Accounts of SpeciesOsteocephalus buckleyi(Boulenger)Hyla buckleyiBoulenger, 1882:362 [Syntypes.—BMNH 1947.2.13.36-39 from Sarayacu, Provincia Pastaza, Ecuador; BMNH 1947.2.13.40-41, 1947.2.13. 43-45 from Canelos, Provincia Pastaza, Ecuador; BMNH 1947.2.13.46 from "Paitanga" (= Pallatanga), Provincia Chimborazo, Ecuador (in error); Mr. Buckley collector; BMNH 1947.2.13.44 here designated as lectotype].Hyla festaePeracca, 1904:39 [Holotype.—MIZS 2950 from "Valle de Santiago" (= lower Río Zamora), Provincia Morona-Santiago, Ecuador; Enrico Festa collector]. New synonymy.Osteocephalus buckleyi—Goin, 1961:13.Hyla carriCochran and Goin, 1970:211 [Holotype.—FMNH 69702 from Acevedo, Río Suaza, Departamento Huila, Colombia; Philip collector]. New synonymy.Hyla cabreraiCochran and Goin, 1970:215 [Holotype.—USNM 152759 from Caño Guacayá, tributary of lower Río Apoporis, Comisaria Amazonas, Colombia; Isadore Cabrera collector]. New synonymy.Justification of Synonymy.—Boulenger (1882:362) listed 11 specimens in his description ofHyla buckleyi. We have examined all of these and conclude that one (BMNH 1947.2.13.42) isO. leprieurii. Cochran and Goin (1970:213) restricted the type locality to Canelos,ProvinciaPastaza, Ecuador; we here select BMNH 1947.2.13.44 from that locality as the lectotype. This specimen is a male having a snout-vent length of 37.9 mm; the diameter of the tympanumis 3.5 mm, 81.4 percent of the diameter of the eye. The type series, exclusive of BMNH 1947.2.13.42 (=O. leprieurii) consists of six males having snout-vent lengths of 37.9-44.6 (mean 40.4) mm, and four females having snout-vent lengths of 50.0-53.9 (mean 51.5) mm. The dorsum in the males bears a mixture of large and small tubercles, whereas the dorsum in females is nearly smooth. The skin on the flanks, especially the axilla, is areolate. The coloration consists of a creamy tan ground color with irregular reddish brown markings on the back and broad transverse bars on the limbs. The dorsal markings are narrowly bordered by creamy white; those on the back consist of an interorbital bar and a pair of longitudinal marks beginning in the scapular region and usually diverging posteriorly in the sacral region or converging into a broad median blotch. One specimen has a middorsal creamy white stripe from the tip of the snout to the vent. In all of the types large dark brown spots are present on the flanks and posterior surfaces of the thighs. The ventral surfaces are pale creamy tan with or without diffuse brown spots on the throat and chest.The holotype ofHyla festaeis a female having a snout-vent length of 75.0 mm; the diameter of the tympanum is 3.9 mm, 57.4 percent of the diameter of the eye. The skin is smooth on the dorsum and areolate on the anterior part of the flanks. The dorsum is pale brown with a large median longitudinal dark brown blotch on the back and broad transverse bars, narrowly outlined by cream, on the limbs. Dark brown spots are present on the flanks; the posterior surfaces of the thighs are dark brown. The throat and belly are grayish white with irregular dark brown spots.The holotype ofHyla carriis a female having a snout-vent length of 66.1 mm; the diameter of the tympanum is 4.7 mm, 81.0 percent of the diameter of the eye. The skin on the dorsum is smooth with scattered small tubercles and areolate on the anterior part of the flanks. The dorsum is tan with irregular dark brown blotches on the back and transverse bars on the limbs; all dorsal markings are narrowly outlined by creamy white. Dark brown spots are present on the flanks; the venter and posterior surfaces of the thighs are tan without dark spots.The holotype ofHyla cabreraiis a female having a snout-vent length of 52.7 mm; the diameter of the tympanum is 4.0 mm, 76.9 percent of the diameter of the eye. The skin on the dorsum is weakly tuberculate and that on the anterior part of the flanks is areolate. The dorsum is creamy tan with dark brown markings (interorbital bar, reticulations on occiput, three longitudinal streaks on back,and broad transverse bars on limbs). Irregular dark brown spots are present on the flanks. The venter is pinkish tan with small reddish brown spots on the throat and darker brown spots on the chest and belly.In their description ofHyla cabrerai, Cochran and Goin (1970:217) stated: "This species, together withbuckleyiandpearsonicertainly make a closely knit groupߪ. Bothbuckleyiandcabreraihave long hind legs, with the extended heel reaching to the tip of the snout, while inpearsonithe extended heel reaches only to the eye.H. buckleyihas the belly dusky, while it is heavily spotted incabreraiand is reticulated inpearsoni.H. cabreraiseems to have the heaviest hands with the most webbing between the fingers; the other two species have the webbing reduced between the fingers." The description ofHyla cabreraiwas based on three specimens. We have examined the holotype and one paratype (WCAB 13284 from Territorio do Amapá, Brasil). Another paratype in the private collection of C. J. Goin from Caño Tuí, between Mitú and Raudal de Yurupari, Comisaria de Vaupés, Colombia, was not examined.Cochran and Goin (1970:211) based their description ofHyla carrion one gravid female and stated: "A largeHylawith the vomerine teeth in two ʌʌ shaped patches between the somewhat squarish choanae; reduced webs between the fingers; and a pattern of dorsal dark blotches bordered by light margins. The species is not similar to any other species known in Colombia. It is perhaps most closely related toHyla claresignataof Brazil, from which it can be differentiated by its more heavily spotted dorsum, larger tympanum, and lack of dark anal spots."Except for the inclusion of the name in checklists,Hyla festaehas not been mentioned in the literature since the original description.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 ofHyla 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 ofHyla festaeis boldly spotted; it isdistinctly spotted incabrerai, uniform tan incarri, and tan, flecked, or spotted in the type series ofbuckleyi. The ventral coloration in series of specimens from Amazonian Ecuador encompasses that observed in all of the types, except that offestae, 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 offestae, and one paratype ofcabreraithe webbing extends to the middle of the penultimate phalanges of the third and fourth fingers. In the holotype ofcabreraithe 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 ofOsteocephalus. 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 ofHyla carri, Cochran and Goin (1970:211) misrepresented the nature of the dentigerous processes of the prevomers, which are angular, not ʌ-shaped. Their suggestion that the ColombianHyla carriis related toHyla claresignatain 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 ofHyla festaeresembles that ofOsteocephalus verrucigerus, but the type differs fromverrucigerusby having areolate skin on the flanks and distinct dark markings on the dorsum. Inverrucigerusthe 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 Perú suggest strongly that the types are representative of one taxon, the oldest name for which isHyla buckleyiBoulenger, 1882. Consequently, we placeHyla festaePeracca, 1904,Hyla carriCochran and Goin, 1970, andHyla cabreraiCochran and Goin, 1970, as junior synonyms ofHyla buckleyiBoulenger, 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 buckleyican 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 Amapá in northeastern Brasil; the upper Amazon Basin from southern Colombia to east-central Bolivia; one locality (Acevedo) in upper Río Magdalena drainage in Colombia (Fig. 8). All localities are at elevations of less than 700 m. Recordsfor 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.Fig. 8.Distribution ofOsteocephalus buckleyi(circles) andO. pearsoni(triangles).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(Duméril and Bibron)Hyla leprieuriiDuméril and Bibron, 1841:553 [Holotype.—MNHN 4629 from "Cayenne"; Mons. Leprieur collector].Hypsiboas leprieurii—Cope, 1867:200.Hyla leprieurii brittiMelin, 1941:42 [Holotype.—NHMG 489 from the Rio Uaupés, 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 ofHyla leprieuriiis 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. Duméril and Bibron (1841:554) described the coloration, as follows: "The loreal regionisblack. 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 ofHyla leprieurii brittiis 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 ofmetalliclustre; 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 onthe 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 ofHyla leprieurii britti"ߪ resembles a good dealH. leprieuriiDum. & 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 ofleprieuriiߪ." The pattern of narrow transverse bars on the backs of the holotypes ofH. leprieuriiandH. brittiis a condition shared only by these two nominal taxa that are placed inOsteocephalus. Melin noted thatbrittidiffered fromleprieuriiin 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 inleprieurii—♀; 0.655 inbritti—♂) 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 forleprieurii; 45 forbritti).