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diff --git a/31293.txt b/31293.txt new file mode 100644 index 0000000..fbb141f --- /dev/null +++ b/31293.txt @@ -0,0 +1,877 @@ +The Project Gutenberg EBook of Systematic Status of a South American Frog, +Allophryne ruthveni Gaige, by John D. Lynch and Howard L. Freeman + +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: Systematic Status of a South American Frog, Allophryne ruthveni Gaige + +Author: John D. Lynch + Howard L. Freeman + +Release Date: February 16, 2010 [EBook #31293] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK SOUTH AMERICAN FROG *** + + + + +Produced by Chris Curnow, Joseph Cooper, Diane Monico, and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + + + + + + +UNIVERSITY OF KANSAS PUBLICATIONS +MUSEUM OF NATURAL HISTORY + +Volume 17, No. 10, pp. 493-502, 3 Figs. +October 27, 1966 + + +Systematic Status of a South American Frog, +Allophryne ruthveni Gaige + + +BY + +JOHN D. LYNCH AND HOWARD L. FREEMAN + + +UNIVERSITY OF KANSAS +LAWRENCE +1966 + + + + +UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + +Editors: E. Raymond Hall, Chairman, Henry S. Fitch, +Frank B. Cross + + +Volume 17, No. 10, pp. 493-502, 3 Figs. +Published October 27, 1966 + + +UNIVERSITY OF KANSAS +Lawrence, Kansas + + +PRINTED BY +ROBERT R. (BOB) SANDERS, STATE PRINTER +TOPEKA, KANSAS +1966 + +31-5378 + + + + +Systematic Status of a South American Frog, +Allophryne ruthveni Gaige + +BY + +JOHN D. LYNCH AND HOWARD L. FREEMAN + + +Gaige (1926) described _Allophryne ruthveni_ as a new genus and species +of diminutive bufonid from British Guiana. Noble (1931) considered _A. +ruthveni_ to be a toothless relative of _Centrolenella_ and placed the +genus in the Hylidae. Gallardo (1965) suggested that _Allophryne_ is a +leptodactylid of uncertain affinities. Other references to the +monotypic genus have consisted only of a listing of the name or of its +inclusion in a key. To date the holotype and one paratype (both +females) have been reported (Gaige, 1926), and the family position of +the genus remains unsettled. + +A male of _Allophryne ruthveni_ is among the amphibians and reptiles +collected in southern British Guiana by William A. Bentley in January, +1962, and deposited in the Museum of Natural History at The University +of Kansas (KU). Four additional specimens (females) are in the American +Museum of Natural History; only one of the latter has definite locality +data. + + _Acknowledgments._--We are grateful to Dr. Ernest E. + Williams, Museum of Comparative Zoology (MCZ) and Dr. + Richard G. Zweifel, American Museum of Natural History + (AMNH) for the loan of specimens. We are further indebted to + Dr. Zweifel for permission to clear and stain one specimen. + Dr. William E. Duellman and Linda Trueb offered many + constructive criticisms. Miss Trueb executed the drawings of + the skull and finger bones. Mr. Martin Wiley provided x-ray + photographs of _Allophryne_. + + +METHODS AND MATERIALS + + Six of the seven known specimens were available for study. + Measurements were taken in the manner described by Duellman + (1956). One specimen was cleared and stained, using the + technique of Davis and Gore (1936), in order to study the + skeleton. X-ray photographs were made of another specimen + for comparison. + + _Specimens examined._--Six, as follows: BRITISH GUIANA, + _Dist. Demarara_: Marudi Creek, AMNH 44749; _Dist. Equibo_: + Tumatumari, MCZ 11790 (paratype); _Dist. Rupununi_ + (_Berbice_): Wai Wai Country, N of Acarahy Mountains, west + of New River (2 deg.N, 58 deg.W), KU 69890. Also, 3 specimens from + "probably British Guiana," AMNH 70108-10 (70110 cleared and + stained). + + +SYSTEMATIC ACCOUNT + +The availability of additional material and the new information +pertaining to osteology permit an amplification