Part 4

The wholesale synonymization of names, which, on the bases of their published diagnoses, seem to apply to distinctly different species, is possible with the application of uniform criteria to the types and series of other specimens. In measurements and proportions the type specimens of the nominal taxa all fall within the range of variation exhibited by a series of 18 males and 15 females from Provincia Pastaza, Ecuador, except the ratio of the diameter of the tympanum to that of the eye in the female holotype of Hyla festae. In that specimen the ratio is 0.574, whereas the ratio in the 15 females from Provincia Pastaza is 0.587-0.905 (mean 0.736).

Ventral coloration is the most variable character among the types. The venter in the type of Hyla festae is boldly spotted; it is distinctly spotted in cabrerai, uniform tan in carri, and tan, flecked, or spotted in the type series of buckleyi. The ventral coloration in series of specimens from Amazonian Ecuador encompasses that observed in all of the types, except that of festae, which has more ventral spotting than any other individual.

The webbing on the hand usually excludes the penultimate phalanges of the fingers, but in some specimens from Amazonian Ecuador the webbing encompasses the proximal parts of the penultimate phalanges of the fingers. In a few of these specimens, the holotype of festae, and one paratype of cabrerai the webbing extends to the middle of the penultimate phalanges of the third and fourth fingers. In the holotype of cabrerai the webbing extends to the middle of the penultimate phalanges of the third and fourth fingers and to the base of the disc of the second finger.

The types of the nominal taxa and series of specimens from Guyana and Amazonian Ecuador display noticeable variation in dorsal coloration. The variety of dorsal patterns of all of the types is included in the variation displayed by the other specimens. All specimens have some amount of dark spotting on the flanks; all have vertically barred lips, on which a pale subocular spot usually is evident. Probably the most unifying physical characteristic of all of the specimens is the nature of the skin on the anterior part of the flank. The skin is elevated amidst an irregular network of depressions. This areolate dermal condition is present in all specimens and does not occur in other species of Osteocephalus. The degree of tubercularity of the skin on the dorsum is variable and sexually dimorphic. All males are tubercular, whereas small females are smooth or have only a few scattered tubercles. Large females usually have pronounced tubercles on the eyelids and supratympanic fold.

In their description of Hyla carri, Cochran and Goin (1970:211) misrepresented the nature of the dentigerous processes of the prevomers, which are angular, not ^-shaped. Their suggestion that the Colombian Hyla carri is related to Hyla claresignata in southeastern Brasil is unfounded. The latter species is smaller (40 mm), has a yellow dorsum and venter, dark brown spots dorsolaterally, oblique dentigerous processes of the prevomers, small tympanum, and smooth skin dorsally.

The ventral coloration of the type of Hyla festae resembles that of Osteocephalus verrucigerus, but the type differs from verrucigerus by having areolate skin on the flanks and distinct dark markings on the dorsum. In verrucigerus the skin on the flanks is smooth, and the dorsum is uniform dark brown, except for a tan snout in females.

Comparisons of the types of the nominal species with series of specimens from Guyana, Colombia, Ecuador, and Peru suggest strongly that the types are representative of one taxon, the oldest name for which is Hyla buckleyi Boulenger, 1882. Consequently, we place Hyla festae Peracca, 1904, Hyla carri Cochran and Goin, 1970, and Hyla cabrerai Cochran and Goin, 1970, as junior synonyms of Hyla buckleyi Boulenger, 1882.

Diagnosis.--1) Size moderate, sexual dimorphism extreme; maximum observed snout-vent length in males 48.1 mm, in females 75.1 mm; 2) skin on dorsum in males bearing a mixture of large and small non-spinous tubercles; 3) skin on flanks, especially anteriorly, areolate; 4) web usually extending only to base of antepenultimate phalange on inner edge of third finger; 5) dorsum pale tan or green with irregular, longitudinal, dark brown blotches, usually narrowly outlined with cream; 6) venter cream or tan, suffused with brown or marked with brown spots in some specimens; 7) lips marked with vertical brown and cream bars; 8) flanks creamy tan with irregular brown spots and/or diagonal marks; 9) dermal roofing bones of skull lacking exostosis; 10) dermal sphenethmoid absent; 11) nasals widely separated medially; 12) anteromedial margin of frontoparietal at mid-level of orbit; 13) frontoparietal fontanelle partially exposed; 14) palatine serrate; 15) parasphenoid bearing odontoids; 16) zygomatic ramus of squamosal extending approximately one-half of distance to maxillary arch; 17) transverse processes of third presacral vertebra narrower than sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width and narrower in males than in females; 18) intermandibularis and submentalis muscles independent; 19) supramandibular portion of interhyoideus extensively developed; associated skin forming broad loose fold.

