Chapter II: Part 2

_Hyla (Trachycephalus) vilarsi_ Melin, 1941:40 [Holotype.--NHMG 488
from Taracua, Rio Uaupes, Territorio do Amazonas, Brasil; Douglas
Melin collector]. (_fide_ Bokermann, 1966:64.)

_Hyla depressa_ Andersson, 1945:73 [Holotype.--NHRM 1966 from the
Rio Pastaza watershed (? Provincia Pastaza), Ecuador; William
Clarke-MacIntyre collector]. New synonymy.

_Justification of Synonymy._--The holotype of _Osteocephalus taurinus_ is 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 of _Osteocephalus flavolineatus_ is a female having a 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 synonymized _flavolineatus_ with _taurinus_. We have observed that a middorsal cream stripe occurs in about 10 percent of the specimens of _taurinus_ and in some specimens of _buckleyi_. This is a common color morph in many species of _Eleutherodactylus_. 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 synonymizing _flavolineatus_ with _taurinus_.

The holotype of _Osteocephalus planiceps_ is 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 distinguishing _planiceps_ from _taurinus_. The coloration and morphometric and structural characters of the type of _planiceps_ all fall within the range of variation displayed by series of _O. taurinus_ from the upper Amazon Basin.

The type of _Hyla vilarsi_ is 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 names _Osteocephalus planiceps_ Cope and _Hyla vilarsi_ Melin was confused by Cochran and Goin (1970:322), who assigned these names to the synonymy of _O. leprieurii_. Bokermann (1966:64) placed _Hyla vilarsi_ in the synonymy of _Osteocephalus taurinus_ without justification. The type specimens of both _planiceps_ and _vilarsi_ have moderately exostosed dermal roofing bones and distinct cranial ridges. The type of _planiceps_ has moderately large tubercles on the dorsum, and the type of _vilarsi_ has spots on the throat, chest, and flanks and longitudinal markings on the back. All of these features are characteristic of _taurinus_ and not of _leprieurii_, 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 of _Hyla depressa_ is 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 of _Hyla depressa_ with _leprieurii_ and _buckleyi_, but he did not compare his specimen with _taurinus_, from which it exhibits no distinguishing features.

_Osteocephalus taurinus_ is a widespread and variable species, and it has received several specific names. We are convinced that _Osteocephalus taurinus_ Steindachner, 1862, is the oldest available name for this large Amazonian species. The following names are junior synonyms: _Osteocephalus flavolineatus_ Steindachner, 1862; _Osteocephalus planiceps_ Cope, 1874; _Hyla (Trachycephalus) vilarsi_ Melin, 1941; _Hyla depressa_ Andersson, 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 distinguish _taurinus_ from 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 included in 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.

_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 Tapirape, 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 Igarape 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: Rio Pastaza drainage, Ecuador 73.8 mm, Surinam 67.7 mm, Rio Ucayali drainage, Peru 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 in _taurinus_, but on the basis of preserved specimens we are unable to detect consistent differences distinguishing two or more taxa.

The coloration and pattern of _taurinus_ are 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 as _O. leprieurii_.

Ontogenetic change in coloration is slight in _taurinus_. 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 as _Hyla palpebrogranulata_ Andersson.

=Osteocephalus verrucigerus= (Werner)

_Hyla verrucigera_ Werner, 1901:601 [Holotype.--ZMB 16589 from
"Ecuador"; Richard Haensch collector].

_Hyla riopastazae_ Andersson, 1945:72 [Holotype.--NHRM 1960 from
Banos, Rio Pastaza, Provincia Tungurahua, Ecuador; William
Clarke-MacIntyre collector].

_Hyla orcesi_ Funkhouser, 1956:78 [Holotype.--CAS-SU 13150 from Rio
Pacayacu, tributary of Rio Cotapino, Provincia Napo, Ecuador;
collector unknown].

_Osteocephalus orcesi_--Cochran and Goin, 1970:317.

_Osteocephalus verrucigerus_--Trueb and Duellman, 1970:601
[Synonymized _Hyla riopastazae_ Andersson, 1945, and _Hyla orcesi_
Funkhouser, 1956, with _Hyla verrucigera_ Werner, 1901].

_Justification of Synonymy._--Trueb and Duellman (1970:605) discussed the assignment of the names in the synonymy of _O. verrucigerus_; only a brief resume is given here.