of Gaige's (1926) +description. + +Genus ~Allophryne~ Gaige + + _Allophryne_ Gaige, Occas. Papers Mus. Zool., Univ. + Michigan, 176:1, Oct. 14, 1926. Crawford, Annals Carnegie + Mus., 21(1):29, 32, Nov. 14, 1931. Noble, The biology of the + amphibia. McGraw-Hill, p. 510, 1931. Ruthven, Herpetologica, + 1:3, July 11, 1936. Gallardo, Papeis Avulsos, 17:79, Jan. 1, + 1965. + + _Type species._--_Allophryne ruthveni_ Gaige. + + _Diagnosis and definition._--A genus of diminutive frogs; + vomers, maxillae, and premaxillae edentate; skin of head + strongly anchored to connective tissue on cranium; + prepollical spine absent in males; disk of third finger + larger than tympanum, smaller than eye; no humeral hook in + either sex; ilia extending anteriorly beyond sacral + expansions; adults attaining snout-vent length of 31 mm.; + male having darkened external subgular vocal sac; skin of + dorsum pustulate. + +~Allophryne ruthveni~ Gaige + + _Allophryne ruthveni_ Gaige, Occas. Papers Mus. Zool., Univ. + Michigan, 176:1-3, pl. I, Oct. 14, 1926. Crawford, Annals + Carnegie Mus., 21(1):32, Nov. 14, 1931. Ruthven, + Herpetologica, 1:3, July 11, 1936. Barbour and Loveridge, + Bull. Mus. Comp. Zool., 96(2):64, Feb., 1946. Peters, Occas. + Papers Mus. Zool., Univ. Michigan, 539:10, Sept. 19, 1952. + + _Holotype._--University of Michigan Museum of Zoology 63419, + adult female, from Tukeit Hill, below Kaiteur Falls, Equibo + District, British Guiana; obtained in May, 1924, by E. N. + Clarke. + + _Diagnosis._--Fingers free; toes two-thirds webbed; no + supernumerary tubercles on soles or palms; no tarsal fold; + elongate anal sheath, anal opening on lower surface of + thighs; head broad, interorbital space 2.5 times width of + upper eyelid; snout subacuminate in dorsal profile, strongly + sloping in lateral profile; tympanum visible in males, + concealed in females; venter areolate. + + _External Morphology._--(Fig. 1) _Additional features not + mentioned in diagnoses_: Head wider than long, about as wide + as body; supratympanic fold present; canthus rostralis + rounded, loreal region slightly concave, nearly vertical; + nostril at tip of snout; pupil horizontal; no teeth on + maxillary, premaxillary, or vomer; tongue small, round, + thick, not notched behind, free posteriorly for one-sixth of + length; choanae large, only partly visible from directly + below; males having darkened subgular vocal sac; vocal slits + present in male. + + Axillary membrane lacking or but slightly developed; no + tubercles or ridge under forearm; two palmar tubercles; + subarticular tubercles small, simple, round, flattened; tips + of fingers slightly expanded, T-shaped, with prominent + transverse groove; first finger shorter than second (stated + as longer than second in diagnosis by Gaige, 1926:2); folds + extending laterally from anus for a short distance, then + downward to venter of thighs; no appendage on heel, no inner + or outer tarsal folds or tubercles; inner metatarsal + tubercle oval, about twice as long as wide; outer metatarsal + tubercle nearly absent; no supernumerary tubercle on sole; + subarticular tubercles on foot small, round, simple, and + diffuse; toes T-shaped, slightly wider than digit; toes + about two-thirds webbed (Fig. 1d). + + Skin of venter coarsely areolate; skin of flanks, throat, + chest, undersurfaces of arms, tibia, tarsi, dorsal surfaces + of thighs, tarsi, hands, and feet smooth; skin of dorsal + surfaces of tibia, forearm, back, and top and sides of head + having large horny pustules (sharply spinous in male). + +[Illustration: FIG. 1. _Allophryne ruthveni_, male (KU 69890); +(_a_) Dorsum. (_b_) Thenar view of right hand. (_c_) Lateral profile of +head. (_d_) Plantar view of right foot. x 3.5.] + + _Color._--Dorsum gray with irregular network of black lines + and elongate