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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/31293-8.txt b/31293-8.txt new file mode 100644 index 0000000..e7965d7 --- /dev/null +++ b/31293-8.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: ISO-8859-1 + +*** 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°N, 58°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, Papéis 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. × 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_. × 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, proötics, and + exoccipitals form a compact central zone. An elongate + frontoparietal fontanelle is present. + +[Illustration: FIG. 3. Dorsal view of skull of _Allophryne_ +(AMNH 70110). × 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 proötic. 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 propósito de los Leptodactylidae (Amphibia Anura). + Papéis 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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Lynch And Howard L. Freeman. + </title> + <style type="text/css"> + +body { + margin-left: 10%; + margin-right: 10%; +} + + h1,h2,h3,h4,h5,h6 { + text-align: center; /* all headings centered */ + clear: both; +} + +p { + margin-top: .75em; + text-align: justify; + margin-bottom: .75em; +} + + p.title { text-align: center; text-indent: 0; + font-weight: bold; + line-height: 1.4; margin-bottom: 3em; } + +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: 92%; + font-size: smaller; + text-align: right; +} /* page numbers */ + +.blockquot { + margin-left: 5%; + margin-right: 10%; +} + +.i4 {display: block; margin-left: 2.5em; + padding-left: 2.5em; text-indent: -2.5em;} + +.center {text-align: center;} + +.smcap {font-variant: small-caps;} + +.u {text-decoration: underline;} + +.caption {font-weight: bold;} + +/* Images */ +.figcenter { + margin: auto; + 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; +} + + </style> + </head> +<body> + + +<pre> + +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: ISO-8859-1 + +*** 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 + + + + + + +</pre> + + + + + +<p class="title"><span class="smcap">University of Kansas Publications</span><br /> +<span class="smcap">Museum of Natural History</span><br /><br /> + +Volume 17, No. 10, pp. 493-502, 3 Figs.<br /> +October 27, 1966</p> +<hr style="width: 25%;" /> + + +<h1>Systematic Status of a South American Frog,<br /> +Allophryne ruthveni Gaige<br /><br /></h1> + + +<p class="title">BY<br /><br /> + +<big>JOHN D. LYNCH AND HOWARD L. FREEMAN</big><br /><br /><br /> + + +<span class="smcap">University of Kansas</span><br /> +<span class="smcap">Lawrence</span><br /> +1966<br /> +</p> +<hr style="width: 65%;" /> + + + +<p class="title"> +<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br /> +<br /> +Editors: E. Raymond Hall, Chairman, Henry S. Fitch,<br /> +Frank B. Cross<br /> +<br /> +<br /> +Volume 17, No. 10, pp. 493-502, 3 Figs.<br /> +Published October 27, 1966<br /> +<br /> +<br /> +<span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas<br /> +<br /> +<br /> +<small>PRINTED BY<br /> +ROBERT R. (BOB) SANDERS, STATE PRINTER<br /> +TOPEKA, KANSAS<br /> +1966<br /> +<br /> +31-5378<br /> +</small></p> +<hr style="width: 65%;" /> + + +<p><span class="pagenum"><a name="Page_495" id="Page_495">[Pg 495]</a></span></p> +<h2> +Systematic Status of a South American Frog,<br /> +Allophryne ruthveni Gaige</h2> + +<p class="center">BY<br /><br /> + +JOHN D. LYNCH AND HOWARD L. FREEMAN</p> + + +<p>Gaige (1926) described <i>Allophryne ruthveni</i> as a new genus and +species of diminutive bufonid from British Guiana. Noble (1931) +considered <i>A. ruthveni</i> to be a toothless relative of <i>Centrolenella</i> +and placed the genus in the Hylidae. Gallardo (1965) suggested +that <i>Allophryne</i> 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.