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diff --git a/32653.txt b/32653.txt new file mode 100644 index 0000000..50597c9 --- /dev/null +++ b/32653.txt @@ -0,0 +1,1527 @@ +The Project Gutenberg EBook of A Review of the Frogs of the Hyla bistincta +Group, by William E. Duellman + +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: A Review of the Frogs of the Hyla bistincta Group + +Author: William E. Duellman + +Release Date: June 2, 2010 [EBook #32653] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK FROGS--HYLA BISTINCTA GROUP *** + + + + +Produced by Chris Curnow, Joseph Cooper and the Online +Distributed Proofreading Team at https://www.pgdp.net + + + + + + + + + + UNIVERSITY OF KANSAS PUBLICATIONS + MUSEUM OF NATURAL HISTORY + + Volume 15, No. 9, pp. 469-491, 4 figs. + March 2, 1964 + + + A Review of the Frogs + Of the Hyla bistincta Group + + BY + + WILLIAM E. DUELLMAN + + + UNIVERSITY OF KANSAS + LAWRENCE + 1964 + + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors: E. Raymond Hall, Chairman, Henry S. Fitch, + Theodore H. Eaton, Jr. + + + Volume 15, No. 9, 469-491, 4 figs. + Published March 2, 1964 + + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + + PRINTED BY THE STATE PRINTER + TOPEKA, KANSAS + 1964 + + 29-8590 + + + + + A Review of the Frogs + Of the Hyla bistincta Group + + BY + + WILLIAM E. DUELLMAN + + + + +CONTENTS + + + PAGE + INTRODUCTION 471 + Acknowledgments 471 + THE HYLA BISTINCTA GROUP 472 + Analysis of Characters 473 + Key to the Species of the Hyla Bistincta Group 474 + ACCOUNTS OF THE SPECIES 475 + _Hyla bistincta_ Cope 475 + _Hyla charadricola_ new species 478 + _Hyla robertsorum_ Taylor 481 + _Hyla pachyderma_ Taylor 485 + _Hyla crassa_ (Brocchi) 486 + RELATIONSHIPS 489 + LITERATURE CITED 491 + + + + +INTRODUCTION + + +In the mountainous regions of Middle America there are several groups +of hylid frogs that inhabit mountain streams. Some of these groups, +such as _Plectrohyla_ and _Ptychohyla_, have been elevated to generic +rank, whereas others are retained in the large and complex genus +_Hyla_. In the mountains of Mexico five species of hylids that seem to +compose a phyletic unit are herein referred to as the _Hyla bistincta_ +group. Since 1955 I have been accumulating specimens of, and data on, +this group with the result that all specimens known to me, including +the types of all named taxa, have been studied. Detailed observations +have been made on the ecology and life histories of three of the +species; the other two species are known to me only from preserved +specimens. + + +Acknowledgments + +For permission to examine specimens in their care I am indebted to +Charles M. Bogert, American Museum of Natural History (AMNH); Doris +M. Cochran, United States National Museum (USNM); Jean Guibe, Museum +National d'Histoire Naturelle, Paris (MNHN); Robert F. Inger, Chicago +Natural History Museum (CNHM); Hobart M. Smith, University of Illinois +Museum of Natural History (UIMNH); Charles F. Walker, University of +Michigan Museum of Zoology (UMMZ). (Abbreviations of institutions +given above in parentheses are used throughout; the Museum of Natural +History, University of Kansas is abbreviated KU.) + +For their willing assistance in the field I am grateful to Ann S. +Duellman, Dale L. Hoyt, and John Wellman. Permits for collecting in +Mexico were generously issued by the late Ing. Luis Macias Arellano, +Departamento de la Fauna Silvestre, Direccion General de Caza. The +drawings in figures 1 and 3 were executed by Gail Selfridge. This +research has been supported by the National Science Foundation (NSF +G-9827). + + + + +THE HYLA BISTINCTA GROUP + + +The five species comprising the _Hyla bistincta_ group are +moderate-sized hylids having rather blunt heads and robust bodies. +The fingers are long and have little webbing (Fig. 1). The skin of +the dorsum is thick and glandular, but not tuberculate. An anal +sheath is present. The skull is rather broad, flat, and solidly +roofed. The ethmoid is broad, curved downward laterally, and solidly +sutured to the frontoparietal. The nasals are broad, sutured for their +entire width with the ethmoid, and broadly in contact medially. The +premaxillaries are in contact medially; each has a long, flat nasal +process. The quadratojugal is absent, and the maxillary tapers to a +point posteriorly. There is no squamosal-maxillary connection. +The maxillary and premaxillary teeth are rather long, bifid, and +moderately spatulate. Some teeth on the premaxillary and anterior +part of the maxillary are hooked. The vomerine teeth are spatulate +and bifid. A broad, flat, ossified prepollex is present but does +not project as a spine. The known tadpoles have ventral mouths, +2/3 tooth-rows, two or more rows of labial papillae, and long tails +with low fins. + + [Illustration: FIG. 1. Palmar view of right hand of: + _a._--_Hyla robertsorum_ (KU 57661), + _b._--_Hyla charadricola_ (KU 58414). x 3.] + +As thus defined the _Hyla bistincta_ group can be distinguished from +all other groups of Middle American frogs by the combination of +absence of the quadratojugal, non-projecting prepollex, long fingers +with little webbing, and stream-inhabiting tadpoles having 2/3 tooth +rows and two or more rows of labial papillae. + +Possibly _Hyla arborescandens_ and _Hyla hazelae_ belong in this +group. Because these species are somewhat different from the included +species and because their tadpoles are as yet unknown, I have +refrained from including these two species in the _Hyla bistincta_ +group. Taylor (1948:261) assigned _Hyla proboscidea_ (= _H. +dalquesti_) and (1949:272) _Hyla cyclomaculata_ to this group, but +because these two species have a quadratojugal and notably different +tadpoles, they are excluded from the group. + +Frogs of the genus _Plectrohyla_ closely resemble species in the _Hyla +bistincta_ group but differ principally in having a projecting +prepollex. In the highlands of Costa Rica a group of species, of which +_Hyla moesta_ is best known, resembles species in the _Hyla bistincta_ +group. At present insufficient information is available on the Costa +Rican species to determine their affinities. + + +Analysis of Characters + +The characters used in the systematic study of the frogs in this group +are those usually employed in anuran systematics. Of the various +measurements and proportions, the snout-vent length and the relative +size of the tympanum to the eye apparently are of more taxonomic +importance than the others (Table 1). In all of the species the +tympanum is at least partially covered by a heavy, dermal +supratympanic fold, and in some specimens of _H. pachyderma_ the +tympanum is completely obscured. In two species (_H. bistincta_ and +_H. charadricola_) the snout is square, whereas in the other species +it is round. + +The fingers are long and slender in _H. crassa_, _pachyderma_, and +_robertsorum_ and somewhat shorter with more webbing in _H. bistincta_ +and _charadricola_. Breeding males of _Hyla pachyderma_ have +moderately large nuptial spines; the other species have small spines, +except _H. charadricola_ in which spines apparently are absent. A +well-defined thoracic fold is present in _H. pachyderma_, and a weak +fold is present in _H. robertsorum_; the other species lack folds. In +all species there is an anal sheath; this sheath is longest in _H. +bistincta_, in which the anal opening is directed ventrally at the +level of the lower edge of the thighs. + + TABLE 1.