HARVARD UNIVERSITY

LIBRARY

OF THE

Museum of Comparative Zoology

lAU^D. Cuivip. ZCUL
'5^(\)f\~ Wajuo^Ovj(Le3 LIBRARY

JAN 21 1972

OCCASIONAL PAPERS

HARVARD
of the UNIVERSITY

MUSEUM OF NATURAL HISTORY
The University of Kansas
Lawrence, Kansas

NUMBER 1 APRIL 29, 1971

A SYNOPSIS OF NEOTROPICAL HYLID FROGS,
GENUS OSTEOCEPHALUS

^^1 ( I; i( ^
Linda Trueb^ and William E. L^uellman^

When we initiated a study of the herpetofauna at Santa Cecilia
in Amazonian Ecuador in 1966, we were immediately confronted
with many kinds of animals that we could not identify with the ex-
isting literature. Comparisons of our specimens with those pre-
served in other museums resolved some of the problems, but many
identifications could be made only after study of type specimens;
even then some determinations remained questionable. We now
find that in order to prepare a meaningful account of the herpeto-
fauna of Santa Cecilia, we must complete several taxonomic studies,
the limits of which extend far beyond eastern Ecuador. Because of
our interests in hylids we have begun our studies on these frogs.

One of us (Trueb, 1970a) studied the cranial osteology of
casque-headed hylid frogs and redefined the genus Osteocephalus
but did not deteiTnine the number of species in the genus. Our work
in Amazonian Ecuador resulted in the discovery of the sympatric
occurrence of three species at each of two localities; one of these
species was found with a fourth species at another locality. Study
of museum specimens confirmed the recognition of these four species
in the Amazon Basin and lower Amazonian slopes of the Andes.
A fifth species from Bolivia and Peru also is included in the genus.
Examination of museum specimens has provided data on the geo-

^ Research Associate, Division of Herpetology, Museum of Natural History,
University of Kansas.

" Curator, Division of Herpetology, Museum of Natural History, University
ol Kansas.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

graphic variation in, and distribution of, the five species. However,
our conclusions pertaining to some populations need substantiation,
because we have been hampered by inadequate material from areas
beyond Ecuador. More than half of the 905 specimens of Osteoceph-
alus are from Ecuador, a relatively small part of the total range
of the genus.

In this paper we are presenting a taxonomic review of the genus
Osteocephalus; of necessity our study has been at the alpha level.
We have utilized all of the usual external characters, as well as
osteological features in our definitions of the species. Tadpoles and
mating calls are available for only one species, O. vernicigerus
( Trueb and Duellman, 1970 ) ; these and other important systematic
characters, such as karyotypes, are not available for the group at
this time. Our tendency has been to take a conservative view of
species; thus it is doubtful that any subsequent worker will recog-
nize fewer species in the genus. Our observations on these frogs in
Amazonian Ecuador are presented in a final section of this paper.

ACKNOWLEDGMENTS

For the loan of specimens or for the provision of working space
in their respective institutions, we are indebted to James E. Bohlke,
Werner C. A. Bokermann, F. W. Braestrup, Nelly Carrillo de
Espinoza, Osvaldo R. da Cunha, Josef Eiselt, M. J. Fouquette, Jr.,
Alice G. C. Grandison, Jean Guibe, Birgitta Hansson, Walter Hell-
mich, M. J. Hoogmoed, Robert F. Inger, Konrad Klemmer, Jean
Lescure, Alan E. Leviton, Clarence J. McCoy, Robert H. Mount,
Charles W. Myers, Umberto Parenti, Giinther Peters, James A.
Peters, William F. Pyburn, Juan A. Rivero, Dorothy M. Smith, Paulo
E. Vanzolini, Greta Vestergren, David B. Wake, Charles F. Walker,
Ernest E. Williams, and Richard G. Zweifel.

