either + with appressed hairs or with microscopic erect + setulae. _Dasyhelea_ Kieff. + + kk. Eyes bare, or otherwise differing from the + foregoing. + + l. Empodium well developed, nearly as long as the + claws and with long hairs at the base; femora + and fifth tarsal segments unarmed, i.e. + without spines or stout setae; fourth tarsal + segment cylindrical. + + m. Wing with erect and microscopic setulae. + Widely distributed. (= Atrichopogon) + _Ceratopogon_ Meig. + + mm. Wing with long and depressed hairs. Widely + distributed. _Forcipomyia_ + + n. Hind metatarsus shorter or not longer than + the following (i.e. the second tarsal) + segment. Subgenus _Prohelea_ Kieff + + nn. Hind metatarsus longer than the following + segment. Subgenus _Forcipomyia_ Meig. + + ll. Empodium short, scarcely reaching the middle + of the claws, or vestigial. + + m. R-m crossvein wanting. + + n. Palpi four segmented; inferior fork of the + media obliterated at the base. Australia. + _Leptoconops_ Skuse + + nn. Palpi three-segmented. + + o. Legs spinulose, tarsal claws of the + female with a basal tooth or strong + bristle, those of the male unequal, the + anterior with a long sinuous tooth, the + posterior with a short arcuate tooth. + Italy. MYCTEROTYPUS Noe + + oo. Legs unarmed; no crossvein between the + branches of the radius (fig. 163e). New + Mexico. TERSESTHES Townsend + + mm. R-m crossvein present. + + n. Fore femora very much swollen, armed with + spines below, fore tibia arcuate and + applied closely to the inferior margin of + the femur. + + o. R_{2+3} present, therefore cell R_1 and + R_2 both present; wing usually fasciate. + United States _Heteromyia_ Say. + + oo. R_{2+3} not distinct from R_{4+5}, hence + cell R_3 obliterated. South America + _Pachyleptus_ Arrib. (Walker) + + nn. Fore femur not distinctly swollen. + + o. R_{2+3} present therefore cells R_1 and + R_3 both present, or if not, then the + branches of the radius more or less + coalescent, obliterating the cells. + + p. At least the tip of the wing with erect + setulae; tip of R_{4+5} scarcely + attaining the middle of the wing, + empodium rather indistinct, not + reaching the middle of the claws, the + claws not toothed, equal, with long + basal bristle; legs without stout + setae. Widely distributed. CULICOIDES + Latr. + + HAEMATOMYIDIUM and OECACTA are probable + synonyms of this. + + pp. Wings bare, if rarely with hair, then + the radius reaches beyond two-thirds + the length of the wing, or the femur + or fifth tarsal segment with stout + black spines. + + q. Media unbranched. Europe. + _Brachypogon_ Kieff + + qq. Media branched. + + r. Hind femur much swollen and spined. + America and Europe. _Serromyia_ + Meg. + + rr. Hind femur not distinctly swollen. + + s. Cell R_1 not longer than high; + fork of the media distad of the + crossvein; wing with microscopic + setulae. _Stilobezzia_ Kieff + + ss. Cell R_1 elongate. + + t. Femora unarmed. Widely + distributed. (= Sphaeromias + Kieff. 1913 not Curtis?). + JOHANNSENIELLA Will. + + tt. Femora, at least in part, with + strong black spines. Widely + distributed. _Palpomyia_ + Megerle + + oo. R_{2+3} coalescent with R_{4+5} hence + cell R_3 is obliterated. + + p. In the female the lower branch of the + media with an elbow near its base + projecting proximad, the petiole of + the media coalescent with the basal + section of the radius, wing long and + narrow, radial sector ending near the + tip of the wing; venation of the male + as in _Bezzia_; front concave. United + States. _Stenoxenus_ Coq. + + pp. Venation otherwise, front not concave. + + q. Subcosta and R_1 more or less + coalescent with the costa; wing + pointed at the apex, much longer + than the body; antennae fourteen + segmented, not plumose. India. + _Haasiella_ Kieff. + + qq. Subcosta and radius distinct from + the costa. + + r. Abdomen petiolate. _Dibezzia_ + Kieff. + + rr. Abdomen not petiolate. + + s. Head semi-globose; hind tarsi + unusually elongate in the + female; antennae of the male not + plumose. Europe. _Macropeza_ + Meigen. + + ss. Head not globose, more or less + flattened in front; antennae of + the male plumose. Widely + distributed. _Bezzia_ Kieff. + + t. Fore femora, at least, armed + with stout spines below. + Subgenus _Bezzia_ Kieff. + + tt. Femora unarmed. Subgenus + _Probezzia_ Kieff. + + ii. Media of the wing simple, and otherwise not as in + "i". To this group belong numerous Chironomid + genera, none of which are known to be noxious to + man. + + ff. The costal vein apparently is continued around the hind + margin of the wing; radius with at least four branches. + + g. Wing ovate pointed, with numerous veins; crossveins, if + evident, before the basal third of the wing; veins very + hairy; very small moth-like flies. PSYCHODIDAE + + h. With elongate biting proboscis; the petiole of the + anterior forked cell of the wing (R_2) arises at or + beyond the middle of the wing (fig. 163d). PHLEBOTOMUS + Rdi. + +[Illustration: 163. Wings of Diptera. (_a_) Anopheles; (_b_) +Prosimulium; (_c_) Johannseniella; (_d_) Phlebotomus (After Doerr and +Russ); (_e_) Tersesthes (after Townsend); (_f_) Tabanus; (_g_) +Symphoromyia; (_h_) Aphiochaeta; (_i_) Eristalis; (_j_) Gastrophilus; +(_k_) Fannia; (_l_) Musca.] + + hh. With shorter proboscis; the petiole of the anterior + forked cell arises near the base of the wing. + _Psychoda_, _Pericoma_, etc. + + gg. The r-m crossvein placed at or beyond the center of the + wing; wings not folded roof-like over the abdomen. + + h. Proboscis short, not adapted for piercing; wings bare + (DIXIDAE); or wings scaled (CULICIDAE, Subf. + CORETHRINAE). + + hh. Proboscis elongate, adapted for piercing; wings + scaled, fringed on the hind margin; antennae of the + male bushy plumose. Mosquitoes. CULICIDAE (exclusive of + CORETHRINAE) + + i. Metanotum without setae. + + j. Proboscis strongly decurved; body with broad, + appressed, metalescent scales; cell R_2 less than + half as long as its petiole; claws of female + simple, some of the claws of the male toothed. + Several large southern species believed to feed + only on nectar of flowers. _Megarhinus_ R. D. + + jj. Proboscis straight or nearly so, or otherwise + different. + + k. Scutellum evenly rounded, not lobed; claws simple + in both sexes. ANOPHELES Meig. + + l. Abdomen with clusters of broad outstanding + scales along the sides; outstanding scales on + the veins of the wing rather narrow, + lanceolate; upper side of the thorax and + scutellum bearing many appressed lanceolate + scales. Florida and southward (CELLIA). + + m. Hind feet from the middle of the second + segment largely or wholly snow white. + + n. With a black band at the base of the last + segment of each hind foot. A. + ALBIMANUS*[H] and TARSIMACULATA* + + nn. Without such a band A. ARGYRITARSIS* + + mm. Hind feet black, mottled with whitish and + with bands of the same color at the sutures + of the segments. W. I. A. MACULIPES + + ll. Abdomen without such a cluster of scales; + outstanding scales of the wing veins rather + narrow, lanceolate; tarsi wholly black. + + m. Deep black, thorax obscurely lined with + violaceous, especially posteriorly; head, + abdomen and legs black; no markings on the + pleura; abdomen without trace of lighter + bindings; wing scales outstanding, uniform, + not forming spots, though little thicker at + the usual points indicating the spottings. + Florida. A. ATROPUS + + mm. Otherwise marked when the wings are + unspotted. + + n. Wings unspotted. + + o. Petiole of the first forked cell (R_2) + more than a third the length of the + cell. Mississippi valley. A. WALKERI + + oo. Petiole of the first forked cell a third + the length of the cell. Md. A. BARBERI + + nn. Wings spotted. + + o. Front margin of the wings with a patch of + whitish and yellow scales at a point + about two-thirds or three-fourths of the + way from base to apex of wing. + + p. Veins of the wing with many broad + obovate outstanding scales; thorax + with a black dot near the middle of + each side. W. I. A. GRABHAMI* + + pp. The outstanding scales of the wings + rather narrow, lanceolate. + + q. Scales of the last vein of the wings + white, those at each end black; + R_{4+5} black scaled, the extreme + apex white scaled. Widely + distributed north and south (fig. + 131). A. PUNCTIPENNIS + + A dark variety from Pennsylvania has + been named A. PERPLEXENS. + + qq. Scales of the last vein of the wing + white, those at its apex black; + R_{4+5} white scaled and with two + patches of black scales. South and + the tropics. A. FRANCISCANUS and + PSEUDOPUNCTIPENNIS* + + oo. Front margin of the wings wholly black + scaled. + + p. Last (anal) vein of the wings white + scaled with three patches of black + scales (fig. 132). New Jersey to + Texas. A. CRUCIANS* + + pp. Last vein of the wings wholly black + scaled. + + q. Widely distributed north and south + (fig. 130), (= MACULIPENNIS). A. + QUADRIMACULATUS* + + qq. Distributed from Rocky Mountains + westward. A. OCCIDENTALIS + + kk. Scutellum distinctly trilobed. + + l. Cell R_2 less than half as long as its petiole; + thorax with metallic blue scales; median lobe + of the scutellum not tuberculate; few small + species which are not common. URANOTAENIA + Arrib. + + ll. Cell R_2 nearly or quite as long as its + petiole, or otherwise distinct. + + m. Femora with erect outstanding scales; occiput + broad and exposed. Large species. P. + CILIATA. P. HOWARDI PSOROPHORA R. D. + + mm. Femora without erect scales. + + n. Clypeus bearing several scales or hairs, + scutellum with broad scales only; back of + head with broad scales; scales along the + sides of the mesonotum narrow; some or the + claws toothed; thorax marked with a pair + of silvery scaled curved stripes; legs + black with white bands at the bases of + some of the segments (fig. 134). Yellow + Fever mosquito. AEDES (= STEGOMYIA) + CALOPUS. + + nn. With another combination of characters. + Numerous species of mosquitoes belonging + to several closely related genera, widely + distributed over the country. (_Culex_, + _Aedes_, _Ochlerotatus_, etc.). CULEX in + the wide sense. + + ii. Metanotum with setae. _Wyeomyia_ (found in the United + States); and related tropic genera. + + bb. Antennae composed of three segments with a differentiated style or + bristle; third segment sometimes complex or annulate, in which + case the empodium is usually developed like the pulvilli, i.e., + pad-like (fig. 161 g). + + c. Empodium developed pad-like (pulvilliform) i.e., three nearly + equal membranous appendages on the underside of the claw (fig. + 161g). + + d. Squamae, head, and eyes large; occiput flattened or concave; + third segment of the antennae with four to eight annuli or + segments, proboscis adapted for piercing; body with fine + hairs, never with bristles; middle tibia with two spurs; wing + venation as figured (fig. 163f); marginal vein encompasses the + entire wing. Horse flies, greenheads, deer flies, gad flies + TABANIDAE[I] + + e. Hind tibia with spurs at tip; ocelli usually present + (PANGONINAE) + + f. Third joint of the antennae with seven or eight segments; + proboscis usually prolonged. + + g. Each section the third antennal segment branched. Central + American species, _P. festae_. _Pityocera_ G. T. + + gg. Sections of the third antennal segment not branched. + + h. Upper corner of the eyes in the female terminating in + an acute angle; wings of both sexes dark anteriorly. + _G. chrysocoma_, a species from the eastern states. + _Goniops_ Ald. + + hh. Upper corner of the eye in the female not so + terminating; wings nearly uniform in color, or + hyaline. + + i. Proboscis scarcely extending beyond the palpi; front + of the female wide; much wider below than above. S. + W. States. _Apatolestes_ Will. + + ii. Proboscis extending beyond the palpi. + + j. Wing with cell M_3 closed. Tropic America. (= + _Diclisa_) _Scione_ Wlk. + + jj. Cell M_3 open; ocelli present or absent. Two or + three eastern species; many south and west + PANGONIA Rdi. + + ff. Third segment of the antenna with five divisions; ocelli + present. + + g. First and second segments of the antenna short, the + second only half as long as the first, three western + species. SILVIUS Rdi. + + gg. First and second segments of the antenna long, the + second distinctly over half as long as the first. Deer + flies. Many species, widely distributed. CHRYSOPS Meig. + + ee. Hind tibia without spurs; ocelli absent. + + f. Third segment of