blotches; flanks and labial region black with + large white ocelli; dorsal surfaces of limbs gray, marked as + follows: two large, elongate white spots on each thigh, + concealed white spot on base of upper arm, black-edged gray + transverse bars on forearms and shanks, white spot on each + knee and elbow; ventral surfaces pale gray; black-edged + white spot on ventral surface of thigh on each side of anal + opening; chin and throat dark gray with white spots; vocal + sac in male black (Fig. 1a and c). + + Gaige (1926) briefly described the color, which conforms to + the above in all particulars. The paratype (MCZ 11790) has + lost the gray color after 40 years in preservation; now + (1966) the ground-color is cream-brown, and the dorsal + spotting, noted by Gaige as being black, is now brown. + + The spots on the feet, tarsi, knees, thighs, flanks and + upper arm are white in preservative, but in life possibly + were red or yellow. These colors usually fade to white in + preservative. Red or yellow spots are common aposematic + colors in frogs. + + _Variation._--Eight measurements were taken on each specimen + and four ratios were computed; these are summarized in Table + 1. Gaige's illustration of the holotype shows that it has a + greatly reduced pattern, whereas the paratype and three of + the other five known specimens have relatively large and + numerous spots. The male (KU 69890) and one female (AMNH + 70108) have a reduced pattern intermediate between that of + the holotype and the four other specimens. + +TABLE I.--Variation in Measurements and Proportions of Allophryne +ruthveni. (Ranges in parentheses below means.) + +--------------------------+----------+----------------- + Character | Male (1) | Females (5) +--------------------------+----------+----------------- +Snout-vent (in mm.) | 20.6 | 23.6 + | | (18.4-31.0)[A] + | | +Tibia/snout-vent | 0.43 | 0.43 + | | (0.41-0.47) + | | +Tympanum/head width | 0.12 | 0.15 + | | (0.14-0.16) + | | +Eyelid/interorbital space | 0.55 | 0.53 + | | (0.49-0.56) + | | +Tympanum/eye length | 0.40 | 0.46 + | | (0.42-0.50) +--------------------------+----------+----------------- + +[Footnote A: Holotype is reported to be 31 mm. snout-vent length +(Gaige, 1926). The largest measured by us was 26.2 mm. snout-vent.] + + The dorsal spinules are most pronounced and extensive on the + male (Fig. 1) and less so in all other specimens examined. + The illustration of the holotype suggests that it has + equally prominent, but fewer, spinules (Gaige, 1926). + + The holotype, a gravid female, is the largest known specimen + (31 mm., snout-vent length). Another gravid female (AMNH + 70108) has a snout-vent length of 26.2 mm. + + _Distribution._--All known specimens have been found in the + foothills of the northeastern face of the Guiana Massif in + British Guiana. + + +FAMILY POSITION + + The following characters of _Allophryne_ are those generally + held to be useful in determining family relationships: + + 1. Presacral vertebrae procoelus, eight in number. + + 2. Parahyoid absent. + + 3. Free ribs lacking. + + 4. Bidder's organ absent. + + 5. Intercalary cartilages present in digits; phalangeal + formulae 3-3-4-4 and 3-3-4-5-4. + + 6. Coccyx articulating with sacrum by two condyles. + + 7. Tarsal bones not fused. + + 8. Pectoral girdle arciferal. + + 9. Epicoracoidal horns present, free. + + 10. Terminal phalanges T-shaped. + + 11. Sacrum procoelus and diapophyses expanded. + + 12. Maxillae, premaxillae, and prevomers edentate. + + 13. Cranial roofing bones well ossified. + +Griffiths (1959) accorded considerable taxonomic weight to the presence +or absence of epicoracoidal horns in showing relationships among the +genera placed in the Brachycephalidae [= Atelopodidae; Dendrobatidae; +and Leptodactylidae (in part)] by Noble (1931). _Allophryne_ possesses +well-developed, free epicoracoidal horns, such as those found in the +Hylidae, Centrolenidae, Leptodactylidae and Bufonidae. + +The presence of intercalary elements in the digits is characteristic of +the Centrolenidae, Hylidae, Phrynomeridae, Pseudidae, and the +rhacophorine ranids (including the Hyperoliidae). This element is bony +in the pseudids and cartilaginous in the other families. Phrynomerids +and rhacophorine ranids lack epicoracoidal horns and have firmisternal +pectoral girdles. Centrolenids are small, delicate, arboreal frogs +having poorly ossified skulls and fused tarsal bones, but agree with +_Allophryne_ in having T-shaped terminal phalanges. + +[Illustration: FIG. 2. Dorsal (_a_) and lateral (_b_) views of +distal phalanges of third finger of _Allophryne_. x 40.] + +Only the presence of intercalary cartilages (Fig. 2) suggests +relationship of _Allophryne_ to the Hylidae. The T-shaped terminal +phalanges suggest affinities with centrolenids, elutherodactyline +leptodactylids, or certain "brachycephalid" frogs. Griffiths (1959) +clearly showed that Noble's Brachycephalidae was a polyphyletic +assemblage. No hylid genus is edentate, and none has either T-shaped +terminal phalanges or the unusual dorsal spinules. Perhaps the presence +of intercalary cartilages is not indicative of relationship but instead +is a parallelism (or convergence) in _Allophryne_ and genera of the +Centrolenidae. + + +CRANIAL OSTEOLOGY + + The skull of _Allophryne_ (Fig. 3) is distinctive among + anurans; it does not closely resemble the skulls of either + hylids or centrolenids, both of which have generally more + delicate (except for casque-headed hylids, such as + _Corythomantis_, _Diaglena_, _Osteocephalus_, _Triprion_) + and generalized skulls. _Allophryne_ on the other hand has a + strongly ossified central region (cranial roofing bones and + sphenethmoid complex) and a weak peripheral zone. The + peripheral elements are reduced (maxilla, pterygoid, and + squamosal) or absent (quadratojugal), whereas the + frontoparietals, nasals, sphenethmoid, prooetics, and + exoccipitals form a compact central zone. An elongate + frontoparietal fontanelle is present. + +[Illustration: FIG. 3. Dorsal view of skull of _Allophryne_ +(AMNH 70110). x 12.] + + Dorsally (Fig. 3), the premaxillae are not visible. The + proportionally gigantic septomaxillae are visible anterior + to the nasals. The moderate-sized nasals are separated + medially and in broad contact with the ethmoid posteriorly. + The palatine process of the nasal does not meet the frontal + process of the maxilla. A large frontoparietal fontanelle is + evident between the frontoparietals. The tegmen tympani are + much reduced and maintain only cartilaginous contact with + the posterior arms of the squamosals. The foramen magnum, + occipital condyles, and exoccipitals show no unusual + features. The _pars facialis_ and frontal process of the + maxilla are greatly reduced. The maxilla and premaxilla are + articulated. The high, narrow alary processes of the + premaxillae extend dorsally about two-thirds of the height + of the snout. A cartilaginous internasal septum is + illustrated (Fig. 3), but sectioning is necessary to + determine the true nature and extent of this element. + + Ventrally, the skull lacks palatines. The maxillae, + premaxillae, and prevomers are edentate. The parasphenoid is + large with relatively short, stout alary (lateral) + processes. The sphenethmoid is extensive in ventral aspect + and forms the major supporting structure in the anterior + part of the skull. The pterygoid has a broad articulation + with the maxilla, a tenuous contact with the squamosal, but + is not attached to the prooetic. The anterior (zygomatic) + process of the squamosal is greatly reduced (only about + one-third