Study of specimens in European museums was made possible
by a grant (No. 5063) from the Penrose Fund of the American
Philosophical Society. Field work in Ecuador was partially sup-
ported by grants from the Watkins Fund of the Museum of Natural
History, University of Kansas. At our base camp at Santa Cecilia,
Ecuador, we enjoyed the hospitality of Ing. Ildefonso Munoz B.
Transportation in Ecuador was generously provided by the Texaco
Petroleum Company. During the course of our field work Stephen
R. Edwards and Thomas H. Fritts contributed directly to our study
of Osteocephalus. Michael J. Tyler of the South Australian Museum
provided infomiation on the vocal sac structure. We extend our

HYLID FROGS, GENUS OSTEOCEPHALUS 3

sincere thanks to all of these persons for their contributions to our
endeavors.

MATERIALS AND METHODS

We have examined 893 preserved frogs, including the type speci-
mens of all included nominal taxa, 8 skeletons, 1 lot of eggs, and 3
lots of tadpoles that we refer to the genus Osteocephalus; in addi-
tion skulls were removed from five preserved specimens, and
radiographs were made of 12 other preserved specimens. We have
been fortunate in seeing living individuals of all species, except
O. pearsoni, but we have colored photographs of a living specimen
of that species. Figures 1 and 2 were drawn from projected colored
transparencies of living frogs. Terminology follows that of Duell-
man ( 1970b ) . On the distribution maps solid symbols indicate lo-
calities from which we have examined specimens ;open symbols rep-
resent additional locality records based on the literature. Through-
out the text specimens are listed by their catalogue numbers pre-
ceded by the appropriate museum abbreviation, as follows:

AMNH American Museum of Natural History

ANSP Academy of Natural Sciences of Philadelphia

ASU Arizona State University

AUM Auburn University Museum

BMNH British Museum (Natural History)

CAS California Academy of Sciences

CAS-SU Stanford University Collection ( In California Academy of

Sciences )

CM Carnegie Museum

FMNH Field Museum of Natural History

KU Uni\'ersity of Kansas Museiuii of Natural History

MCZ Museum of Comparative Zoology, Harvard University

MIZS Museo ed Istituto di Zoologi Sistematico, Universita di Torino

MJP Museo Javier Prado, Lima

MNHN Museum National d'Histoire Naturelle, Paris

MPEG Museu Paraense Emiliano Goeldi, Belem

MVZ Museum of Vertebrate Zoology, University of California,

Berkeley

MZUSP Museu de Zoologia, Universidade da Sao Paulo

NHMG Naturhistoriska Museet Goteborg

NHMW Naturhistorisches Museum, Wien

NHRM Naturhistoriska Riksmuseet, Stockholm

RMNH Rijksniuseum van Natiuirlijke Histoire, Leiden

SMF Senckenbergische Museum, Frankfurt

UIMNH University of Illinois, Museiun of Natural History

UMMZ University of Michigan Museum of Zoology

UP Universite de Paris

UPR-M University of Puerto Rico, Mayagiiez

UTA University of Texas, Arlington

4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

USNM United States National Museum

UZM Uni\ersitets Zoologiske Museum, Copenhagen

WCAB Werner C. A. Bokennann, Sao Paulo, Brasil

ZMB Zoologisches Museum Berlin

ZSM Zoologisches Sammlung Miinchen

HISTORICAL RESUME

Because of the taxonomic confusion that has surrounded the
generic name Osteocephalus and two of the species (and their
synonyms), we present a brief resume of the taxonomic history of
the group.

Among the amphibians sent to the Museum National d'Histoire
Naturelle in Paris by a Monsieur Leprieur in French Guiana was a
single female specimen of a moderately large hylid having a well-
ossified skull and smooth dorsal skin. This specimen escaped from
the covetous eyes of Johann Tschudi, who prematurely named sev-
eral species on the basis of specimens in Paris, and survived without
an epithet until Dumeril and Bibron (1841) proposed for it the
name Hijla leprieurii. The description of the species is fairly de-
tailed, but the specimen was not illustrated. This is the earliest
trivial name now associated with Osteocephalus.