antenna with four divisions, no tooth or + angulation; wings marked with rings and circles of darker + coloring; front of the female very wide. Widely + distributed. _H. americana_, _H. punctulata_. HAEMATOPOTA + Meig. + + ff. Third segment of the antenna with five divisions (fig. + 161b). + + g. Third segment of the antenna not furnished with a tooth + or distinct angular projection. + + h. Body covered with metallic scales; front of female of + normal width; front and middle tibiae greatly dilated. + _L. lepidota_. _Lepidoselaga_ Macq. + + hh. Body without metallic scales; antennae not very long, + the third segment not cylindrical, not situated on a + projecting tubercle; front of the female narrow. + South. _D. ferrugatus._ (= _Diabasis_) _Diachlorus_ O. + S. + + gg. Third segment of the antenna furnished with a tooth or a + distinct angular projection. + + h. Hind tibiae ciliate with long hairs. S. W. and tropics. + _Snowiella_ and _Stibasoma_. + + hh. Hind tibiae not ciliate. + + i. Species of slender build, usually with a banded + thorax and abdomen; third segment of the antenna + slender, the basal prominence long; wings mostly + with brownish markings. Tropic America. + _Dichelacera_ Macq. + + ii. Species of a stouter build; third segment of the + antenna stout, its basal process short (fig. 161b). + Many species, widely distributed TABANUS L. + + dd. With another group of characters. + + e. Squamae small, antennae variable, thinly pilose or nearly + bare species, without distinct bristles; wing veins not + crowded anteriorly, R_4 and R_5 both present, basal cells + large; middle tibiae at least with spurs. LEPTIDAE + + f. Flagellum of the antenna more or less elongated, composed + of numerous more or less distinct divisions. XYLOPHAGINAE + and ARTHROCERATINAE. + + ff. Antennae short, third segment simple, with arista or style; + face small, proboscis short LEPTINAE + + g. Front tibiae each with one or two spurs, or if absent, + then no discal cell. _Triptotricha_, _Pheneus_, + _Dialysis_, _Hilarimorpha_. + + gg. Front tibiae without terminal spurs, discal cell + present. + + h. Hind tibiae each with a single spur. + + i. Anal cell open (fig. 163g); third antennal segment + kidney-shaped with dorsal or subdorsal arista; first + antennal segment elongate and thickened. About a + dozen species have been described from the United + States, of which at least one (S. PACHYCERAS) is + known to be a vicious blood sucker. SYMPHOROMYIA + Frauenf. + + ii. Anal cell closed; third antennal segment not + kidney-shaped. _Chrysopila_, _Ptiolina_, _Spania_. + + hh. Hind tibiae each with two spurs. + + i. Third segment kidney-shaped, the arista subdorsal; + anal cell closed. _Atherix_ Meig. + + ii. Third segment of the antenna short and with terminal + arista; anal cell open _Leptis_ Fabr. Two European + species of this genus have been accused of blood + sucking habits, but the record seems to have been + based upon error in observation. + + ee. With another combination of characters. STRATIOMYIIDAE, + CYRTIDAE, etc. + + cc. Empodium bristlelike or absent. + + d. Antennae apparently two segmented, with three-segmented arista, + wings (rarely wanting) with several stout veins anteriorly, + the weaker ones running obliquely across the wing (fig. 163h); + small, quick running, bristly, humpbacked flies. Several + genera; APHIOCHAETA, PHORA, TRINEURA, etc. PHORIDAE + + dd. Flies with other characters. + + e. No frontal lunule above the base of the antennae; both R_4 + and R_5 often present; third segment of the antenna often + with a terminal bristle. ASILIDAE, MYDAIDAE, + APIOCERIDAE, THEREVIDAE, SCENOPINIDAE, BOMBYLIIDAE, + EMPIDIDAE, DOLICHOPODIDAE, LONCHOPTERIDAE. + + ee. A frontal lunule above the base of the antennae; third + segment of the antenna always simple, i.e., not ringed, + usually with a dorsal arista; R_4 and R_5 coalesced into a + simple vein. + + f. A spurious vein or fold between the radius and the media, + rarely absent; the cell R_{4+5} closed at the apex by vein + M_1; few or no bristles on the body, none on the head; + flies frequently with yellow markings. ERISTALIS (fig. + 163i), HELOPHILUS, and many other genera. SYRPHIDAE + + ff. No spurious vein present. + + g. Body without bristles; proboscis elongate and slender, + often folding; front of both male and female broad. + CONOPIDAE + + gg. Bristles almost always present on head, thorax, abdomen + and legs. + + h. Arista terminal; hind metatarsus enlarged, sometimes + ornamented, hind tarsus more or less flattened beneath. + PLATYPEZIDAE + + hh. Flies having a different combination of characters. + + i. Head large, eyes occupying nearly the entire head; + cell R_{4+5} narrowed in the margin; small flies. + PIPUNCULIDAE + + ii. Head and eyes not unusually large. + + j. Squamae (tegulae, or calyptrae, or alulae) + not large, often quite small, the lower one + lacking, or at most barely projecting from below + the upper one (antisquama); front of both male and + female broad, the eyes therefore widely separated; + posthumeral and intraalar macrochaeta not + simultaneously present; thorax usually without a + complete transverse suture; postalar callus + usually absent; the connectiva adjoining the + ventral sclerites always visible; hypopleural + macrochaetae absent; last section of R_{4+5} + and M_{1+2} with but few exceptions nearly + parallel; subcostal vein often wanting or + vestigial or closely approximated to R_1; the + latter often short, basal cells small, the + posterior ones often indistinct or wanting; + vibrissae present or absent. ACALYPTRATE + MUSCOIDEA + + k. Subcosta present, distinctly separated from R_1 + at the tip; R_1 usually ends distad of the + middle of the wing; the small basal cells of the + wing distinct. + + l. A bristle (vibrissa) on each side of the face + near the margin of the mouth. CORDYLURIDAE, + SEPSIDAE, PHYCODROMIDAE, HETERONEURIDAE, + HELOMYZIDAE. + + ll. No vibrissae present. + + m. Head nearly spherical, cheeks broad and + retreating; proboscis short; the cell R_5 + closed or narrowed in the margin; legs very + long; tarsi shorter than the tibiae. CALOBATA + and other genera. MICROPEZIDAE + + mm. Flies with another combination of + characters. RHOPALOMERIDAE, TRYPETIDAE, + ORTALIDAE, SCIOMYZIDAE. + + kk. Subcosta absent or vestigial, or if present, + then apparently ending in the costa at the point + where R_1 joins it; R_1 usually ends in the + costa at or before the middle of the wing. + + l. Arista long plumose, or pectinate above; oral + vibrissae present; anal cell complete; costa + broken at the apex of R_1. DROSOPHILA, + PHORTICA, and other genera. DROSOPHILIDAE + + ll. With another combination of characters. + + m. The cell M and first M_2 not separated by a + crossvein; anal cell absent; front bare or + only bristly above; usually light colored + flies. HIPPELATES, OSCINUS, and other + genera. (See also m m m.) OSCINIDAE + + mm. Cell M and cell first M_2 often separated by + a crossvein; anal cell present, complete, + though frequently small; scutellum without + spines or protuberances; oral vibrissae + present; arista bare or short plumose; front + bristly at vertex only; small dark flies. + PIOPHILA (fig. 99), SEPSIS and other genera. + SEPSIDAE + + mmm. The GEOMYZIDAE, AGROMYZIDAE, + PSILIDAE, TRYPETIDAE, RHOPALOMERIDAE, + BORBORIDAE and DIOPSIDAE differ in + various particulars from either the + OSCINIDAE and the SEPSIDAE noted above. + + jj. Squamae well developed, usually large, the lower + one frequently projecting from below the upper + one; both posthumeral and intraalar macrochaetae + present; thorax with a complete transverse suture; + postalar callus present and separated by a + distinct suture from the dorsum of the thorax; + front of the female broad, of the male frequently + narrow, the eyes then nearly or quite contiguous; + the connectiva adjoining the ventral sclerites + either visible or not; hypopleural macrochaetae + present or absent; subcosta always distinct in its + whole course, R_1 never short. CALYPTRATE + MUSCOIDEA[J] + + k. Oral opening small, mouth parts usually much + reduced or vestigial. This family is undoubtedly + of polyphyletic origin but for convenience it is + here considered as a single family. OESTRIDAE. + + l. The costal vein ends at the tip of R_{4+5}, + M_{1+2} straight, not reaching the wing + margin, hence cell R_5 wide open (fig. 163j); + squamae small; arista bare; ovipositor of the + female elongate. Larvae in the alimentary canal + of horses, etc. GASTROPHILUS + + m. Posterior crossvein (m-cu) wanting; wings + smoky or with clouds. Europe. G. PECORUM + + mm. Posterior crossvein (m-cu) present, at least + in part. + + n. Wing hyaline with smoky median cross band, + and two or three spots; posterior + trochanters with hook in the male and a + prominence in the female. World wide + distribution. G. EQUI. + + nn. Wings without spots. + + o. Posterior crossvein (m-cu) distad of the + anterior crossvein (r-m); legs, + particularly the femora, blackish brown. + Europe and North America. G. + HAEMORRHOIDALIS + + oo. Posterior crossvein opposite or proximad + of the anterior crossvein. Europe and + North America. G. NASALIS + + ll. The costal vein ends at the tip of M_{1+2}, + M_{1+2} with a bend, the cell R_5 hence much + narrowed in the margin, or closed. + + m. Proboscis geniculate, inserted in a deep + slit; female without extricate ovipositor; + arista either bare or plumose; squamae large; + facial grooves approximated below. + + n. Arista bare, short. Larvae in rodents. + Tropic America. _B. princeps_. _Bogeria_ + Austen + + nn. Arista pectinate above. + + o. Tarsi broadened and flattened, hairy, + anal lobe of the wing large. Larvae in + rodents. A number of American species. + _Cuterebra_. + + oo. Tarsi slender, not hairy; anal lobe of + the wing moderate. Larvae in man and + other mammals. Tropic America. D. + CYANIVENTRIS. DERMATOBIA Br. + + mm. Mouth parts very small, vestigial; arista + bare. + + n. Facial grooves approximated below, leaving + a narrow median depression or groove. + + o. Cell R_5 closed and petiolate, body + nearly bare. Larvae in the nasal cavities + of the smaller Ungulates. The sheep bot + fly. O. OVIS. Widely distributed OESTRUS + L. + + oo. Cell R_5 narrowly open, body hairy. + Larvae parasitic on deer. Europe and + America. _Cephenomyia_ Latr. + + nn. Facial grooves far apart, enclosing + between them a broad shield-shaped + surface; squamae large; female with + elongate ovipositor. Larvae hypodermatic on + Ungulates HYPODERMA Clark + + o. Palpi wanting; tibiae thickened in the + middle. + + p. Hair at apex of the abdomen yellow; + legs including femora yellowish brown. + H. DIANA + + pp. Hair at the apex of the abdomen + reddish yellow. Europe and America. + + q. Tibiae and tarsi yellow; femora black. + H. LINEATA + + qq. Legs black with black hair; tips of + hind tibia and tarsi yellowish + brown. H. BOVIS + + oo. Palpi small, globular; tibiae + cylindrical, straight. On reindeer. _O. + tarandi_ _Oedemagena_ Latr. + + kk. Oral opening of the usual size; mouth parts not + vestigial. + + l. Hypopleurals wanting; if three sternopleurals + are present the arrangement is 1:2; + conjunctiva (fig. 161c) of the venter usually + present; if the terminal section of M_{1+2} is + bent it has neither fold nor appendage + (ANTHOMYIIDAE of Girschner). + + m. Sternopleurals wanting; M_{1+2} straight + toward the apex, costa ends at or slightly + beyond the tip of R_{4+5}; mouth parts + vestigial. GASTROPHILINAE. See OESTRIDAE + + mm. Sternopleurals present, if rarely absent + then differing in other characters. + + n. Caudal margin of the fifth ventral + abdominal sclerite of the male deeply + notched on the median line usually to + beyond the middle; abdomen often + cylindrical or linear; abdomen often with + four to eight spots; eyes of the male + usually widely separated; sternopleurals + three, arranged in an equilateral + triangle; subapical seta of the hind tibia + placed very low; M_{1+2} straight; anal + vein abbreviated; wings not rilled. + _Caenosia_, _Caricea_, _Dexiopsis_, + _Hoplogaster_, _Schoenomyia_, etc. + (COENOSINAE)[K]. ANTHOMYIIDAE in part + + nn. Caudal margin of the fifth ventral + abdominal sclerite of the male incurved, + rarely deeply cleft, rarely entire, in a + few genera deeply two or three notched; + M_{1+2} straight or curved; abdomen + usually short or elongate oval; + sternopleurals, if three are present, + arranged