Osteocephalus buckleyi can be distinguished readily from all other species in the genus by the presence of areolate skin anteriorly on the flanks and by the rather boldly contrasting dorsal pattern. Furthermore, females are distinctive in having tubercles on the eyelids and supratympanic folds.

Distribution.--The periphery of the Amazon Basin, in the Guianas and Territorio do Amapa in northeastern Brasil; the upper Amazon Basin from southern Colombia to east-central Bolivia; one locality (Acevedo) in upper Rio Magdalena drainage in Colombia (Fig. 8). All localities are at elevations of less than 700 m. Records for Pallatanga and Santiago in Provincia Chimborazo, Ecuador (high on the Pacific slopes of the Andes), are considered to be erroneous. 78 specimens from 40 localities.

Remarks.--In life the dorsum is green with dark markings. A male (KU 123171) from Santa Cecilia, Ecuador, was: "Dorsum green with dark brown blotches. Anterior and posterior surfaces of thighs dull blue. Venter brown, flecked with white. Iris greenish bronze with brown horizontal triangles and ventromedian brown line." (W. E. Duellman, field notes, 16 June 1968.) A female (KU 126646) from Lago Agrio, Ecuador, was: "Dorsum pale green with darker green blotches and creamy yellow middorsal stripe. Lateral blotches bronze-tan. Flanks tan with black blotches. Anterior surfaces of thighs dark brown. Dorsal and posterior surfaces of thighs and shanks tan with dark brown blotches. Webbing brown. Suborbital spot green. Postorbital bar black. Belly grayish brown in appearance--tips of granules white; intergranular spaces brown. Iris golden bronze with black flecks peripherally and median, horizontal, reddish brown streak." (W. E. Duellman, field notes, 12 May 1969.)

No ontogenetic change in coloration has been noted.

=Osteocephalus leprieurii= (Dumeril and Bibron)

Hyla leprieurii Dumeril and Bibron, 1841:553 [Holotype.--MNHN 4629 from "Cayenne"; Mons. Leprieur collector].

Hypsiboas leprieurii--Cope, 1867:200.

Hyla leprieurii britti Melin, 1941:42 [Holotype.--NHMG 489 from the Rio Uaupes, north of the Rio Japu, Territorio do Amazonas, Brasil; Douglas Melin collector]. New synonymy.

Hyla leprieurii leprieurii--Melin, 1941:42.

Osteocephalus britti--Goin, 1961:13.

Osteocephalus leprieurii--Goin, 1961:13.

Justification of Synonymy.--The holotype of Hyla leprieurii is a female having a snout-vent length of 46.6 mm. The diameter of the tympanum is 3.7 mm, 69.8 percent of the diameter of the eye. The dorsal roofing bones are smooth, and the skin on the dorsum is smooth. The penultimate phalanges of the fingers are not included in the webbing. When we examined the specimen on 2 July 1969, it was slightly soft and somewhat faded to a peculiar grayish green color with faint darker transverse bars on the limbs. Dumeril and Bibron (1841:554) described the coloration, as follows: "The loreal region is black. A stripe of the same color extends from the posterior border of the orbit to the corner of the mouth, passing through the tympanum. All of the dorsal parts are grayish white with large transverse brown bands, which are more expanded and less regularly outlined on the back than on the limbs. There is one of these on the occiput that is in a triangular shape. All of the venter is white." (Free translation from French.)

The holotype of Hyla leprieurii britti is a male having a snout-vent length of 48.1 mm. The diameter of the tympanum is 3.6 mm, 65.5 percent of the diameter of the eye. The skin on the dorsum is tubercular; the tubercles are small on head and on the dorsal surfaces of the limbs and slightly larger on the back. The penultimate phalanges of the fingers are not included in the webbing. Melin (1941:43) stated: "Above blackish brown with a very indistinct band between the eyes; iris with mottle of metallic lustre; hinder parts of upper jaw whitish; sides of body mottled with blackish brown; hind limbs (especially tibiae and tarsi) with narrow, diffuse cross bars; beneath whitish with slight brown mottle along jaw." We examined the type on 17 February 1969; at that time it was dull brown above with faint, narrow, dark brown, transverse bars on the back and dorsal surfaces of the limbs. A cream subocular spot was evident, and the venter was creamy white.