Fitzinger ( 1843 ) in his generic synopsis of amphibians and rep-
tiles proposed the generic name Osteocephalus but did not associate
a specific name with the genus. Consequently, Osteocephalus
Fitzinger, 1843, is a nomen nudum. Franz Steindachner followed
Leopoldo Fitzinger at the Naturhistorisches Museum in Vienna,
where he had access to Fitzinger's notes and, of course, the im-
portant collections housed in that museum. Steindachner (1862)
named two species of Osteocephahis on the basis of Brasilian
specimens collected by Johann Natterer. Both species were named
in the same publication; O. taurinus appeared on page 77, and
O. flavolineatus, on p. 80. This is the earliest association of the
generic name Osteocephalus with a specific name and a description,
both of which satisfy the Code of Zoological Nomenclature for
generic availability. Therefore, Steindachner is the authority for
the generic name Osteocephahis, which has O. taurinus as the type
species by original designation. It is not possible to determine
whether or not Steindachner's usage of Osteocephalus was the same
as that intended by Fitzinger 19 years earlier.

Steindachner (1862) gave reasonably good descriptions of his
two new species and provided excellent illustrations of the two
specimens, both large females. Apparently impressed by the simi-
larities between Trachycephahis nigromaculatus Tschudi, 1838, and

HYLID FROGS, GENUS OSTEOCEPHALUS 5

Osteocephalus taurinus, Steindachner (1867) used the combination
Trachycephalus (Osteocephalus) taurinus. This ambiguous usage
for the LSGOs prechidcs our determining if Steindachner was in
effect synonymizing OsteocepJiahis witli Trachycephalus or whether
he was placing Osteocephahis in a subgeneric status. Steindachner
(1867) did not mention O. flavoUneatus; perhaps by that time he
had conchided that flavoUneatus was only a color morph of taurinus.
Cope (1867) placed Hyla leprieurii in the genus Hypsihoas
Wagler, 1830. Cope (1874) named Osteocephahis pkiniceps from
Nauta, Peru. The single specimen was among the collections made
by the Orton Expedition to the upper Amazon Basin and was de-
posited in the Academy of Natural Sciences in Philadelphia.

Boulenger ( 1882 ) placed both Osteocephalus and Trachyce-
phahis is the synonymy of Hyla; he recognized Hyla taurina (with
O. flavoUneatus as a synonym), H. leprieurii, and H. planiceps. In
the same publication Boulenger named Hyla buckleyi on the basis
of 10 specimens in the British Museum from Ecuador; in the descrip-
tion he stated that Ijuckleyi was like leprieurii and taurinus in hav-
ing paired lateral vocal sacs. Boulenger held a lasting influence on
taxonomic herpetology, and his generic synonymy of Osteocephalus
was unchallenged until only a decade ago.

Coin ( 1961 ) presented a generic synopsis of the genera of hylid
frogs, in which he recognized Osteocephalus and stated: "There are
perhaps eight or ten species of this genus in South America. Cer-
tainly taurinus, britti, leprieuri, buckleyi and pearsoni belong here.
O. planiceps is surely a synonym of leprieuri and I believe that
garbei is as well. The status of such forms as macrotis, riopastazae,
and depressa has not yet been settled." Coin defined Osteocephalus
as follows: "Males with paired vocal pouches, one at each angle of
the jaw; derm of head not co-ossified with skull but roof of skull
exostosed." Trueb ( 1970a ) elaborated on Coin's definition and
assuredly included only O. taurinus and O. leprieurii in the genus.

Coin's inclusion of buckley, britti, and pearsoni in Osteocephalus
was the first association of any of these names with that genus.
Duellman (1970a) demonstrated that Garbeana garbei Miranda-
Ribeiro, 1926, was a member of the Hyla rubra group. Hyla
macrotis Andersson, 1945, is a Phrynohyas. Trueb and Duellman
(1970) determined that Hyla verrucigera Werner, 1901, is the
earliest name for an Osteocephalus displaying striking sexual di-
morphism in coloration and texture of the dorsal skin; Hyla
riopastazae Andersson, 1945 (female holotype), and Hyla orcesi

6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Funkhouser, 1956 ( male holotype ) , were placed in the synonymy of
Osteocephalus verrucigerus.