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 considerHyla leprieuriiDuméril and Bibron, 1841, to be a monotypic species withHyla leprieurii brittiMelin, 1941, as a junior synonym.In their account ofOsteocephalus leprieurii, Cochran and Goin (1970:323) stated: "The specimen described and illustrated (MCZ 28042) has been directly compared with the types ofleprieurii,planiceps, andvilarsiby 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 ofplanicepsandvilarsiand these, likewise, are considered conspecific." With this justification Cochran and Goin (1970:322) includedOsteocephalus planicepsCope, 1874, andHyla vilarsiMelin, 1941, in the synonymy ofOsteocephalus leprieurii.We do not concur with Cochran and Goin's synonymy and contend thatplanicepsandvilarsiare synonyms ofOsteocephalus taurinus; we give our reasons in the account of that species. We have examined the specimens listed asO. leprieuriiby Cochran andGoin; several of them, including CNHM (= FMNH) 69716, aretaurinus. 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 includeHyla leprieurii brittiin their synonymy ofOsteocephalus leprieuriibut did discuss the name in their account ofOsteocephalus orcesi(=O. verrucigerus), as follows (1970:319): "When we first examined one of the specimens we felt sure that we had Melin'sHyla brittiat hand, but on direct comparison with the type ofbrittithe two proved to be different. After studying the type oforcesi(SUNHM 13150) we have no doubt that the specimens at hand areorcesiand thatbrittiis 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 leprieuriidiffers from all other members of the genus by having transverse dark bars on the back. Two other hylids (Hyla lanciformisandmultifasciata) in the Amazon Basin have transverse dark marks on the dorsum. Both of these differ fromleprieuriiby having pointed snouts, much longer hind limbs, and smooth skin dorsally.Distribution.—The periphery of the Amazon Basin, in the Guianas and the upper part of the basin in southern Colombia, Ecuador, Perú, and extreme western Brasil (Fig. 9). Most localities are at elevations of less than 500 m, but the species ascends the lowerAndean slopes to elevations of 1100 m. 265 specimens from 31 localities.Fig. 9.Distribution ofOsteocephalus leprieurii(circles) andO. verrucigerus(triangles).Remarks.—Most adults ofleprieuriihave distinct transverse markings on the back; these are variable in width, extent, and arrangement. In some specimens, such as USNM 166557, some of the transverse bars are fragmented into spots; in a few specimens the dorsal pattern consists solely of small dark spots arranged in transverse rows. Such specimens have a dorsal pattern resembling that of sometaurinus. The transverse nature of the dorsal markings is further modified in some specimens, such as USNM 166555, in which the dark bars are fragmented and oblique.Extreme ontogenetic change in color pattern is exhibited by this species (Fig. 10). Juveniles having snout-vent lengths of less than 28 mm have an olive-brown dorsum with a pale cream stripe across the head and broad, cream, dorsolateral stripes; transverse dark bars are absent on the body and limbs. Individuals having snout-vent lengths of 30-35 mm have dark brown transverse bars onthe back and limbs but still retain the light dorsolateral stripes, whereas the stripes are lost in larger individuals.Fig. 10.Ontogenetic change in color pattern inOsteocephalus leprieurii:a. KU 126644; b. KU 126640; c. KU 126625. ×2.Coloration in life of specimens from Lago Agrio, Ecuador: "In males the dorsal ground color varies from dark brown to ochre-tan; dorsal markings uniformly dark brown. Most specimens have dark brown and cream anal stripes; labial area cream-colored. Flanks vary from tan to white. Ventral coloration varies from salmon to tan to white. The iris is bronze with a greenish cast and black reticulations. In females the dorsal coloration is the same as in males, except that dark marks tend to be outlined with cream; venter tannish salmon." (W. E. Duellman, field notes, 12 May 1969).Osteocephalus pearsoni(Gaige)Hyla pearsoniGaige, 1929:3 [Holotype.—UMMZ 57548 from the upper Río Beni, below mouth of Río Mapiri, Departamento El Beni, Bolivia; N. E. Pearson collector].Osteocephalus pearsoni—Goin, 1961:13.Justification of Synonymy.—Goin (1961:13) suggested thatHyla pearsoniGaige was anOsteocephalus, but Cochran and Goin (1970:217)consideredpearsonito be aHyla. The presence of exostosed dermal roofing bones, angulate prevomerine dentigerous processes, and the structure of the vocal sacs are characters which place the species inOsteocephalus.Diagnosis.—1) Size moderate, sexual dimorphism evident; maximum observed snout-vent length in males 46.2 mm, in females 54.7 mm; 2) skin on dorsum in males bearing a few, small, scattered non-spinous tubercles; 3) skin on flanks smooth; 4) web extending to base of antepenultimate phalange on inner edge of third finger; 5) dorsum tan with irregular brown blotches; 6) venter cream with fine brown reticulations; 7) lips dark with pale vertical bar below eye; 8) flanks pale tan with round, brown spots; 9) dermal roofing bones of skull slightly exostosed; 10) dermal sphenethmoid absent; 11) nasals narrowly separated medially; 12) anteromedial margin of frontoparietal between mid- and anterior levels of orbit; 13) frontoparietal fontanelle covered; 14) palatine not serrate; 15) parasphenoid lacking odontoids; 16) zygomatic ramus of squamosal extending about one-half distance to maxillary arch; 17) transverse processes of third presacral vertebra approximately equal in width to sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus extensively developed; associated skin forming broad loose fold.Osteocephalus pearsonican be distinguished most readily from other members of the genus by the brown reticulate pattern on the venter, round brown spots on the flanks, and smooth skin on the flanks. Also, it is the least tuberculate species in the genus.Distribution.—Upper Amazon Basin and Amazonian slopes of the Andes in central Perú (1620 m in Río Ucayali drainage) and northern Bolivia (less than 500 m in Río Beni drainage) (Fig. 8). 6 specimens from 3 localities.Remarks.—The specimen from Yaupi, Perú (KU 136312) is a subadult female having a snout-vent length of 39.8 mm. In life the coloration was: "Dorsum light pinkish brown with large rich chocolate brown blotch from eyes to anterior tips of ilia; numerous small chocolate blotches on flanks; dorsal surfaces of thighs and shanks, canthus, and supraorbital region to insertion of forearm chocolate brown; supralabial border and short bar from eye to lip bronze-white; venter bronze-white with numerous tiny chocolate brown flecks [tending to form reticulations on throat and chest]; anterior and posterior surfaces of thighs light olive-brown; iris largely black with gold flecks." (Thomas H. Fritts, field notes, 23March 1970.) On the basis of this one subadult, it seems likely that reticulations on the venter develop with age.Osteocephalus taurinusSteindachnerOsteocephalus taurinusSteindachner, 1862:77 [Holotype.—NHMW 16492 from Barra do Río Negro, Manáus, Territorio do Amazonas, Brasil; Johann Natterer collector].Osteocephalus flavolineatusSteindachner, 1862:80 [Holotype.—NHMW 16495 from Cucuí, Territorio do Amazonas, Brasil; Johann Natterer collector].Trachycephalus (Osteocephalus) taurinusSteindachner, 1867:64.Osteocephalus planicepsCope. 1874:122 [Holotype.—ANSP 11399 from Nauta, Departamento de Loreto, Perú; James Orton collector]. New synonymy.Hyla taurina—Boulenger, 1882:363 [synonymizedOsteocephalus flavolineatusSteindachner, 1862, withO. taurinusSteindachner, 1862].Hyla planiceps—Boulenger, 1882:364.Hyla (Trachycephalus) vilarsiMelin, 1941:40 [Holotype.—NHMG 488 from Taracuá, Río Uaupés, Territorio do Amazonas, Brasil; Douglas Melin collector]. (fideBokermann, 1966:64.)Hyla depressaAndersson, 1945:73 [Holotype.—NHRM 1966 from the Río Pastaza watershed (? Provincia Pastaza), Ecuador; William Clarke-MacIntyre collector]. New synonymy.Justification of Synonymy.