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Title: A Review of the Frogs of the Hyla bistincta Group

Author: William E. Duellman

Release Date: June 2, 2010 [EBook #32653]

Language: English

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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

[Pg 471]

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 México 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

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, ⅔ tooth-rows, two or more rows of labial papillae, and long tails with low fins.

Fig. 1. Palmar view of right hand of:
a.—Hyla robertsorum (KU 57661),
b.—Hyla charadricola (KU 58414). × 3.

[Pg 473] 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 ⅔ 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.[Pg 474]

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.)

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

[Pg 475]

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

[Pg 477]

Remarks.—Shannon (1951:470) named Hyla bistincta labeculata on the basis of a single male from San Lucas Camotlán, 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 México have reticulate mottling on the flanks as compared with the marbling on the flanks in specimens from eastern México. 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 ⅔ 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.

[Pg 478]

Fig. 2. Map showing locality records for Hyla bistincta and Hyla charadricola.]

Hyla charadricola new species

Fig. 3. Holotype of Hyla charadricola (KU 58414). × 1.5.

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.

Hyla robertsorum Taylor

[Pg 482]

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° 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° 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 Zacualtipán.

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 Zacualtipán. Four [Pg 484] 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.

Fig. 4. Map showing locality records for Hyla crassa, Hyla pachyderma, and Hyla robertsorum.

[Pg 485]

Hyla pachyderma Taylor

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 Tezuitlán.

Hyla crassa (Brocchi)

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, tronqués a leur extrémité antérieure." 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 [Pg 489] 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.

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. [Pg 490] 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 México. Hyla charadricola also lives in the Sierra Madre Oriental, whereas Hyla bistincta is widespread in the mountains of México southeastward to the Isthmus of Tehuantepec.

[Pg 491]

LITERATURE CITED

Barbour, T.

Brocchi, P.

Duellman, W. E.

Hartweg, N.

Kellogg, R.

Rabb, G. B. and Mosimann, J. E.

Shannon, F. A.

Stuart, L. C.

Taylor, E. H.

Taylor, E. H. and Smith, H. M.

    Transmitted June 24, 1963.

29-8590

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