in the order 1:2 in a right + triangle. (MUSCINAE-ANTHOMYIINAE of + Girschner) + + o. M_{1+2} straight, hence cell R_5 not + narrowed in the margin. ANTHOMYIIDAE in + part + + p. Underside of the scutellum more or less + sparsely covered with fine hairs; anal + vein nearly always reaches the hind + margin of the wing; extensor surface + of the hind tibiae with a number of + stout setae; squamae often small and + equal. ANTHOMYIA, _Chortophila_, + _Eustalomyia_, _Hammomyia_, + _Hylemyia_, _Prosalpia_, _Pegomyia_, + etc. HYLEMYINAE-PEGOMYINAE + + pp. Underside of the scutellum bare; anal + vein does not reach the wing margin. + + q. First anal vein short, second anal + suddenly flexed upwards; hind tibiae + each with one or two strong setae on + the extensor surface. FANNIA (= + HOMALOMYIA), _Coelomyia_, + _Choristoma_, _Euryomma_, _Azelia_, + etc. FANNINAE-AZELINAE + + qq. Anal veins parallel or divergent. + + r. Setae on the exterior surface of the + hind tibiae wanting (except in + _Limnaricia_ and _Coenosites_), + lower squama not broadened to the + margin of the scutellum. + _Leucomelina_, _Limnophora_, + _Limnospila_, _Lispa_, _Mydaea_, + _Spilogaster_, etc. + MYDAEINAE-LIMNOPHORINAE + + rr. One (rarely more) seta on the + extensor surface of the hind + tibia; squamae usually large and + unequal. HYDROTAEA, _Aricia_, + _Drymeia_, _Ophyra_, _Phaonia_ (= + _Hyetodesia_), _Pogonomyia_, + _Trichophthicus_, etc. ARICINAE + + oo. M_{1+2} curved or bent, hence the cell + R_5 more or less narrowed in the margin. + (MUSCINAE). MUSCIDAE in part. See + page 303 for generic synopsis. + + ll. Hypopleurals present; when three + sternopleurals are present the arrangement is + 2:1 or 1:1:1. (TACHINIDAE of Girschner) + + m. Conjunctiva of the ventral sclerites of the + abdomen present, frequently well developed, + surrounding the sclerites. + + n. Mouth parts vestigial. OESTRIDAE. See page + 297 for generic synopsis. + + nn. Mouth parts well developed. + + o. M_{1+2} straight, hence cell R_5 wide + open in the margin; costa ending at the + tip of R_5; three sternopleurals + present; antennal arista plumose. + _Syllegoptera_. Europe. + (SYLLEGOPTERINAE). DEXIIDAE in part + + oo. M_{1+2} bent, hence cell R_5 narrowed in + the margin; sternopleurals rarely + wanting, usually 1:1 or 0:1; facial + plate strongly produced below vibrissal + angle like the bridge of the nose; + antennal arista bare. Parasitic on + Hemiptera and Coleoptera. _Allophora_, + _Cistogaster_, _Clytia_, _Phasia_, etc. + (PHASIINAE) TACHINIDAE in part. + + mm. Conjunctiva of the ventral sclerites + invisible (fig. 161a). + + n. Second ventral sclerite of the abdomen + lying with its edges either upon or in + contact with the ventral edges of the + corresponding dorsal sclerite. + + o. Outermost posthumeral almost always lower + (more ventrad) in position than the + presutural macrochaeta; fifth ventral + abdominal sclerite of the male cleft + beyond the middle, often strongly + developed; body color very frequently + metallic green or blue, or yellow; + arista plumose. (CALLIPHORINAE) + MUSCIDAE in part. + + See page 303 for generic synopsis. + + oo. Outermost posthumeral macrochaeta on + level or higher (more dorsad) than the + presutural macrochaeta; arista bare, + pubescent, or plumose only on the basal + two-thirds; body coloring usually + grayish (fig. 106). SARCOPHAGIDAE + + p. Fifth ventral sclerite of the male + either wanting or with the caudal + margin straight; presutural intraalar + rarely present. (SARCOPHAGINAE) + + q. Fifth ventral abdominal sclerite of + the male much reduced, the remaining + segments with straight posterior + margin, overlapping scale-like; in + the female only segment one and two + scale-like, the others wholly or in + part covered; sternopleurals usually + three or more. SARCOPHAGA and + related genera. + + qq. Fifth ventral sclerite of the male + plainly visible; sternopleurals + usually two. SARCOPHILA, + WOHLFAHRTIA, _Brachycoma_, + _Hilarella_, _Miltogramma_, + _Metopia_, _Macronychia_, _Nyctia_, + _Paramacronychia_, _Pachyphthalmus_, + etc. + + pp. Fifth ventral abdominal sclerite of + the male cleft to beyond the middle; + ventral sclerites usually visible, + shield-like. _Rhinophora_, _Phyto_, + _Melanophora_ RHINOPHORINAE + +[Illustration: 164. Glossina palpalis. (x4.) After Austen.] + + nn. Second ventral abdominal sclerite as well + as the others more or less covered, + sometimes wholly, by the edges of the + dorsal sclerite. + + o. The presutural intraalar wanting; ventral + sclerites two to five nearly or quite + covered by the edges of the + corresponding dorsal sclerites; base of + the antennae usually at or below the + middle of the eye; arista usually + plumose; legs usually elongate; + abdominal segments with marginal and + often discal macrochaetae. DEXIIDAE + + oo. Presutural intraalar present, if absent, + then the ventral sclerites broadly + exposed or the fifth ventral sclerite + vestigial; base of the antennae usually + above the middle of the eye; arista + bare; at least two posthumerals and + three posterior intraalars present. + Parasitic on caterpillars, etc. + TACHINIDAE + + +SYNOPSIS OF THE PRINCIPAL GENERA OF THE MUSCIDAE OF THE WORLD + +a. Proboscis long, directed forward, adapted for piercing, or oral + margin much produced, snout-like. + + b. Oral margin produced snout-like; vibrissa placed high above the + oral margin; antennal arista either pectinate or more or less + plumose. + + c. Antennal arista short or long-plumose; neither sex with distinct + orbital bristles. + + d. No facial carina between the antennae. RHYNCHOMYIINAE + + e. Arista short-plumose. _R. speciosa._ Europe. _Rhynchomyia_ R. + D. + + ee. Arista long-plumose. _I. phasina_. Europe and Egypt. + _Idiopsis_. B. B. + + dd. With flattened carina, the bases of the antennae separated; no + abdominal macrochaetae. COSMININAE _C. fuscipennis_. South + Africa. _Cosmina_ + + cc. Antennal arista pectinate; bases of the antennae separated by a + flattened carina. RHINIINAE R. D. + + d. Cell R_5 open, or closed at the margin. + + e. Third segment of the antenna twice as long as the second; + claws of both sexes short; cell R_5 open. _I. lunata._ + Eastern Hemisphere. _Idia_ Meigen + + ee. Third segment of the antenna three times as long as the + second; cell R_5 open or closed; claws of the male long and + slender, of the female shorter than the last tarsal joint. + _I. mandarina_, China. _Idiella_ B. B. + + dd. Cell R_5 petiolate. _Rhinia_; and _Beccarimyia_ Rdi. + + bb. Proboscis long, directed forward, adapted for piercing. STOMOXINAE + + c. Arista flat, pectinate above with plumose rays; sternopleurals + 1:2; bases of the veins R_1 and R_{4+5} without setae; base of + the media bowed down; apical cell opens before the apex of the + wing. African species. GLOSSINA Wied. + + d. Species measuring over twelve mm. in length. _G. longipennis_ + and FUSCA. + + dd. Species less than twelve mm. in length. + + e. All segments of the hind tarsi black. + + f. The fourth and fifth segments of the fore tarsi black; + antennae black (fig. 164). G. PALPALIS R. D. + + ff. Otherwise marked. _G. bocagei_, _tachinoides_, + _pallicera._ + + ee. First three segments of the hind tarsi are yellow, the + fourth and fifth segments are black. + + f. Fourth and fifth segments of the first and second pair of + tarsi are black. + + g. The yellow bands of the abdominal segments occupy a third + of the segment (fig. 165). G. MORSITANS Westw. + + gg. The yellow band on each segment of the abdomen occupies + a sixth of the segment. G. LONGIPALPIS Wied. + + ff. Tarsi of the first and second pairs of legs wholly yellow. + _G. pallidipes_ Austen + + cc. Rays of the arista not plumose; only one or two sternopleurals; + base of the media not strongly bowed down; apical cell opens at + or very near the apex of the wing. + + d. Vein R_{4+5} without setae at the base; palpi about as long as + the proboscis. + + e. Arista pectinate (i.e. rays on one side only), the rays often + undulate; two yellow sternopleurals often difficult to + detect; vein M_{1+2} only slightly bent, the apical cell + hence wide open. The horn fly, H. IRRITANS (= _Lyperosia + serrata_) and related species. Widely distributed (figs. + 167, 168). HAEMATOBIA R. D. not B. B. + +[Illustration: 165. Glossina morsitans. (x4.) After Austen.] + + ee. Arista also with rays below; vein M_{1+2} more strongly + bent, the apical cell hence less widely open. + + f. Palpi strongly spatulate at the tips, lower rays of the + arista about six in number, B. SANGUINOLENTUS. South Asia. + BDELLOLARYNX Austen + + ff. Palpi feebly spatulate; apical cell of the wing narrowly + open slightly before the tip; sternopleurals black, + anterior bristle sometimes absent. H. ATRIPALPIS. Europe. + HAEMATOBOSCA Bezzi + + dd. Vein R_{4+5} with setae at the base.[L] + + e. Veins R_1 and R_{4+5} with setae at the base; two equally + prominent sternopleural macrochaetae; arista with rays both + above and below; palpi as long as the proboscis; apical cell + of the wing wide open. L. TIBIALIS. (_Haematobia_ B. B. not + R. D.). LYPEROSIOPS Town. + + ee. Only vein R_{4+5} with basal setae; anterior sternopleural + macrochaeta wanting; arista pectinate. + + f. Palpi as long as the proboscis, the latter stout, with + fleshy terminal labellae; apical cell narrowly open; + sternopleural macrochaetae black. S. MACULOSA from Africa + and related species from Asia. STYGEROMYIA Austen + + ff. Palpi much shorter than the proboscis, the latter pointed + at the apex, without fleshy labellae; apical cell of the + wing wide open. S. CALCITRANS, the stable fly and related + species. Widely distributed in both hemispheres (fig. + 110). STOMOXYS Geof. + +aa. Proboscis neither slender nor elongate, the labellae fleshy and not + adapted for piercing. + + b. Hypopleurae without a vertical row of macrochaetae. MUSCINAE + + c. Arista bare; distal portion of M_{1+2} broadly curved at the end; + hypopleurae with a sparse cluster of fine hairs. _S. + braziliana_, Southern States and southward. _Synthesiomyia_ B. + B. + + cc. Arista pectinate or plumose. + + d. Arista pectinate. _H. vittigera_, with the posterior half of + the abdomen metallic blue. Mexico. _Hemichlora_ V. d. W. + + dd. Arista plumose. + + e. Middle tibia with one or more prominent setae on the inner + (flexor) surface beyond the middle, or inner surface very + hairy. + + f. R_1 ends distad of the m-cu crossvein; M_{1+2} with a broad + curve near its apical end. (= _Neomesembrina_ Schnabl. = + _Metamesembrina_ Town). _M. meridiana._ Europe. + _Mesembrina_ Meigen + + ff. R_1 ends proximad of the m-cu crossvein. + + g. Eyes pilose, sometimes sparsely in the female. + + h. Female with two or three stout orbital setae; the hind + metatarsus of the male thickened below at the base and + penicillate. _D. pratorum._ Europe. _Dasyphora_ R. + D.