The extant type of _Hyla verrucigera_ is 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 of _Hyla riopastazae_ is 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 of _Hyla orcesi_ is 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 of _H. verrucigera_, _riopastazae_, and _orcesi_ are a juvenile, adult female, and adult male, respectively, of one species, the earliest available name for which is _Hyla verrucigera_ Werner, 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) nasals widely 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 verrucigerus_ can 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 Peru; one locality (Acevedo) in upper Rio 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.

GENERIC RELATIONSHIPS

Among the 33 genera currently recognized in the family Hylidae, there are two basic types of vocal sac structure (Duellman, 1970b), namely the subgular type and the lateral type. Only four hylid genera, all Neotropical lowland groups, are known to possess paired lateral vocal sacs; these are _Osteocephalus_, _Argenteohyla_, _Phrynohyas_, and _Trachycephalus_. The geographical distributions and morphological characteristics of these four genera suggest that they are more closely related to one another than with any other hylid genera.

Of the four genera, _Osteocephalus_ is the most generalized in morphology, and, like _Phrynohyas_, has no specialized habits. _Osteocephalus_ and _Argenteohyla_ are similarly distinguished from _Phrynohyas_ and _Trachycephalus_ on the basis of vocal sac structure. The vocal sacs of _Osteocephalus_ and _Argenteohyla_ are posterior and protrude posterolateral to the angles of the jaws when they are inflated, whereas those of _Phrynohyas_ and _Trachycephalus_ are more lateral and protrude posterior to the angles of the jaws when inflated.

Although _Osteocephalus_ and _Argenteohyla_ have similar vocal sac structure, they are obviously distinct. The monotypic _Argenteohyla_ is a rather specialized, semifossorial frog (Trueb, 1970b), characterized by smooth skin, moderate-sized digital discs, and a large inner metatarsal tubercle. The general architecture of the skull is not unlike that of _Osteocephalus_; the skulls of both are well roofed, broader than long, and characterized by posterolaterally oriented parasphenoid alae. _Argenteohyla_ bears small, slightly curved prevomerine dentigerous processes in contrast to the large, angular processes of _Osteocephalus_. The skull of _Argenteohyla_ shows specializations, apparently adaptations to its semifossorial mode of existence, which further distinguish the genus from _Osteocephalus_. In comparison with _Osteocephalus_, the cranium of _Argenteohyla_ is slightly depressed anteriorly, the roofing bones extensively casqued, and the palatines robust.

Osteologically, _Osteocephalus_ more closely resembles _Phrynohyas_ than either of the other two genera, but _Osteocephalus_ and _Phrynohyas_ are clearly distinct on the basis of their respective vocal sac structure. Like _Osteocephalus_, skulls of the members of the genus _Phrynohyas_ are broader than long, have extensive dermal roofing bones, and have posterolaterally oriented parasphenoid alae. In contrast to _Osteocephalus_, the dentigerous processes of the prevomers are curved, rather than angular in _Phrynohyas_. Furthermore, the latter genus is singularly distinguished from _Osteocephalus_, _Argenteohyla_, and _Trachycephalus_ by having extensively developed parotoid glands that produce a viscous, milky volatile secretion.

_Trachycephalus_ is the most readily identifiable of the four genera under discussion. Members of this genus are large frogs with heavily casqued and co-ossified skulls (Trueb, 1970a). The dermal roofing bones bear ornate and characteristic patterns of sculpturing. The medial ramus of the pterygoid does not articulate with the otic capsule, and the parasphenoid alae are laterally, rather than posterolaterally, oriented. A dermal sphenethmoid is present, and the parasphenoid bears odontoids. The basic structure of the skull has many characters in common with both _Osteocephalus_ and _Phrynohyas_. The obvious modifications of dermal roofing bones and of palatal and suspensory elements seem to be specializations adapting members of the genus _Trachycephalus_ to their peculiar phragmotic habits. The vocal sac structure of _Trachycephalus_ is like that of _Phrynohyas_ and therefore further distinguishes it from _Osteocephalus_.

Morphologically, _Osteocephalus_ seems to be sufficiently diverse and generalized so as to represent a modern derivative of an ancestral type which might have given rise to _Phrynohyas_, _Trachycephalus_, and _Argenteohyla_. The specialized vocal sac structure in _Phrynohyas_ and _Trachycephalus_ suggests that these two genera may be rather closely allied and represent a single phyletic line from an ancestral stock similar to _Osteocephalus_. _Argenteohyla_ is quite distinct from _Phrynohyas_ and _Trachycephalus_ and apparently represents a distinct phyletic line from the ancestral stock.