—The holotype ofOsteocephalus taurinusis a female having a snout-vent length of 103.9 mm. The diameter of the tympanum is 6.8 mm, 77.3 percent of the diameter of the eye. The skull is strongly exostosed, and the lateral edges of the frontoparietals are elevated so as to form distinct ridges. The skin on the dorsum is smooth. When we examined the type on 5 August 1969, the specimen was soft and badly faded to a pale creamy tan with pale brown transverse bars on the hind limbs and spots on the flanks. Steindachner (1862:79) described the coloration of the type: "In the preserved specimen the dorsum of the entire body, including fore and hind limbs, is a light yellow-brown color, which becomes lighter towards the venter. The belly is whitish, as are the undersides of the arms and legs. The throat is indistinctly marbled with brown. Roundish dark brown flecks are randomly distributed in a considerable number along the side of the body up to the eye; the tympanum is more or less fully surrounded by brown. A few discrete spots, always more or less drawn out in length, on the sides of the body, are also found on the posterior part of the back. The dorsal surfaces of the fore and hind feet are marked with somewhat obliquely arranged brown transverse bands, which are more intensively colored near the margin than in the middle of the band." (free translation from German.)The holotype ofOsteocephalus flavolineatusis a female havinga snout-vent length of 81.8 mm. The diameter of the tympanum is 6.0 mm, 71.4 percent of the diameter of the eye. The skull is strongly exostosed, and the lateral edges of the frontoparietals are elevated so as to form a ridge on each side. The skin on the dorsum is very weakly tuberculate. We examined the type on 9 August 1969 and found it to be in excellent condition. The color pattern is unchanged from that described by Steindachner (1862:81). The dorsum is tan with irregular brown blotches on the back, spots on the flanks, and transverse bars on the limbs. A narrow creamy white, middorsal stripe extends from the snout to the vent. The subocular area is creamy tan, and the venter is tan. Boulenger (1882:363) questionably synonymizedflavolineatuswithtaurinus. We have observed that a middorsal cream stripe occurs in about 10 percent of the specimens oftaurinusand in some specimens ofbuckleyi. This is a common color morph in many species ofEleutherodactylus. In the absence of distinguishing morphological characteristics we can only conclude that the middorsal stripe is a pattern variant and that Boulenger was correct in synonymizingflavolineatuswithtaurinus.The holotype ofOsteocephalus planicepsis a male having a snout-vent length of 58.5 mm. The diameter of the tympanum is 4.9 mm, 77.8 percent of the diameter of the eye. The skull is moderately exostosed, and the lateral edges of the frontoparietals are distinctly elevated. The skin on the dorsum is tuberculate. Cope (1874:122) described the coloration of the type as follows: "Color above uniform dark brown, concealed surfaces on the limbs similar and without any markings. Sides a little varied with the white of the belt. A light border to the upper lip, and lighter line from the orbit to the angle of the mouth; dermal scapular fold pale edged. Femur and tibia with dark crossbands on the exposed surfaces." We examined the holotype on 25 September 1969, and found it to be soft and rubbed. The coloration remains much the same as described by Cope, who provided no means of distinguishingplanicepsfromtaurinus. The coloration and morphometric and structural characters of the type ofplanicepsall fall within the range of variation displayed by series ofO. taurinusfrom the upper Amazon Basin.The type ofHyla vilarsiis a gravid female having a snout-vent length of 62.7 mm. The diameter of the tympanum is 4.8 mm, 73.8 percent of the diameter of the eye. The dorsal roofing bones of the skull are moderately exostosed, and the lateral edges of the frontoparietals are distinctly elevated. The skin on the dorsum is smooth.Melin (1941:42) described the coloration of the holotype as follows: "Above uniform reddish brown; upper eyelids and sides of head darkish brown; below the rostral edge a narrow dark band, continuing as a broader light-edged one through the eye and tympanum towards the base of the forelimb and then farther on continuing along the sides as a line of black spots; sides of upper jaw whitish with traces of dark cross bars (one distinct under the eye); sides of body darkish with black spots and marble, often on a whitish ground; thighs, tibiae, and tarsi each with two broad light-edged, dark cross bars on a brownish ground (less distinct on thighs); sides of thighs finely mottled with brown; beneath whitish with small, sparse spots along jaw, on the chest and sides." We examined the type on 17 February 1969, at which time the specimen was somewhat desiccated, especially the hands and feet. The coloration remains much the same as described by Melin, except that he failed to note the presence of four elongate spots on the back.The status of the namesOsteocephalus planicepsCope andHyla vilarsiMelin was confused by Cochran and Goin (1970:322), who assigned these names to the synonymy ofO. leprieurii. Bokermann (1966:64) placedHyla vilarsiin the synonymy ofOsteocephalus taurinuswithout justification. The type specimens of bothplanicepsandvilarsihave moderately exostosed dermal roofing bones and distinct cranial ridges. The type ofplanicepshas moderately large tubercles on the dorsum, and the type ofvilarsihas spots on the throat, chest, and flanks and longitudinal markings on the back. All of these features are characteristic oftaurinusand not ofleprieurii, which lacks exostosis and cranial ridges and has transverse markings on the back, no spots on the throat, chest, and flanks, and in males has small dorsal tubercles.The type ofHyla depressais a male having a snout-vent length of 69.8 mm. The diameter of the tympanum is 5.2 mm, 77.6 percent of the diameter of the eye. The dorsal roofing bones of the skull are moderately exostosed, and the lateral edges of the frontoparietals are elevated. The skin on the dorsum is tuberculate. The dorsum is dull brown with a broad darker brown longitudinal mark having indistinct lateral edges from the snout to the post-sacral area. A narrow cream middorsal line extends from the snout to the vent. The side of the head is dark brown, palest posteroventral to the orbit. The posterior surfaces of the thighs are dull brown; the flanks are pale brown, and the ventral surfaces are pale creamy tan. Dark brown transverse bars are present on the limbs. When we examined the type on 3 January 1969, it was in excellent condition.Andersson (1945:75) contrasted the type ofHyla depressawithleprieuriiandbuckleyi, but he did not compare his specimen withtaurinus, from which it exhibits no distinguishing features.Osteocephalus taurinusis a widespread and variable species, and it has received several specific names. We are convinced thatOsteocephalus taurinusSteindachner, 1862, is the oldest available name for this large Amazonian species. The following names are junior synonyms:Osteocephalus flavolineatusSteindachner, 1862;Osteocephalus planicepsCope, 1874;Hyla (Trachycephalus) vilarsiMelin, 1941;Hyla depressaAndersson, 1945.Diagnosis.—1) Size large; sexual dimorphism evident; maximum observed snout-vent length in males 84.6 mm, in females 104 mm; 2) skin on dorsum in males bearing many moderately large, spinous tubercles; 3) skin on flanks smooth; 4) web extending to middle of antepenultimate phalange on inner edge of third finger; 5) dorsum brown usually with a large medial dark brown blotch or, less frequently, several dark spots; narrow middorsal yellow line present in some; 6) venter cream or tan with or without small, irregular brown flecks; 7) lips brown with vertical cream bar below eye in some, expanded into pale labial stripe posteriorly in some females; 8) flanks tan or cream with or without small, irregular brown spots; 9) dermal roofing bones of skull exostosed, casqued, and co-ossified (in large adults); 10) dermal sphenethmoid present; 11) nasals juxtaposed medially; 12) anteromedial margin of frontoparietals at mid-level of orbit; 13) frontoparietal fontanelle covered; 14) palatine serrate; 15) parasphenoid bearing odontoids; 16) zygomatic ramus of squamosal usually articulating with maxillary arch; 17) transverse processes of third presacral vertebra approximately equal in width to sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus extensively developed; associated skin forming everted pouch.The moderately rugose dorsum (in males), large size, extensive webbing on the hand, and frontoparietal flanges in adults serve to distinguishtaurinusfrom other members of the genus.Distribution.—The Amazon Basin, the upper Orinoco Basin, and the Guianas. Most localities are below 500 m, but the species ascends the lower Amazonian slopes of the Andes to elevations of about 1000 m (Fig. 11). A record from Caracas, Venezuela, and those from Provincia Carchi and Provincia Esmeraldas, Ecuador, are considered to be erroneous. The latter specimens were includedin a collection sold to the University of Illinois; contained in the collection are many common Amazonian species unknown from the Pacific lowlands. 