--COMPARISON OF CERTAIN MEASUREMENTS AND PROPORTIONS + IN THE SPECIES OF THE HYLA BISTINCTA GROUP. (DATA FROM ADULT + MALES; MEANS ARE GIVEN IN PARENTHESES BELOW THE RANGES.) + + A: Snout-vent length + B: Tibia length / Snout-vent length + C: Head width / Snout-vent length + D: Tympanum / Eye + + ==================================================================== + SPECIES | N | [A] | [B] | [C] | [D] + ---------------+----+-----------+-----------+-----------+----------- + _H. bistincta_ | 38 | 43.0-53.8 | 0.47-0.52 | 0.32-0.37 | 0.35-0.48 + | | (46.3) | (0.49) | (0.34) | (0.42) + | | | | | + _H. | 10 | 35.3-44.4 | 0.50-0.54 | 0.31-0.33 | 0.30-0.37 + charadricola_ | | (40.4) | (0.52) | (0.32) | (0.34) + | | | | | + _H. | 26 | 39.9-47.9 | 0.48-0.51 | 0.30-0.36 | 0.36-0.47 + robertsorum_ | | (43.1) | (0.49) | (0.32) | (0.41) + | | | | | + _H. pachyderma_| 1 | 39.9 | 0.53 | 0.32 | ... + | | | | | + _H. crassa_ | 1 | 53.7 | 0.50 | 0.33 | 0.28 + ---------------+----+-----------+-----------+-----------+----------- + +Frogs in this group are rather drab in appearance. The dorsal color +varies from dull green to various shades of brown. The most distinct +aspect of the coloration is the different color patterns on the flanks +and posterior surfaces of the thighs. The flanks in all species are +marked with spots or reticulations. + +_Hyla bistincta_ differs from other members of the group in having +vocal slits and a distensible vocal sac. Only this species has been +reported to call (Shannon, 1951:473). Insofar as is known, the other +species are mute. Examination of skeletal preparations of _H. +bistincta_, _charadricola_, and _robertsorum_ and X-rays of the other +species shows no notable specific differences in the osteology. Since +the tadpoles of only _H. bistincta_ (Duellman, 1961:47) and _H. +robertsorum_ (Rabb and Mosimann, 1955) are known, larval characters +are of limited use in intra-group systematics. + + +Key to the Species of the _Hyla bistincta_ Group + + 1. Snout, in dorsal profile, short and bluntly rounded; + canthus rounded, sometimes indistinct, vocal slits absent 2 + + Snout, in dorsal profile, longer, squared; canthus distinct, + vocal slits present or absent 4 + + 2. Feet webbed to base of discs; dorsum dull olive green; + flanks having cream-colored spots _H. crassa_, p. 486 + + Feet not webbed to base of discs; dorsum brown 3 + + 3. Tarsal fold strong; thoracic fold heavy; webbing on feet + extending to middle of penultimate phalanx of fourth toe; + distinct white stripe above anus; cluster of largo spines on + thumb in breeding males, _H. pachyderma_, p. 485 + + Tarsal fold weak and short; thoracic fold absent or barely + indicated; webbing of feet extending to base of penultimate + phalanx of fourth toe; no distinct white stripe above anus; + nuptial tuberosities in breeding males consisting of small + spines _H. robertsorum_, p. 481 + + 4. Tarsal fold strong; anal flap elongate; anus opening at + lower edge of femur; dorsum tan or brown; flanks mottled with + cream and brown; venter immaculate creamy white; no anal + stripe; vocal slits present, _H. bistincta_, p. 475 + + Tarsal fold faint and short; anal flap not elongate; anus + opening at middle of femur; dorsum olive-green with black + reticulations; flanks greenish gray with brown or black + spots; a row of white flecks above and below anus, vocal + slits absent _H. charadricola_, p. 478 + + + + +ACCOUNTS OF THE SPECIES + + +In the following accounts complete synonymies are given for each +species. In each account one specimen is described in detail; after +this description the variation is discussed. In the list of specimens +examined, localities are arranged alphabetically within the states, +which also are given in alphabetical order. Localities given in +italics are not shown on the accompanying maps (Figs. 2 and 4) due +to crowding of symbols. + + +#Hyla bistincta# Cope + + _Hyla bistincta_ Cope, Proc. Amer. Philos. Soc, 17:87, + August, 1877 [Holotype.--USNM 32261 from "most probably + Veracruz," Mexico; Francis Sumichrast collector]. Brocchi, + Etude des batraciens de l'Amerique Centrale, p. 43, 1881. + Boulenger, Catalogue Batrachia Salientia, 2nd Ed., p. 401, + February 1, 1882. Sumichrast, La Naturaleza, 6:81, 1882. + Cope, Bull. U. S. Natl. Mus., 32:14, 1887; Bull. U. S. Natl. + Mus., 34:351, 1889. Guenther, Biologia Centrali-Americana, + Reptilia and Batrachia, pp. 265-6, June, 1901. Diaz de Leon, + Indice de los batracios que se encuentran en la Republica + Mexicana, p. 17, June, 1904. Nieden, Das Tierreich, Amphibia, + Anura I, p. 247, June, 1923. Kellogg, Bull. U. S. Natl. Mus., + 160:163-164, March 31, 1932. Taylor, Proc. Biol. Soc. + Washington, 50:50-53, April 21, 1937; Univ. Kansas Sci. + Bull., 26:389, November 27, 1940. Taylor and Smith, Proc. U. + S. Natl. Mus., 194:87, June 7, 1948. Taylor, Univ. Kansas + Publ. Mus. Nat. Hist., 1:261, August 16, 1948; Copeia, no. + 4:272-273, December 15, 1949. Smith and Taylor, Univ. Kansas + Sci. Bull., 33:346, March 20, 1950 [Type locality restricted + to Acultzingo, Veracruz, Mexico]. Rabb and Mosimann, Occas. + Papers Mus. Zool. Univ. Michigan, 563:6-9, March 29, 1955. + Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 15:47-49, + December 20, 1961. + + _Hyla bistincta labeculata_ Shannon, Proc. U. S. Natl. Mus., + 101:470-473, figs. 92a, 93d, May 17, 1951 [Holotype.--USNM + 123689 from San Lucas Camotlan, Oaxaca, Mexico; Walter S. + Miller collector]. Smith and Williams, Herpetologica, 19:23, + April 11, 1963. + + _Hyla bistincta bistincta_, Shannon, Proc. U. S. Natl. Mus., + 101:472, May 17, 1951. Shannon and Werler, Herpetologica, + 11:85, July 15, 1955. Smith and Williams, Herpetologica, + 19:23, April 11, 1963. + +_Diagnosis._--Maximum snout-vent length in males 54 mm.; snout in +dorsal profile truncate; tarsal fold strong; inner metatarsal tubercle +large, high, and elongate; outer metatarsal tubercle absent; webbing +on foot extending to middle of antepenultimate phalanx of fourth toe; +nuptial spines on thumb small; thoracic fold absent; anal opening at +level of lower edge of femur; dorsum brown or tan; belly cream-color; +flanks creamy yellow with brown reticulations or spots; anal stripe +absent; vocal slits present. + +_Description._--The following description is based on KU 68078 from +Uruapan, Michoacan, Mexico. Adult male having a snout-vent length of +43.2 mm.; tibia length, 22.1 mm., 51.1 per cent of snout-vent length; +foot length (measured from proximal edge of inner metatarsal tubercle +to tip of longest toe), 20.8 mm.; greatest width of head, 15.4 mm., +35.6 per cent of snout-vent length; head length, 13.9 mm., 32.1 per +cent of snout-vent length; diameter of eye, 5.1 mm.; diameter of +tympanum, 2.4 mm., 47.1 per cent of diameter of eye. Snout in lateral +profile bluntly rounded, in