OCCURRENCE OF _OSTEOCEPHALUS_ IN AMAZONIAN ECUADOR

All of our observations on members of this genus have been made at four localities: 1) Santa Cecilia at an elevation of 340 meters on the Rio Aguarico, a tributary of the Rio Napo, 2) Lago Agrio, 330 meters, about 14 kilometers east of Santa Cecilia, 3) Puerto Libre, 570 meters, on the Rio Aguarico just east of its formation by the confluence of the Rio Cofanes and Rio Chingua, and 4) south slope of the Cordillera del Due, above the Rio Coca, 1150 meters. _Osteocephalus leprieurii_ was found at all four localities, and _buckleyi_ was found at all but the last; _taurinus_ was found at Santa Cecilia and Lago Agrio, and _verrucigerus_ was found only in the Cordillera del Due. Our data are based on collections of 113 frogs and three lots of tadpoles, as well as observations on calling sites and young. The observations are summarized by species, as follows:

_Osteocephalus buckleyi._--No breeding activity was observed. Males were found only at night in March, June, and July. One was perched on a _Heliconia_ leaf in a swamp at Puerto Libre, and two were on bushes in the forest at Santa Cecilia. A gravid female was found on a recently felled tree at Lago Agrio on the night of 12 May 1969.

_Osteocephalus leprieurii._--Males were heard calling sporadically at Puerto Libre in July 1968, and at Santa Cecilia in May 1969. A small chorus was found on the night of 12 May 1969 at Lago Agrio, where the frogs were perched on branches of fallen trees over a temporary pool. The call is a soft rattling chuckle. In late April and May many gravid females and males with well-developed nuptial excrescences were obtained from trees as they were felled at Lago Agrio. The reproductive condition of the frogs indicates that they probably breed in May. One individual called nearly every night from a large tree at Puerto Libre between 4-17 July 1968. The tree was felled on the latter date, but no frog was found. Two nights later apparently the same individual called from a bromeliad at a height of about 10 m on a large bamboo adjacent to the felled tree; the frog was collected when the bamboo was cut down.

Throughout the rainy months that we have worked in Ecuador (April-August) we have found occasional individuals perched on bushes or low trees at night. Large numbers of adults were observed only during a clearing operation which resulted in the felling of many large trees. Thus, it seems likely that _leprieurii_ is a tree-top inhabitant. A partially digested adult male was removed from the stomach of a _Hemiphractus proboscideus_.

At Santa Cecilia many recently metamorphosed young and juveniles were found in June and July 1968. Most of these were on low bushes or herbs in swamp forest at night; some were found in unfolded _Heliconia_ leaves by day, and one was observed on the forest floor by day. Snout-vent lengths of 18 specimens are 12.3-17.0 (mean 15.1) mm. The smaller frogs were recently metamorphosed as evidenced by the melanophore deposits above the vent. The coloration of the young is strikingly different from that of the adults (see account of _O. leprieurii_), so the association of the young and adults was not made until individuals with intermediate patterns were obtained at Lago Agrio in May 1969. Probably juveniles obtained in June and July are the offspring of an April or May breeding. We have been unable to associate tadpoles with this species.

_Osteocephalus taurinus._--A small chorus occurred at Lago Agrio on 12 May 1969. Males were calling from the ground adjacent to a small pool amidst recently felled trees. The males were very wary and, when approached, jumped onto limbs and ran up branches; this behavior was noted by Bokermann (1964). The call consists of a series of low-pitched, short notes--like a slow trill--four to six notes per call group. Call groups are repeated two, three, or four times followed by a lapse of several minutes. Although no amplectant pairs were found, several gravid females were collected at Lago Agrio in May, so it can be safely assumed that the species breeds in May. From April through July occasional individuals were observed on bushes and trees at night. During clearing operations at Lago Agrio several individuals were obtained from the tops of trees as they were felled.

_Osteocephalus verrucigerus._--Observations were made in a broad, shallow ravine, in which there was a small stream. On 2-4 August 1968, males were observed calling from low bushes and rocks at the edge of a quiet pool in the stream. The call consists of a series of well-pulsed, low-pitched, guttural notes produced at the rate of 5-10 per minute. One amplectant pair was found at the base of a bush adjacent to the pool on 3 August. Another female was found on a branch of a tree 2 m above the ground and 10 m from the stream. Tadpoles of this species were found in the quiet silt-bottomed pool.