Hyla pearsoni Gaige, 1929, is a small species of Osteocephalus.
Our findings substantiate Coin's suggestions relative to two other
taxa. Hyla lepheurii britti Melin, 1941, from the Rio Uaupes, Brasil,
and Hyla depressa Andersson, 1945, from the Rio Pastaza watershed,
Ecuador, are members of the genus Osteocephalus, but both are
synonyms of earlier names— leprieiirii and taurinus, respectively.
Another name proposed by Melin (1941), Hyla (Trachycephahis)
vilarsi from Taracua, Brasil, also is placed in the synonymy of
O. taurinus.

Cochran and Coin ( 1970) were unaware of the identities of Hyla
verrucigera and riopastazae- they used the later name Osteocephalus
orcesi for Colombian frogs that are correctly referred to O. ver-
rucigerus. Although Coin (1961) placed Hyla buckleyi and H.
pear.soni in Osteocephalus, Cochran and Coin (1970) recognized a
"buckleyi group" in Hyla that included these two species plus a new
species, Hyla cabrerai from Amazonian Colombia and Brasil (total
of three specimens). Also, these authors named Hyla carri from a
single Colombian specimen. Study of the types of Hyla cabrerai,
H. carri, and //. festae Peracca, 1904, from Ecuador, reveal that all
of these names are synonyms of Osteocephalus buckleyi.

Much of the taxonomic confusion and multiplicity of trivial
names is due to the great amount of color variation in taurinus and
to the sexual dimoi-phism in the texture of the dorsal skin in all of
the species. The details of variation in these and other characters
and our justifications for the synonymies are given in the accounts
of the species. All of the trivial names that apply to species herein
recognized as members of the genus Osteocephalus are listed in
table 1.

Osteocephalus Steindachner, 1862

Osteocephalus Steindachner, 1862:77 [Type species. — Osteocephalus taurinus
Steindachner, 1862, by original designation]. Not Osteocephalus Fitzinger,
1843:50 (nomen nudum).

Diagnostic Definition.— I) Skull broader than long; 2) deiTnal
roofing bones of skull well ossified, exostosed, and/or co-ossified in
some species; 3) prenasal and internasal bones absent; 4) parasphe-
noid alae posterolaterally oriented; 5) dentigerous processes of
prevomers angular (/ \); 6) vocal sacs paired, posterior, and
when inflated protruding posteroventral or posterolateral to angles
of jaws; 7) submentalis muscle moderate in size and araphic; 8) in-
termandibularis muscle undifferentiated and bearing an elongate

HYLID FROGS, GENUS OSTEOCEPHALUS
Table 1. — Alphabetical Synonymy of the Species of Osteocephalus.