516 specimens from 151 localities.Fig. 11.Distribution ofOsteocephalus taurinus.Remarks.—This widespread species is highly variable in size and coloration. Striking differences in snout-vent length are evident in series from various parts of the range. The smallest calling males (CAS-SU 12351-6 from Rio Tapirapé, Brasil) have snout-vent lengths of 46.5-60.3 (mean 53.3) mm, whereas the largest (FMNH 140254, KU 92243-6, WCAB 9997, 10001, 10003-4 from Igarapé Marmelo, Brasil) have snout-vent lengths of 71.5-84.6 (mean 77.6) mm. Mean values of snout-vent lengths of males from other localities are: Río Pastaza drainage, Ecuador 73.8 mm, Surinam 67.7 mm, Río Ucayali drainage, Perú 57.6 mm, and Guyana 55.5 mm. Although the difference between the smallest and largest adults is highly significant, populations bridging the gap do exist. Furthermore, the geographic arrangement of small versus large frogs is a confusing mosaic. We have entertained the thought that we have included more than one species intaurinus, but on the basis of preservedspecimens we are unable to detect consistent differences distinguishing two or more taxa.The coloration and pattern oftaurinusare so variable that no one series of statements can describe samples drawn from the entire range of species. We have been unable to determine geographic trends in color pattern; instead the variation within a given sample can encompass the variety known in most other samples. Two minor exceptions do exist. A narrow middorsal light stripe is present in some individuals from throughout the range, but striped specimens are most common in the upper Amazon Basin. The absence of dorsal markings is uncommon in the entire species, but it is most frequent in individuals from the Guianas. A few individuals, such as KU 105230, have scattered white spots on the dorsum.The coloration of four males in life from Lago Agrio, Ecuador (KU 126652-5) was: "Dorsal ground color tan to dark brown with darker brown markings. Flanks creamy tan to yellow with brown or black flecks or mottling. Venter uniform creamy yellow or yellow with brown spots or reticulations. Iris greenish yellow with radiating black streaks and a median, horizontal reddish brown streak." (W. E. Duellman, field notes, 12 May 1969.) A female from Santa Cecilia, Ecuador (KU 123173), was: "Dorsum mottled olive-green and tan. Flanks tan with brown spots. Belly and throat creamy white, becoming tan posteriorly. Edge of upper jaw olive-green." (W. E. Duellman, field notes, 16 June 1968.) Another female from Santa Cecilia (KU 123175), was: "Brown dorsally with cream-colored mottling. Transverse bars on legs darker brown with cream-colored edges. Margin of upper lip creamy yellow. Anterior and posterior surfaces of thighs tan. Flanks white with brown spots. Venter creamy white. Iris greenish bronze with heavy radiating reticulations of black." (W. E. Duellman, field notes, 22 July 1968.)The tendency for females to have a labial stripe posteriorly and the absence of dorsal tubercles in females has resulted in the identification of many such specimens asO. leprieurii.Ontogenetic change in coloration is slight intaurinus. Most juveniles (less than 40 mm in snout-vent length) can be identified readily. There is a tendency for the dorsal markings of juveniles to consist of several small spots. Apparently with growth the spots usually coalesce, forming a large median blotch, but some adults retain the juvenile pattern. Cochran and Goin (1970:251) erroneously identified several juveniles from Colombia asHyla palpebrogranulataAndersson.Osteocephalus verrucigerus(Werner)Hyla verrucigeraWerner, 1901:601 [Holotype.—ZMB 16589 from "Ecuador"; Richard Haensch collector].Hyla riopastazaeAndersson, 1945:72 [Holotype.—NHRM 1960 from Baños, Río Pastaza, Provincia Tungurahua, Ecuador; William Clarke-MacIntyre collector].Hyla orcesiFunkhouser, 1956:78 [Holotype.—CAS-SU 13150 from Río Pacayacu, tributary of Río Cotapino, Provincia Napo, Ecuador; collector unknown].Osteocephalus orcesi—Cochran and Goin, 1970:317.Osteocephalus verrucigerus—Trueb and Duellman, 1970:601 [SynonymizedHyla riopastazaeAndersson, 1945, andHyla orcesiFunkhouser, 1956, withHyla verrucigeraWerner, 1901].Justification of Synonymy.—Trueb and Duellman (1970:605) discussed the assignment of the names in the synonymy ofO. verrucigerus; only a brief resumé is given here.The extant type ofHyla verrucigerais a juvenile male having a snout-vent length of 32.0 mm. The dorsum is smooth except for tubercles on the eyelids; the skin is loose, and the body is soft. The specimen is faded to a pale brown; indistinct dark spots are present on the back, and transverse bars are evident on the limbs.The holotype ofHyla riopastazaeis a gravid female having a snout-vent length of 64.7 mm. The dorsum is smooth. The dorsal ground color is pale brown with indistinct brown transverse bars on the limbs. The throat, chest, and belly are cream with brown spots and mottling.The holotype ofHyla orcesiis an adult male having a snout-vent length of 52.6 mm. The dorsum is heavily tuberculate. The dorsum is dark brown with faint transverse bars on the forearms and feet; the ventral surfaces are creamy brown.Trueb and Duellman (1970) provided conclusive evidence that the types ofH. verrucigera,riopastazae, andorcesiare a juvenile, adult female, and adult male, respectively, of one species, the earliest available name for which isHyla verrucigeraWerner, 1901.Diagnosis.—1) Size moderate, sexual dimorphism evident; maximum observed snout-vent length in males 54.3 mm, in females 65.8 mm; 2) skin on dorsum in males bearing large, keratinized tubercles; 3) skin on flanks smooth; 4) web extending to base of antepenultimate phalange on inner edge of third finger; 5) dorsum uniformly dark brown or black, with tan snout in females; 6) venter creamy white, heavily mottled with black or dark brown, especially in females; 7) lips marked with pale tan labial stripe and suborbital bar; 8) flanks dull reddish brown; 9) dermal roofing bones of skull lacking exostosis; 10) dermal sphenethmoid absent; 11) nasalswidely separated medially; 12) anteromedial margin of frontoparietals at anterior border of orbit; 13) frontoparietal fontanelle covered; 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 approximately equal in width to sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus forming simple, tubular, posterolateral extension; associated skin unmodified.Osteocephalus verrucigeruscan be distinguished from other members of the genus by its uniformly dark dorsum, heavily mottled venter, and large, spinous tubercles on the dorsum in males.Distribution.—Lower Amazonian slopes (500-1840 m) of the Andes and on the western fringe of the Amazon Basin in Ecuador and Perú; one locality (Acevedo) in upper Río Magdalena drainage in Colombia (Fig. 9). 40 specimens from 13 localities.Remarks.—In life the dorsum in males is dull olive-green; the groin, anterior and posterior surfaces of the thighs, inner surfaces of limbs, and upper arms are dark brown. The ventral surfaces of the limbs are pinkish tan; the other ventral surfaces are pale creamy tan with reddish brown spots. The suborbital spot is pale greenish tan, and the iris is deep reddish brown. In females the dorsum is dull olive-brown; the anterior part of the head is tan, and the suborbital spot is yellowish tan. The groin and hidden surfaces of the limbs are dark reddish brown. The ventral surfaces of the limbs are brown; the throat and chest are creamy white, and the belly is reddish tan, both with dark brown mottling.Considerable ontogenetic change occurs in coloration. Juveniles are pale above with a dark median dorsal blotch and dark transverse bars on the limbs. The venter is white. The change consists principally of an increase in dark pigment and subsequent obliteration of the juvenile pattern.Tadpoles of this species have moderately long tails with low fins, robust bodies, two rows of labial papillae with median part of the upper lip bare, and two upper and five lower rows of teeth. Trueb and Duellman (1970) described the eggs, tadpoles, mating call, and variation in the adults.
Accounts of Species
Osteocephalus buckleyi(Boulenger)
Hyla buckleyiBoulenger, 1882:362 [Syntypes.—BMNH 1947.2.13.36-39 from Sarayacu, Provincia Pastaza, Ecuador; BMNH 1947.2.13.40-41, 1947.2.13. 43-45 from Canelos, Provincia Pastaza, Ecuador; BMNH 1947.2.13.46 from "Paitanga" (= Pallatanga), Provincia Chimborazo, Ecuador (in error); Mr. Buckley collector; BMNH 1947.2.13.44 here designated as lectotype].