SPECIMENS EXAMINED

The localities for each of the specimens examined are given in the following paragraphs. The arrangement of the data is as follows: alphabetically by country, state (department or province), and locality; alphabetically by the first letter in the abbreviations for the museums, and numerically after each museum abbreviation. Specimens lacking precise locality data are listed first in the most restricted political unit possible; localities which have not been found on maps or the positions of which are not known to us are given in quotation marks. Where more than one specimen is included under one museum number, the number of specimens is given in parentheses after the museum number. Unless noted otherwise, all specimens are alcoholics.

_Osteocephalus buckleyi_

BOLIVIA: _El Beni_: Ivon, BMNH 1967.2070-1. _Santa Cruz_:
Buenavista, CM 4333, 4339, UMMZ 66563-5.

BRASIL: _Amapa_: No specific locality, WCAB 13284.

COLOMBIA: _Amazonas_: Rio Guacaya, USNM 152759. _Huila_: Acevedo,
Rio Suaza, FMNH 69702. _Narino_: Rumiyacu, FMNH 54756. _Meta_:
Rio Guejar, Campamento La Macarena, USNM 152199.

ECUADOR: No specific locality, NHMW 6209, WCAB 35499. _Chimborazo_:
Pallatanga, BMNH 1947.2.13.46; Santiago, FMNH 42529.
_Morona-Santiago_: "Rio Santiago" (= Rio Zamora), MIZS 2950.
_Napo_: Lago Agrio, KU 126646; Puerto Libre, Rio Aguarico, KU
123172; Santa Cecilia, AUM 8138, KU 105208-9, 109506, 123171.
_Pastaza_: Alpayacu, BMNH 1912.11.1.64; Canelos, BMNH
1947.2.13.40-1, 1947.2.13.43-5; Colonia Mena, Rio Conambo, ZSM
33/1962; Don Tomas, USNM 166014; Guache, Rio Pastaza, AMNH 79986;
Rio Bobonaza, USNM 166005; Rio Capahuari, USNM 166554; Rio Conambo
at Rio Shiona-yacu, USNM 166018; Rio Copataza, upper Rio Pastaza,
USNM 166007-13; Rio Pastaza, NHRM 1946; Rio Pucyacu, USNM 165997
(skeleton), 165998-6001; Rio Rutuno, USNM 166006; Rio Villano,
USNM 166002-4; Sarayacu, BMNH 1947.2.13.36-9, MCZ 26090, ZMB 10166.

GUYANA: _Mazaruni-Potaro_: Kartabo, AMNH 70971; Membaru River,
upper Mazaruni River, UMMZ 85168; Oko Mountains, FMNH 26722-3.
_North West_: Amakura River, Haulover, UMMZ 83558-9. _Rupununi_:
Marudi River, AMNH 46233; Shudi-kar-wau, AMNH 49252. _West
Demerara_: Dunoon, UMMZ 52449, 52508.

PERU: _Junin_: Chanchamayo, BMNH 1911.12.13.79-80. _Loreto_:
Andoas, AMNH 79984-5; Cashiboya, AMNH 43454; San Antonio, Rio
Itaya, AMNH 43218. _Puno_: Yahuaramayo, BMNH 1913.2.25.7.

SURINAM: _Suriname_: Powakka, CM 44217.

SOUTH AMERICA: No specific locality, NHMW 6208.

_Osteocephalus leprieurii_

BRASIL: _Acre_: Tarauaca, FMNH 83247. _Amazonas_: Rio Javari,
Benjamin Constant, CAS-SU 12620; Rio Uaupes, north of Rio Japu,
NHMG 489.

COLOMBIA: _Amazonas_: Gino-goje, lower Rio Apoporis, MCZ 28038,
28040-2, 28044, USNM 152136-8.