Trivial Name, Original Generic Name, Author, Date Current Name

britti (Hyla leprieurii) Melin, 1941 O. leprieurii

bitcklcyi (Hyla) Boulenger, 1882 O. buckleyi

cabrcrai (Hyla) Cochran and Coin, 1970 O. buckleyi

cam (Hyla) Cochran and Coin, 1970 .._ O. buckleyi

depressa (Hyla) Andersson, 1945 O. tatirinus

festae (Hyla) Peracca, 1904 O. buckleyi

flavoliueatii.s- (Osteocephalus) Steindachner, 1862 O. tauiinus

leprieurii (Hyla) Dumeril and Bibron, 1841 _— O. lei)rieurii

orcesi (Hyla) Funkhouser, 1956 O. verrucigerus

pearsoni (Hyla) Caige, 1929 O. pearsoni

planiceps (Osteocephalus) Cope, 1874 O. taurinus

riopastazae (Hyla) Andersson, 1945 O. verrucigerus

taurinus (Osteocephalus) Steindachner, 1862 O. taurinus

verrucigera (Hyla) Werner, 1901 O. verrucigerus

vilarsi (Hyla) Melin, 1941 O. taurinus

median aponeurosis; 9) parotoid glands absent or poorly developed,
skin not producing viscous secretion characteristic of Phrynohyas;
10) skin on dorsum tuberculate in males, smooth in females; 11)
tympanum large, 60 percent or more of diameter of eye; 12) fingers
about one-third, toes more than three-fourths webbed; 13) discs
large, round; 14) nuptial excrescences present in breeding males;
15) inner metatarsal tubercle not modified for digging; 16) outer
metatarsal tubercle absent; 17 ) tarsal fold weak or absent; 18 ) pupil
horizontal; 19) palpebrum clear; 20) known tadpoles having two
upper and five lower rows of teeth.

Content— As defined here, the genus contains five known
species: O. buckleyi (Boulenger), O. leprieurii (Dumeril and
Bibron), O. pearsoni (Gaige), O. taurinus Steindachner, and O.
verrucigerus ( Werner ) .

Distrilmtion.— The Guianas and Amazon Basin; also in the upper
Orinoco and Magdalena drainages. Most localities are at elevations
below 500 m, but the genus ascends the Amazonian slopes of the
Andes to elevations of about 1800 m.

Analysis of Characters

Size and Proportions— Frogs of the genus Osteocephalus are
moderate to large hylids. The largest species is taurinus, attaining
a snout- vent length of 103.1 mm; the smallest is pearsoni, which at-
tains a length of 54.7 mm. Considerable intraspecific geographic
variation occurs in adult size, especially in taurinus. Females of all
species attain a noticeably larger size than males, but no significant
differences are apparent in proportions ( Table 2 ) .

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

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HYLID FROGS, GENUS OSTEOCEPHALUS 9

Coloration.— AW Osteocephalus are predominantly brown frogs
usually with some darker dorsal markings (Figs. 1 and 2). Osteo-
cephalus vernicigerus has a nearly uniform dark brown dorsum and
no distinct transverse bars on the limbs, whereas all of the other
species have distinct bars on the limbs. The dorsal markings on the
body consist of irregular blotches in buckleyi, pearsoni, and taurinus
but are narrow transverse marks in leprieiirii. A narrow middorsal
cream or yellow stripe is present in some individuals of buckleyi
and taurinus but absent in all individuals of the other species. The
flanks are uniform pale tan in leprieurii and uniform reddish brown
in verruciiierus; in the other species the flanks are cream to brown
with dark brown or black spots ( also dark diagonal marks in some
buckleyi ) . A creamy white anal stripe is present in some specimens
of leprieurii but absent in all individuals of other species.

The postocular region, encompassing the tympanum, is dark
brown in most specimens. In adults of pearsoni and taurinus the
upper lips are dark brown. A pale cream or tan suborbital spot is
present in pearsoni and in some taurinus; in some specimens of
taurinus the suborbital spot is expanded posteriorly forming a labial
stripe on the posterior part of the lip. The labial markings of
verruci'^erus are similar to the latter pattern, except that in females
a distinct, light labial stripe extends the length of the lip. Osteoceph-
alus leprieurii has a distinct, broad, pale labial stripe. The lips
are barred cream and dark brown in buckleyi.

The venter is uniform creamy white or pale tan in leprieurii,
unifonn white in some buckleyi (most males), and uniform tan in
some taurinus. The other species and some individuals of taurinus
and buckleyi (most females) have dark ventral markings. These
markings are most distinctive in verrucigerus, in which the venter
is white with bold black mottling and spots ( Fig. 3c ) . Those indi-
viduals of taurinus having ventral markings usually have indistinct,
diffuse brown spots on the throat and chest ( Fig. 3b ) . Osteocepha-
lus pearsoni is characterized by a fine brown reticulation on the
venter and on the anterior and posterior surfaces of the thighs in
adults ( Fig. 3a). Individuals of buckleyi that have ventral markings
vary between the patterns illustrated for pearsoni and taurinus
(Figs. 3b and c).