Hyla festaePeracca, 1904:39 [Holotype.—MIZS 2950 from "Valle de Santiago" (= lower Río Zamora), Provincia Morona-Santiago, Ecuador; Enrico Festa collector]. New synonymy.
Osteocephalus buckleyi—Goin, 1961:13.
Hyla carriCochran and Goin, 1970:211 [Holotype.—FMNH 69702 from Acevedo, Río Suaza, Departamento Huila, Colombia; Philip collector]. New synonymy.
Hyla cabreraiCochran and Goin, 1970:215 [Holotype.—USNM 152759 from Caño Guacayá, tributary of lower Río Apoporis, Comisaria Amazonas, Colombia; Isadore Cabrera collector]. New synonymy.
Justification of Synonymy.—Boulenger (1882:362) listed 11 specimens in his description ofHyla buckleyi. We have examined all of these and conclude that one (BMNH 1947.2.13.42) isO. leprieurii. Cochran and Goin (1970:213) restricted the type locality to Canelos,ProvinciaPastaza, Ecuador; we here select BMNH 1947.2.13.44 from that locality as the lectotype. This specimen is a male having a snout-vent length of 37.9 mm; the diameter of the tympanumis 3.5 mm, 81.4 percent of the diameter of the eye. The type series, exclusive of BMNH 1947.2.13.42 (=O. leprieurii) consists of six males having snout-vent lengths of 37.9-44.6 (mean 40.4) mm, and four females having snout-vent lengths of 50.0-53.9 (mean 51.5) mm. The dorsum in the males bears a mixture of large and small tubercles, whereas the dorsum in females is nearly smooth. The skin on the flanks, especially the axilla, is areolate. The coloration consists of a creamy tan ground color with irregular reddish brown markings on the back and broad transverse bars on the limbs. The dorsal markings are narrowly bordered by creamy white; those on the back consist of an interorbital bar and a pair of longitudinal marks beginning in the scapular region and usually diverging posteriorly in the sacral region or converging into a broad median blotch. One specimen has a middorsal creamy white stripe from the tip of the snout to the vent. In all of the types large dark brown spots are present on the flanks and posterior surfaces of the thighs. The ventral surfaces are pale creamy tan with or without diffuse brown spots on the throat and chest.
The holotype ofHyla festaeis a female having a snout-vent length of 75.0 mm; the diameter of the tympanum is 3.9 mm, 57.4 percent of the diameter of the eye. The skin is smooth on the dorsum and areolate on the anterior part of the flanks. The dorsum is pale brown with a large median longitudinal dark brown blotch on the back and broad transverse bars, narrowly outlined by cream, on the limbs. Dark brown spots are present on the flanks; the posterior surfaces of the thighs are dark brown. The throat and belly are grayish white with irregular dark brown spots.
The holotype ofHyla carriis a female having a snout-vent length of 66.1 mm; the diameter of the tympanum is 4.7 mm, 81.0 percent of the diameter of the eye. The skin on the dorsum is smooth with scattered small tubercles and areolate on the anterior part of the flanks. The dorsum is tan with irregular dark brown blotches on the back and transverse bars on the limbs; all dorsal markings are narrowly outlined by creamy white. Dark brown spots are present on the flanks; the venter and posterior surfaces of the thighs are tan without dark spots.
The holotype ofHyla cabreraiis a female having a snout-vent length of 52.7 mm; the diameter of the tympanum is 4.0 mm, 76.9 percent of the diameter of the eye. The skin on the dorsum is weakly tuberculate and that on the anterior part of the flanks is areolate. The dorsum is creamy tan with dark brown markings (interorbital bar, reticulations on occiput, three longitudinal streaks on back,and broad transverse bars on limbs). Irregular dark brown spots are present on the flanks. The venter is pinkish tan with small reddish brown spots on the throat and darker brown spots on the chest and belly.
In their description ofHyla cabrerai, Cochran and Goin (1970:217) stated: "This species, together withbuckleyiandpearsonicertainly make a closely knit groupߪ. Bothbuckleyiandcabreraihave long hind legs, with the extended heel reaching to the tip of the snout, while inpearsonithe extended heel reaches only to the eye.H. buckleyihas the belly dusky, while it is heavily spotted incabreraiand is reticulated inpearsoni.H. cabreraiseems to have the heaviest hands with the most webbing between the fingers; the other two species have the webbing reduced between the fingers." The description ofHyla cabreraiwas based on three specimens. We have examined the holotype and one paratype (WCAB 13284 from Territorio do Amapá, Brasil). Another paratype in the private collection of C. J. Goin from Caño Tuí, between Mitú and Raudal de Yurupari, Comisaria de Vaupés, Colombia, was not examined.
Cochran and Goin (1970:211) based their description ofHyla carrion one gravid female and stated: "A largeHylawith the vomerine teeth in two ʌʌ shaped patches between the somewhat squarish choanae; reduced webs between the fingers; and a pattern of dorsal dark blotches bordered by light margins. The species is not similar to any other species known in Colombia. It is perhaps most closely related toHyla claresignataof Brazil, from which it can be differentiated by its more heavily spotted dorsum, larger tympanum, and lack of dark anal spots."
Except for the inclusion of the name in checklists,Hyla festaehas not been mentioned in the literature since the original description.
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 ofHyla 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 ofHyla festaeis boldly spotted; it isdistinctly spotted incabrerai, uniform tan incarri, and tan, flecked, or spotted in the type series ofbuckleyi. The ventral coloration in series of specimens from Amazonian Ecuador encompasses that observed in all of the types, except that offestae, 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 offestae, and one paratype ofcabreraithe webbing extends to the middle of the penultimate phalanges of the third and fourth fingers. In the holotype ofcabreraithe 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 ofOsteocephalus. 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 ofHyla carri, Cochran and Goin (1970:211) misrepresented the nature of the dentigerous processes of the prevomers, which are angular, not ʌ-shaped. Their suggestion that the ColombianHyla carriis related toHyla claresignatain 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 ofHyla festaeresembles that ofOsteocephalus verrucigerus, but the type differs fromverrucigerusby having areolate skin on the flanks and distinct dark markings on the dorsum. Inverrucigerusthe 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 Perú suggest strongly that the types are representative of one taxon, the oldest name for which isHyla buckleyiBoulenger, 1882. Consequently, we placeHyla festaePeracca, 1904,Hyla carriCochran and Goin, 1970, andHyla cabreraiCochran and Goin, 1970, as junior synonyms ofHyla buckleyiBoulenger, 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 buckleyican 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 Amapá in northeastern Brasil; the upper Amazon Basin from southern Colombia to east-central Bolivia; one locality (Acevedo) in upper Río Magdalena drainage in Colombia (Fig. 8). All localities are at elevations of less than 700 m. Recordsfor 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.
Fig. 8.Distribution ofOsteocephalus buckleyi(circles) andO. pearsoni(triangles).
Fig. 8.Distribution ofOsteocephalus buckleyi(circles) andO. pearsoni(triangles).
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(Duméril and Bibron)
Hyla leprieuriiDuméril and Bibron, 1841:553 [Holotype.—MNHN 4629 from "Cayenne"; Mons. Leprieur collector].
Hypsiboas leprieurii—Cope, 1867:200.
Hyla leprieurii brittiMelin, 1941:42 [Holotype.—NHMG 489 from the Rio Uaupés, 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 ofHyla leprieuriiis 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. Duméril and Bibron (1841:554) described the coloration, as follows: "The loreal regionisblack. 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 ofHyla leprieurii brittiis 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 ofmetalliclustre; 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 onthe 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 ofHyla leprieurii britti"ߪ resembles a good dealH. leprieuriiDum. & 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 ofleprieuriiߪ." The pattern of narrow transverse bars on the backs of the holotypes ofH. leprieuriiandH. brittiis a condition shared only by these two nominal taxa that are placed inOsteocephalus. Melin noted thatbrittidiffered fromleprieuriiin 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 inleprieurii—♀; 0.655 inbritti—♂) 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 forleprieurii; 45 forbritti).