dorsal profile truncate; canthus +pronounced, rounded, not angular; loreal region slightly concave; lips +thick, round, not flaring; nostrils slightly protuberant; internarial +distance, 3.4 mm.; interorbital distance, 4.5 mm., somewhat broader +than width of eyelid, 3.5 mm. A heavy dermal fold from posterior +corner of eye above tympanum and curved downward to insertion of +forearm; tympanum round, its diameter slightly more than its distance +from eye. Forearm not robust; row of small pustules on ventral surface +of forearm; fold on wrist; prepollex moderately enlarged, covered with +small, horny, nuptial spines continuous on edge of digit; row of +spines on inner surface of second finger; subarticular tubercles +large, round, none bifid; supernumerary tubercles rather large, round; +palmar tubercles rather small, flat, elliptical; fingers long, +moderately slender; length of fingers from shortest to longest, +1-2-4-3; discs moderately large, that on third finger slightly larger +than tympanum; rudimentary web between first and second and between +second and third fingers; web between third and fourth fingers +extending about one-fourth length of fourth finger. Heels overlap by +about one-third length of shank when hind limbs adpressed; tibiotarsal +articulation extends to anterior edge of eye; tarsal fold strong, +extending to heel; inner metatarsal tubercle large, high, and +elongate; outer metatarsal tubercle absent; subarticular tubercles +moderately large, round; supernumerary tubercles small, in single rows +on proximal segments of digits; toes moderately short; length of toes +from shortest to longest, 1-2-3-5-4, third and fifth toes about equal +in length; toes about two-thirds webbed; web extending to middle of +antepenultimate phalanx of fourth toe, to discs of first, second, and +fifth toes, and to base of penultimate phalanx of third toe; discs +rather small, about two-thirds size of those on fingers. Anal opening +at level of lower edge of thighs; anal sheath elongate, deeply creased +medially; transverse dermal fold above anus. Skin of dorsal surfaces +of head, body, and limbs faintly areolate; skin of chin, belly, and +ventral surfaces of thighs granular, that of ventral surfaces of +limbs, except thighs, areolate; thoracic fold absent. Tongue nearly +round, slightly longer than wide, shallowly notched behind and barely +free posteriorly. Vomerine teeth 4-4, situated on rounded vomerine +ridges between rather small ovoid inner nares; vocal slits present, +situated along posterior edge of each ramus. + +Color (in alcohol) pale brown on dorsal surfaces of head, body, and +limbs; flanks and anterior surfaces of thighs creamy white with dark +brown reticulations; posterior surfaces of thighs tan with creamy +white spots; belly cream-color; anal stripe absent. + +Color (in life) pale tan on dorsal surfaces; flanks and anterior +surfaces of thighs pale creamy yellow with purplish brown +reticulations; posterior surfaces of thighs tan with yellow spots; +ventral surfaces yellow; iris pale copper-color. + +_Variation._--There is little variation in structure. The total number +of vomerine teeth varies from 6 to 14. In some individuals the +supratympanic fold covers the upper part of the tympanum, but at least +the lower part of the tympanum is always visible. The extent of the +webbing between the toes varies from three-fourths to two-thirds +complete. Usually the web extends to some point on the antepenultimate +phalanx of the fourth toe, but in some specimens the web extends to +the base of the penultimate phalanx. + +In the large series of specimens from Uruapan, Michoacan, the +coloration of the flanks and anterior surfaces of the thighs varies +from nearly uniform creamy yellow with only fine dark reticulations to +bold reticulations enclosing yellow spots. Some specimens from Oaxaca +and Veracruz have slightly different markings on the flanks; in these +the dark pigment is in the form of irregular spots or dashes, instead +of reticulations. + +There is considerable variation in color in the living frogs. The +dorsum varies from greenish tan and pale yellowish tan to reddish +brown, and some individuals are dark chocolate brown. + +_Remarks._--Shannon (1951:470) named _Hyla bistincta labeculata_ on +the basis of a single male from San Lucas Camotlan, Oaxaca; he +diagnosed the subspecies as differing from _Hyla bistincta bistincta_ +by having "the gray reticulation of the sides entirely broken up into +elongate black blotches; tarsal fold moderately elevated." The +condition of the tarsal fold is characteristic of the species. The +dispersion of dark pigment on the flanks is variable. The type of +_Hyla bistincta labeculata_ (USNM 123689) is extreme in the +development of dark dashes on the flanks, but this condition is +approached in several specimes from Oaxaca and Veracruz. For example, +in some specimens from Cumbres de Acultzingo, Veracruz, the mottling +on the flanks is bold; in others the flanks are reticulated. The +specimen from San Vicente, Oaxaca, has black dashes on the flanks +(Smith and Williams, 1963:23), whereas a specimen from Cerro San +Felipe, Oaxaca, has no pattern on the flanks. In general, specimens +from western Mexico have reticulate mottling on the flanks as compared +with the marbling on the flanks in specimens from eastern Mexico. On +the basis of available data, the recognition of subspecies in _Hyla +bistincta_ is unwarranted. + +The tadpoles of this species described by Duellman (1961:47) are like +those of _Hyla robertsorum_ in having 2/3 tooth-rows, peglike +serrations on the beaks, and long, rounded tails. At Uruapan tadpoles +were found in a rocky stream on April 24, 1956, and metamorphosing +young were found there on August 2, 1956. A completely metamorphosed +juvenile has a snout-vent length of 24.8 mm. + +_Hyla bistincta_ is found only along streams, where individuals can be +seen clinging to vines and other vegetation closely over-hanging +fast-moving parts of the stream. + +_Distribution._--_Hyla bistincta_ occurs at elevations from 1400 to +2600 meters in the mountains of the Sierra Madre Occidental in western +Jalisco southward through the Cordillera Volcanica in Michoacan, +Mexico, and Morelos, the Sierra de Coalcoman in Michoacan, and the +Sierra Madre del Sur in Guerrero and Oaxaca, and thence northward in +the Sierra Madre Oriental to central Veracruz (Fig. 2). + +Specimens examined.