Ontogenetic change in coloration is slight or non-existent in
buckleyi, pearsoni, and taurinus, except that juveniles lack ventral
markings. A dark blotch on the back and distinct transverse bars on
the limbs are evident in juveniles of verrucigerus; these markings
are obscured in the adults. Juveniles of leprieurii are olive-brown

10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

«<;

Fic;. 1. Species of Osteocephalus: Top. O. pearsoni, KU 136312, $ ; Middle.
O. huckleiji, KU 123172, i ; Bottom. O. verrucigerus, KU 123177, 6. Xl.5.

HYLID FROGS, GENUS OSTEOCEPHALUS

11

.K^

%;

%

/

Fig. 2. Species of Osteocephalits: Top. O. leprieuni, KU 126611, $; Bottom.

O. taiirinus, KU 126648, S . Xl.

12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 3. Diagrammatic views of ventral color patterns in Osteocephalus: a. O.
pearsoni, UMMZ 57533, $ ; b. O. taurinus, USNM 166037, $ ; c. O. ver-

nicigenis, KU 123185, 9 .

with yellow dorsolateral stripes; the transverse dark marks, char-
acteristic of the adults, appear before the stripes are lost.

Skin.— The dorsal skin of all males of Osteocephalus is tubercu-
late to varying degrees, whereas the dorsal skin of females is smooth,
or nearly so ( Fig. 4 ) . Osteocephalus verrucigerus differs from other
members of the genus by the presence of niunerous, large tubercles
bearing keratinized tips. The tubercles of leprieurii are numerous
and spinous but much smaller than those of verrucigerus; those of
taurinus are spinous but less numerous than in leprieurii. Os-
teocephalus buckleiji has a mixture of large and small, non-spinous
tubercles, and pearsoni has only a few, small, scattered, non-spinous
tubercles. Fleshy tubercles occur on the eyelids and supratympanic
fold in females of buckleyi; a few small tubercles are present on the
back of females of pearsoni, whereas the dorsal skin in females of
the other species is smooth. The skin on the flanks of both sexes of
buckleyi is weakly areolatc; in tlie other species the flanks are
smooth. The skin on the top of the head in taurinus is rugose as a
consequence of co-ossification. In all species the anal opening is
directed posteriorly at the upper level of the thighs.

Hands and Feet.— The feet of Osteocephalus are fully webbed or

HYLID FROGS, GENUS OSTEOCEPHALUS

13

Lnar

*

( ■■

mHH

:.^ -s— i

Fig. 4. Segments of dorsal skin of males of Osteocephalus showing size and
arrangement of tubercles: a. O. verrucigeius, KU 123183; b. O. tauriniis,
USNM 166033; c. O. leprieurii, KU 126616; d. O. bticklcyi, USNM 165999.

Each square = 1 sq. cm.

nearly so. Webbing between fingers one and two is basal in all
species. Webbing between fingers two, three, and four is most ex-
tensive in taurimis, in which the three fingers are about one-half
webbed (Fig. 5). Osteocephalus buckleyi, pearsoni, and ver-
rucigerus have reduced webbing between fingers two and three,
and leprieurii has reduced webbing between fingers two, three, and
four. All members of the genus have well-developed subconical
subarticular tubercles on the fingers and toes; there is a tendency
for the distal tubercle on the fourth finger to be weakly bifid.
Palmar and plantar supernumerary tubercles are well developed in
taurimis, moderately developed in buckleyi, leprieurii, and pearsoni,
and barely evident in verrucigerus. All of the species have a notice-
able fold on the wrist and enlarged prepollices, bearing horny
nuptial excrescences in breeding males. The prepollex is least en-
larged in buckleyi. Outer metatarsal tubercles are absent. The
inner metatarsal tubercle is moderately well developed and ovoid
in leprieurii and pearsoni; it is elliptical and flat in the other species.
Tarsal folds are absent in all species except verrucigerus. in which
the folds are barely evident.