ECUADOR: No specific locality, WCAB 35452-3; "Napo-Pastaza," USNM
166571. _Napo_: Avila, UMMZ 92093; south slope Cordillera del Due,
KU 123170; Lago Agrio, KU 125961-2 (skeletons), 126611-44, UMMZ
129326 (2); Limon Cocha, Rio Napo, KU 99210-6, UIMNH 63087-9,
63098, 63106-9, 63118-9, 64802-4, 64858, 87998-9, 88001-30,
88437-8, 88580, 88604-5, 89852-97, 89999-90000; Loreto, CAS-SU
11439, WCAB 36526; Puerto Libre, Rio Aguarico, KU 123190-1; Puerto
Napo, UIMNH 55818-20; Rio Cotapino, UMMZ 92094; Rio Napo, UMMZ
92078; Santa Cecilia, AUM 8099, 8102, 8113-5, 8127-9, 8131, 8137,
8139-46, 8148, KU 105210-20, 109509-11, 111971, 122964-87, 123169,
126645. _Pastaza_: Canelos, BMNH 1947.2.13.42, KU 120915; Rio
Alpayacu, UMMZ 92079; Rio Arajuno, USNM 166560-2, WCAB 40176; Rio
Oglan, USNM 16655203, 166558; Rio Rutuno, USNM 166559; Rio
Shilcayacu, below Puyo, USNM 166557; Rio Villano, USNM 166551.

FRENCH GUIANA: No specific locality, MNHN 4629. _Inini_: Lunier
River, MNHN 98/217.

GUYANA: _Mazaruni-Potaro_: Kartabo, AMNH 70967-8, 70972, 70976.
_Rupununi_: Shudi-kar-wau, AMNH 49255. _West Demerara_: Demerara
Falls, BMNH 72.10.16.23, 72.10.16.37-8.

PERU: _Loreto_: Estiron, Rio Ampiyacu, MZUSP 31033-4; Pebas, CAS-SU
3158, 3160; Roaboya, AMNH 43064.

SURINAM: No specific locality, MCZ 2036, RMNH 11468. _Marowijne_:
Camp 3, RMNH 13045-6; Wane Creek North, RMNH 11469-70. _Saramacca_:
Right Coppename River, RMNH 11467.

_Osteocephalus pearsoni_

BOLIVIA: _El Beni_: upper Rio Beni, below mouth of Rio Mapiri, MCZ
15565, UMMZ 57548, 67464-5; Rurrenbaque, UMMZ 57533.

PERU: _Pasco_: Yaupi, KU 136312.

_Osteocephalus taurinus_

BOLIVIA: _El Beni_: Ivon, BMNH 1967.2040; Reyes, UMMZ 57532. _La
Paz_: San Ernesto, Mapiri District, BMNH 1901.8.2.54. _Santa Cruz_:
Buenavista, AMNH 33951-2, 33958, BMNH 1927.8.1.19, 1927.8.1.118,
FMNH 27091, UMMZ 63319-21, 63959(2), 63961(2), 66566(2), 66567,
66568(2), 66569 (2), 66570, 66571(2), 66575-6, 68196; Rio Mamore,
2 km N Boca Chapare, AMNH 79324; Sara, CM 3840-1; Surutu,
CM 3814-5.

BRASIL: No specific locality: "Interior," BMNH 74.7.16.8-9. _Acre_:
Placido de Castro, MZUSP 6518; Tarauaca, WCAB 2496. _Amazonas_:
Cucui, NHMW 16495; Manacapuru, ZMB 28492, ZSM 278/1925; Manaus,
MCZ 56281, NHMW 16492; Maues, AMNH 69623, 76177; Taracua, NHMG 488,
WCAB 18463-4. _Mato Grosso_: Mabuca, MZUSP 4272; Posto Coluene, Rio
Xingu, WCAB 812; "Puerto Cabello," AMNH 3154; Tapirape, AMNH
73647-62, CAS-SU 12351-6, MNHN 46/324. _Para_: No specific
locality, MPEG 623-6; Belem, KU 129866; Cachimbo, FMNH 175876,
UIMNH 42149, WCAB 813; Cameta, NHMW 15892; Gurupa, BMNH 96.6.29.13;
Ilha de Marajo, BMNH 1923.11.9.20-4; Ilha Mexicana, ZSM 111/1911,
112/1912; "Ponto Dois Indios," BMNH 1939.1.5.5; Santarem, BMNH
75.10.22.1-4, MCZ 354. _Rondonia_: Abuna, CAS 49773-4, FMNH 64239;
Forte Principe da Beira, WCAB 10230; Igarape Marmelo, FMNH 140254,
KU 84725 (skeleton), 92243-6, 92247-8 (skeletons), WCAB 9997,
10001, 10003-4; Porto Velho, MZUSP 16343.