[M] + + hh. Neither sex with orbital setae. + + i. Abdomen without macrochaetae; arista plumose. _C. + asiatica._ Eastern Hemisphere. _Cryptolucilia_ B. B. + + ii. Abdomen with strong macrochaetae; arista very + short-plumose, nearly bare. _B. tachinina._ Brazil. + _Reinwardtia_ B. B. + + gg. Eyes bare. + + h. Body densely pilose; thoracic macrochaetae wanting; + middle tibiae much elongate and bent; last section of + M_{1+2} with a gentle curve. H. (_Mesembrina_) + _mystacea, et al._, Europe and _H. solitaria_, N. + America. _Hypodermodes_ Town. + + hh. Body not densely pilose. + + i. Dorsocentrals six; last section of M_{1+2} with a + gentle curve. + + j. Inner dorsocentrals ("acrostichals") wanting; + sternopleurals arranged 1:3. _P. cyanicolor_, + _cadaverina_, etc. Europe and America _Pyrellia_ + R. D. + + jj. Inner dorsocentrals ("acrostichals") present; + sternopleurals arranged 1:2. E. _latreillii._ + North America. _Eumesembrina_ Town. + + ii. Dorsocentrals five; inner dorsocentrals present; + last section of M_{1+2} with a rounded angle; + sternopleurals arranged 1:2. _P. cornicina_ Europe + and America. (_Pseudopyrellia_ Girsch.) _Orthellia_ + R. D. + + ee. Middle tibia without a prominent bristle on the inner + surface beyond the middle. + +[Illustration: 166. Pycnosoma marginale. (x4.) After Graham-Smith.] + + f. Squamula thoracalis broadened mesad and caudad as far as + the scutellum. + + g. Sternopleural macrochaetae arranged in an equilateral + triangle; front of both sexes broad; genae bare; + dorsocentrals six, small; wing not rilled. (To + COENOSINAE). _Atherigona_ Rdi. + + gg. Sternopleural macrochaetae when three are present, + arranged in a right triangle. + + h. Last section of M_{1+2} with a more or less rounded + angle (fig. 163l). + + i. Eyes of the male pilose or pubescent, of the female + nearly bare; m-cu crossvein usually at or proximad + of the mid-distance between the r-m crossvein and + the bend of M_{1+2}. P. (= _Placomyia_ R. D.) + _vitripennis_. _Plaxemyia_ R. D. + + ii. Eyes bare; the m-cu crossvein always nearer to the + bend of M_{1+2} than to the r-m crossvein. + + j. Apex of the proboscis when extended reveals a + circlet of stout chitinous teeth. P. INSIGNIS + Austen, of India, bites both man and animals. (= + _Pristirhynchomyia_.) PHILAEMATOMYIA Austen + + jj. Apex of the proboscis without black teeth. + + k. Eyes of male separated by a distance equal to a + fourth the width of the head. House or typhoid + fly. M. DOMESTICA L. Widely distributed. MUSCA + L. + + kk. Eyes of the male contiguous. E. CORVINA. Europe. + EUMUSCA Town + + hh. Last section of M_{1+2} with a gentle curve (fig. + 102). + + i. Eyes pilose. + + j. Claws in the male somewhat elongated; no orbitals + in either sex; antennae separated at the base by a + flat carina; abdomen marked with red or yellow. + _G. maculata._ Europe and America. _Graphomyia_ R. + D. + + jj. Claws short and equal in the two sexes; two or + three stout orbital macrochaetae in the + female; R_1 scarcely produced beyond the r-m + crossvein; eyes contiguous in the male. _P. + obsoleta._ Brazil. _Phasiophana_ Br. + + ii. Eyes bare; fronto-orbital macrochaetae in a double + row, antennae contiguous at the base. + + j. One or more pairs of well developed anterior inner + dorsocentral (acrostichal) macrochaetae; seta on + extensor surface of hind tibia. M. ASSIMILIS, + STABULANS, etc. Europe and America. MUSCINA R. D. + + jj. Anterior inner dorsocentrals and the setae; on the + extensor surface of the hind tibia wanting. _M. + micans_, etc. Europe and North America. _Morellia_ + R. D. + + ff. Squamula thoracalis not broadened mesad and caudad, not + reaching the margin of the scutellum; macrochaetae on + extensor surface of the hind tibia wanting. + + g. Eyes pubescent. _M. meditabunda._ Europe and America. + _Myiospila_ Rdi. + + gg. Eyes bare; R_1 ends before the middle of the wing. A + number of species from the tropics of both hemispheres. + _Clinopera_ V. d. W. + + bb. Hypopleurae with a vertical row of macrochaetae. + + c. Eyes pubescent. + + d. R_1 ends about opposite the r-m crossvein; basal section of + R_{4+5} bristly nearly to the crossvein; _S. enigmatica_. + Africa. _Somalia_ Hough + + dd. R_1 ends distad of the r-m crossvein. + + e. Eastern hemisphere. Australasia. _N. ochracea_, + _dasypthalma_. _Neocalliphora_ Br. + + ee. Western Hemisphere. _T. muscinum._ Mexico. _Tyreomma_ V. d. + W. + + cc. Eyes bare. + + d. The vibrissal angle situated at a noticeable distance above the + level of the margin of the mouth. + + e. Sternopleural macrochaetae arranged in the order 1:1. + + f. Genae with microchaetae. + + g. Body grayish, with depressed yellow woolly hair among the + macrochaetae; wings folded longitudinally over the body + when at rest. Cluster flies. _P. rudis_ and related + species, widely distributed. _Pollenia_ R. D.[N] + + gg. Body metallic blue or green. Eastern Hemisphere. + + h. Vibrissal angle placed very high above the oral margin; + a carina between the antennae; outer posthumeral + wanting; anterior intraalar present. _T. viridaurea_. + Java. _Thelychaeta_ Br. + +[Illustration: 167. Horn fly. (_a_) egg; (_b_) larva; (_c_) puparium; +(_d_) adult. (x4). Bureau of Entomology] + + hh. Vibrissal angle moderately high above the oral margin; + carina small or wanting; no post humeral macrochaeta; + lower squamae hairy above. (= _Paracompsomyia_ Hough) + (fig. 166). _Pycnosoma_ Br. + + ff. Genae bare. _S. terminata_. Eastern Hemisphere. + _Strongyloneura_ Bigot + + ee. Sternopleurals arranged 2:1. + + f. Body metallic green or blue, with gray stripes; genae hairy + to the lower margin; post humerals often wanting; lower + squamae bare above. (= _Compsomyia_ Rdi.). CHRYSOMYIA R. D. + + g. With one or two orbitals; height of bucca less than half + the height of the eye. South and east U. S. (fig. 107). + C. MARCELLARIA + + gg. No orbitals; height of bucca about a third less than + height of eye. West U. S. _C. wheeleri_ Hough + + ff. Body black or sordidly metallic greenish gray, usually + yellow pollinose or variegate; genae at most hairy above. + _N. stygia_. Eastern Hemisphere. _Neopollenia_ Br. + + dd. Vibrissal angle situated nearly on a level of the oral margin. + + e. Species wholly blackish, bluish, or greenish metallic in + color. + + f. First section of R_{4+5} with at most three or four small + bristles at the immediate base. + + g. The bend of M_{1+2} a gentle curve; costal spine present; + cell R_5 closed, ending before the apex of the wing. _S. + cuprina._ Java. _Synamphoneura_ Bigot + + gg. Bend of M_{1+2} angular; or the insect differs in other + characters; dorsal surface of the squamula thoracalis + hairy (except in _Melinda_); arista plumose only on the + basal two-thirds (except usually in _Calliphora_ and + _Eucalliphora_). + +[Illustration: 168. Head of horn-fly (Lyperosia irritans); (_a_) female; +(_b_) male; (_c_) lateral aspect of female. Bureau of Entomology] + + h. Arista plumose only on the basal two-thirds. + + i. Base of the antennae ventrad of the middle of the + eye; eyes of the male nearly contiguous; genae + hairy; second abdominal segment with median marginal + macrochaetae; two, rarely three, postsutural + intraalar macrochaetae. + + j. Squamula thoracalis dorsally with long black hairs; + male hypopygium two-segmented, large, projecting; + claws and pulvilli of the male elongate; three + strong sternopleural macrochaetae; genae at + least half the width of the eye; buccae (cheeks) + half the height of the eyes; oviviparous. _O. + sepulcralis._ Europe. _Onesia_ R. D. + + jj. Dorsal surface of the squamula thoracalis bare; + male hypopygium small, scarcely projecting below; + claws and pulvilli not elongate; two stout + sternopleural macrochaetae, sometimes with a + delicate one below the anterior; genae nearly + linear in the male; buccae about a third of the eye + height; oviparous. _M. caerulea_. Europe. + _Melinda_. R. D. + +[Illustration: 169. Lateral and dorsal aspects of the thorax, and +frontal aspect of the head of a muscoidean fly, with designations of the +parts commonly used in taxonomic work.] + + ii. Base of the antennae dorsad of the middle of the eye; + eyes of both sexes distinctly separated; dorsal + surface of the squamula thoracalis with black hairs; + two postsutural intraalar macrochaetae. + + j. Hypopygium of the male large, with a pair of + slightly curved forceps whose ends are concealed + in a longitudinal slit in the fifth ventral + sclerite; third posterior inner dorso-central + (acrostichal) macrochaetae absent; anterior + intraalar rarely present; abdomen usually not + pollinose; the second segment without median + marginal macrochaetae; face yellow. _C. + mortuorum_, _cadaverina_, and related species. + Both hemispheres. _Cynomyia_ R. D.[O] + +[Illustration: 170. Sepsis violacea; puparium and adult. (See page 297.) +After Howard.] + + jj. Three pairs of posterior inner dorsocentrals + (acrostichals) present; second abdominal segment + with a row of marginal macrochaetae; genae + hairy, at least above. + + k. Hypopygium of the male with a projecting style. + _S. stylifera_. Europe. _Steringomyia_ Pok. + + + kk. Hypopygium of the male without style. _A. + stelviana_ B. B. _Acrophaga_ B. B. + + hh. Arista usually plumose nearly to the tip; posterior + dorsocentrals and inner dorsocentrals (acrostichals) + well developed; dorsal surface of the squamula + thoracalis hairy; abdomen metallic and usually + pollinose; genae hairy. + + i. With one pair of ocellar macrochaetae. C. VOMITORIA, + ERYTHROCEPHALA, VIRIDESCENS, and related species. + Both hemispheres. CALLIPHORA R. D. + + ii. With two strong pairs of ocellar macrochaetae. _E. + latifrons._ Pacific slope of the U. S. + _Eucalliphora_ Town. + + ff. First section of R_{4+5} bristly near or quite half way to + the small crossvein; dorsal surface of the squamula + thoracalis is bare; the hypopygium of the male is + inconspicuous. + + g. Genae bare; posterior inner and outer dorsocentrals + distinct and well developed. _L. caesar_, _sericata_, + _sylvarum_, and related species. Widely distributed in + both hemispheres (fig. 103). LUCILIA R. D. + + gg. Genae with microchaetae, at least down to the + level of the base of the arista. + + h. Mesonotum flattened behind the transverse suture. + + i. Posterior dorsocentrals inconstant and unequally + developed; one pair of posterior inner + dorsocentrals. _P. terraenovae._ North America. + _Protophormia_ Town. + + ii. Posterior dorsocentrals well developed, the inner + dorsocentrals (acrostichals) unequally developed. + _P. azurea_, _chrysorrhoea_, etc. Europe and America. + _Protocalliphora_ Hough + + hh. Mesonotum not flattened behind the transverse suture; + posterior inner and outer dorsocentrals inconstant and + unequally developed. _P. regina._ Europe and America. + _Phormia_ R. D. + + ee. Species more or less rufous or yellow in color. + + f. Anterior dorsocentrals wanting; first section of the + R_{4+5} at most only bristly at the base, bend near apex + of M_{1+2} rectangular, R_1 ends over the crossvein; + fronto-orbital macrochaeta absent; eyes of the male + contiguous. _C. semiviridis._ Mexico. _Chloroprocta_ + V. d. W + + ff. With another combination of characters. + + g. Body robust, of large size, abdomen elongate, not round; + genae with several ranges of microchaetae; + vibrissal ridges strongly convergent; abdomen with well + developed macrochaetae; costal spine usually absent; + eyes of the male widely separated. + +[Illustration: 171. Stigmata of the larvae of Muscoidea. Third instar. +(_a_) Cynomyia cadaverina; (_b_) Phormia regina; (_c_) Chrysomyia +macellaria; (_d_) Musca domestica; (_e_) Sarcophaga sp.; (_f_) Oestris +ovis; (_g_) Gastrophilus equi; (_h_) Sarcophaga sp.; (_i_) Pegomyia +vicina; (_j_) Protocalliphora azurea; (_k_) Hypoderma lineata; (_l_) +Muscina stabulans. Magnification for f, g, and k, x25; all others, x50.] + + h. Peristome broad, pteropleural macrochaetae + distinct; one or two sternopleurals; in the female a + single orbital macrochaeta; last abdominal segment + without discal macrochaetae; hypopygial processes + of the male with a long stylet; second abdominal + segment of the female sometimes much elongate. A. + LUTEOLA (fig. 86). Africa. The sub-genus + _Choeromyia_ Roub. is included here. AUCHMEROMYIA B. + B. + + hh. Peristome narrow; no pteropleurals, two + sternopleurals; two orbitals in the female; second + segment not elongate; the fourth with two well + developed discal macrochaetae. B. DEPRESSA. Africa. + BENGALIA R. D + + gg. With another combination of characters. + + h. Costal spine present; body in part black; antennae + noticeably shorter than the epistome, inserted above + the middle of the eye and separated from each other by + a carina; abdominal segments with marginal + macrochaetae; sternopleurals 2:1 or 1:1. + _Paratricyclea_ Villen. + + hh. Costal spine not distinct, or if present, insect + otherwise different. + + i. Genae with several ranges of microchaetae; + vibrissal ridges strongly converging; peristome + broad; arista moderately plumose; sternopleurals + usually 1:1; color entirely testaceous. C. + ANTHROPOPHAGA (fig. 87) and GRUNBERGI. Africa. + CORDYLOBIA Grunb. + + ii. Genae bare or with but one range of setae; + vibrissal ridges less converging; peristome narrow; + arista long plumose. + + j. Genae with a single row of microchaetae. + + k. Sternopleurals 2:1; color entirely testaceous. + _Ochromyia_ Macq.