--GUERRERO: Omiltemi, UIMNH 38023-5. JALISCO: 25 +km. SE Autlan, UMMZ 102076. MEXICO: 19 km. W Villa Victoria, UIMNH +28162, USNM 114513. MORELOS: Cuernavaca, USNM 121523; _3 km. N +Cuernavaca_, UIMNH 28168-70. MICHOACAN: Cerro San Andres, UMMZ 102075; +Dos Aguas, UMMZ 119193; _12.5 km. ENE Dos Aguas_, UMMZ 119194; _Los +Conejos_, UMMZ 94238-40; Uruapan, KU 68077-8, 69093 (skeleton), UIMNH +20457, 28167, UMMZ 85452-3, 112838, 112839 (16), 115231 (tadpoles), +115232, 115233 (12), 121515, S-1699 (skeleton), S-1826 (skeleton), +USNM 114514-5, 114517-24. OAXACA: Cerro San Felipe, UIMNH 28163; Pluma +Hidalgo, AMNH 13447; San Lucas Camotlan, USNM 123689; San Vincente, +UIMNH 51346 (Smith and Williams, 1963:23). VERACRUZ: no specific +locality, USNM 32261; Cumbres de Acultzingo, CNHM 105482-3, UIMNH +28164-6, 49133-4, USNM 114525. + + [Illustration: FIG. 2. Map showing locality records for + _Hyla bistincta_ and _Hyla charadricola_.] + + +#Hyla charadricola# new species + +_Holotype._--University of Kansas Museum of Natural History No. 58414 +from the Rio Totolapa, 14.4 kilometers by road west of Huachinango, +Puebla, Mexico, 2280 meters; obtained by John Wellman on June 8, 1960. + +_Paratypes._--KU 58415-58423, same data as holotype, and UIMNH 50966, +obtained at the type locality by William E. Duellman on February 11, +1961. + +_Diagnosis._--Maximum snout-vent length in males, 45 mm.; snout in +dorsal profile truncate; tarsal fold short and weak; inner metatarsal +tubercle moderately large, elliptical, and flat; outer metatarsal +tubercle minute, round, and indistinct; webbing extending to base of +terminal phalanx of fourth toe; nuptial spines on thumb absent; +thoracic fold absent; anal opening at level of middle of thigh; dorsum +olive-green with black reticulations; venter cream color with brown +flecks on chin; flanks pale grayish green with brown spots; anal +stripe absent; row of white flecks above and a row below anus; vocal +slits absent. + +_Description of Holotype._--Adult male having a snout-vent lenth of +44.4 mm.; tibia length, 22.2 mm., 50 per cent of snout-vent length; +foot length (measured from proximal edge of inner metatarsal tubercle +to tip of longest toe), 20.4 mm.; greatest width of head, 14.0 mm., +31.5 per cent of snout-vent length; head length, 13.6 mm., 30.6 per +cent of snout-vent length; diameter of eye, 5.0 mm.; diameter of +tympanum, 1.6 mm., 32.0 per cent of diameter of eye. Snout in lateral +profile bluntly rounded, in dorsal profile truncate (Fig. 3); canthus +pronounced, rounded, not angular; loreal region slightly concave; lips +thick, rounded, not flaring; nostrils slightly protuberant; +internarial distance, 3.7 mm.; interorbital distance, 4.1 mm., +somewhat broader than width of eyelid, 3.3 mm. A heavy dermal fold +from posterior corner of eye above tympanum and then to insertion of +forearm; tympanum round, its diameter three-fourths its distance from +eye. Forearm rather slender; a short axillary membrane; no fold on +wrist; no nuptial excrescence or spines on enlarged prepollex; +subarticular tubercles moderately small, round; none bifid; few +supernumerary tubercles on proximal segments of digits; large, flat +palmar tubercle present; fingers long and slender; length of fingers +from shortest to longest, 1-2-4-3; discs moderately large, that on +third finger about equal to diameter of tympanum; rudimentary web +between first and second fingers; web between third and fourth fingers +extending to about one-fifth length of fourth finger (Fig. 1). Heels +overlap by about one-third length of tibia when hind limbs adpressed; +tibiotarsal articulation extends to middle of eye; tarsal fold weak, +extending from moderately large, flat, elliptical inner metatarsal +tubercle to middle of tarsus; outer metatarsal tubercle minute, round, +indistinct; subarticular tubercles round; single row of small, round +supernumerary tubercles on proximal segments of each digit; toes +moderately short and slender; length of toes from shortest to longest, +1-2-3-5-4, third and fifth about equal in length; toes about +three-fourths webbed; web extending to base of terminal phalanx of +fourth toe; discs small, about two-thirds size of those on fingers. +Anal opening directed posteroventrally at middle of thighs; two +transverse dermal folds above anus; short, thin anal sheath present; +many small tubercles lateral and ventral to anal opening. Skin of +dorsum, chin, and ventral surfaces of limbs smooth; belly moderately +granular; posterior surfaces of thighs heavily granular; no thoracic +fold. Tongue nearly round, shallowly notched behind, free posteriorly +for about one-fourth its length; vomerine teeth, 3-4, long, situated +on posteroventral edges of narrow transverse vomerine ridges between +moderately large, round inner nares; no vocal slits. + +Color (in alcohol) purplish brown on dorsal surfaces of head, body, +and limbs; fine darker reticulations on dorsum; flanks pale tan with +dark brown spots; posterior surfaces of thighs tan; chin creamy white +with brown spots; belly dusty white; undersides of thighs and shanks +yellow; webbing grayish brown; undersides of first two fingers dusty +white, of third and fourth fingers and of feet brown; anal stripe +absent; small white flecks above and below anal opening. + +Color (in life) dark green on dorsal surfaces of head, body, and +limbs; darker green reticulations on back; flanks dusty white with +dark olive-gray mottling; dark olive-gray stripe from nostril through +eye and then to insertion of forelimb; upper lips pale green; inguinal +region, anterior and posterior surfaces of thighs dark yellowish +orange; ventral surfaces of tarsi and shanks, and webbing of feet +dusty yellow; belly white; iris silvery gold. + + [Illustration: FIG. 3. Holotype of _Hyla charadricola_ + (KU 58414). x 1.5.] + +_Variation._--Adult males have snout-vent lengths from 35.3 to 44.4 +mm., and adult females from 43.4 to 50.9 mm. No notable variation in +structure is displayed in the type series. In some individuals the +dark reticulation on the dorsum is faint. Juveniles in life had dorsal +colorations varying from rich brown with darker reticulations to pale +green or gray with dark green reticulations. Some adults when +collected were pale green with faint or no dorsal reticulations; later +these individuals darkened. In all specimens the anal stripe is +absent, and the flanks are heavily spotted. + +_Comparisons.--Hyla charadricola_ differs from all other members of +the _Hyla bistincta_ group in the following combination of characters: +truncate snout, green dorsum, and absence of a thoracic fold. From +other hylids that occur in the same area, _Hyla charadricola_ differs +from _Hyla miotympanum_ and _H. arborescandens_, both of which are +green dorsally, by having a truncate snout and longer fingers with +less webbing. _Hyla eximia_, though green, has brown dorsal spots, +shorter fingers, and a round snout. _Hyla robertsorum_ differs in +having a round snout and brown dorsum, and _Hyla taeniopus_ is much +larger, has transverse bands on the limbs, and has extensive webbing +between fingers. + +_Remarks._--At the type locality, a shallow rocky stream in pine +forest, _Hyla charadricola_ was found beneath rocks at the edge of +fast moving sections of the stream and beneath rocks in shallow +riffles in the stream. Most of the frogs were in water. At night +they were found sitting on rocks in the stream. _Hyla miotympanum_, +which is abundant at the type locality, lives in bushes and beneath +rocks along the stream but usually is not found in the riffles inhabited +by _Hyla charadricola_. At Lago de Tejocotal _Hyla charadricola_ +was found beneath rocks near the shore of the lake and by a +stream in the pine forest. Individuals were found on low vegetation +over-hanging a small stream in pine-oak forest four kilometers southwest +of Tianguistengo. + +Five recently metamorphosed young (KU 58424-9) found at the +type locality on June 8, 1960, have snout-vent lengths of 22.4 to +24.0 (average 23.2) mm. The young are colored like the adults, +except that in life the dorsum is a brighter green and the flanks are +more yellow than tan and have less dark spotting than in adults. + +_Distribution.