Cranium.— As a genus, the cranial structure is remarkably uni-
form and quite generalized when viewed in the context of the family
Hylidae. The skulls are broad and relatively flat, each being only
slightly more broad than long and about one-third as high as long.
In dorsal aspect the snouts are broadly rounded; the snout of
buckleyi is somewhat less rounded and appears to be slightly longer
than the snouts of other species. This subtle difference relates to
the relative narrowness of the premaxillaries in buckleyi.

The genus is characterized by well-developed dermal roofing
bones and an unusually large exposure of the sphenethmoid dorsally
(Fig. 6). The conformation of the sphenethmoid exposed dorsally
is determined by the marginal positions of the adjacent, overlapping

14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 5. Palmar views of hands of males of Osteocephahis: a. O. htickleyi, KU

109506; b. O. leprieurii, KU 126627; c. O. pcarsoni, MCZ 15565; d. O. taurinus,

KU 126653; e. O. venucigems, KU 123177. X4.

HYLID FROGS, GENUS OSTEOCEPHALUS

15

Fig. 6. Skulls of hvo species of Osteocephalus: a and b. O. leprieurii, KU
125961; c and d. O. pcarsoni, UMMZ 67465. x3.

elements— the nasals and frontoparietals. Medially the nasals over-
lap the lateral margins of the sphenethmoid. Anteromedially, the
nasals converge in leprieurii and taurinus, are narrowly separated
in buckleyi and pearsoni, or are more widely separated in ver-
rucigerus. In all species the nasals terminate at the anterodorsal
corner of the orbit. The frontoparietals of buckleyi, leprieurii, and
taurinus have an anterolateral extension, which marginally overlaps
the dorsolateral edge of the sphenethmoid and articulates with the
posterodorsal corner of the nasal in buckleyi and taurinus; the bones
are narrowly separated in leprieurii. The frontoparietals of pearsoni
and verrucigerus have more extensive median ossification and less
extensive anterolateral ossification. Thus, in those species the
posteromedian portion of the sphenethmoid is obscured, and the

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

lateral margins are partly exposed. The frontoparietal fontanelle is
completely covered in all species, except buckleyi and Jeprieiirii, in
which an irregular, median separation of the frontoparietals exposes
a small portion of the fontanelle. The posterolateral margins of the
frontoparietals lie medial to the epiotic eminences.

Dermal ornamentation, involving the nasals, frontoparietals, and
sphenethmoid, occurs in taurinus and, to a limited extent, in peor-
soni. In the latter species marginal portions of the frontoparietals,
the dorsal surfaces of the nasals, and the posteromedial part of the
exposed sphenethmoid are slightly exostosed, resulting in an open,
reticulate pattern of dermal sculpturing of exceedingly low relief
( Fig. 6c ) . Osteocephalus taurinus is characterized by casquing,
co-ossification, and a rather intricate pattern of dermal sculpturing,
which was described in detail and illustrated by Trueb ( 1970a ) .

The squamosals of all species are moderately large structures
having otic plates that overlie the lateral portion of the cristae
paroticae. The posterior rami are short; the zygomatic rami of all
species, except taurinus, extend slightly more than one-half of the
distance to the maxillary. In taurinus the zygomatic ramus extends
nearly to, or articulates with, the maxillary.

The maxillary arches are complete and relatively robust. The
alary processes of the premaxillaries are vertically oriented in
leprieurii, pearsoni, and taurinus and very slightly inclined pos-
teriorly in buckleyi and verrucigerus. The maxillaries are uniformly
characterized by the absence of postorbital processes and by the
presence of preorbital processes that articulate with the maxillary
processes of the nasals. The partes facialae of the maxillaries are
moderately developed in all species, except taurinus, in which the
pars fascialis tends to articulate with the lateral margin of the nasal
in well-ossified individuals. The partes palatinae are poorly de-
veloped in all species, except buckleyi, in which the pars palatina
of the premaxillaiy is moderately robust.