[P] + + kk. Sternopleurals 1:1. _P. varia_ Hough. Africa. + _Parochromyia_ Hough + + jj. Genae bare. + + k. Basal section of R_{4+5} bristly only at the + immediate base, distally M_{1+2} with a broad + curve; distal portion of the abdomen metallic; + sternopleurals usually 1:1, rarely 2:1. _M. + aeneiventris_ Wd. Tropic America. + _Mesembrinella._ G. T. + + kk. R_{4+5} bristly at least nearly half way to the + small crossvein; sternopleurals 1:1. + + l. Macrochaetae of the abdomen marginal; + neither sex with orbitals; no carina between + the base of the antennae; three pairs of + presutural inner dorsocentrals. Eastern + hemisphere. _T. ferruginea._ _Tricyclea V. d. + W_. (= _Zonochroa_ B. B. according to + Villeneuve 1914). + + ll. Abdomen without macrochaetae; wing + usually with a marginal streak and gray + markings. Brazil. Hemilucilia B. B. + +[Illustration: 172. Left hand stigmata of the larvae of muscoidea. Third +instar. (_a_) Lucilia caesar; (_b_) Calliphora vomitoria; (_c_) Stomoxys +calcitrans; (_d_) Orthellia cornicina; (_e_) Pyrellia cadavarina; +(_f_) Haematobia irritans; (_g_) Mesembrina mystacea; (_h_) Mesembrina +meridiana; (_i_) Myospila meditabunda; (_j_) Mydaea urbana; (_k_) +Polietes albolineata; (_l_) Polietes lardaria; (_m_) Morellia hortorum; +(_n_) Hydrotaea dentipes; (_o_) Hebecnema umbratica; (_p_) H. vespertina; +(_q_) Limnophora septemnotata; (_r_) Muscina stabulans. (_a_ and _b_) +after MacGregor; (_d_) after Banks; all others after Portchinsky. +Magnification varies. The relative distance to the median line is +indicated in each figure.] + + +SIPHONAPTERA. Fleas + +Adapted from a table published by Oudemans. + +a. Elongated fleas, with jointed (articulated) head, with combs + (ctenidia) on head and thorax; with long, oval, free-jointed + flagellum of the antenna (fig. 92d). Suborder FRACTICIPATA + + b. With ctenidia in front of the antennae and on the genae (cheeks); + maxillae with acute apices; labial palpi five-segmented, + symmetrical; eyes poorly developed or wanting. On rodents. + HYSTRICHOPSYLLIDAE + + c. Abdominal segments without ctenidia. + + d. Post-tibial spines in pairs and not in a very close set row; + head with ctenidia. _Ctenophthalmus_ Kol. + + dd. Post-tibial spines mostly single and in a close set row. + _Ctenopsyllus_ and _Leptopsyllus_. The last genus has recently + been erected for _L. musculi_, a widely distributed species + occurring on rats and mice. + + cc. Abdominal segments with one or more ctenidia; post-tibial spines + in numerous, short, close-set transverse rows on posterior + border with about four spines in each row. _H. americana._ + _Hystrichopsylla_ Taschenb. + + bb. With only two pairs of subfrontal ctenidia; labial palpi + five-segmented, symmetrical; eyes vestigial or wanting. On bats. + (= ISCHNOPSYLLIDAE). NYCTERIDIPSYLLIDAE + + With more or less blunt maxilla; all tibiae with notch; a single + antepygidial bristle; metepimeron without ctenidium. _N. + crosbyi_ from Missouri was found on bats. Rothschild suggests + that this is probably the same as _N. insignis_. (= + _Ischnopsyllus_ = _Ceratopsyllus_), _Nycteridiphilus_ + +aa. Head not jointed, i.e. the segments coalescent, traces of the + segmentation still being visible in the presence of the vertex + tubercle, the falx (sickle-shaped process), and a suture. Suborder + INTEGRICIPITA + + b. Flagellum of the antennae long and oval. + + c. Usually elongate fleas, with a free-segmented flagellum of the + antenna; thorax not shorter than the head, longer than the first + tergite. + + d. Genae of the head and the pronotum with ctenidia. NEOPSYLLIDAE + + e. Labial palpi four or five-segmented; symmetrical; hind coxae + with patch of spines inside; row of six spatulate spines on + each side in front of the antennae. _C. ornate_ found on a + California mole. _Corypsylla_ + + ee. Labial palpi two-segmented, transparent, membranous. On + hares. _Spilopsyllus_ Baker + + dd. No ctenidium on the head. + + e. Pronotum with ctenidium. DOLICHOPSYLLIDAE + + f. Labial palpi five-segmented, symmetrical. + + g. Antepygidial bristles one to three; eyes present. + + h. Inner side of hind coxae distally with a comb of minute + teeth; falx present. On rodents and carnivores. + _Odontopsyllus_ Baker + + hh. Inner side of hind coxae without comb or teeth. Many + North American species on rodents. CERATOPHYLLUS + Curtis + + gg. Antepygidial bristles five on each side; eyes absent; + suture white. _D. stylosus_ on rodents. _Dolichopsyllus_ + Baker + + ff. Labial palpi four or five-segmented; asymmetrical + (membranous behind), apex acute. _Hoplopsyllus anomalus_ + found on Spermophiles in Colorado. HOPLOPSYLLIDAE + + ee. Pronotum without ctenidium. _Anomiopsyllus californicus_ and + _nudatus_ on rodents. ANOMIOPSYLLIDAE + + cc. Very short fleas; flagellum of the antenna with pseudo-segments + coalescent; thorax much shorter than the head and than the first + tergite. HECTOPSYLLIDAE + + Flagellum of the antenna with six coalescent pseudo-segments; + maxilla blunt. The chigger on man (fig. 93). D. PENETRANS. (= + RHYNCHOPRION = SARCOPSYLLA) DERMATOPHILUS Guerin + + bb. Flagellum short, round, free portion of the first segment shaped + like a mandolin. + + c. Thorax not shorter than the head, longer than the first tergite; + flagellum either with free segments or in part with the segments + coalescent. + + d. Head and pronotum with ctenidium; labial palpi asymmetrical. + ARCHAEOPSYLLIDAE + + With four subfrontal, four genal, and one angular ctenidia. + Widely distributed. CTENOCEPHALUS Kol. + + e. Head rounded in front (fig. 92a). Dog flea. C. CANIS + + ee. Head long and flat (fig. 92b). Cat flea. C. FELIS + + dd. Neither head nor pronotum with ctenidium. Labial palpi + asymmetrical, membranous behind. PULICIDAE + + e. Mesosternite narrow, without internal rod-like thickening + from the insertion of the coxae upwards. Human flea, etc. + PULEX L. + + ee. Mesosternite broad with a rod-like internal thickening from + the insertion of the coxae upwards (fig. 89). X. + (LOEMOPSYLLA) CHEOPIS, plague or rat flea. XENOPSYLLA + + cc. Thorax much shorter than the head and than the first tergite. + ECHIDNOPHAGIDAE. E. GALLINACEA, the hen flea also attacks man + (fig. 96). (= ARGOPSYLLA = XESTOPSYLLA) ECHIDNOPHAGA Olliff. + + +FOOTNOTES: + +[E] Adapted from Banks, Nuttall, Warburton, Stiles, _et al._ + +[F] Dr. C. W. Stiles considers the species which is responsible for +spotted fever distinct from the _venustus_ of Banks, separating it as +follows: + +Goblet cells about 75 in the male or 105 in the female. Texas. _D. +venustus._ + +Goblet cells 157 in the male, or 120 in the female; stigmal plate shaped +as shown in the figure (figs. 150 a, b). Montana, etc. D. ANDERSONI. + +[G] Professor C. R. Crosby who has been working upon certain capsids +states that he and his assistant have been bitten by LYGUS PRATENSIS, +the tarnished plant bug, by CHLAMYDATUS ASSOCIATUS and by ORTHOTYLUS +FLAVOSPARSUS, though without serious results. + +[H] Species marked with an * are known to transmit malaria. Species +found only in tropical North America and not known to carry malaria have +been omitted from this table, but all found in the United States are +included. + +[I] This table to the North American genera of the Tabanidae is adapted +from one given by Miss Ricardo. + +[J] The classification of the Muscoidea as set forth by Schiner and +other earlier writers has long been followed, although it is not +satisfactory, being admittedly more or less artificial. Within the last +two or three decades several schemes have been advanced, that of Brauer +and Bergenstamm and of Girschner, with the modifications of Schnabl and +Dziedzicki having obtained most favor in Europe. Townsend, in 1908, +proposed a system which differs from Girschner's in some respects, but +unfortunately it has not yet been published in sufficient detail to +permit us to adopt it. From considerations of expediency we use here the +arrangement given in Aldrich's Catalogue of North American Diptera, +though we have drawn very freely upon Girschner's most excellent paper +for taxonomic characters to separate the various groups. + +It may sometimes be found that a species does not agree in all the +characters with the synopsis; in this case it must be placed in the +group with which it has the most characters in common. + +[K] There are several genera of flies of the family _Cordyluridae_; (i.e. +_Acalyptratae_) which might be placed with the _Anthomyiidae_ (i.e. +_Calyptratae_), owing to the relatively large size of their squamae. As +there is no single character which will satisfactorily separate all +doubtful genera of these two groups we must arbitrarily fix the limits. +In general those forms on the border line having a costal spine, or +lower squama larger than the upper, or the lower surface of the +scutellum more or less pubescent, or the eyes of the male nearly or +quite contiguous, or the eyes hairy, or the frontal setae decussate in +the female; or any combination of these characters may at once be placed +with the _Anthomyiidae_. Those forms which lack these characteristics and +have at least six abdominal segments (the first and second segments +usually being more or less coalescent) are placed with the Acalyptrates. +There are other acalyptrates with squamae of moderate size which have +either no vibrissae, or have the subcosta either wholly lacking or +coalescent in large part with R_1 or have spotted wings; they, therefore +will not be confused with the calyptrates. + +[L] _Pachymyia_ Macq. is closely related to _Stomoxys_. It differs in +having the arista rayed both above and below. _P. vexans_, Brazil. + +[M] The genus _Eudasyphora_ Town. has recently been erected to contain +_D. lasiophthalma_. + +[N] _Nitellia_, usually included in this genus has the apical cell +petiolate. _Apollenia_ Bezzi, has recently been separated from +_Pollenia_ to contain the species _P. nudiuscula_. Both genera belong to +the Eastern hemisphere. + +[O] The following three genera are not sufficiently well defined to +place in this synopsis. In color and structural characters they are +closely related to _Cynomyia_ from which they may be distinguished as +follows. _Catapicephala_ Macq., represented by the species _C. +splendens_ from Java, has the setae on the facial ridges rising to the +base of the antennae and has median marginal macrochaetae on the +abdominal segments two to four: _Blepharicnema_ Macq., represented by +_B. splendens_ from Venezuela has bare genae, oral setae not +ascending; tibiae villose; claws short in both sexes; _Sarconesia_ +Bigot with the species _S. chlorogaster_ from Chile, setose genae; +legs slender, not villose; claws of the male elongate. + +[P] _Plinthomyia_ Rdi. and _Hemigymnochaeta_ Corti are related to +_Ochromyia_, though too briefly described to place in the key. + + + + +APPENDIX + +HYDROCYANIC ACID GAS AGAINST HOUSEHOLD INSECTS + + +The following directions for fumigating with hydrocyanic acid +gas are taken from Professor Herrick's circular published by the +Cornell Reading Course: + +Hydrocyanic acid gas has been used successfully against household +insects and will probably be used more and more in the future. +It is particularly effective against bed-bugs, and cockroaches, but +because _it is such a deadly poison it must be used very carefully_. + +The gas is generated from the salt potassium cyanid, by treating +it with sulfuric acid diluted with water. Potassium cyanid is a +most poisonous substance and the gas emanating from it is also +deadly to most, if not all, forms of animal life. The greatest care +must always be exercised in fumigating houses or rooms in buildings +that are occupied. Before fumigation a house should be vacated. +It is not necessary to move furniture or belongings except brass or +nickel objects, which may be somewhat tarnished, and butter, milk, +and other larder supplies that are likely to absorb gas. If the nickel +and brass fixtures or objects are carefully covered with blankets +they will usually be sufficiently protected. + +There may be danger in fumigating one house in a solid row of +houses if there is a crack in the walls through which the gas may find +its way. It also follows that the fumigation of one room in a house +may endanger the occupants of an adjoining room if the walls between +the two rooms are not perfectly tight. It is necessary to keep +all these points in mind and to do the work deliberately and thoughtfully. +The writer has fumigated a large college dormitory of 253 +rooms, once a year for several years, without the slightest accident +of any kind. In order to fumigate this building about 340 pounds +of cyanid and the same amount of sulfuric acid were used each time. +In addition to this, the writer has fumigated single rooms and smaller +houses with the gas. In one instance the generating jars were too +small; the liquid boiled over and injured the floors and the rugs. +Such an accident should be avoided by the use of large jars and