--Hyla charadricola_ inhabits streams in pine and +pine-oak forests at elevations of 2000 to 2300 meters in northern +Puebla and eastern Hidalgo (Fig. 2). + +Specimens examined.--HIDALGO: Lago de Tejocotal, 11 km. E +Acaxochitlan, KU 58438, UMMZ 104032, 118165; 4 km. SW Tianguistengo, +KU 53811-2. PUEBLA: _11.7 km. W Huachinango_, UMMZ 121567 (5); Rio +Totolapa, 14.4 km. W Huachinango, KU 53813-5, 55624, 58414-37, 59813 +(skeleton), 59886 (skeleton), MCZ 34964-5, UIMNH 50966, UMMZ 118166 +(5), S-2242 (skeleton). + + +#Hyla robertsorum# Taylor + + _Hyla robertsorum_ Taylor, Univ. Kansas Sci. Bull., + 26:393-396, figs. 5-6, November 27, 1940 [Holotype.--CNHM + 100124 (formerly EHT-HMS 16264) from El Chico Parque + Nacional, Hidalgo, Mexico; Mr. and Mrs. Radclyffe Roberts and + Edward H. Taylor collectors]; Univ. Kansas Sci. Bull., + 28:310, November 15, 1942. Taylor and Smith, Proc. U. S. + Natl. Mus., 95:589, June 30, 1945. Smith and Taylor, Bull. U. + S. Natl. Mus., 194:87, June 17, 1948; Univ. Kansas Sci. + Bull., 33:333, March 20, 1950. Rabb and Mosimann, Occas. + Papers Mus. Zool. Univ. Michigan, 563:1-9, March 29, 1955. + Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 15:48, December + 20, 1961. + +_Diagnosis._--Maximum snout-vent length in males, 48 mm.; snout in +dorsal profile rounded; tarsal fold short and weak; inner metatarsal +tubercle moderate in size and elliptical; outer metatarsal tubercle +small, round, and indistinct; webbing extending to base of penultimate +phalanx of fourth toe; nuptial spines on thumb small; weak thoracic +fold present; anal opening above level of middle of thighs; dorsum +dark brown with dark reticulations; venter brown with cream-colored +flecks; flanks brown with creamy white spots; anal stripe absent; +small white spots in anal region; vocal slits absent. + +_Description._--The following description is based on KU 57651 from El +Chico Parque Nacional, Hidalgo. Adult male having a snout-vent length +of 45.1 mm.; tibia length, 22.1 mm., 49.0 per cent of snout-vent +length; foot length (measured from proximal edge of inner metatarsal +tubercle to tip of longest toe), 21.5 mm.; greatest width of head, +13.7 mm., 30.4 per cent of snout-vent length; head length, 12.6 mm., +27.9 per cent of snout-vent length; diameter of eye, 4.0 mm.; diameter +of tympanum, 1.8 mm., 45 per cent of diameter of eye. Snout short, in +lateral profile blunt, in dorsal profile round; canthus rounded; +loreal region slightly concave; lips thick, round, and not flaring; +nostrils slightly protuberant; internarial distance, 3.6 mm.; +interorbital distance, 4.0 mm., slightly broader than width of eyelid, +3.5 mm. A moderately heavy dermal fold from posterior corner of eye +above tympanum and curving downward towards insertion of forearm; +tympanum nearly round, covered by dermal fold above, its diameter +slightly less than its distance from eye. Forearm moderately robust; +distinct fold on wrist; prepollex much enlarged with patch of small +nuptial spines continuous on side of digit; similar line of nuptial +spines on inner edge of second finger; subarticular tubercles round, +moderate in size, none bifid; supernumerary tubercles small and +present only proximally; fingers long and slender; length of fingers +from shortest to longest, 1-2-4-3; discs moderately large, that on +third finger about size of tympanum; no web between first and second +fingers; rudimentary web between other fingers. Legs robust; heels +over-lap by about one-fourth length of shank when hind limbs +adpressed; tibiotarsal articulation extending to posterior corner of +eye; tarsal fold weak, extending to about middle of tarsus; inner +metatarsal tubercle moderately large, flat, and elliptical; outer +metatarsal tubercle small, round, and indistinct; subarticular +tubercles round; supernumerary tubercles small, in single row on +proximal segments of each digit; toes moderately long and slender; +length of toes from shortest to longest, 1-2-5-3-4, the fifth nearly +as long as third; toes nearly fully webbed; web extending to base of +penultimate phalanx of fourth toe and to discs on other toes; discs +small, about two-thirds size of those on fingers. Anal opening above +middle of thighs; anal sheath short, deeply creased medially; heavy +transverse dermal fold above anus; no large anal tubercles. Skin of +all dorsal surfaces, chin, and ventral surfaces of limbs, except +proximal parts of thighs, smooth; belly and proximal parts of thighs +areolate; thoracic fold present, weak. Tongue elliptical, slightly +longer than wide, not notched behind, and free posteriorly for about +one-fourth of its length; vomerine teeth 3-3, situated on small, +widely separated, transverse ridges between rather small elliptical +inner nares; no vocal slits. + +Color (in alcohol) dark brown with irregular darker reticulations on +dorsal surfaces of head, body, and limbs; flanks brown with small +creamy white spots; posterior surfaces of thighs dark brown; chin +creamy tan; belly grayish brown with cream-colored flecks; ventral +surfaces of limbs pale brown; webbing on feet gray; small white spots +in anal region. + +Color (in life) chocolate brown with darker brown reticulations and +irregular blotches above; flanks brown with yellow spots; belly gray +to grayish brown with faint cream-colored spots; iris a deep bronze +color. + +_Variation._--In males the total number of vomerine teeth varies from +4 to 7. In many specimens the vomerine ridges are larger and more +closely approximated medially than in the specimen described above. +Females attain snout-vent lengths of 51 mm., have as many as 9 +vomerine teeth, and have a proportionately larger tympanum than males. +Some of the largest specimens of both sexes have indistinct +cream-colored pustules scattered on the ventral surface of the +forearm. Some individuals have nearly uniform grayish brown ventral +surfaces; in others the chin, as well as the abdomen, is brown with +cream-colored spots. The dorsal surfaces of some specimens are nearly +uniform dark brown with no reticulations. In others the dorsum is +paler brown with distinct darker mottling; in some of these there is +little mottling laterally, so that there is the effect of an +irregular, pale brown, dorsolateral stripe. + +_Tadpoles._--The tadpoles of this species were described by Rabb and +Mosimann (1955). Tadpoles obtained from streams at 3.3 kilometers +north and at 8.5 kilometers southeast of Zacualtipan, Hidalgo, are +like those described by Rabb and Mosimann in having 2/3 tooth rows, +peglike serrations on the beaks, and long rounded tails. The largest +tadpole (KU 60078) has small hind legs, a body length of 22 mm., and a +total length of 61 mm. + +_Remarks._--Taylor (1940:393) found frogs of this species in plants +along small spring-fed rivulets