Territorio do Amapa in northeastern Brasil; the upper Amazon Basin from
southern Colombia to east-central Bolivia; one locality (Acevedo) in
upper Rio Magdalena drainage in Colombia (Fig. 8). All localities are at
elevations of less than 700 m. Records for Pallatanga and Santiago in
Provincia Chimborazo, Ecuador (high on the Pacific slopes of the Andes),
are considered to be erroneous. 78 specimens from 40 localities.
[Illustration: FIG. 8. Distribution of _Osteocephalus buckleyi_
(circles) and _O. 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= (Dumeril and Bibron)
_Hyla leprieurii_ Dumeril and Bibron, 1841:553 [Holotype.--MNHN
4629 from "Cayenne"; Mons. Leprieur collector].
_Hypsiboas leprieurii_--Cope, 1867:200.
_Hyla leprieurii britti_ Melin, 1941:42 [Holotype.--NHMG 489 from
the Rio Uaupes, north of the Rio Japu, Territorio do Amazonas,
Brasil; Douglas Melin collector]. New synonymy.
_Hyla leprieurii leprieurii_--Melin, 1941:42.
_Osteocephalus britti_--Goin, 1961:13.
_Osteocephalus leprieurii_--Goin, 1961:13.
_Justification of Synonymy._--The holotype of _Hyla leprieurii_ is a
female having a snout-vent length of 46.6 mm. The diameter of the
tympanum is 3.7 mm, 69.8 percent of the diameter of the eye. The dorsal
roofing bones are smooth, and the skin on the dorsum is smooth. The
penultimate phalanges of the fingers are not included in the webbing.
When we examined the specimen on 2 July 1969, it was slightly soft and
somewhat faded to a peculiar grayish green color with faint darker
transverse bars on the limbs. Dumeril and Bibron (1841:554) described
the coloration, as follows: "The loreal region is black. A stripe of the
same color extends from the posterior border of the orbit to the corner
of the mouth, passing through the tympanum. All of the dorsal parts are
grayish white with large transverse brown bands, which are more expanded
and less regularly outlined on the back than on the limbs. There is one
of these on the occiput that is in a triangular shape. All of the venter
is white." (Free translation from French.)
The holotype of _Hyla leprieurii britti_ is a male having a snout-vent
length of 48.1 mm. The diameter of the tympanum is 3.6 mm, 65.5 percent
of the diameter of the eye. The skin on the dorsum is tubercular; the
tubercles are small on head and on the dorsal surfaces of the limbs and
slightly larger on the back. The penultimate phalanges of the fingers
are not included in the webbing. Melin (1941:43) stated: "Above blackish
brown with a very indistinct band between the eyes; iris with mottle of
metallic lustre; hinder parts of upper jaw whitish; sides of body
mottled with blackish brown; hind limbs (especially tibiae and tarsi)
with narrow, diffuse cross bars; beneath whitish with slight brown
mottle along jaw." We examined the type on 17 February 1969; at that
time it was dull brown above with faint, narrow, dark brown, transverse
bars on the back and dorsal surfaces of the limbs. A cream subocular
spot was evident, and the venter was creamy white.
Melin (1941:42) stated that the holotype of _Hyla leprieurii britti_
"... resembles a good deal _H. leprieurii_ Dum. & Bibr. As, however, it
differs from the latter species by its very concave loreal region, small
tympanum, and almost uniformly brownish colour, it may at least form a
subspecies of _leprieurii_...." The pattern of narrow transverse bars on
the backs of the holotypes of _H. leprieurii_ and _H. britti_ is a
condition shared only by these two nominal taxa that are placed in
_Osteocephalus_. Melin noted that _britti_ differed from _leprieurii_ in
the depth of the loreal concavity and in the size of the tympanum.
Neither of these differences is noteworthy in comparison with series of
specimens. The depth of the loreal concavity is a highly subjective
character, and we note no differences between the types. The ratio of
the diameter of the tympanum to the diameter of the eye is relatively
smaller in both holotypes (0.698 in _leprieurii_--[F]; 0.655 in
_britti_--[M]) than in series of fresh specimens from Lago Agrio,
Ecuador (0.652-0.884, mean 0.785 in 17 males; 0.700-0.909, mean 0.790 in
20 females). The smaller proportions in the types may be due to
geographic variation or to shrinkage as a result of many years in
preservative (130+ years for _leprieurii_; 45 for _britti_).
Comparisons of the holotypes with series of specimens from Ecuador,
Guyana, and Surinam indicate that one morphological species occurs
throughout the upper Amazon Basin and the Guianas and that both type
specimens are representatives of one species. Consequently, we consider
_Hyla leprieurii_ Dumeril and Bibron, 1841, to be a monotypic species
with _Hyla leprieurii britti_ Melin, 1941, as a junior synonym.
In their account of _Osteocephalus leprieurii_, Cochran and Goin
(1970:323) stated: "The specimen described and illustrated (MCZ 28042)
has been directly compared with the types of _leprieurii_, _planiceps_,
and _vilarsi_ by the junior author and there seems to be no doubt that
all are conspecific. Another specimen (CNHM 69716) has been directly
compared with the types of _planiceps_ and _vilarsi_ and these,
likewise, are considered conspecific." With this justification Cochran
and Goin (1970:322) included _Osteocephalus planiceps_ Cope, 1874, and
_Hyla vilarsi_ Melin, 1941, in the synonymy of _Osteocephalus
leprieurii_.
