Part 3

Vertebral Column.--The cervical cotyles are uniformly widely displaced. The neural arches are low and non-imbricate. The transverse processes of the third presacral vertebrae are approximately equal in width to the sacral diapophyses in all species, except buckleyi, in which the processes of the third presacral are slightly narrower than the diapophyses. Osteocephalus buckleyi is further distinguished by the presence of narrow transverse processes on presacrals five through eight (Fig. 7b); males have narrower processes than do females. The processes are moderately wide but subequal in width in pearsoni, taurinus, and verrucigerus, whereas they are nearly equivalent in width to one another and to the sacral diapophyses in leprieurii (Fig. 7a). The sacral diapophyses of all species are moderately dilated and posterolaterally inclined. The coccyx bears a distinct dorsal ridge and has a bicondylar articulation with the sacrum.

Pectoral Girdle.--The pectoral girdles are fully arciferal and bear small, cartilaginous omosterna and moderately large cartilaginous sterna. The coracoids are robust, and the clavicles are strongly arched. Procoracoid cartilage seems to be absent. The scapulae are large, longer than the clavicles, and bicapitate proximally. The suprascapulae are moderately large and well ossified in leprieurii and taurinus. The suprascapula of verrucigerus is poorly ossified, and that of buckleyi is not ossified.

Pelvic Girdle.--The ilia of buckleyi, taurinus, and verrucigerus lack any indication of a crest on the shaft, whereas leprieurii has a low crest. The dorsal acetabular expansion of the ilia is moderately low in taurinus and verrucigerus, but distinctly lower in buckleyi and leprieurii. The ilia of all species bear low dorsal protuberances. The ischia of leprieurii, taurinus, and verrucigerus are moderately expanded; that of buckleyi is somewhat less expanded dorsally. The pubis of leprieurii, taurinus, and verrucigerus are calcified, whereas that of buckleyi remains cartilaginous.

Throat Musculature and Vocal Sac Structure.--Tyler (1971) described the throat myology of Osteocephalus. The genus is characterized by a moderate-sized araphic submentalis muscle and an undifferentiated intermandibularis having an elongate median aponeurosis. The intermandibularis and submentalis are completely independent in buckleyi, whereas in the other species there is a small attachment between these muscles.

According to Tyler (pers. com.), Osteocephalus has three distinctive types of vocal sac structure which result from differences in the development of the interhyoideus muscle and the overlying skin. In leprieurii and verrucigerus the supramandibular portions of the interhyoideus form a simple tubular, posterolateral extension; there is no modification of the associated skin. Osteocephalus buckleyi and pearsoni have more extensive development of the supramandibular portions of the interhyoideus; furthermore, the associated skin forms a broad, loose fold extending from the ventromedial surface of the throat dorsolaterally to the base of the supratympanic fold. Like buckleyi and pearsoni, the supramandibular portion of the interhyoideus is much expanded in taurinus. The vocal sac structure of taurinus differs from that of other members of the genus in that the skin of taurinus forms an everted pouch, which dangles loosely beneath the supratympanic fold.

KEY TO THE SPECIES OF Osteocephalus

1. Inner edge of third finger webbed to mid-length of antepenultimate phalange; dorsum brown with dark brown spots or median blotch; skull in adults casqued and co-ossified with prominent supraorbital flanges O. taurinus

Inner edge of third finger webbed to base of antepenultimate phalange; dorsum plain or marked with dark blotches or transverse bars; skull in adults smooth or slightly exostosed, lacking supraorbital flanges 2

2. Skin on flanks areolate; dorsum in males bearing a mixture of large and small non-spinous tubercles; lips distinctly barred O. buckleyi

Skin on flanks smooth; dorsum in males bearing tubercles of uniform size; lips not barred 3

3. Dorsal pattern consisting of narrow transverse dark bars; dorsum in males bearing numerous small spinous tubercles O. leprieurii

Dorsal pattern not consisting of transverse bars; dorsal tubercles large or few in number 4

4. Dorsum uniformly dark brown; venter heavily mottled with black, especially in females; dorsum in males bearing large, keratinized tubercles O. verrucigerus

Dorsum tan with irregular dark brown blotches; venter cream with fine brown reticulations; dorsum in males bearing few, small non-spinous tubercles O. pearsoni

ACCOUNTS OF SPECIES

=Osteocephalus buckleyi= (Boulenger)

Hyla buckleyi Boulenger, 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 festae Peracca, 1904:39 [Holotype.--MIZS 2950 from "Valle de Santiago" (= lower Rio Zamora), Provincia Morona-Santiago, Ecuador; Enrico Festa collector]. New synonymy.

Osteocephalus buckleyi--Goin, 1961:13.

Hyla carri Cochran and Goin, 1970:211 [Holotype.--FMNH 69702 from Acevedo, Rio Suaza, Departamento Huila, Colombia; Philip collector]. New synonymy.

Hyla cabrerai Cochran and Goin, 1970:215 [Holotype.--USNM 152759 from Cano Guacaya, tributary of lower Rio Apoporis, Comisaria Amazonas, Colombia; Isadore Cabrera collector]. New synonymy.

Justification of Synonymy.--Boulenger (1882:362) listed 11 specimens in his description of Hyla buckleyi. We have examined all of these and conclude that one (BMNH 1947.2.13.42) is O. leprieurii. Cochran and Goin (1970:213) restricted the type locality to Canelos, Provincia Pastaza, 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 tympanum is 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 of Hyla festae is 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 of Hyla carri is 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 of Hyla cabrerai is 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 of Hyla cabrerai, Cochran and Goin (1970:217) stated: "This species, together with buckleyi and pearsoni certainly make a closely knit group.... Both buckleyi and cabrerai have long hind legs, with the extended heel reaching to the tip of the snout, while in pearsoni the extended heel reaches only to the eye. H. buckleyi has the belly dusky, while it is heavily spotted in cabrerai and is reticulated in pearsoni. H. cabrerai seems 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 of Hyla cabrerai was based on three specimens. We have examined the holotype and one paratype (WCAB 13284 from Territorio do Amapa, Brasil). Another paratype in the private collection of C. J. Goin from Cano Tui, between Mitu and Raudal de Yurupari, Comisaria de Vaupes, Colombia, was not examined.

Cochran and Goin (1970:211) based their description of Hyla carri on one gravid female and stated: "A large Hyla with the vomerine teeth in two ^^ 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 to Hyla claresignata of 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 festae has not been mentioned in the literature since the original description.