Melin (1941:42) stated that the holotype of Hyla leprieurii britti "... resembles a good deal H. leprieurii Dum. & Bibr. As, however, it differs from the latter species by its very concave loreal region, small tympanum, and almost uniformly brownish colour, it may at least form a subspecies of leprieurii...." The pattern of narrow transverse bars on the backs of the holotypes of H. leprieurii and H. britti is a condition shared only by these two nominal taxa that are placed in Osteocephalus. Melin noted that britti differed from leprieurii in the depth of the loreal concavity and in the size of the tympanum. Neither of these differences is noteworthy in comparison with series of specimens. The depth of the loreal concavity is a highly subjective character, and we note no differences between the types. The ratio of the diameter of the tympanum to the diameter of the eye is relatively smaller in both holotypes (0.698 in leprieurii--; 0.655 in britti--) than in series of fresh specimens from Lago Agrio, Ecuador (0.652-0.884, mean 0.785 in 17 males; 0.700-0.909, mean 0.790 in 20 females). The smaller proportions in the types may be due to geographic variation or to shrinkage as a result of many years in preservative (130+ years for leprieurii; 45 for britti).

Comparisons of the holotypes with series of specimens from Ecuador, Guyana, and Surinam indicate that one morphological species occurs throughout the upper Amazon Basin and the Guianas and that both type specimens are representatives of one species. Consequently, we consider Hyla leprieurii Dumeril and Bibron, 1841, to be a monotypic species with Hyla leprieurii britti Melin, 1941, as a junior synonym.

In their account of Osteocephalus leprieurii, Cochran and Goin (1970:323) stated: "The specimen described and illustrated (MCZ 28042) has been directly compared with the types of leprieurii, planiceps, and vilarsi by the junior author and there seems to be no doubt that all are conspecific. Another specimen (CNHM 69716) has been directly compared with the types of planiceps and vilarsi and these, likewise, are considered conspecific." With this justification Cochran and Goin (1970:322) included Osteocephalus planiceps Cope, 1874, and Hyla vilarsi Melin, 1941, in the synonymy of Osteocephalus leprieurii.

We do not concur with Cochran and Goin's synonymy and contend that planiceps and vilarsi are synonyms of Osteocephalus taurinus; we give our reasons in the account of that species. We have examined the specimens listed as O. leprieurii by Cochran and Goin; several of them, including CNHM (= FMNH) 69716, are taurinus. Thus, due to Cochran and Goin's confusion of two taxa, their comparisons of certain specimens with types has little meaning.

Cochran and Goin did not include Hyla leprieurii britti in their synonymy of Osteocephalus leprieurii but did discuss the name in their account of Osteocephalus orcesi (= O. verrucigerus), as follows (1970:319): "When we first examined one of the specimens we felt sure that we had Melin's Hyla britti at hand, but on direct comparison with the type of britti the two proved to be different. After studying the type of orcesi (SUNHM 13150) we have no doubt that the specimens at hand are orcesi and that britti is a different, probably valid species."

Diagnosis.--1) Size moderate, sexual dimorphism evident; maximum observed snout-vent length in males 48.4 mm, in females, 61.5 mm; 2) skin on dorsum in males bearing numerous, minute, spinous tubercles; 3) skin on flanks smooth; 4) web extending to base of antepenultimate phalange on inner edge of third finger; 5) dorsum tan or olive-brown with transverse brown or olive bars; 6) venter creamy white or pale tan without markings; 7) lips marked with creamy tan labial stripe and suborbital spot; 8) flanks pale tan with no markings; 9) dermal roofing bones of skull lacking exostosis; 10) dermal sphenethmoid absent; 11) nasals juxtaposed medially; 12) anteromedial margin of frontoparietal between mid- and anterior levels of orbit; 13) frontoparietal fontanelle partially exposed; 14) palatine not serrate; 15) parasphenoid lacking odontoids; 16) zygomatic ramus of squamosal extending about one-half of distance to maxillary arch; 17) transverse processes of presacral vertebrae 3-8 about equal in width to one another and to sacral diapophyses; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus forming simple tubular posterolateral extension; associated skin unmodified.

Osteocephalus leprieurii differs from all other members of the genus by having transverse dark bars on the back. Two other hylids (Hyla lanciformis and multifasciata) in the Amazon Basin have transverse dark marks on the dorsum. Both of these differ from leprieurii by having pointed snouts, much longer hind limbs, and smooth skin dorsally.