We do not concur with Cochran and Goin's synonymy and contend that
_planiceps_ and _vilarsi_ are synonyms of _Osteocephalus taurinus_; we
give our reasons in the account of that species. We have examined the
specimens listed as _O. leprieurii_ by Cochran and Goin; several of
them, including CNHM (= FMNH) 69716, are _taurinus_. Thus, due to
Cochran and Goin's confusion of two taxa, their comparisons of certain
specimens with types has little meaning.
Cochran and Goin did not include _Hyla leprieurii britti_ in their
synonymy of _Osteocephalus leprieurii_ but did discuss the name in their
account of _Osteocephalus orcesi_ (= _O. verrucigerus_), as follows
(1970:319): "When we first examined one of the specimens we felt sure
that we had Melin's _Hyla britti_ at hand, but on direct comparison with
the type of _britti_ the two proved to be different. After studying the
type of _orcesi_ (SUNHM 13150) we have no doubt that the specimens at
hand are _orcesi_ and that _britti_ is a different, probably valid
species."
_Diagnosis._--1) Size moderate, sexual dimorphism evident; maximum
observed snout-vent length in males 48.4 mm, in females, 61.5 mm; 2)
skin on dorsum in males bearing numerous, minute, spinous tubercles; 3)
skin on flanks smooth; 4) web extending to base of antepenultimate
phalange on inner edge of third finger; 5) dorsum tan or olive-brown
with transverse brown or olive bars; 6) venter creamy white or pale tan
without markings; 7) lips marked with creamy tan labial stripe and
suborbital spot; 8) flanks pale tan with no markings; 9) dermal roofing
bones of skull lacking exostosis; 10) dermal sphenethmoid absent; 11)
nasals juxtaposed medially; 12) anteromedial margin of frontoparietal
between mid- and anterior levels of orbit; 13) frontoparietal fontanelle
partially exposed; 14) palatine not serrate; 15) parasphenoid lacking
odontoids; 16) zygomatic ramus of squamosal extending about one-half of
distance to maxillary arch; 17) transverse processes of presacral
vertebrae 3-8 about equal in width to one another and to sacral
diapophyses; 18) intermandibularis and submentalis muscles connected;
19) supramandibular portion of interhyoideus forming simple tubular
posterolateral extension; associated skin unmodified.
_Osteocephalus leprieurii_ differs from all other members of the genus
by having transverse dark bars on the back. Two other hylids (_Hyla
lanciformis_ and _multifasciata_) in the Amazon Basin have transverse
dark marks on the dorsum. Both of these differ from _leprieurii_ by
having pointed snouts, much longer hind limbs, and smooth skin dorsally.
_Distribution._--The periphery of the Amazon Basin, in the Guianas and
the upper part of the basin in southern Colombia, Ecuador, Peru, and
extreme western Brasil (Fig. 9). Most localities are at elevations of
less than 500 m, but the species ascends the lower Andean slopes to
elevations of 1100 m. 265 specimens from 31 localities.
[Illustration: FIG. 9. Distribution of _Osteocephalus leprieurii_
(circles) and _O. verrucigerus_ (triangles).]
_Remarks._--Most adults of _leprieurii_ have 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 some _taurinus_.
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 on the back and limbs but still retain
the light dorsolateral stripes, whereas the stripes are lost in larger
individuals.
[Illustration: FIG. 10. Ontogenetic change in color pattern in
_Osteocephalus leprieurii_: a. KU 126644; b. KU 126640;
c. KU 126625. x2.]
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 pearsoni_ Gaige, 1929:3 [Holotype.--UMMZ 57548 from the upper
Rio Beni, below mouth of Rio Mapiri, Departamento El Beni,
Bolivia; N. E. Pearson collector].
_Osteocephalus pearsoni_--Goin, 1961:13.
_Justification of Synonymy._--Goin (1961:13) suggested that _Hyla
pearsoni_ Gaige was an _Osteocephalus_, but Cochran and Goin (1970:217)
considered _pearsoni_ to be a _Hyla_. The presence of exostosed dermal
roofing bones, angulate prevomerine dentigerous processes, and the
structure of the vocal sacs are characters which place the species in
_Osteocephalus_.
_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 pearsoni_ can 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 Peru (1620 m in Rio Ucayali drainage) and northern Bolivia (less
than 500 m in Rio Beni drainage) (Fig. 8). 6 specimens from 3
localities.
_Remarks._--The specimen from Yaupi, Peru (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, 23 March 1970.) On the basis of this one subadult, it
seems likely that reticulations on the venter develop with age.
=Osteocephalus taurinus= Steindachner
_Osteocephalus taurinus_ Steindachner, 1862:77 [Holotype.--NHMW
16492 from Barra do Rio Negro, Manaus, Territorio do Amazonas,
Brasil; Johann Natterer collector].
_Osteocephalus flavolineatus_ Steindachner, 1862:80
[Holotype.--NHMW 16495 from Cucui, Territorio do Amazonas, Brasil;
Johann Natterer collector].
_Trachycephalus (Osteocephalus) taurinus_ Steindachner, 1867:64.
_Osteocephalus planiceps_ Cope. 1874:122 [Holotype.--ANSP 11399
from Nauta, Departamento de Loreto, Peru; James Orton collector].
New synonymy.
_Hyla taurina_--Boulenger, 1882:363 [synonymized _Osteocephalus
flavolineatus_ Steindachner, 1862, with _O. taurinus_ Steindachner,
1862].
_Hyla planiceps_--Boulenger, 1882:364.
_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.