by +placing old rugs or a quantity of newspapers beneath the jars. + + +THE PROPORTIONS OF INGREDIENTS + +Experiments and experience have shown that the potassium +cyanid should be ninety-eight per cent pure in order to give satisfactory +results. The purchaser should insist on the cyanid being of +at least that purity, and it should be procurable at not more than +forty cents per pound. The crude form of sulfuric acid may be used. +It is a thickish, brown liquid and should not cost more than four or +five cents a pound. If a room is made tight, one ounce of cyanid for +every one hundred cubic feet of space has been shown to be sufficient. +It is combined with the acid and water in the following proportions: + + Potassium cyanid 1 ounce + Commercial sulfuric acid 1 fluid ounce + Water 3 fluid ounces + + +A SINGLE ROOM AS AN EXAMPLE + +Suppose a room to be 12 by 15 by 8 feet. It will contain 12 x 15 x 8, or +1440 cubic feet. For convenience the writer always works on the basis of +complete hundreds; in this case he would work on the basis of 1500 cubic +feet, and thus be sure to have enough. The foregoing room, then, would +require 15 ounces of cyanid, 15 ounces of sulfuric acid, and 45 ounces +of water. The room should be made as tight as possible by stopping all +the larger openings, such as fireplaces and chimney flues, with old rags +or blankets. Cracks about windows or in other places should be sealed +with narrow strips of newspaper well soaked in water. Strips of +newspaper two or three inches wide that have been thoroughly soaked in +water may be applied quickly and effectively over the cracks around the +window sash and elsewhere. Such strips will stick closely for several +hours and may be easily removed at the conclusion of the work. + +While the room is being made tight, the ingredients should be measured +according to the formula already given. The water should be measured and +_poured first_ into a stone jar for holding at least two gallons. The +jar should be placed in the middle of the room, with an old rug or +several newspapers under it in order to protect the floor. + +The required amount of sulfuric acid should then be poured rather slowly +into the water. _This process must never be reversed; that is, the acid +must never be poured into the jar first._ The cyanid should be weighed +and put into a paper bag beside the jar. All hats, coats, or other +articles that will be needed before the work is over should be removed +from the room. When everything is ready the operator should drop the bag +of cyanid gently into the jar, holding his breath, and should walk +quickly out of the room. The steam-like gas does not rise immediately +under these conditions, and ample time is given for the operator to walk +out and shut the door. If preferred, however, the paper bag may be +suspended by a string passing through a screw eye in the ceiling and +then through the keyhole of the door. In this case the bag may be +lowered from the outside after the operator has left the room and closed +the door. + +The writer has most often started the fumigation toward evening and left +it going all night, opening the doors in the morning. The work can be +done, however, at any time during the day and should extend over a +period of five or six hours at least. It is said that better results +will be obtained in a temperature of 70^o F., or above, than at a lower +degree. + +At the close of the operation the windows and doors may be opened from +the outside. In the course of two or three hours the gas should be +dissipated enough to allow a person to enter the room without danger. +The odor of the gas is like that of peach kernels and is easily +recognized. The room should not be occupied until the odor has +disappeared. + + +FUMIGATING A LARGE HOUSE + +The fumigation of a large house is merely a repetition, in each room and +hall, of the operations already described for a single room. All the +rooms should be made tight, and the proper quantities of water and +sulfuric acid should be measured and poured into jars placed in each +room with the cyanid in bags besides the jars. When all is ready, the +operator should _go to the top floor and work downward_ because the gas +is lighter than air and tends to rise. + + +PRECAUTIONS + +The cyanid should be broken up into small pieces not larger than small +eggs. This can best be done on a cement or brick pavement. It would be +advantageous to wear gloves in order to protect the hands, although the +writer has broken many pounds of cyanid without any protection on the +hands. Wash the hands thoroughly at frequent intervals in order to +remove the cyanid. + +The operations of the work must be carried out according to directions. + +The work should be done by a calm, thoughtful and careful person--best +by one who has had some experience. + +Conspicuous notices of what has been done should be placed on the doors, +and the doors should be locked so that no one can stray into the rooms. + +The gas is lighter than air, therefore one should always begin in the +rooms at the top of the house and work down. + +After fumigation is over the contents of the jar should be emptied into +the sewer or some other safe place. The jars should be washed thoroughly +before they are used again. + +_It must be remembered that cyanid is a deadly poison_; but it is very +efficient against household insects, if carefully used, and is not +particularly dangerous when properly handled. + + +LESIONS PRODUCED BY THE BITE OF THE BLACK-FLY + +While this text was in press there came to hand an important paper +presenting a phase of the subject of black fly injury so different from +others heretofore given that we deem it expedient to reproduce here the +author's summary. The paper was published in _The Journal of Cutaneous +Diseases_, for November and December, 1914, under the title of "A +Clinical, Pathological and Experimental Study of the Lesions Produced by +the Bite of the Black Fly (_Simulium venustum_)," by Dr. John Hinchman +Stokes, of the University of Michigan. + + +RESUME AND DISCUSSION OF EXPERIMENTAL FINDINGS + +The principal positive result of the work has been the experimental +reproduction of the lesion produced by the black-fly in characteristic +histological detail by the use of preserved flies. The experimental +lesions not only reproduced the pathological pictures, but followed a +clinical course, which in local symptomatology especially, tallied +closely with that of the bite. This the writer interprets as +satisfactory evidence that the lesion is not produced by any living +infective agent. The experiments performed do not identify the nature of +the toxic agent. Tentatively they seem to bring out, however, the +following characteristics. + +1. The product of alcoholic extraction of flies do not contain the toxic +agent. + +2. The toxic agent is not inactivated by alcohol. + +3. The toxic agent is not destroyed by drying fixed flies. + +4. The toxic agent is not affected by glycerin, but is, if anything, +more active in pastes made from the ground fly and glycerin, than in the +ground flies as such. + +5. The toxic agent is rendered inactive or destroyed by hydrochloric +acid in a concentration of 0.25%. + +6. The toxic agent is most abundant in the region of the anatomical +structures connected with the biting and salivary apparatus (head and +thorax). + +7. The toxic agent is not affected by a 0.5% solution of sodium +bicarbonate. + +8. The toxic agent is not affected by exposure to dry heat at 100^o C. +for two hours. + +9. The toxic agent is destroyed or rendered inactive in alkaline +solution by a typical hydrolytic ferment, pancreatin. + +10. Incomplete experimental evidence suggests that the activity of the +toxic agent may be heightened by a possible lytic action of the blood +serum of a sensitive individual, and that the sensitive serum itself may +contain the toxic agent in solution. + +These results, as far as they go (omitting No. 10), accord with Langer's +except on the point of alcoholic solubility and the effect of acids. The +actual nature of the toxic agent in the black-fly is left a matter of +speculation. + +The following working theories have suggested themselves to the writer. +First, the toxin may be, as Langer believes in the case of the bee, an +alkaloidal base, toxic as such, and neutralized after injection by +antibodies produced for the occasion by the body. In such a case the +view that a partial local fixation of the toxin occurs, which prevents +its immediate diffusion, is acceptable. Through chemotactic action, +special cells capable of breaking up the toxin into harmless elements +are attracted to the scene. Their function may be, on the other hand, to +neutralize directly, not by lysis. This would explain the role of the +eosinophiles in the black-fly lesion. If their activities be essential +to the destruction or neutralization of the toxin, one would expect them +to be most numerous where there was least reaction. This would be at the +site of a bite in an immune individual. A point of special interest for +further investigation, would be the study of such a lesion. + +Second, it is conceivable that the injected saliva of the fly does not +contain an agent toxic as such. It is possible, that like many foreign +proteins, it only becomes toxic when broken down. The completeness and +rapidity of the breaking down depends on the number of eosinophiles +present. In such a case immunity should again be marked by intense +eosinophilia. + +[Illustration: 173. Fifth day mature lesion. Lower power drawing showing +papillary oedema and infiltrate in the region of the puncture. After Dr. +J. H. Stokes] + +Third, lytic agents in the blood serum may play the chief role in the +liberation of the toxic agent from its non-toxic combination. An immune +individual would then be one whose immunity was not the positive one of +antibody formation, but the negative immunity of failure to metabolize. +An immune lesion in such a case might be conceived as presenting no +eosinophilia, since no toxin is liberated. If the liberation of the +toxin is dependent upon lytic agents present in the serum rather than in +any cellular elements, a rational explanation would be available for the +apparent results (subject to confirmation) of the experiment with +sensitive and immune sera. In this experiment it will be recalled that +the sensitive serum seemed to bring out the toxicity of the ground +flies, and the serum itself seemed even to contain some of the dissolved +or liberated toxin. The slowness with which a lesion develops in the +case of the black-fly bite supports the view of the initial lack of +toxicity of the injected material. The entire absence of early +subjective symptoms, such as pain, burning, etc., is further evidence +for this view. It would appear as if no reaction occurred until lysis of +an originally non-toxic substance had begun. Regarding the toxin itself +as the chemotactic agent which attracts eosinophiles, its liberation in +the lytic process and diffusion through the blood stream attracts the +cells in question to the point at which it is being liberated. Arriving +upon the scene, these cells assist in its neutralization. + +The last view presented is the one to which the author inclines as the +one which most adequately explains the phenomena. + +A fourth view is that the initial injection of a foreign protein by the +fly (i.e., with the first bite) sensitizes the body to that protein. Its +subsequent injection at any point in the skin gives rise to a local +expression of systematic sensitization. Such local sensitization +reactions have been described by Arthus and Breton, by Hamburger and +Pollack and by Cowie. The description of such a lesion given by the +first named authors, in the rabbit, however, does not suggest, +histopathologically at least, a strong resemblance to that of the +black-fly. Such an explanation of many insect urticariae deserves further +investigation, however, and may align them under cutaneous expressions +of anaphylaxis to a foreign protein injected by the insect. Depending on +the chemical nature of the protein injected, a specific chemotactic +reaction like eosinophilia may or may not occur. Viewed in this light +the development of immunity to insect bites assumes a place in the +larger problem of anaphylaxis. + +[Illustration: 174. Experimental lesion produced from alcohol-fixed +flies, dried and ground into a paste with