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 considerHyla leprieuriiDuméril and Bibron, 1841, to be a monotypic species withHyla leprieurii brittiMelin, 1941, as a junior synonym.
In their account ofOsteocephalus leprieurii, Cochran and Goin (1970:323) stated: "The specimen described and illustrated (MCZ 28042) has been directly compared with the types ofleprieurii,planiceps, andvilarsiby 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 ofplanicepsandvilarsiand these, likewise, are considered conspecific." With this justification Cochran and Goin (1970:322) includedOsteocephalus planicepsCope, 1874, andHyla vilarsiMelin, 1941, in the synonymy ofOsteocephalus leprieurii.
We do not concur with Cochran and Goin's synonymy and contend thatplanicepsandvilarsiare synonyms ofOsteocephalus taurinus; we give our reasons in the account of that species. We have examined the specimens listed asO. leprieuriiby Cochran andGoin; several of them, including CNHM (= FMNH) 69716, aretaurinus. 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 includeHyla leprieurii brittiin their synonymy ofOsteocephalus leprieuriibut did discuss the name in their account ofOsteocephalus orcesi(=O. verrucigerus), as follows (1970:319): "When we first examined one of the specimens we felt sure that we had Melin'sHyla brittiat hand, but on direct comparison with the type ofbrittithe two proved to be different. After studying the type oforcesi(SUNHM 13150) we have no doubt that the specimens at hand areorcesiand thatbrittiis 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 leprieuriidiffers from all other members of the genus by having transverse dark bars on the back. Two other hylids (Hyla lanciformisandmultifasciata) in the Amazon Basin have transverse dark marks on the dorsum. Both of these differ fromleprieuriiby having pointed snouts, much longer hind limbs, and smooth skin dorsally.
Distribution.—The periphery of the Amazon Basin, in the Guianas and the upper part of the basin in southern Colombia, Ecuador, Perú, and extreme western Brasil (Fig. 9). Most localities are at elevations of less than 500 m, but the species ascends the lowerAndean slopes to elevations of 1100 m. 265 specimens from 31 localities.
Fig. 9.Distribution ofOsteocephalus leprieurii(circles) andO. verrucigerus(triangles).
Fig. 9.Distribution ofOsteocephalus leprieurii(circles) andO. verrucigerus(triangles).
Remarks.—Most adults ofleprieuriihave distinct transverse markings on the back; these are variable in width, extent, and arrangement. In some specimens, such as USNM 166557, some of the transverse bars are fragmented into spots; in a few specimens the dorsal pattern consists solely of small dark spots arranged in transverse rows. Such specimens have a dorsal pattern resembling that of sometaurinus. The transverse nature of the dorsal markings is further modified in some specimens, such as USNM 166555, in which the dark bars are fragmented and oblique.
Extreme ontogenetic change in color pattern is exhibited by this species (Fig. 10). Juveniles having snout-vent lengths of less than 28 mm have an olive-brown dorsum with a pale cream stripe across the head and broad, cream, dorsolateral stripes; transverse dark bars are absent on the body and limbs. Individuals having snout-vent lengths of 30-35 mm have dark brown transverse bars onthe back and limbs but still retain the light dorsolateral stripes, whereas the stripes are lost in larger individuals.
Fig. 10.Ontogenetic change in color pattern inOsteocephalus leprieurii:a. KU 126644; b. KU 126640; c. KU 126625. ×2.
Fig. 10.Ontogenetic change in color pattern inOsteocephalus leprieurii:a. KU 126644; b. KU 126640; c. KU 126625. ×2.
Coloration in life of specimens from Lago Agrio, Ecuador: "In males the dorsal ground color varies from dark brown to ochre-tan; dorsal markings uniformly dark brown. Most specimens have dark brown and cream anal stripes; labial area cream-colored. Flanks vary from tan to white. Ventral coloration varies from salmon to tan to white. The iris is bronze with a greenish cast and black reticulations. In females the dorsal coloration is the same as in males, except that dark marks tend to be outlined with cream; venter tannish salmon." (W. E. Duellman, field notes, 12 May 1969).
Osteocephalus pearsoni(Gaige)
Hyla pearsoniGaige, 1929:3 [Holotype.—UMMZ 57548 from the upper Río Beni, below mouth of Río Mapiri, Departamento El Beni, Bolivia; N. E. Pearson collector].
Osteocephalus pearsoni—Goin, 1961:13.
Justification of Synonymy.—Goin (1961:13) suggested thatHyla pearsoniGaige was anOsteocephalus, but Cochran and Goin (1970:217)consideredpearsonito be aHyla. The presence of exostosed dermal roofing bones, angulate prevomerine dentigerous processes, and the structure of the vocal sacs are characters which place the species inOsteocephalus.
Diagnosis.—1) Size moderate, sexual dimorphism evident; maximum observed snout-vent length in males 46.2 mm, in females 54.7 mm; 2) skin on dorsum in males bearing a few, small, scattered non-spinous tubercles; 3) skin on flanks smooth; 4) web extending to base of antepenultimate phalange on inner edge of third finger; 5) dorsum tan with irregular brown blotches; 6) venter cream with fine brown reticulations; 7) lips dark with pale vertical bar below eye; 8) flanks pale tan with round, brown spots; 9) dermal roofing bones of skull slightly exostosed; 10) dermal sphenethmoid absent; 11) nasals narrowly separated medially; 12) anteromedial margin of frontoparietal between mid- and anterior levels of orbit; 13) frontoparietal fontanelle covered; 14) palatine not serrate; 15) parasphenoid lacking odontoids; 16) zygomatic ramus of squamosal extending about one-half distance to maxillary arch; 17) transverse processes of third presacral vertebra approximately equal in width to sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus extensively developed; associated skin forming broad loose fold.
Osteocephalus pearsonican be distinguished most readily from other members of the genus by the brown reticulate pattern on the venter, round brown spots on the flanks, and smooth skin on the flanks. Also, it is the least tuberculate species in the genus.
Distribution.—Upper Amazon Basin and Amazonian slopes of the Andes in central Perú (1620 m in Río Ucayali drainage) and northern Bolivia (less than 500 m in Río Beni drainage) (Fig. 8). 6 specimens from 3 localities.
Remarks.—The specimen from Yaupi, Perú (KU 136312) is a subadult female having a snout-vent length of 39.8 mm. In life the coloration was: "Dorsum light pinkish brown with large rich chocolate brown blotch from eyes to anterior tips of ilia; numerous small chocolate blotches on flanks; dorsal surfaces of thighs and shanks, canthus, and supraorbital region to insertion of forearm chocolate brown; supralabial border and short bar from eye to lip bronze-white; venter bronze-white with numerous tiny chocolate brown flecks [tending to form reticulations on throat and chest]; anterior and posterior surfaces of thighs light olive-brown; iris largely black with gold flecks." (Thomas H. Fritts, field notes, 23March 1970.) On the basis of this one subadult, it seems likely that reticulations on the venter develop with age.
Osteocephalus taurinusSteindachner
Osteocephalus taurinusSteindachner, 1862:77 [Holotype.—NHMW 16492 from Barra do Río Negro, Manáus, Territorio do Amazonas, Brasil; Johann Natterer collector].
Osteocephalus flavolineatusSteindachner, 1862:80 [Holotype.—NHMW 16495 from Cucuí, Territorio do Amazonas, Brasil; Johann Natterer collector].
Trachycephalus (Osteocephalus) taurinusSteindachner, 1867:64.
Osteocephalus planicepsCope. 1874:122 [Holotype.—ANSP 11399 from Nauta, Departamento de Loreto, Perú; James Orton collector]. New synonymy.
Hyla taurina—Boulenger, 1882:363 [synonymizedOsteocephalus flavolineatusSteindachner, 1862, withO. taurinusSteindachner, 1862].
Hyla planiceps—Boulenger, 1882:364.
Hyla (Trachycephalus) vilarsiMelin, 1941:40 [Holotype.—NHMG 488 from Taracuá, Río Uaupés, Territorio do Amazonas, Brasil; Douglas Melin collector]. (fideBokermann, 1966:64.)