</p> + +<p>A male of <i>Allophryne ruthveni</i> 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.</p> + +<div class="blockquot"><p><i>Acknowledgments.</i>—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 <i>Allophryne</i>.</p></div> + + +<h3>METHODS AND MATERIALS</h3> + +<div class="blockquot"><p>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.</p> + +<p><i>Specimens examined.</i>—Six, as follows: BRITISH GUIANA, <i>Dist. Demarara</i>: +Marudi Creek, AMNH 44749; <i>Dist. Equibo</i>: Tumatumari, MCZ 11790 (paratype); +<i>Dist. Rupununi</i> (<i>Berbice</i>): Wai Wai Country, N of Acarahy Mountains, +west of New River (2°N, 58°W), KU 69890. Also, 3 specimens from "probably +British Guiana," AMNH 70108-10 (70110 cleared and stained).</p></div> + + +<h3>SYSTEMATIC ACCOUNT</h3> + +<p>The availability of additional material and the new information +pertaining to osteology permit an amplification of Gaige's (1926) +description.</p> +<p><span class="pagenum"><a name="Page_496" id="Page_496">[Pg 496]</a></span></p> + +<h4><big><span style="font-weight: normal">Genus</span> Allophryne <span style="font-weight: normal">Gaige</span></big></h4> + +<div class="blockquot"><p><i>Allophryne</i> 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, Papéis Avulsos, +17:79, Jan. 1, 1965.</p> + +<p><i>Type species.</i>—<i>Allophryne ruthveni</i> Gaige.</p> + +<p><i>Diagnosis and definition.</i>—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.</p></div> + +<h4><big>Allophryne ruthveni <span style="font-weight: normal">Gaige</span></big></h4> + +<div class="blockquot"><p><i>Allophryne ruthveni</i> 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.</p> + +<p><i>Holotype.</i>—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.</p> + +<p><i>Diagnosis.</i>—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.</p> + +<p><i>External Morphology.</i>—(<a href="#fig1">Fig. 1</a>) <i>Additional features not mentioned in diagnoses</i>: +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.</p> + +<p>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 +(<a href="#fig1">Fig. 1d</a>).</p> + +<p>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).</p></div> + +<p><span class="pagenum"><a name="Page_497" id="Page_497">[Pg 497]</a></span></p> + +<p class="figcenter" style="width: 415px;"> +<a name="fig1" id="fig1"></a><img src="images/image001.jpg" width="415" height="600" alt="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. × 3.5." title="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. × 3.5." /> +<span class="caption">Fig. 1. Allophryne ruthveni, male (KU 69890); (a) Dorsum.<br /> (b) Thenar +view of right hand. (c) Lateral profile of head.<br /> (d) Plantar view of right +foot. × 3.5.</span> +</p> + +<p><span class="pagenum"><a name="Page_498" id="Page_498">[Pg 498]</a></span></p> +<div class="blockquot"><p><i>Color.</i>—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 (<a href="#fig1">Fig. 1</a>a and c).</p> + +<p>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.</p> + +<p>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.</p> + +<p><i>Variation.</i>—Eight measurements were taken on each specimen and four +ratios were computed; these are summarized in <a href="#table1">Table 1</a>. 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.</p></div> + +<p class="center"><a name="table1" id="table1"></a><b>TABLE I.—Variation in Measurements and Proportions of Allophryne +ruthveni. (Ranges in parentheses below means.)</b></p> + +<div class="center"> +<table border="1" cellpadding="8" cellspacing="0" summary="table1"> +<tr><td align="center"><b>Character</b></td><td align="center"><b>Male (1)</b></td><td align="center"><b>Females (5)</b></td></tr> +<tr><td align="left">Snout-vent (in mm.)