in an open meadow at El Chico +Parque Nacional. Also, he noted that active frogs dove into the +streams and took refuge in the mud on the bottom. Rabb and +Mosimann (1955:1) found this species along banks of tiny streams +in open meadows and noted that the frogs sought refuge in the +water. At El Chico Parque Nacional on June 8, 1960, I found _Hyla +robertsorum_ under rocks along small rivulets by day; at night, when +the temperature was 14 deg. C., frogs were sitting on rocks and in +junipers overhanging a small stream. At the same locality on June +23 and 24, 1962, frogs of this species were found on rainy nights, +when the temperature varied from 10 to 12 deg. C. At this time the +frogs were sitting on the grassy banks of rivulets in the meadow. +During the day _Hyla robertsorum_ was found on the earthen banks +of the rivulets in places where dense growths of grass overhung +the streams. On December 23, 1959, one specimen of _Hyla robertsorum_ +was found beneath a rock in a small stream in pine forest at +3.3 kilometers north of Zacualtipan. + +Rabb and Mosimann (1955:1) obtained tadpoles of _Hyla robertsorum_ +from quiet pools of a stream at El Chico Parque Nacional. I +found tadpoles in pools in rocky streams in pine forest at 3.3 kilometers +north and at 8.5 kilometers southeast of Zacualtipan. Four +completely metamorphosed juveniles obtained on June 8, 1960, at +El Chico Parque Nacional have snout-vent lengths of 30.6 to +32.0 mm. Gravid females were found at the same locality on June +8, 1960, and June 23, 1962. + + [Illustration: FIG. 4. Map showing locality records for + _Hyla crassa_, _Hyla pachyderma_, and _Hyla robertsorum_.] + +_Distribution.--Hyla robertsorum_ inhabits streams in the pine and fir +forests in the higher parts (2250 to 3050 meters) of the Sierra Madre +Oriental in extreme northern Puebla and eastern Hidalgo (Fig. 4). + +Specimens examined.--HIDALGO: 16 km. W Agua Blanca, UMMZ 106432 (6); +El Chico Parque Nacional, CNHM 75786, 100124, KU 57650-71, 59824-5 +(skeletons), 59914-5 (skeletons), 71269-95, 71757 (skeleton), UIMNH +10349-64, 27022-35, 39434-49, UMMZ 92462, 106401 (5), 106443 +(tadpoles), USNM 114762-85, 134268; 3.3 km. N Zacualtipan, KU 53810, +60078 (tadpoles); 8.5 km. SE Zacualtipan, KU 60079 (tadpoles). PUEBLA: +Honey, UMMZ 95245. + + +#Hyla pachyderma# Taylor + + _Hyla pachyderma_ Taylor, Univ. Kansas Sci. Bull., + 28:308-310, pl. 27, figs. 1-4, November 12, 1942 + [Holotype.--USNM 115029 from Pan de Olla, Veracruz, south of + Tezuitlan, Puebla, Mexico; Hobart M. Smith collector]. Taylor + and Smith, Proc. U. S. Natl. Mus., 95:588, June 30, 1945. + Smith and Taylor, Bull. U. S. Natl. Mus., 194:86, June 17, + 1948; Univ. Kansas Sci. Bull, 33:350, March 20, 1950. Rabb + and Mosimann, Occ. Papers Mus. Zool. Univ. Michigan, 563:7-8, + March 29, 1955. + +_Diagnosis._--Maximum snout-vent length in males 40 mm.; snout in +dorsal profile round; tarsal fold strong; inner metatarsal tubercle +round and moderate in size; outer metatarsal tubercle small and +indistinct; webbing on foot extending to middle of penultimate phalanx +of fourth toe; nuptial spines on thumb large; thoracic fold present; +anal opening at level of middle of thighs; dorsum dull grayish brown +with scattered indistinct dark flecks; venter cream-color mottled with +brown on throat and chest; flanks grayish brown with cream-colored +reticulations; anal stripe distinct, creamy white, sometimes extending +outward on thighs; white spots or line below anus; vocal slits absent. + +_Description._--The following description is based on USNM 115028 from +Pan de Olla, Veracruz. Adult male having a snout-vent length of 39.9 +mm.; tibia length, 21.0 mm., 52.6 per cent of snout-vent length; foot +length (measured from proximal edge of inner metatarsal tubercle to +tip of longest toe), 20.5 mm.; greatest width of head, 12.8 mm., 32.1 +per cent of snout-vent length; head length, 12.3 mm., 30.8 per cent of +snout-vent length. Snout short, in lateral profile bluntly rounded, in +dorsal profile rounded; canthus rounded; loreal region slightly +concave; lips thick, round, and not flaring; nostrils slightly +protuberant; internarial distance, 2.7 mm.; interorbital distance, 3.7 +mm., somewhat broader than eyelid, 2.9 mm. A heavy dermal fold from +posterior corner of eye above tympanic region and then to insertion of +forearm; tympanum completely concealed. Forearm moderately robust; +distinct fold on wrist; prepollex enlarged bearing cluster of +moderate-sized, horny, nuptial spines continuous on edge of digit; row +of spines present on second finger; subarticular tubercles round, +small proximally and slightly larger distally; supernumerary tubercles +small and indistinct; three palmar tubercles, median and outer partly +fused; fingers long, moderately slender; discs moderately large; +length of fingers from shortest to longest, 1-2-4-3; second and fourth +fingers subequal in length; webbing between fingers rudimentary. Heels +overlap by about one-fourth length of shank when hind limbs adpressed; +tibiotarsal articulation extends to anterior edge of eye; tarsal fold +thick, low, extending nearly to heel; inner metatarsal tubercle +moderately large and round; outer metatarsal tubercle small and +indistinct; subarticular tubercles small and round; supernumerary +tubercles small, present on proximal segments of digits; toes +moderately long and slender; length of toes from shortest to longest, +1-2-3-5-4; third and fifth toes subequal in length; toes about +three-fourths webbed; web extending to middle of penultimate phalanx +of fourth toe; discs rather small, about two-thirds size of those on +fingers. Anal opening directed posteroventrally at level of middle of +thighs; anal flap slightly elongate; thick, transverse dermal fold +above anus. Skin of dorsum and ventral surfaces of limbs, except +thighs, smooth; skin of chin, belly, and ventral surfaces of thighs +granular; thoracic fold present. Tongue nearly round, slightly notched +behind, and barely free posteriorly; vomerine teeth 3-3, situated on +posteroventral edges of small, transverse vomerine ridges between +rather large triangular inner nares; no vocal slits. + +Color (in alcohol) of dorsal surfaces of head, body, and limbs dull +grayish brown with indistinct scattered darker flecks; flanks grayish +brown with cream-colored reticulations; posterior surfaces of thighs +tan; chin cream-color, mottled with brown; belly creamy yellow; anal +stripe cream-color. + +_Variation._--In addition to the specimen described above three others +are known. One is a juvenile having a snout-vent length of 29.5 mm., +and two are females having snout-vent lengths of 46.9 and 41.6 mm. +Variation in structure and coloration between the four specimens is +slight. In the females the tympani are partly visible and are about +one-third the diameter of the eye; the chest is mottled with brown; +the anal stripe extends laterally in the form of a row of +cream-colored dashes and spots onto the posterodorsal surfaces of the +thighs. + +_Remarks._--On the basis of the four specimens available for study, +_Hyla pachyderma_ seems to be closely related to _Hyla crassa_ and +_Hyla robertsorum_. In the _Hyla bistincta_ group, _Hyla pachyderma_ +is unique in having enlarged nuptial spines. + +Taylor and Smith (1945:588) stated that the frogs were found on +bushes and weeds beside a small, bounding stream near Pan de Olla. +I have searched unsuccessfully for this species in the area around +Pan de Olla and Tezuitlan. + +_Distribution._--This species is known only from a stream at an +elevation of about 1600 meters on the Atlantic slopes of the Sierra +Madre Oriental in central Veracruz (Fig. 4). + +Specimens examined.