Hyla depressaAndersson, 1945:73 [Holotype.—NHRM 1966 from the Río Pastaza watershed (? Provincia Pastaza), Ecuador; William Clarke-MacIntyre collector]. New synonymy.
Justification of Synonymy.—The holotype ofOsteocephalus taurinusis a female having a snout-vent length of 103.9 mm. The diameter of the tympanum is 6.8 mm, 77.3 percent of the diameter of the eye. The skull is strongly exostosed, and the lateral edges of the frontoparietals are elevated so as to form distinct ridges. The skin on the dorsum is smooth. When we examined the type on 5 August 1969, the specimen was soft and badly faded to a pale creamy tan with pale brown transverse bars on the hind limbs and spots on the flanks. Steindachner (1862:79) described the coloration of the type: "In the preserved specimen the dorsum of the entire body, including fore and hind limbs, is a light yellow-brown color, which becomes lighter towards the venter. The belly is whitish, as are the undersides of the arms and legs. The throat is indistinctly marbled with brown. Roundish dark brown flecks are randomly distributed in a considerable number along the side of the body up to the eye; the tympanum is more or less fully surrounded by brown. A few discrete spots, always more or less drawn out in length, on the sides of the body, are also found on the posterior part of the back. The dorsal surfaces of the fore and hind feet are marked with somewhat obliquely arranged brown transverse bands, which are more intensively colored near the margin than in the middle of the band." (free translation from German.)
The holotype ofOsteocephalus flavolineatusis a female havinga snout-vent length of 81.8 mm. The diameter of the tympanum is 6.0 mm, 71.4 percent of the diameter of the eye. The skull is strongly exostosed, and the lateral edges of the frontoparietals are elevated so as to form a ridge on each side. The skin on the dorsum is very weakly tuberculate. We examined the type on 9 August 1969 and found it to be in excellent condition. The color pattern is unchanged from that described by Steindachner (1862:81). The dorsum is tan with irregular brown blotches on the back, spots on the flanks, and transverse bars on the limbs. A narrow creamy white, middorsal stripe extends from the snout to the vent. The subocular area is creamy tan, and the venter is tan. Boulenger (1882:363) questionably synonymizedflavolineatuswithtaurinus. We have observed that a middorsal cream stripe occurs in about 10 percent of the specimens oftaurinusand in some specimens ofbuckleyi. This is a common color morph in many species ofEleutherodactylus. In the absence of distinguishing morphological characteristics we can only conclude that the middorsal stripe is a pattern variant and that Boulenger was correct in synonymizingflavolineatuswithtaurinus.
The holotype ofOsteocephalus planicepsis a male having a snout-vent length of 58.5 mm. The diameter of the tympanum is 4.9 mm, 77.8 percent of the diameter of the eye. The skull is moderately exostosed, and the lateral edges of the frontoparietals are distinctly elevated. The skin on the dorsum is tuberculate. Cope (1874:122) described the coloration of the type as follows: "Color above uniform dark brown, concealed surfaces on the limbs similar and without any markings. Sides a little varied with the white of the belt. A light border to the upper lip, and lighter line from the orbit to the angle of the mouth; dermal scapular fold pale edged. Femur and tibia with dark crossbands on the exposed surfaces." We examined the holotype on 25 September 1969, and found it to be soft and rubbed. The coloration remains much the same as described by Cope, who provided no means of distinguishingplanicepsfromtaurinus. The coloration and morphometric and structural characters of the type ofplanicepsall fall within the range of variation displayed by series ofO. taurinusfrom the upper Amazon Basin.
The type ofHyla vilarsiis a gravid female having a snout-vent length of 62.7 mm. The diameter of the tympanum is 4.8 mm, 73.8 percent of the diameter of the eye. The dorsal roofing bones of the skull are moderately exostosed, and the lateral edges of the frontoparietals are distinctly elevated. The skin on the dorsum is smooth.Melin (1941:42) described the coloration of the holotype as follows: "Above uniform reddish brown; upper eyelids and sides of head darkish brown; below the rostral edge a narrow dark band, continuing as a broader light-edged one through the eye and tympanum towards the base of the forelimb and then farther on continuing along the sides as a line of black spots; sides of upper jaw whitish with traces of dark cross bars (one distinct under the eye); sides of body darkish with black spots and marble, often on a whitish ground; thighs, tibiae, and tarsi each with two broad light-edged, dark cross bars on a brownish ground (less distinct on thighs); sides of thighs finely mottled with brown; beneath whitish with small, sparse spots along jaw, on the chest and sides." We examined the type on 17 February 1969, at which time the specimen was somewhat desiccated, especially the hands and feet. The coloration remains much the same as described by Melin, except that he failed to note the presence of four elongate spots on the back.
The status of the namesOsteocephalus planicepsCope andHyla vilarsiMelin was confused by Cochran and Goin (1970:322), who assigned these names to the synonymy ofO. leprieurii. Bokermann (1966:64) placedHyla vilarsiin the synonymy ofOsteocephalus taurinuswithout justification. The type specimens of bothplanicepsandvilarsihave moderately exostosed dermal roofing bones and distinct cranial ridges. The type ofplanicepshas moderately large tubercles on the dorsum, and the type ofvilarsihas spots on the throat, chest, and flanks and longitudinal markings on the back. All of these features are characteristic oftaurinusand not ofleprieurii, which lacks exostosis and cranial ridges and has transverse markings on the back, no spots on the throat, chest, and flanks, and in males has small dorsal tubercles.
The type ofHyla depressais a male having a snout-vent length of 69.8 mm. The diameter of the tympanum is 5.2 mm, 77.6 percent of the diameter of the eye. The dorsal roofing bones of the skull are moderately exostosed, and the lateral edges of the frontoparietals are elevated. The skin on the dorsum is tuberculate. The dorsum is dull brown with a broad darker brown longitudinal mark having indistinct lateral edges from the snout to the post-sacral area. A narrow cream middorsal line extends from the snout to the vent. The side of the head is dark brown, palest posteroventral to the orbit. The posterior surfaces of the thighs are dull brown; the flanks are pale brown, and the ventral surfaces are pale creamy tan. Dark brown transverse bars are present on the limbs. When we examined the type on 3 January 1969, it was in excellent condition.Andersson (1945:75) contrasted the type ofHyla depressawithleprieuriiandbuckleyi, but he did not compare his specimen withtaurinus, from which it exhibits no distinguishing features.
Osteocephalus taurinusis a widespread and variable species, and it has received several specific names. We are convinced thatOsteocephalus taurinusSteindachner, 1862, is the oldest available name for this large Amazonian species. The following names are junior synonyms:Osteocephalus flavolineatusSteindachner, 1862;Osteocephalus planicepsCope, 1874;Hyla (Trachycephalus) vilarsiMelin, 1941;Hyla depressaAndersson, 1945.
Diagnosis.—1) Size large; sexual dimorphism evident; maximum observed snout-vent length in males 84.6 mm, in females 104 mm; 2) skin on dorsum in males bearing many moderately large, spinous tubercles; 3) skin on flanks smooth; 4) web extending to middle of antepenultimate phalange on inner edge of third finger; 5) dorsum brown usually with a large medial dark brown blotch or, less frequently, several dark spots; narrow middorsal yellow line present in some; 6) venter cream or tan with or without small, irregular brown flecks; 7) lips brown with vertical cream bar below eye in some, expanded into pale labial stripe posteriorly in some females; 8) flanks tan or cream with or without small, irregular brown spots; 9) dermal roofing bones of skull exostosed, casqued, and co-ossified (in large adults); 10) dermal sphenethmoid present; 11) nasals juxtaposed medially; 12) anteromedial margin of frontoparietals at mid-level of orbit; 13) frontoparietal fontanelle covered; 14) palatine serrate; 15) parasphenoid bearing odontoids; 16) zygomatic ramus of squamosal usually articulating with maxillary arch; 17) transverse processes of third presacral vertebra approximately equal in width to sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus extensively developed; associated skin forming everted pouch.
The moderately rugose dorsum (in males), large size, extensive webbing on the hand, and frontoparietal flanges in adults serve to distinguishtaurinusfrom other members of the genus.