the length of the posterior process). + + +DISCUSSION + +The skull of _Allophryne_ is definitely non-hylid. Most of the +post-cranial features do not help to clarify relationships. +_Allophryne_ shares several osteological features with the +Dendrobatidae: T-shaped terminal phalanges, general cranial morphology +and procoelus vertebrae. But, the dendrobatids possess firmisternal +pectoral girdles and lack epicoracoidal horns. Also, no dendrobatid has +intercalary elements in the digits. We are, therefore, left with a +taxonomic enigma. In one or more characters generally regarded as +important, _Allophryne_ differs from all presently defined families of +frogs. The Hylidae and Dendrobatidae are the only currently recognized +families in which the genus might be placed. + +The function and taxonomic importance of the large septomaxillae are +unknown and are probably associated with the modification of the +sphenethmoid-prevomer area. A more detailed study of the cranial +osteology of _Allophryne_, especially the structural relationships of +the sphenethmoid-prevomer area may elucidate the relationships of +_Allophryne_. + +The relationships of _Allophryne_ cannot be understood without a +re-analysis of some of the features used as major criteria in frog +classification (the nature of an intercalated cartilage; the nature of +the sternal complex; the relative value of cranial osteology; the +vertebral structure; and the thigh musculature). Some of these features +have been investigated by other workers, most notably Griffiths, but +others have not and need re-examination. A re-analysis of some of the +major criteria used in frog classification is in progress (Callison, +Lynch, and Trueb) and upon completion of that study we think the +relationships of _Allophryne_ will become apparent. + +A more comprehensive study of the cranial anatomy of certain hylids, +leptodactylids, dendrobatids, and atelopodids along with that of +_Allophryne_ is needed to clarify the relationships of _Allophryne_, +and might indicate that the recognition of a fifth family is necessary. + + +CONCLUSION + +Among currently recognized families of frogs, _Allophryne_ is least +different from the Hylidae although it is our opinion that inclusion of +this genus in the Hylidae probably represents an unnatural +classification. However, the present evidence suggesting that +_Allophryne_ should be in another family is less convincing than +evidence suggesting it should be in the Hylidae. We tentatively place +_Allophryne_ in the Hylidae. + + +LITERATURE CITED + +DAVIS, D. D. and GORE, U. R. + + 1936. Clearing and staining skeletons of small vertebrates. + Fieldiana: Technique, 4:1-16. + +DUELLMAN, W. E. + 1956. The frogs of the hylid genus _Phrynohyas_ Fitzinger, + 1843. Misc. Publs. Mus. Zool., Univ. Michigan, 96:1-47, + February 21. + +GAIGE, H. T. + 1926. A new frog from British Guiana. Occas. Papers Mus. + Zool., Univ. Michigan, 176:1-3, October 14. + +GALLARDO, J. M. + 1965. A proposito de los Leptodactylidae (Amphibia Anura). + Papeis Avulsos, 17:77-87, January 1. + +GRIFFITHS, I. + 1959. The phylogeny of _Sminthillus limbatus_ and the status + of the Brachycephalidae (Amphibia: Salientia). Proc. + Zool. Soc. London, 132:457-87, May. + +NOBLE, G. K. + 1931. The biology of the amphibia. McGraw-Hill, New York, + vii + 577 pp. + + +_Transmitted August 2, 1966._ + + +31-5378 + + + + * * * * * + +Transcriber's Notes + +Italicized text is shown within _underscores_. + +Bold text is shown within ~tildes~. + +Table 1 and Figs. 2 and 3 have been moved slightly to avoid breaking +up the paragraphs of text. + + + + + + + + +End of the Project Gutenberg EBook of Systematic Status of a South American +Frog, Allophryne ruthveni Gaige, by John D. Lynch and Howard L. 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