--VERACRUZ: Pan de Olla, south of Tezuitlan, +Puebla, USNM 115026-9. + + +#Hyla crassa# (Brocchi) + + _Cauphias crassus_ Brocchi, Bull. Soc. Philom. Paris, ser. 7, + 1:130, 1877 [Holotype.--MNHN 6331 from "Mexico;" Adolphe + Boucard collector]. + + _Cauphias crassum_ Brocchi, Etudes des batraciens de + l'Amerique Centrale, p. 64, pl. 12, fig. 4, 1882. Diaz de + Leon, Indice de los batracios que se encuentran en la + Republica Mexicana, p. 21, June, 1904. Kellogg, Bull. U. S. + Nat. Mus., 160:118-120, March 31, 1932. Taylor, Univ. Kansas + Sci. Bull., 26:392, November 15, 1940. Rabb and Mosimann, + Occas. Papers Mus. Zool. Univ. Michigan, 563:7, March 29, + 1955. + + _Hyla crassa_, Boulenger, Catalogue Batrachia Salientia, + 2nd. Ed., p. 396, February 1, 1882. Guenther, Biologia + Centrali-Americana, Reptilia and Batrachia, p. 281, + September, 1901. Nieden, Das Tierreich, Amphibia, Anura 1, + p. 248, June, 1923. Smith and Taylor, Bull. U. S. Natl. Mus., + 194:86, June 17, 1948. Taylor, Amer. Mus. Novitates, 1437:20, + December 7, 1949. + + _Hypsiboas crassus_, Cope, Bull. U. S. Natl. Mus., 32:14, + 1887. + + _Hyla robustofemora_ Taylor, Univ. Kansas Sci. Bull., + 26:389-393, figs. 3-4, November 27, 1940 [Holotype.--UIMNH + 25050 (formerly EHT-HMS 16314) from Cerro San Felipe, 15 + kilometers northeast of Oaxaca, Oaxaca, Mexico; Edward H. + Taylor collector]; Univ. Kansas Sci. Bull., 28:310, November + 15, 1942. Smith and Taylor, Bull. U. S. Natl. Mus., 194:86, + June 17, 1948. Taylor, Amer. Mus. Novitates, 1437:20, + December 7, 1949. Smith and Taylor, Univ. Kansas Sci. Bull., + 33:339, March 20, 1950. Rabb and Mosimann, Occas. Papers Mus. + Zool. Univ. Michigan, 563:7, March 29, 1955. + + _Plectrohyla crassa_, Hartweg, Occas. Papers Mus. Zool. Univ. + Michigan, 437:1, June 30, 1941. Stuart, Occas. Papers Mus. + Zool. Univ. Michigan, 455:6, January 5, 1942. + +_Diagnosis._--Maximum snout-vent length in males 54 mm.; snout in +dorsal profile round; tarsal fold strong; inner metatarsal tubercle +small and elliptical; outer metatarsal tubercle small, flat, and +indistinct; foot fully webbed; nuptial spines on thumb small; thoracic +fold absent; anal opening at level of middle of femur; dorsum dull +olive-green; belly creamy yellow; chin gray with yellow flecks; flanks +dull olive-green with scattered cream-colored spots; and stripe faint, +cream-color; vocal slits absent. + +_Description._--The following description is based on UIMNH 25050 from +Cerro San Felipe, Oaxaca. Adult male having a snout-vent length of +53.7 mm.; tibia length, 26.9 mm., 50.1 per cent of snout-vent length; +foot length (measured from proximal edge of inner metatarsal tubercle +to tip of longest toe), 25.4 mm.; greatest width of head, 17.6 mm., +32.8 per cent of snout-vent length; head length, 16.0 mm., 29.8 per +cent of snout-vent length; diameter of eye, 5.4 mm.; diameter of +tympanum, 1.5 mm., 27.8 per cent of diameter of eye. Snout short, in +lateral profile bluntly rounded, in dorsal profile broadly round; +canthus absent; loreal region nearly flat; lips thick and not flaring; +nostrils barely protuberant; internarial distance, 3.8 mm.; +interorbital distance, 4.7 mm., somewhat broader than width of eyelid, +3.8 mm. Heavy dermal fold from posterior corner of eye above tympanum +and then to insertion of forearm; tympanum concealed above, its +diameter about equal to its distance from eye. Forearm thick; distinct +fold on wrist; prepollex enlarged bearing patch of small nuptial +spines continuous on side of digit; similar patch on second finger; +subarticular tubercles small and round, none bifid; few supernumerary +tubercles on proximal segments of digits; large, flat palmar tubercle; +fingers long and slender; length of fingers from shortest to longest, +1-2-4-3; discs moderately large; rudimentary web between second and +third fingers and between third and fourth. Legs thick; heels overlap +by about one-fourth length of shank when hindlimbs adpressed; +tibiotarsal articulation extends to posterior corner of eye; tarsal +fold thick, extending to heel; inner metatarsal tubercle small and +elliptical; outer metatarsal tubercle small, flat, and indistinct; +subarticular tubercles small and round; single row of supernumerary +tubercles on proximal segments of each digit; toes moderately short +and slender; length of toes from shortest to longest, 1-2-3-5-4; toes +fully webbed; flap of skin on inner surface of first toe; discs about +same size as those on fingers. Anal opening directed posteroventrally +at middle of thighs; anal sheath moderately elongate; small tubercles +below anal opening. Skin of dorsum rather smooth, somewhat granular +on dorsal surfaces of limbs; skin of chin and belly moderately +granular; that of posterior surfaces of thighs smooth; no thoracic +fold. Tongue nearly round, shallowly notched posteriorly, and free for +about one-fourth its length; vomerine teeth 5-5, situated on rounded +ridges between small inner nares; no vocal slits. + +Color (in alcohol) dull olive-green on dorsal surfaces of head, body, +and limbs; flanks dull olive-green with scattered cream-colored spots; +posterior surfaces of thighs grayish brown with faint creamy mottling; +chin gray with cream-colored spots; belly creamy yellow, suffused with +gray posteriorly; undersides of feet and webbing gray; anal stripe +faint, pale cream-color. + +_Variation._--The only other known specimen (MNHN 6331) is a female +having a snout-vent length of 53.7 mm. and resembling the specimen +described above in most details of morphology. In MNHN 6331 the +tympanum is completely concealed, and the 8-7 vomerine teeth are +arranged in two irregular rows. The female has more cream-colored +mottling on the flanks and posterior surfaces of the thighs and more +distinct mottling on the throat than the male described above. + +_Remarks._--The systematic status of _Cauphias crassus_ Brocchi has +been in doubt since the time of the original description. Brocchi +(1877:130) stated: "Les dernieres phalanges sont obtuses, tronques a +leur extremite anterieure." Brocchi placed the species in his genus +_Cauphias_ (type species, _C. guatemalensis_), which he considered to +be related to _Hylodes_ ( = _Eleutherodactylus_ in the sense used by +Brocchi); he thereby placed _Cauphias_ in his Hylodidae ( = +Leptodactylidae, in part). This idea of relationships was perpetuated +by Barbour (1927:96), who reported on the second known specimen of +_Cauphias guatemalensis_ and stated: "When I dissected the sternum I +was at once struck by its similarity to Noble's figures of +transitional types between arciferal and firmisternal forms. The +_Cauphias_ sternum recalls some of his figures for _Sminthillus_ and +_Eleutherodactylus_. This genus is probably most closely related to +the latter and has probably become highly modified to meet some +peculiar environmental condition or on account of some specialized +habits as yet unknown." Kellogg (1932:118) placed _Cauphias_ in the +Leptodactylidae and stated that the terminal phalanges are T-shaped. +Hartweg (1941:1) considered _Plectrohyla_ to be the correct generic +name for _Cauphias guatemalensis_; he thereby relegated _Cauphias_ to +the synonomy of _Plectrohyla_. Hartweg (1941:9) further showed that +the terminal phalanges of _Plectrohyla guatemalensis_ were not +T-shaped and that intercalary cartilages were present. Thus, he +correctly concluded that _Plectrohyla guatemalensis_ (and _P. crassa_ +by implication) was a member of the family Hylidae. Stuart (1942:6) +followed Hartweg's allocations and further suggested that _Plectrohyla +crassa_ might be the same species as _Hyla robustofemora_ Taylor. In +his description of _H. robustofemora_ Taylor (1940:392), who had not +examined the type of _Cauphias crassus_, stated that were it not for +the statements of Brocchi and Kellogg that _C. crassus_ has T-shaped +terminal phalanges, "I might suspect I had before me a specimen of +_Cauphias_ closely related to _crassum_." + +I have compared the type of _Cauphias crassus_ with that of _Hyla +robustofemora_. With the exception of the minor differences mentioned +in the preceding section on variation, the specimens are alike, +leaving little doubt that they represent the same species. The +statements of Brocchi and Kellogg to the contrary, the type of +_Cauphias crassus_ possesses intercalary cartilages between the +penultimate and terminal phalanges; the latter are not T-shaped, but +as in the type of _Hyla robustofemora_, resemble those typical of +_Hyla_. On the basis of the morphological characters, as pointed out +for _Hyla robustofemora_ by Taylor (1940:392), _Hyla crassa_ is a +member of the _Hyla bistincta_ group. + +_Distribution._--This species is definitely known only from a small +stream at an elevation of 2300 meters in the mountains of central +Oaxaca (Fig. 4). + +Specimens examined.--OAXACA: Cerro San Felipe, UIMNH 25050. "Mexico," +MNHN 6331. + + + + +RELATIONSHIPS + + +The evolutionary trend in the members of the _Hyla bistincta_ group +is towards aquatic habits. _Hyla bistincta_, the least specialized +species in the group, has relatively short fingers, webbing between +the fingers, a truncate, high snout, and relatively large subarticular +and supernumerary tubercles. _Hyla charadricola_ resembles _bistincta_ +in having relatively short fingers, a slight amount of webbing, and a +truncate snout. Apparently these two species are more closely related +to one another than either is to the other species in the group. _Hyla +robertsorum_, _pachyderma_, and _crassa_ are the most aquatic members +of the group. These species are closely related, possibly conspecific. +All have round, sloping snouts, robust forearms, long, unwebbed +fingers, and large webbed feet. Both _H. pachyderma_ and _H. crassa_ +seem to be advanced beyond _H. robertsorum_. If small nuptial spines, +moderately webbed feet, and absence of a well-defined thoracic fold +are considered to be less advanced than large nuptial spines and a +strong thoracic fold, as in _H. pachyderma_, or fully webbed feet, as +in _H. crassa_, then _H. robertsorum_ must be considered to be less +advanced than _H. pachyderma_ or _H. crassa_. + +Members of the _Hyla bistincta_ group inhabit mountain streams. The +frogs can be found along these streams throughout the year. Since in +most stream-breeding hylids there is no migration to breeding sites, +the breeding call does not function to attract females to the breeding +site. Apparently voices are lacking in all members of the _Hyla +bistincta_ group, except in _Hyla bistincta_. The presence of vocal +slits and the ability to call further indicate that _Hyla bistincta_ +is the primitive member of this group. + +Members of the _Hyla bistincta_ group and the species of _Plectrohyla_ +closely resemble each other in osteology and body form of the adults +and in structure of the tadpoles. This resemblance suggests a close +relationship between the two groups. _Plectrohyla_ apparently evolved +from an ancestral stock resembling the extant _Hyla bistincta_. +Probably this stock gave rise independently to _Plectrohyla_ and +to the _Hyla robertsorum-pachyderma-crassa_ complex. In the former +the voice was retained, and a projecting prepollex spine developed, +whereas in the latter the voice was lost, and the prepollex spine +did not project. _Plectrohyla_ lives in mountain streams in the +Chiapan-Guatemalan highlands; the _Hyla robertsorum-pachyderma-crassa_ +complex inhabits similar environments in the Sierra Madre Oriental in +Mexico. _Hyla charadricola_ also lives in the Sierra Madre Oriental, +whereas _Hyla bistincta_ is widespread in the mountains of Mexico +southeastward to the Isthmus of Tehuantepec. + + + + +LITERATURE CITED + + +BARBOUR, T. + + 1927. _Cauphias_ rediscovered. Copeia, no. 165:96-98, + December 23. + +BROCCHI, P. + + 1877. Note sur quelques batraciens hylaeformes recueillis au + Mexique et au Guatemala. Bull. Soc. Philom. Paris, ser. + 7, 1 (3):122-132. + +DUELLMAN, W. E. + + 1961. The amphibians and reptiles of Michoacan, Mexico. Univ. + Kansas Publ. Mus. Nat. Hist., 15:1-148, pls. 1-6, December + 20. + +HARTWEG, N. + + 1941. Notes on the genus _Plectrohyla_, with descriptions of + new species. Occas. Papers Mus. Zool. Univ. Michigan, + 437:1-10, pl. 1, June 30. + +KELLOGG, R. + + 1932. Mexican tailless amphibians in the United States National + Museum. Bull. U. S. Natl. Mus., 160:iv + 224 pp., pl. 1, + March 31. + +RABB, G. B. and MOSIMANN, J. E. + + 1955. The tadpole of _Hyla robertsorum_, with comments on the + affinities of the species. Occas. Papers Mus. Zool. Univ. + Michigan, 563:1-9, March 29. + +SHANNON, F. A. + + 1951. Notes on a herpetological collection from Oaxaca and + other localities in Mexico. Proc. U. S. Natl. Mus., + 101:465-484, May 17. + +STUART, L. C. + + 1942. Descriptions of two new species of _Plectrohyla_ Brocchi, + with comments on several forms of tadpoles. Occas. Papers + Mus. Zool. Univ. Michigan, 455:1-14, January 5. + +TAYLOR, E. H. + + 1940. New species of Mexican Anura. Univ. Kansas Sci. Bull., + 41:385-405, November 27. + + 1948. A new hylid frog from eastern Mexico. Univ. Kansas + Publ. Mus. Nat. Hist., 1:259-262, August 16. + + 1949. A new hylid frog from central Veracruz. Copeia, 1949 + (4):272-274, pl. 1, December 15. + +TAYLOR, E. H. and SMITH, H. M. + + 1945. Summary of the collections of amphibians made in Mexico + under the Walter Rathbone Bacon Traveling Scholarship. + Proc. U. S. Natl. Mus., 95:521-613, pls. 18-32, June 30. + + _Transmitted June 24, 1963._ + + + 29-8590 + + + + + +End of the Project Gutenberg EBook of A Review of the Frogs of the Hyla +bistincta Group, by William E. 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