glycerin. After Dr. J. H. +Stokes] + + +SUMMARY + +In order to bring the results of the foregoing studies together, the +author appends the following resume of the clinical data presented in +the first paper. + +The black-fly, _Simulium venustum_, inflicts a painless bite, with +ecchymosis and haemorrhage at the site of puncture. A papulo-vesicular +lesion upon an urticarial base slowly develops, the full course of the +lesion occupying several days to several weeks. Marked differences in +individual reaction occur, but the typical course involves four stages. +These are, in chronological order, the papular stage, the vesicular or +pseudovesicular, the mature vesico-papular or weeping papular stage and +the stage of involution terminating in a scar. The papule develops in +from 3 to 24 hours. The early pseudovesicle develops in 24 to 48 hours. +The mature vesico-papular lesion develops by the third to fifth day and +may last from a few days to three weeks. Involution is marked by +cessation of oozing, subsidence of the papule and scar-like changes at +the site of the lesion. The symptoms accompanying this cycle consist of +severe localized or diffused pruritus, with some heat and burning in the +earlier stages if the oedema is marked. The pruritus appears with the +pseudovesicular stage and exhibits extraordinary persistence and a +marked tendency to periodic spontaneous exacerbation. The flies tend to +group their bites and confluence of the developing lesions in such cases +may result in extensive oedema with the formation of oozing and +crusted plaques. A special tendency on the part of the flies to attack +the skin about the cheeks, eyes and the neck along the hair line and +behind the ears, is noted. In these sites inflammation and oedema may +be extreme. + +A distinctive satellite adenopathy of the cervical glands develops in +the majority of susceptible persons within 48 hours after being bitten +in the typical sites. This adenopathy is marked, discrete and painful, +the glands often exquisitely tender on pressure. It subsides without +suppuration. + +Immunity may be developed to all except the earliest manifestations, by +repeated exposures. Such an immunity in natives of an infested locality +is usually highly developed. There are also apparently seasonal +variations in the virulence of the fly and variations in the reaction of +the same individual to different bites. + +Constitutional effects were not observed but have been reported. + + + + +BIBLIOGRAPHY + + +ALDRICH, J. M. 1905. A catalogue of North American Diptera. Washington, + D. C. 1-680. + +ALESSANDRI, G. 1910. Studii ed esperienze sulle larve della Piophila + casei. Arch. Parasit. xiii, p. 337-387. + +ANDERSON, J. F. and FROST, W. H. 1912. Transmission of poliomyelitis by + means of the stable-fly (Stomoxys calcitrans). Public Health Reports. + Washington. xxvii, p. 1733-1735. + +---- 1913. Further attempts to transmit the disease through the agency + of the stable-fly (Stomoxys calcitrans). Public Health Repts., + Washington. xxviii, p. 833-837. + +ANDERSON, J. F. and GOLDBERGER, J. 1910. 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Science, n. s. xxxviii, p. 513. + +HONEIJ, J. A. and PARKER, R. R. 1914. Leprosy: flies in relation to the + transmission of the disease. Journ. Med. Research, Boston, xxx, p. + 127-130. + +HOOKER, W. A. 1908 a. Life history, habits, and methods of study of the + Ixodoidea. Jour. Econ. Ent. i, p. 34-51. + +---- 1908 b. A review of the present knowledge of the role of ticks in + the transmission of disease. ibid., i, p. 65-76. + +HOPE, F. W. 1837. On insects and their larvae occasionally found in the + human body. Trans. Ent. Soc., London, ii, p. 256-271. + +HOUGH, G. DE N. 1899 a. Synopsis of the Calliphorinae of the United + States. Zoological Bulletin, ii, p. 283-290. + +---- 1899 b. Some Muscinae of North America, Biological Bulletin i, p. + 19-33. + +---- 1899 c. Some North American Genera of Calliphorinae. Entom. News, + x, p. 62-66. + +HOVARTH, G. 1912. Revision of the American Cimicidae. Ann. Mus. Nat. + Hungarici, x, p. 257-262. + +HOWARD, C. W. 1908. 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A general + consideration of mosquitoes, their habits, and their relations to the + human species. 4^o. Carnegie Institution of Washington (vii + 520 + p.). + +HOWARD, L. O. and MARLATT, C. L. 1902. The principal household insects + of the United States. U. S. Dept. Agric., Bur. Ent. Bul. 4. + +HUEBNER, W. 1907. Ueber das Pfeilgift der Kalahari. Arch. exper. Path. + und Pharm., lvii, p. 358-366. + +HUNTER, S. J. 1913. Pellagra and the sand-fly. Jour. econ. Ent. vi, p. + 96-99. + +HUNTER, W. D. 1913. American interest in medical entomology. Jour. econ. + Ent. vi, p. 27-39. + +HUNTER, W. D. and BISHOPP, F. C. 1910. Some of the more important ticks + of the United States. U. S. Dept. Agric. Yearbook 1910, p. 219-230, + pls. xv-xvi. + +---- 1911. The Rocky Mountain spotted fever tick. With special reference + to the problem of its control in the Bitter Root Valley in Montana. + U. S. Dept. Agric., Bur. Ent. Bul. 105, p. 1-47. + +HUTCHISON, R. H. 1914. 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Observations on the mechanism of infection in + tick fever and on the hereditary transmission of _Spirochaeta duttoni_ + in the tick. Trans. Soc. Trop. Med. Hyg., iii, p. 77-95. Abstr. in + Bul. Inst. Pasteur, viii, p. 312-313. + +---- 1910 b. On the hereditary transmission and mechanism of infection + in tick fever and on the hereditary transmission of _Spirochaeta + duttoni_ in the tick. Lancet., clxxvii, p. 11. + +LINNELL, R. McC. 1914. Notes on a case of death following the sting of a + scorpion. Lancet, 1914, p. 1608-1609. + +LIVINGSTONE, D. 1857. Missionary travels and researches in South Africa. + +LUCAS, H. 1843. (note) _Latrodectus malmignatus_ Bul. Soc. Ent., France, + 1843, p. viii. + +LUDLOW, C. S. 1914. Disease bearing mosquitoes of North and Central + America, the West Indies and the Philippine Islands. War Dept., + Office of Surgeon General. Bul. No. 4, 1-96. + +LUGGER, 1896. Insects injurious in 1896. Agr. Exp. Sta. Bul. 48. p. 33 + to 270. + +MacCALLUM, W. C. 1898. 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The mechanical transmission of surra. ibid., viii, sec. B., + p. 223-229. + +---- 1914 a. Experimental insect transmission of anthrax. U. S. Public + Health Repts. xxix, p. 75-77. + +---- 1914 b. I. Collected studies on the insect transmission of + _Trypanosoma evansi_. II. Summary of experiments in the transmission + of anthrax by biting flies. U. S. Pub. Health Service, Hyg. Lab. + Bul., 94, p. 1-48. + +MIYAKE, H. and SCRIBA, J. 1893. Vorlaufige Mitteilung uber einen neuen + Parasit des Menschen. Berl. klin. Wochenschr., xxx, p. 374. + +MOLLERS, B. 1907. Experimentelle Studien uber die Uebertragung des + Ruckfallfiebers durch Zecken. Zeitschr. fur Hyg. u. + Infektionskrankheiten, lviii, p. 277-286. + +MOTE, D. C. 1914. The cheese-skipper (_Piophila casei_). Ohio Naturalist + xiv, p. 309-310. + +NEIVA, A. 1910. Beitrage zur Biologie der _Conorhinus megistus_ Burm. + Memorias de Institute Oswaldo Cruz., ii, p. 206-212. + +NEVEU-LEMAIRE, M. 1907. 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Egyptian Gov't + School of Medicine, Cairo, ii, p. 7-44. + + + + +INDEX + + +Abscess, 178 + +Acanthia, 87 + +Acariasis, 58 + +Acarina, 23, 58, 131, 259 + +Acarus dysenteriae, 132 + +Accidental parasites, 131, 132, 134 + +Aedes, 194, 293 + +Aedes calopus, 182, 201, 205, 206, 208 + +Aedes cantator, 101 + +Aedes sollicitans, 101 + +Aedes taeniorhynchus, 101 + +Aerobic bacteria, 152 + +Aestivo-autumnal, 186 + +African Relapsing Fever, 230 + +Akis spinosa, 177 + +Alternation of Generations, 175 + +Amblyomma, 264 + +Amblyomma americanum, 67 + +Amblyomma cajennense, 67 + +American dog tick, 228 + +Amoeboid organism, 189 + +Anisolabis annulipes, 177 + +Anterior poliomyelitis, 241 + +Anopheles, 194, 291 + +Anopheles crucians, 199 + +Anopheles maculipennis, 182 + +Anopheles punctipennis, 198 + +Anopheles quadrimaculatus, 197 + +Anopheline, 192 + +Anthocoris, 279 + +Anthomyiidae, 300 + +Anthomyia, 138 + +Anthrax, 165 + +Antipruritic treatment, 72 + +Ants, 42 + +Aphiochaeta, 295 + +Apis mellifica, 36 + +Arachnida, 258 + +Araneida, 6 + +Argas, 64 + +Argas persicus, 63, 235, 237 + +Argasidae, 62 + +Argopsylla, 317 + +Argus, 259 + +Arilus, 284 + +Arthropods, poisonous, 6 + +Asopia farinalis, 177 + +Assassin-bugs, 31, 219 + +Auchmeromyia, 117 + +Automeris io, 47 + +Avicularoidea, 12 + + +Babesia, 226 + +Babesia bovis, 223 + +Babesia ovis, 225 + +Babesiosis, 221-222 + +Bacilli, 170 + +Bacillus icteroides, 202, 205 + +Bacillus pestis, 166 + +Bacillus typhosus, 153 + +Back swimmers, 30 + +Bdellolarynx, 304 + +Beauperthuy, Louis Daniel, 2 + +Bed-bug, 86, 88, 90, 173, 219-220 + +Bed-bug, cone-nosed, 92 + +Blister beetles, 54 + +Belostoma, 28, 277 + +Belostoma americana, 31 + +Belostomatidae, 30 + +Bengalia, 314 + +Bird-spiders, 10 + +Black death, 1, 166 + +Black flies, 33, 104, 247 + +Black heads, 80 + +Blaps mortisaga, 134 + +Blepharoceridae, 286 + +Boophilus, 264 + +Boophilus annulatus, 67, 223-225 + +Bot-flies, 112 + +Blue bottle flies, 140 + +Brill's disease, 238 + +Brown-tailed moth, 48 + +Bruck, 34 + +Buthus quinquestriatus, 21 + + +Cabbage butterfly, 56 + +Calliphora, 136, 140, 312 + +Calliphora erythrocephala, 141 + +Calobata, 296 + +Camponotinae, 43 + +Cancer, 254 + +Cantharidin, 54 + +Cantharidin poison, 55 + +Canthariasis, 134 + +Capsidae, 280 + +Carriers, simple, 4, 144 + +Carriers of disease, 144 + +Carrion's fever, 253 + +Caterpillar rash, 45 + +Cat flea, 172 + +Cattle ticks, 222 + +Causative organism, 170 + +Cellia, 291 + +Centipedes, 25, 257 + +Ceratophyllus, 120, 316 + +Ceratophyllus acutus, 123 + +Ceratophyllus fasciatus, 122, 172, 213 + +Ceratopogon, 108 + +Cheese-fly, 137 + +Cheyletus eruditus, 271 + +Chigger, 60, 70 + +Chigoes, 126 + +Chilopoda, 25, 257 + +Chiracanthium nutrix, 18 + +Chironomidae, 107 + +Chorioptes, 270 + +Chrysomelid, 55 + +Chrysomyia, 136, 308 + +Chrysomyia macellaria, 117, 140 + +Chrysops, 294 + +Chylous dropsy, 179 + +Chyluria, 178 + +Cicadidae, 55 + +Cimex L., 278 + +Cimex boueti, 92 + +Cimex columbarius, 92 + +Cimex hemipterus, 91, 220 + +Cimex hirundinis, 92 + +Cimex inodorus, 92 + +Cimex lectularius, 87, 219 + +Citheronia regalis, 44 + +Clinocoris, 87 + +Coleoptera, 134, 274 + +Comedons, 80 + +Complete metamorphosis, 80 + +Compressor muscle, 20 + +Compsomyia, 117 + +Cone-nosed bed-bug, 92 + +Conjunctivitis, nodular, 52 + +Conorhinus, 282 + +Conorhinus megistus, 93, 219-220 + +Conorhinus rubrofasciatus, 220 + +Conorhinus sanguisugus, 32, 92 + +Copra itch, 72 + +Cordylobia, 118 + +Coriscus, 280 + +Coriscus subcoleoptratus, 32 + +Creeping myasis, 112 + +Crustacea, 257 + +Cryptocystis, 176 + +Cryptotoxic, 54-55 + +Cteniza sauvagei, 13 + +Ctenocephalus, 120, 172, 213, 317 + +Culex, 194, 201, 293 + +Culex pipiens, 35, 98 + +Culex quinquefasciatus, 180 + +Culex sollicitans, 200 + +Culex territans, 101 + +Culicidae, 33, 97 + +Culicin, 34 + +Culicoides, 109, 288 + +Cyclops, 183, 257 + +Cynomyia, 136, 311 + + +Dance, St. Vitus, 8 + +Dancing mania, 8 + +Deer-flies, 110 + +Definitive host, 192 + +Demodecidae, 78 + +Demodex, 259 + +Demodex folliculorum, 78 + +Dermacentor, 262 + +Dermacentor andersoni, 67, 228 + +Dermacentor occidentalis, 227 + +Dermacentor variabilis, 67 + +Dermacentor venustus, 24, 228 + +Dermanyssidae, 68 + +Dermanyssus, 266 + +Dermanyssus gallinae, 68 + +Dermatitis, 72, 77, 85 + +Dermatobia, 115, 298 + +Dermatobia cyaniventris, 163 + +Dermatophilus, 317 + +Dermatophilus penetrans, 60, 126 + +Diamphidia simplex, 55 + +Dimorphism, 65 + +Direct inoculators, 4 + +Diplopoda, 25, 257 + +Diptera, 33, 94, 274 + +Dipterous Larvae, 135 + +Dipylidium, 175, 221 + +Dipylidium caninum, 4, 175-176 + +Dog flea, 172 + +Dracunculus, 257 + +Dracunculus medinensis, 182 + +Drosophila, 296 + +Dum-dum fever, 220 + +Dysentery, 154 + + +Ear-flies, 110 + +Earwig, 177 + +Echidnophaga, 317 + +Echinorhynchus, 185 + +Elephantiasis, 178-179 + +Empoasca mali, 33 + +Empretia, 46 + +English Plague Commission, 171 + +Epeira diadema, 18 + +Epizootic, 170 + +Eristalis, 137, 295 + +Essential hosts, 4, 165 + +Eumusca, 307 + +European Relapsing Fever, 233 + +Euproctis chrysorrhoea, 48 + +Eusimulium, 286 + + +Facultative parasites, 131 + +Fannia, 136, 138, 145, 300 + +Federal Health Service, 169 + +Fever, lenticular, 237 + African Relapsing, 230, 234 + Carrion's, 253 + dum-dum, 154 + European