Distribution.—The Amazon Basin, the upper Orinoco Basin, and the Guianas. Most localities are below 500 m, but the species ascends the lower Amazonian slopes of the Andes to elevations of about 1000 m (Fig. 11). A record from Caracas, Venezuela, and those from Provincia Carchi and Provincia Esmeraldas, Ecuador, are considered to be erroneous. The latter specimens were includedin a collection sold to the University of Illinois; contained in the collection are many common Amazonian species unknown from the Pacific lowlands. 516 specimens from 151 localities.
Fig. 11.Distribution ofOsteocephalus taurinus.
Fig. 11.Distribution ofOsteocephalus taurinus.
Remarks.—This widespread species is highly variable in size and coloration. Striking differences in snout-vent length are evident in series from various parts of the range. The smallest calling males (CAS-SU 12351-6 from Rio Tapirapé, Brasil) have snout-vent lengths of 46.5-60.3 (mean 53.3) mm, whereas the largest (FMNH 140254, KU 92243-6, WCAB 9997, 10001, 10003-4 from Igarapé Marmelo, Brasil) have snout-vent lengths of 71.5-84.6 (mean 77.6) mm. Mean values of snout-vent lengths of males from other localities are: Río Pastaza drainage, Ecuador 73.8 mm, Surinam 67.7 mm, Río Ucayali drainage, Perú 57.6 mm, and Guyana 55.5 mm. Although the difference between the smallest and largest adults is highly significant, populations bridging the gap do exist. Furthermore, the geographic arrangement of small versus large frogs is a confusing mosaic. We have entertained the thought that we have included more than one species intaurinus, but on the basis of preservedspecimens we are unable to detect consistent differences distinguishing two or more taxa.
The coloration and pattern oftaurinusare so variable that no one series of statements can describe samples drawn from the entire range of species. We have been unable to determine geographic trends in color pattern; instead the variation within a given sample can encompass the variety known in most other samples. Two minor exceptions do exist. A narrow middorsal light stripe is present in some individuals from throughout the range, but striped specimens are most common in the upper Amazon Basin. The absence of dorsal markings is uncommon in the entire species, but it is most frequent in individuals from the Guianas. A few individuals, such as KU 105230, have scattered white spots on the dorsum.
The coloration of four males in life from Lago Agrio, Ecuador (KU 126652-5) was: "Dorsal ground color tan to dark brown with darker brown markings. Flanks creamy tan to yellow with brown or black flecks or mottling. Venter uniform creamy yellow or yellow with brown spots or reticulations. Iris greenish yellow with radiating black streaks and a median, horizontal reddish brown streak." (W. E. Duellman, field notes, 12 May 1969.) A female from Santa Cecilia, Ecuador (KU 123173), was: "Dorsum mottled olive-green and tan. Flanks tan with brown spots. Belly and throat creamy white, becoming tan posteriorly. Edge of upper jaw olive-green." (W. E. Duellman, field notes, 16 June 1968.) Another female from Santa Cecilia (KU 123175), was: "Brown dorsally with cream-colored mottling. Transverse bars on legs darker brown with cream-colored edges. Margin of upper lip creamy yellow. Anterior and posterior surfaces of thighs tan. Flanks white with brown spots. Venter creamy white. Iris greenish bronze with heavy radiating reticulations of black." (W. E. Duellman, field notes, 22 July 1968.)
The tendency for females to have a labial stripe posteriorly and the absence of dorsal tubercles in females has resulted in the identification of many such specimens asO. leprieurii.
Ontogenetic change in coloration is slight intaurinus. Most juveniles (less than 40 mm in snout-vent length) can be identified readily. There is a tendency for the dorsal markings of juveniles to consist of several small spots. Apparently with growth the spots usually coalesce, forming a large median blotch, but some adults retain the juvenile pattern. Cochran and Goin (1970:251) erroneously identified several juveniles from Colombia asHyla palpebrogranulataAndersson.
Osteocephalus verrucigerus(Werner)
Hyla verrucigeraWerner, 1901:601 [Holotype.—ZMB 16589 from "Ecuador"; Richard Haensch collector].
Hyla riopastazaeAndersson, 1945:72 [Holotype.—NHRM 1960 from Baños, Río Pastaza, Provincia Tungurahua, Ecuador; William Clarke-MacIntyre collector].
Hyla orcesiFunkhouser, 1956:78 [Holotype.—CAS-SU 13150 from Río Pacayacu, tributary of Río Cotapino, Provincia Napo, Ecuador; collector unknown].
Osteocephalus orcesi—Cochran and Goin, 1970:317.
Osteocephalus verrucigerus—Trueb and Duellman, 1970:601 [SynonymizedHyla riopastazaeAndersson, 1945, andHyla orcesiFunkhouser, 1956, withHyla verrucigeraWerner, 1901].
Justification of Synonymy.—Trueb and Duellman (1970:605) discussed the assignment of the names in the synonymy ofO. verrucigerus; only a brief resumé is given here.
The extant type ofHyla verrucigerais a juvenile male having a snout-vent length of 32.0 mm. The dorsum is smooth except for tubercles on the eyelids; the skin is loose, and the body is soft. The specimen is faded to a pale brown; indistinct dark spots are present on the back, and transverse bars are evident on the limbs.
The holotype ofHyla riopastazaeis a gravid female having a snout-vent length of 64.7 mm. The dorsum is smooth. The dorsal ground color is pale brown with indistinct brown transverse bars on the limbs. The throat, chest, and belly are cream with brown spots and mottling.
The holotype ofHyla orcesiis an adult male having a snout-vent length of 52.6 mm. The dorsum is heavily tuberculate. The dorsum is dark brown with faint transverse bars on the forearms and feet; the ventral surfaces are creamy brown.
Trueb and Duellman (1970) provided conclusive evidence that the types ofH. verrucigera,riopastazae, andorcesiare a juvenile, adult female, and adult male, respectively, of one species, the earliest available name for which isHyla verrucigeraWerner, 1901.
Diagnosis.—1) Size moderate, sexual dimorphism evident; maximum observed snout-vent length in males 54.3 mm, in females 65.8 mm; 2) skin on dorsum in males bearing large, keratinized tubercles; 3) skin on flanks smooth; 4) web extending to base of antepenultimate phalange on inner edge of third finger; 5) dorsum uniformly dark brown or black, with tan snout in females; 6) venter creamy white, heavily mottled with black or dark brown, especially in females; 7) lips marked with pale tan labial stripe and suborbital bar; 8) flanks dull reddish brown; 9) dermal roofing bones of skull lacking exostosis; 10) dermal sphenethmoid absent; 11) nasalswidely separated medially; 12) anteromedial margin of frontoparietals at anterior border of orbit; 13) frontoparietal fontanelle covered; 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 approximately equal in width to sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus forming simple, tubular, posterolateral extension; associated skin unmodified.
Osteocephalus verrucigeruscan be distinguished from other members of the genus by its uniformly dark dorsum, heavily mottled venter, and large, spinous tubercles on the dorsum in males.
Distribution.—Lower Amazonian slopes (500-1840 m) of the Andes and on the western fringe of the Amazon Basin in Ecuador and Perú; one locality (Acevedo) in upper Río Magdalena drainage in Colombia (Fig. 9). 40 specimens from 13 localities.
Remarks.—In life the dorsum in males is dull olive-green; the groin, anterior and posterior surfaces of the thighs, inner surfaces of limbs, and upper arms are dark brown. The ventral surfaces of the limbs are pinkish tan; the other ventral surfaces are pale creamy tan with reddish brown spots. The suborbital spot is pale greenish tan, and the iris is deep reddish brown. In females the dorsum is dull olive-brown; the anterior part of the head is tan, and the suborbital spot is yellowish tan. The groin and hidden surfaces of the limbs are dark reddish brown. The ventral surfaces of the limbs are brown; the throat and chest are creamy white, and the belly is reddish tan, both with dark brown mottling.
Considerable ontogenetic change occurs in coloration. Juveniles are pale above with a dark median dorsal blotch and dark transverse bars on the limbs. The venter is white. The change consists principally of an increase in dark pigment and subsequent obliteration of the juvenile pattern.
Tadpoles of this species have moderately long tails with low fins, robust bodies, two rows of labial papillae with median part of the upper lip bare, and two upper and five lower rows of teeth. Trueb and Duellman (1970) described the eggs, tadpoles, mating call, and variation in the adults.