<br /> </td><td align="center">20.6<br /> </td><td align="center">23.6<br />(18.4-31.0)<a name="FNanchor_A_1" id="FNanchor_A_1"></a><a href="#Footnote_A_1" class="fnanchor">[A]</a></td></tr> +<tr><td align="left">Tibia/snout-vent<br /> </td><td align="center">0.43<br /> </td><td align="center">0.43<br />(0.41-0.47)</td></tr> +<tr><td align="left">Tympanum/head width<br /> </td><td align="center">0.12<br /> </td><td align="center">0.15<br />(0.14-0.16)</td></tr> +<tr><td align="left">Eyelid/interorbital space<br /> </td><td align="center">0.55<br /> </td><td align="center">0.53<br />(0.49-0.56)</td></tr> +<tr><td align="left">Tympanum/eye length<br /> </td><td align="center">0.40<br /> </td><td align="center">0.46<br />(0.42-0.50)</td></tr> +</table></div> + + +<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>Holotype is reported to be 31 mm. snout-vent length +(Gaige, 1926). The largest measured by us was 26.2 mm. snout-vent.</p></div> + +<div class="blockquot"><p>The dorsal spinules are most pronounced and extensive on the male (<a href="#fig1">Fig. +1</a>) and less so in all other specimens examined. The illustration of the holotype +suggests that it has equally prominent, but fewer, spinules (Gaige, 1926).</p> + +<p>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.</p> + +<p><i>Distribution.</i>—All known specimens have been found in the foothills of the +northeastern face of the Guiana Massif in British Guiana.</p></div> + +<p><span class="pagenum"><a name="Page_499" id="Page_499">[Pg 499]</a></span></p> + +<h3>FAMILY POSITION</h3> + +<p>The following characters of <i>Allophryne</i> are those generally held to be useful +in determining family relationships:</p> + + +<ol><li>Presacral vertebrae procoelus, eight in number.</li> + +<li>Parahyoid absent.</li> + +<li>Free ribs lacking.</li> + +<li>Bidder's organ absent.</li> + +<li>Intercalary cartilages present in digits; phalangeal formulae 3-3-4-4 and 3-3-4-5-4.</li> + +<li>Coccyx articulating with sacrum by two condyles.</li> + +<li>Tarsal bones not fused.</li> + +<li>Pectoral girdle arciferal.</li> + +<li>Epicoracoidal horns present, free.</li> + +<li>Terminal phalanges T-shaped.</li> + +<li>Sacrum procoelus and diapophyses expanded.</li> + +<li>Maxillae, premaxillae, and prevomers edentate.</li> + +<li>Cranial roofing bones well ossified.</li> +</ol> + +<p>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). <i>Allophryne</i> possesses well-developed, +free epicoracoidal horns, such as those found in the Hylidae, Centrolenidae, +Leptodactylidae and Bufonidae.</p> + +<p>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 <i>Allophryne</i> in having T-shaped terminal phalanges.</p> + +<p class="figcenter" style="width: 600px;"> +<a name="fig2" id="fig2"></a><img src="images/image002.png" width="600" height="326" alt="Fig. 2. Dorsal (a) and lateral (b) +views of distal phalanges of third +finger of Allophryne. × 40." title="Fig. 2. Dorsal (a) and lateral (b) +views of distal phalanges of third +finger of Allophryne. × 40." /> +<span class="caption">Fig. 2. Dorsal (a) and lateral (b) +views of distal phalanges of third +finger of Allophryne. × 40.</span> +</p> + +<p>Only the presence of intercalary cartilages (<a href="#fig2">Fig. 2</a>) suggests relationship of +<i>Allophryne</i> 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<span class="pagenum"><a name="Page_500" id="Page_500">[Pg 500]</a></span> +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 <i>Allophryne</i> and genera of the Centrolenidae.</p> + + +<h3>CRANIAL OSTEOLOGY</h3> + +<div class="blockquot"><p>The skull of <i>Allophryne</i> (<a href="#fig3">Fig. 3</a>) 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 <i>Corythomantis</i>, +<i>Diaglena</i>, <i>Osteocephalus</i>, <i>Triprion</i>) and generalized skulls. <i>Allophryne</i> +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, proötics, and exoccipitals +form a compact central zone. An elongate frontoparietal fontanelle is +present.