COLOMBIA: _Amazonas_: Gino-goje, lower Rio Apoporis, USNM 152139;
Leticia, USNM 152010-1; Raudal de la Playa, lower Rio Apoporis, MCZ
28050; Rio Apoporis, MCZ 28060. _Boyaca_: Sutatenza, USNM 152054-6.
_Cundinamarca_: Medina, MCZ 16269-71, USNM 152089-90, 152092-7,
152757. _Meta_: El Mico, Rio Guejar, USNM 152203; Rio Duda, Sierra
de Macarena, AMNH 79914; Rio Guapaya, Sierra de Macarena, FMNH
81332; Rio Guaviari, Casa de Piedra, UTA No number. _Putumayo_: Rio
Mecaya, FMNH 69711-4, 69716. _Vaupes_: Gomogoje, lower Rio
Apoporis, MCZ 28048.

ECUADOR: No specific locality, WCAB 35451, 35785; "Oriente," UMMZ
90418. _Carchi_: below Salinas, USNM 166059. _Esmeraldas_:
Carondelet, UIMNH 53560-9; Lagartera, Rio Caoni, UIMNH 53441,
53458-79. _Morona-Santiago_: Macuma, UIMNH 63142-3, 63145, 63147,
63151, 63154, 63157, USNM 166060. _Napo_: Avila, UMMZ 92077;
Cuyabeno, UIMNH 63158, 90111; Lago Agrio, KU 126647-55; Limon
Cocha, Rio Napo, AUM 8132-4, KU 99207-8, 99421-3, 99424 (skeleton),
99425, UIMNH 64801, 87798, 87800, 88032-5, 88576, 90066, 90082,
90102, 90104, 90314, 90984; Loreto, WCAB 35352; Rio Cotapino, UMMZ
92080; Rio Napo, UMMZ 84120; San Jose Abajo, AMNH 1295, 1449,
22180, 79990; Santa Cecilia, AUM 8117, 8150, KU 105230-3; south
slope Volcan Sumaco, USNM 166570. _Pastaza_: No specific locality,
ZSM 31/1956; Arajuno, USNM 165995; Bufeo, lower Rio Bobonaza, USNM
166046-8; Canelos, BMNH 80.12.5.179, 1947.2.13.48, UMMZ 89066; Don
Tomas, Rio Bobonaza, USNM 166049-50; Montalvo, CAS-SU 10320, USNM
165987-9, 166058, 166566; 2.5 km SE Puyo, USNM 166051; Rio Arajuno,
USNM 166043-5; Rio Arajuno (headwaters), USNM 166053; Rio Bobonaza,
WCAB 3613-4, 35504; Rio Capahuari, USNM 165990, 166555-6; Rio
Capahuari (headwaters), USNM 166057; Rio Conambo, USNM 166569, ZSM
28/1962, 35/1962; Rio Conambo at Rio Ollaguanga, USNM 166568; Rio
Conambo at Rio Shiona-yacu, USNM 166019, 166563-5; Rio Corrientes,
USNM 195994, 166020-38, WCAB 3841-2; Rio Huiyo-yacu, Pico de
Conambo, USNM 166052; Rio Pastaza, MCZ 19697; Rio Pastaza
(drainage), NHRM 1966, USNM 165996; Rio Pindo, USNM 166039-41; Rio
Pindo at Rio Tigre (village), USNM 165992-3, 166042; Rio Pucayacu,
USNM 166054, 166056; Rio Rutuno, USNM 166055; Rio Solis, upper Rio
Bobonaza, WCAB 39914; Rio Villano, USNM 165991, 166567; Sarayacu,
BMNH 80.12.5.213, 80.12.5.239-40, MZUSP 323; Shell Mera, KU 99420.
_Zamora-Chinchipe_: "Yani-Inzari," AMNH 43259, 43394; Zamora, AMNH
78928.

FRENCH GUIANA: _Cayenne_: Crique Gregoire, UP 40; Maripa, Oyapok
River, UP 72; Oyapok River, UZM 1473. _Inini_: Crique Gabrielle,
UP 118-20.