Relapsing, 233 + pappatici, 96 + red water, 220 + Rocky Mt. Spotted, 226 + three day, 96 + Typhus, 237 + +Filaria, 178, 221 + immitis, 182 + +Filariasis, 178 + +Flannel-moth larvae, 44 + +Fleas, 119, 166, 213 + cat, 172 + dog, 172 + human, 172, 176 + rodent, 123, 172 + rat, 171 + +Flesope, 125 + +Formaldehyde, 91 + +Fomites, 199, 204 + +Fulgoridae, 28 + +Fumigation, 320 + + +Gamasid, 68 + +Gangrene, 129 + +Gastrophilus, 113, 297 + +Giant crab spiders, 13 + +Giant water bugs, 30 + +Gigantorhynchus, 185 + +Glossina, 117, 297, 303 + +Glossina morsitans, 214, 217 + palpalis, 215, 217, 218 + +Glyciphagus, 267 + +Grain moth, 69 + +Grocer's itch, 72 + +Guinea-worm, 182 + + +Habronema muscae, 156, 183 + +Haematobia, 166, 304 + irritans, 146 + +Haematobosca, 304 + +Haematomyidium, 288 + +Haematopinus spinulosus, 213 + +Haematopota, 294 + +Haematosiphon, 279 + +Haemoglobinuria, 222 + +Haemozoin, 189 + +Harpactor, 284 + +Harvest mites, 60 + effect of, 59 + +Head-louse, 173 + +Helminthiasis, 138 + +Helophilus, 295 + +Hemiptera, 27, 86, 273-275 + +Heteropodidae, 13 + +Heuchis sanguinea, 55 + +Hexapod larvae, 58 + +Hexapoda, 27, 80, 258 + +Hippelates, 297 + +Hippobosca, 285 + +Histiogaster, 269 + spermaticus, 132 + +Homalomyia, 136, 138, 300 + +Honey bee, 36 + poison of, 37 + +Hornets, 43 + +Horn-fly, 137, 304, 308 + +Horse-fly, 110, 165 + +House-fly, 137-139, 144, 183 + control of, 156, 160 + +Human flea, 124 + +Host, definitive, 175 + intermediate, 175 + primary, 175 + +Hyalomma, 264 + aegypticum, 224-225 + +Hydrocyanic Acid Gas, 318 + +Hydrotaea, 300 + +Hymenolepis diminuta, 176 + +Hymenoptera, 36, 275 + +Hypoderma, 113, 298 + diana, 113 + lineata, 113 + +Hypopharynx, 80 + + +Immunity from stings, 39 + +Incomplete metamorphosis, 80 + +Infantile paralysis, 162, 241 + splenic, 220 + Direct inoculation, 164 + +Insects, 258 + blood-sucking, 170 + +Intermediate host, 192, 203 + +Intestinal infestation, 112, 133 + myasis, 137 + +Isosoma, 69 + +Itch, 73-74 + mite, 73 + Norwegian, 77 + +Ixodes, 260 + ricinus, 66, 225 + scapularis, 66 + +Ixodidae, 64-65 + +Ixodoidea, 62 + + +Janthinosoma lutzi, 116 + +Jigger, 60 + +Johannseniella, 110, 288 + +Journal of Tropical Medicine and Hygiene, 36 + +Julus terrestris, 25 + +June bug, 185 + + +Kala-azar, 220 + +Karakurte, 14 + +Katipo, 14 + +King, A. F. A., 3 + +Kircher, Athanasius, 1, 8 + +Kissing-bug, 31 + + +Labium, 29, 80 + +Labrum, 28, 80 + +Lachnosterna, 185 + +Laelaps, 266 + +Lagoa crispata, 45 + +Lamblia intestinalis, 154 + +Langer, Josef, 37 + +Larder beetles, 135 + +Latrodectus, 12, 14, 17 + mactans, 15 + +Leishmanioses, 220 + +Lenticular fever, 237 + +Lepidoptera, 274 + +Lepidopterous larvae, 134 + +Leprosy, 252 + +Leptidae, 112 + +Leptis, 295 + +Leptus, 60, 273 + +Lice, 80 + +Linguatulina, 258 + +Liponyssus, 265 + +Loemopsylla, 172, 317 + +Lone star tick, 228 + +Louse, body, 84 + crab, 85 + dog, 176 + head, 82 + pubic, 85 + +Lucilia, 136, 312 + +Lycosa tarantula, 10 + +Lycosidae, 10 + +Lyctocoris, 279 + +Lygus pratensis, 33 + +Lymphangitis, 67 + +Lymph scrotum, 178 + +Lyperosia, 304 + +Lyperosiops, 305 + + +Macloskie, 34 + +Maggots, rat-tail, 137 + +Magnes sive de Arte Magnetica, 8 + +Malaria, 186 + +Malmigniatte, 14 + +Mandibles, 28, 80 + +Mange, 73-75 + +Margaropus, 237, 264 + annulatus, 223 + +Masked bed-bug hunter, 32 + +Mastigoproctus giganteus, 19, 80 + +Maxillae, 28 + +Meal infesting species, 135 + +Melanin granules, 189 + +Melanolestes, 280 + picipes, 32 + +Mena-vodi, 14 + +Mercurialis, 1 + +Merozoites, 190 + +Metamorphosis, 80 + +Miana bug, 63 + +Microgametoblast, 192 + +Midges, 107 + +Migratory ookinete, 192 + +Millipedes, 25, 257 + +Mites, 23, 58 + +Monieziella, 269 + +Mosquitoes, 33, 97, 178, 196, 250 + treatment for bites of, 34, 36, 102 + +Musca, 137, 307 + domestica, 139, 145, 146, 157, 162 + +Muscidae, 117 + +Muscina, 137, 146, 307 + stabulans, 140 + +Mutualism, 57 + +Myasis, 112, 135 + intestinal, 135-140 + nasal, 141 + +Mycterotypus, 287 + +Myiospila, 146, 307 + +Myriapoda, 225, 132, 257 + + +Nagana, 165, 214 + +Nasal infestation, 114, 133 + +Necrobia, 135 + +Nematode parasite, 182 + +Nepa, 28 + +Nephrophages sanguinarius, 132 + +Nettling insects, 43 + larvae, poison of, 53 + +Neurasthenia, 89 + +Nits, 86 + +North African Relapsing Fever, 234 + +Norwegian itch, 77 + +No-see-ums, 109 + +Notoedres, 269 + cati, 78 + +Notonecta, 28, 277 + +Notonectidae, 30 + +Nott, Dr. Josiah, 2 + +Nuttall, 34 + + +Occipital headaches, 138 + +Oecacta, 288 + +Oeciacus, 279 + +Oesophageal diverticula, 35 + +Oestridae, 112, 136 + +Oestris ovis, 113 + +Oestrus, 298 + oocyst, 192 + ookinete, 192 + +Opsicoetes personatus, 32 + +Opthalmia, 155 + nodosa, 52 + +Oriental sore, 221 + +Ornithodoros, 65, 260 + moubata, 220, 230 + +Orthotylus flavosparsus, 33 + +Ornithomyia, 286 + +Oroya, 253 + +Oscinus, 297 + +Otiobius, 259 + megnini, 65 + +Otodectes, 271 + + +Pangonia, 294 + +Pappatici fever, 96 + +Parasimulium, 286 + +Parasite, 3, 57, 131, 134, 182 + accidental, 3, 131, 134 + facultative, 3, 57, 131 + nematode, 182 + stationary, 57 + temporary, 57 + true, 3 + +Parasitism, accidental, 134 + +Pathogenic bacteria, 152 + organisms, 144, 164 + +Pawlowsky, 81 + +Pediculoides, 267 + ventricosus, 69, 72 + +Pediculosis, 81 + +Pediculus, 275 + corporis, 84, 233, 238 + humanus, 82, 173 + +Pellagra, 162, 246 + +Pernicious fever, 186 + +Pest, 166 + +Phidippus audax, 19 + +Philaematomyia, 306 + +Phisalix, 13, 43 + +Phlebotomus, 289 + papatasii, 94 + verrucarum, 254 + vexator, 95 + +Phora, 295 + +Phormia, 136 + +Phormictopus carcerides, 13 + +Phthirus pubis, 85, 275 + +Phortica, 296 + +Pieris brassicae, 56 + +Piophila, 297 + +Piophila casei, 136, 137 + +Piroplasmosis, 222 + +Plague, 166 + bubonic, 166, 169, 170 + pneumonic, 167 + +Plasmodium, 186 + +Platymetopius acutus, 33 + +Plica palonica, 83 + +Pneumonic, 166 + plague, 167, 173 + +Poisoning by nettling larvae, 53 + +Poison of spiders, 7 + +Pollenia, 308 + rudis, 146, 147 + +Primary gland, 28 + +Prionurus citrinus, 20 + +Prosimulium, 286 + +Protocalliphora, 136, 312 + +Protozoan blood parasite, 165 + +Pseudo-tubercular, 52 + +Psorophora, 293 + +Psoroptes, 270 + +Psychodidae, 94 + +Pulex, 120, 124, 126, 172, 317 + cheopis, 172 + irritans, 124 + penetrans, 126 + serraticeps, 120 + +Pulvillus, 150 + +Punkies, 109 + +Pycnosoma, 308 + + +Rasahus, 280 + thoracicus, 32 + +Rat fleas, 120, 124, 171 + +Rat louse, 213 + +Red bugs, 70-72 + +Reduviidae, 31 + +Reduviolus, 280 + +Reduvius, 282 + personatus, 32 + +Redwater fever, 222 + +Relapsing fever, 230, 233 + +Rhinoestrus nasalis, 115 + +Rhipicentor, 264 + +Rhipicephalus, 264 + +Rhizoglyphus, 269 + +Rhodnius, 281 + +Rocky Mountain Spotted Fever, 226 + spotted fever tick, 67 + +Russian gad-fly, 115 + + +St. Vitus's or St. John's dance, 8 + +Salivary syringe, 28 + +Sand-flies, 109, 250 + +Sanguinetti, 11 + +Sarcophaga, 136, 142, 143 + +Sarcophila, 302 + +Sarcopsylla, 317 + penetrans, 126 + +Sarcoptes, 270 + minor, 78 + scabiei, 73 + +Sarcoptidae, 72 + +Scabies, 72, 73, 74, 75 + +Scaurus striatus, 177 + +Schaudinn, 34 + +Schizont, 189, 190 + +Scholeciasis, 134 + +Scolopendra morsitans, 26 + +Scorpions, 20 + poison of, 21 + +Screw worm fly, 140 + +Sepsidae, 296 + +Sepsis, 136, 297 + +Shipley, 34 + +Sibine, 46 + +Silvius, 294 + +Simple carriers, 4, 144 + +Simuliidae, 33, 104 + +Simulium, 247, 249, 286, 321 + pictipes, 104 + +Siphonaptera, 119, 274, 316 + +Siphunculata, 80, 275 + +Sitotroga cerealella, 69 + +Skippers, 137 + +Sleeping sickness, 166, 215 + +Snipe-flies, 112 + +Solpugida, 22 + +Spanish fly, 54 + +Spermatozoa, 192 + +Spinose ear-tick, 65 + +Spirochaeta, 35 + berberi, 234 + duttoni, 234 + +Spirochaetosis, 235 + +Sporozoite, 189 + +Spotted fever, 67, 226 + +Squirrel flea, 123 + +Stable-fly, 137, 160, 163, 165 + +Stegomyia, 182, 293 + calopus, 206 + fasciata, 206 + +Stomoxys, 137, 305 + calcitrans, 117, 146, 160, 161, 165, 242 + +Straw-worm, 69 + +Stygeromyia, 305 + +Sucking stomach, 35 + +Sulphur ointment, 77 + +Surra, 165 + +Symbiosis, 57 + +Symphoromyia, 112, 295 + + +Tabanidae, 110 + +Tabanus, 110, 166, 294 + striatus, 165 + +Taenia, 175 + +Tapeworm, 4, 176 + +Tarantella, 8 + +Tarantism, 8 + +Tarantula, 10 + +Tarsonemidae, 69 + +Tarsonemus, 267 + +Tenebrionid beetles, 127 + +Tersesthes, 110, 288 + +Tetanus, 129 + +Tetranychus, 273 + +Texas fever, 220-223 + +Three-day fever, 96 + +Tick, 23, 226 + bites, Treatment of, 68 + fever, 230 + paralysis, 67 + +Treatment, + Bee stings, 36, 41 + Bites of, + Bed-bugs, 90, 93 + Blackflies, 107 + Buffalo flies, 107 + Bugs, 31, 33 + Centipedes, 26, 27 + Chiggers, 127 + Chigoes, 127 + Fleas, 127 + Harvest mites, 61 + Jiggers, 129 + Lice, 83, 85 + Mosquitoes, 34, 36, 102 + Phlebotomus flies, 97 + Sand flies, 96, 107, 109 + Scorpions, 22, 23 + Spiders, 19 + Ticks, 61, 68, 72 + Ticks, ear, 65 + Blister beetle poison, 55 + Brown-tail moth rash, 45 + Cantharidin poison, 55 + Caterpillar rash, 45 + Ear ticks, 65 + House fly control, 156, 160 + Itch, 77 + Itch, grocer's, 72 + Lice, 85 + Nasal myasis, 143 + Rocky Mt. spotted fever, 228, 229 + Rash, caterpillar, 45 + Scabies, 77 + Sleeping sickness control, 218 + Spotted fever, 228, 229 + Stings, bee, 36, 41 + Typhus fever, prophylaxis, 239 + +Trichodectes canis, 176 + +Trichoma, 82 + +Trineura, 295 + +Trochosa singoriensis, 11 + +Trombidium, 60, 273 + +True insects, 80 + +Trypanosoma, 35 + +Trypanosoma, brucei, 165 + +Trypanosoma cruzi, 219 + +Trypanosoma lewisi, 213 + +Trypanosomiases, 212 + +Trypanosomiasis, 165, 219 + +Tsetse flies, 117, 166, 214, 219 + +Tsetse flies disease, 165 + +Tuberculosis, 155 + +Tumbu-fly, 118 + +Tydeus, 271 + +Typhoid, 155 + +Typhoid fever, 154 + +Typhus, 237 + +Typhus fever, 237 + +Tyroglyphus, 72, 131, 268 + +Dr. Tyzzer, 49 + + +Uranotaenia, 292 + + +Vancoho, 14 + +Varicose groin glands, 178 + +Verruga peruviana, 253 + +Vescicating insects, 54 + + +Wanzenspritze, 29 + +Warble-flies, 112 + +Wasps, 43 + +Whip-scorpions, 19 + +Wohlfahrtia, 143, 302 + +Wolf-spiders, 10 + +Wyeomyia smithii, 101, 293 + + +Xenopsylla, 172, 317 + +Xenopsylla cheopis, 171, 124 + +Xestopsylla, 317 + + +Yaws, 2 + +Yellow fever, 196, 203, 205 + + + + +TRANSCRIBER'S NOTES + +The following discrepancies in the text are as in the original: + + The inconsistent hyphenation of the following: + assassin-bugs/assassin bugs; + bedbug/bed-bug (and bedbugs); + beekeeper/bee-keeper (and beekeepers); + blackflies/black-flies; + blow-flies/blow flies; + bluebottles/blue-bottles; + bot-flies/bot flies; + bristlelike/bristle-like; + browntail/brown-tail; + coextensive/co-extensive; + deer-flies/deer flies; + dorsocentral/dorso-central; + ectoparasites/ecto-parasites; + endoparasites/endo-parasites; + flesh-fly/flesh fly (and flesh flies); + hotbed/hot-bed; + housefly/house-fly (and houseflies); + horsefly/horse-fly (and horse flies); + horse-manure/horse manure; + midsummer/mid-summer; + preeminently/pre-eminently; + sandfly/sand-fly (and sandflies); + screw-worm fly/screw worm fly; + stable-fly/stable fly; + subequal/sub-equal; + subfamily/sub-family; + subtropical/sub-tropical; + tapeworm/tape-worm (and tapeworms); + today/to-day; + tsetse-flies/tsetse flies; + widespread/wide-spread; + wormlike/worm-like. + + Inconsistent spelling of the following: + defence/defense; + disc/disk; + hemolysis/haemolysis; + hemolytic/haemolytic; + hexapod/hexopod; + Levaditi/Lavaditi; + metalescent/metallescent; + Mitzmain/Mitzman; + Neveau-Lemaire/Neveau-Lamaire; + offence/offense; + Phthirus/Phthirius + Portschinsky/Portchinsky/Portchinski; + travelled/traveled; + ventra-/ventro-; + Villot/Villet; + Wohlfartia/Wohlfahrtia; + +toxine, insiduous, efficaceous, cyanid are spelt as in the original + +In the first paragraph of chapter XII, "the student might not be lead" +is as in the original. Lead perhaps should be led + +In the bibliography, the entry for Vaughan without a text specified is +as in the original + + + + + +End of the Project Gutenberg EBook of Handbook of Medical Entomology, by +William Albert Riley and Oskar Augustus Johanssen + +*** END OF THIS PROJECT GUTENBERG EBOOK HANDBOOK OF MEDICAL ENTOMOLOGY *** + +***** This file should be named 34279.txt or 34279.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/4/2/7/34279/ + +Produced by Bryan Ness, Iris Schimandle, Brownfox and the +Online Distributed Proofreading Team at http://www.pgdp.net +(This file was produced from images generously made +available by The Internet Archive/American Libraries.) + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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