</p> + +<p class="figcenter" style="width: 600px;"> +<a name="fig3" id="fig3"></a><img src="images/image003.png" width="600" height="567" alt="Fig. 3. Dorsal view of skull of Allophryne (AMNH 70110). × 12." title="Fig. 3. Dorsal view of skull of Allophryne (AMNH 70110). × 12." /> +<span class="caption">Fig. 3. Dorsal view of skull of Allophryne (AMNH 70110). × 12.</span> +</p> + +<p>Dorsally (<a href="#fig3">Fig. 3</a>), 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.<span class="pagenum"><a name="Page_501" id="Page_501">[Pg 501]</a></span> +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 <i>pars facialis</i> +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 (<a href="#fig3">Fig. 3</a>), but sectioning is necessary to determine +the true nature and extent of this element.</p> + +<p>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 proötic. The anterior (zygomatic) +process of the squamosal is greatly reduced (only about one-third the length +of the posterior process).</p></div> + + +<h3>DISCUSSION</h3> + +<p>The skull of <i>Allophryne</i> is definitely non-hylid. Most of the post-cranial +features do not help to clarify relationships. <i>Allophryne</i> +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, <i>Allophryne</i> 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.</p> + +<p>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 <i>Allophryne</i>, especially the structural relationships +of the sphenethmoid-prevomer area may elucidate the relationships +of <i>Allophryne</i>.</p> + +<p>The relationships of <i>Allophryne</i> 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 <i>Allophryne</i> will become apparent.</p> +<p><span class="pagenum"><a name="Page_502" id="Page_502">[Pg 502]</a></span></p> +<p>A more comprehensive study of the cranial anatomy of certain +hylids, leptodactylids, dendrobatids, and atelopodids along with +that of <i>Allophryne</i> is needed to clarify the relationships of <i>Allophryne</i>, +and might indicate that the recognition of a fifth family is +necessary.</p> + + +<h3>CONCLUSION</h3> + +<p>Among currently recognized families of frogs, <i>Allophryne</i> 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 <i>Allophryne</i> +should be in another family is less convincing than evidence suggesting +it should be in the Hylidae. We tentatively place <i>Allophryne</i> +in the Hylidae.</p> + + +<h3>LITERATURE CITED</h3> + +<p><span class="smcap">Davis, D. D.</span> and <span class="smcap">Gore, U. R.</span></p> + +<p class="i4">1936. Clearing and staining skeletons of small vertebrates. Fieldiana: +Technique, 4:1-16.</p> + +<p><span class="smcap">Duellman, W. E.</span></p> + +<p class="i4">1956. The frogs of the hylid genus <i>Phrynohyas</i> Fitzinger, 1843. Misc. +Publs. Mus. Zool., Univ. Michigan, 96:1-47, February 21.</p> + +<p><span class="smcap">Gaige, H. T.</span></p> + +<p class="i4">1926. A new frog from British Guiana. Occas. Papers Mus. Zool., Univ. +Michigan, 176:1-3, October 14.</p> + +<p><span class="smcap">Gallardo, J. M.</span></p> + +<p class="i4">1965. A propósito de los Leptodactylidae (Amphibia Anura). Papéis +Avulsos, 17:77-87, January 1.</p> + +<p><span class="smcap">Griffiths, I.</span></p> + +<p class="i4">1959. The phylogeny of <i>Sminthillus limbatus</i> and the status of the Brachycephalidae +(Amphibia: Salientia). Proc. Zool. Soc. London, 132:457-87, +May.</p> + +<p><span class="smcap">Noble, G. K.</span></p> + +<p class="i4">1931. The biology of the amphibia. McGraw-Hill, New York, vii + 577 pp.</p> + + +<p><i>Transmitted August 2, 1966.</i></p> + + +<p class="center"><small>31-5378</small></p> + + + +<hr style="width: 65%;" /> + +<h3>Transcriber's Notes</h3> + +<p><a href="#table1">Table 1</a> and Figs. <a href="#fig2">2</a> and <a href="#fig3">3</a> have been moved slightly to avoid breaking +up the paragraphs of text.</p> + + + + + + + + + +<pre> + + + + + +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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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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