GUYANA: No specific locality: RMNH 1873(3), ZMB 3102(2). _East
Demerara_: Atkinson Field, ASU 11622. _Mazaruni-Potaro_: Chinapora
River, upper Potaro River, BMNH 1905.11.1.20-1; Kamakusa, AMNH
21416, 21418-9, 21422; Kartabo, AMNH 11689, 11691, 11697-9, 11703,
11706-8, 23107, 39730, 70966, 70969-70, 70973-5, USNM 118057;
Moraballi Creek, Essequibo River, BMNH 1930.10.10.47-51; Oko
Mountains, FMNH 26692-705; upper Potaro River, Tung District, BMNH
1905.11.1.40; Rockstone, FMNH 26591. _North West_: Amakura River,
Haulover, UMMZ 83735. _Rupununi_: north of Acaray River, west of
New River, KU 69747-8; Kuyuwini Landing, AMNH 46283; Pakaraima
Mountains, BMNH 1933.6.19.49; Shudi-kar-wau, AMNH 10665, 39637,
49256(2). _West Demerara_: Demerara, CAS 54773-4; Demerara Falls,
BMNH 72.10.16.16-22, 72.10.16.25-32; Dunoon, MCZ 4834, UMMZ 46736,
52493-4, 52502, 52504-5, 57271; Vryheid, BMNH 78.12.13.18.

PERU: _Amazonas_: Rio Cenepa, AMNH 43400. _Huanuco_: Monte Alegre,
Rio Pachitea, AMNH 43014, 43019. _Loreto_: Achinamisa, Rio
Huallaga, AMNH 42178, 42502; Andoas, Rio Pastaza, AMNH 79991;
Cashiboya, AMNH 43388, 43453; Estiron, Rio Ampiyacu, CAS 93264-74,
93276, 93278-9, 93281, 93283-6, 93289, 93311, 93327; Igarape
Champuia, upper Rio Curiuja, MZUSP 10339; Iquitos, AMNH 42204,
42442, 43468, NHMW 6118; Lago de Mirano, mouth of Rio Napo, AMNH
42712, 43186; Nauta, ANSP 11399; Ollanta, AMNH 42865; Pampa
Hermosa, Rio Cushabatay, AMNH 43124, 43146; Pebas, CAS-SU 6375;
Pucallpa, MJP 101(2), 140(3); Punga, Rio Tapiche, AMNH 43194;
"Rancho de Indiana, Iquitos District," MVZ 16890; upper Rio Abujao,
AMNH 42908; Rio Itaya, AMNH 42755; upper Rio Pisqui, AMNH 43536;
Rio Tapiche at Rio Contaya, AMNH 42983; Rio Utoquinia at Brasilian
frontier, AMNH 43137; Sobral, Rio Tamaya, AMNH 43242; Yurimaguas,
BMNH 84.2.18.50. _San Martin_: Cainarachi, AMNH 42763; Moyobamba,
ZSM 19/1914.

SURINAM: No specific locality, BMNH 70.3.10.67, NHMW 18433.3.
_Brokopondo_: Afobaka, RMNH 16536; Brownsweg, RMNH 16537; Railway
km. 121, RMNH 16534. _Marowijne_: Djai Creek, RMNH 16513-4; Maroni
River, ZMB 8240, 8531; Nassaugebergte, RMNH 16517-33; Paloemeu,
USNM 159025; Swamp Camp, RMNH 16515. _Nickerie_: Sipaliwini, RMNH
16538. _Saramacca_: Left Coppename River, RMNH 16535; Tibiti, RMNH
16516. _Suriname_: Berlijn, RMNH 15064; Powakka, CM 44226;
Zanderij, CM 50568.

VENEZUELA: _Amazonas_: Cerro Duida, UPR-M 2875; Cerro Marahuaca,
UPR-M 114-5; Esmeralda, AMNH 23174; Iniridi, SMF 2640; La Culebra,
MCZ 28572, UPR-M 117; Laguna, between Tama Tama and Esmeralda,
UPR-M 2760; Rio Pescado, AMNH 23177; Tapara, UPR-M 113. _Distrito
Federal_: Caracas, BMNH 51.7.17.182.

_Osteocephalus verrucigerus_

COLOMBIA: _Huila_: Acevedo, Rio Suaza, FMNH 69709-10.

ECUADOR: No specific locality, ZMB 16589. _Napo_: Avila, UMMZ
90413; south slope Cordillera del Due, KU 123176-88, 123189
(skeleton), 124208 (eggs), 124209-11 (tadpoles); L'Alegria, USNM
167472-3; Rio Pacayacu, tributary of Rio Cotapino, CAS-SU 13150;
southeast slope Volcan Sumaco, CAS-SU 11442. _Pastaza_: Abitagua,
CAS-SU 5067, FMNH 25791, 27619, UMMZ 90414, 92092; Alpayaca, Rio
Pastaza, BMNH 1912.11.1.64; Mera, UMMZ 90412(4). _Tungurahua_:
Banos, NHRM 1960.

PERU: _Ayacucho_: La Mar, Sivia, Rio Apurimac, FMNH 39853.
_Huanuco_: Rio Pachitea, midway between Puerto Victoria and Puerto
Inca, CAS-SU 17745. _Junin_: Satipo, MJP 38.

LITERATURE CITED

ANDERSSON, L. G.

1945. Batrachians from east Ecuador collected 1937, 1938 by Win.
Clarke-MacIntyre and Rolf Blomberg. Arkiv Zool., 37A(2):1-88.

BOKERMANN, W. C. A.

1964. Field observations on the hylid frog _Osteocephalus taurinus_
Fitz. Herpetologica, 20:252-255.

1966. Lista anotada das localidades tipo de anfibios Brasileiros.
Sao Paulo, 183 pp.

BOULENGER, G. A.

1882. Catalogue of the Batrachia Salientia s. Ecaudata in the
collection of the British Museum, ed. 2, London, xvi+503 pp.

COCHRAN, D. M. and C. J. GOIN

1970. Frogs of Colombia. Bull. U.S. Natl. Mus., 288:xii+655 pp.

COPE, E. D.

1867. On the families of the raniform Anura. Jour. Acad. Nat. Sci.
Philadelphia, 2:189-206.

1874. On some Batrachia and Nematognathi brought from the upper
Amazon by Prof. Orton. Proc. Acad. Nat. Sci. Philadelphia,
25:120-137.

DUELLMAN, W. E.

1970a. Identity of the South American hylid frog _Garbeana garbei_.
Copeia, (3):534-538.

1970b. The hylid frogs of Middle America. Monog. Mus. Nat. Hist.,
Univ. Kansas, 1:xi+753 pp.

DUMERIL, A. M. C. and G. BIBRON

1841. Erpetologie generale ou histoire naturelle complete des
reptiles, vol. 8. Paris, 792 pp.

FITZINGER, L.

1843. Systema reptilium. Vienna, ix+106 pp.

FUNKHOUSER, J.

1956. New frogs from Ecuador and southwestern Colombia. Zoologica,
91:73-80.

GAIGE, H. T.

1929. Three new tree-frogs from Panama and Bolivia. Occas. Papers
Mus. Zool. Univ. Michigan, 207:1-6.

GOIN, C. J.

1961. Synopsis of the genera of hylid frogs. Ann. Carnegie Mus.,
36:5-18.

MELIN, D.

1941. Contribution to the knowledge of Amphibia of South America.
Goeteborgs Kungl. Vetensk.-och Vitterh.-Sam. Handl., Ser. B,
1(4):1-71.

PERACCA, M. G.

1904. Viaggio del Dr. Enrico Festa nell' Ecuador e regioni vicine.
Reptile ed amfibii. Boll. Mus. Zool. Anat. Comp., Univ. Torino,
19:1-41.

STEINDACHNER, F.

1862. Ueber zwei noch unbeschriebene Batrachier. Arch. Zool. Anat.
Fisiol., 2:77-82.

1867. Amphibien. Novara Expedition. Zool. Theil, I, Vienna, 70 pp.

TRUEB, L.

1970a. The evolutionary relationships of casque-headed treefrogs
with co-ossified skulls (family Hylidae). Univ. Kansas
Publ. Mus. Nat. Hist., 18:547-716.

1970b. The generic status of _Hyla siemersi_ Mertens.
Herpetologica, 26:254-267.

TRUEB, L. and W. E. DUELLMAN

1970. The systematic status and life history of _Hyla verrucigera_
Werner. Copeia (4):601-610.

TYLER, M.

1971. The phylogenetic significance of vocal sac structure in hylid
frogs. Univ. Kansas Publ. Mus. Nat. Hist., 19:319-360.

WERNER, F.

1901. Ueber Reptilien und Batrachier aus Ecuador und Neu-Guinea.
Verh. Zool.-Bot. Gesell. Wien, 50:593-614.

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Transcriber's Notes

Except for the list of corrections below and minor corrections not listed, the text presented here is that of the original printed version.

Typographical Corrections

Page Correction
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5 is => in
5 buckley => buckleyi
17 scaral => sacral
19 Provicia => Provincia
25 in => is
25 metalic => metallic
41 bromeiad => bromeliad

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