The pterygoids are uniformly tri-radiate structures. The anterior
rami terminate at about the mid-level of the orbit, and the medial
rami articulate firmly with the anterolateral corner of the otic cap-
sule. The palatines are well-developed elements bearing ventral
ridges; the ridges are somewhat irregular in buckleyi, taurinus, and
verrucigerus but smooth in leprieurii and pearsoni. The parasphe-
noids are large elements characterized by acuminate cultrifonn
processes and posterolaterally inclined alary processes. The basal
areas of the cultrifonn processes bear small odontoid protuberances
in buckleyi, taurinus, and verrucigerus, whereas they are smooth in

HYLID FROGS, GENUS OSTEOCEPHALUS

17

Fig. 7. Dorsal views of vertebral columns of two species of OsteocepJialus:
a. O. leprieurii, KU 125962, ? ; h. O. huckleyi, USNM 165997, 9 . x2.

leprieurii and pearsoni. Tlie prevomers are remaikalDly uniform,
moderately well-developed structures. In each species the anterior
ramus lies adjacent to the premaxillary, and the lateral wings form
the anterior, medial, and posteromedial margins of the internal
nares. The dentigerous processes are characteristically large and
angular and bear numerous teeth. The sphenethmoid and otoccipi-
tals are well ossified; a dermal sphenethmoid is present only in
taurinus.

Vertebral Column.— The cervical cotyles are uniformly widely
displaced. The neural arches are low and non-imbricate. The trans-
verse processes of the third presacral vertebrae are approximately
equal in width to the sacral diapophyses in all species, except huck-
leyi, in which the processes of the third presacral are slightly nar-
rower than the diapophyses. Osteocephahis huckleyi is further dis-
tinguished by the presence of narrow transverse processes on pre-
sacrals 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 scaral diapophyses of all
species are moderately dilated and posterolaterally inclined. The
coccyx bears a distinct dorsal ridge and has a bicondylar articula-
tion with the sacrum.

Pectoral Girdle.— The pectoral girdles are fully arciferal and
bear small, cartilaginous omosterna and moderately large car-
tilaginous sterna. The coracoids are robust, and the clavicles are
strongly arched. Procoracoid cartilage seems to be absent. The

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

scapulae are large, longer than the clavicles, and bicapitate prox-
imally. 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) de-
scribed the throat myology of Osteocephalus. The genus is charac-
terized by a moderate-sized araphic submentalis muscle and an un-
differentiated intermandibularis having an elongate median apo-
neurosis. The intermandibularis and submentalis are completely in-
dependent in buckleyi, whereas in the other species there is a small
attacliment between these muscles.

According to Tyler (pers. com.), Osteocephalus has three dis-
tinctive 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 fomi a simple tubular, posterolateral extension; there
is no modification of the associated skin. OsteocepJmlus buckleyi
and pearsoni have more extensive development of the supramandib-
ular 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 in-
terhyoideus 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 fomis an everted pouch, which dangles loosely
beneath the supratympanic fold.

Key to the Species of 0.steocephalus

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

HYLID FROGS, GENUS OSTEOCEPHALUS 19

Inner edge of third finger webbed to base of antepenultimate
phalange; dorsum plain or marked with dark blotches or trans-
verse 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. buckleiji

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 ])rown 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 Hershkovitz

collector]. New synonymy.
Hyla cabrerai Cochran and Goin, 1970:215 [Holotype.— USNM 152759 from

Caiio Guacaya, tributary of lower Rio Apoporis, Comisaria Amazonas,

Colombia; Isadore Cabrera collector]. New synonymy.

Justification of Sy no mjimj.— Boulenger (1882:362) listed 11 spec-
imens 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, Provicia 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 tym-

20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

panum 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 pos-
teriorly 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 pos-
terior 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 cahrerai 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,

HYLID FROGS, GENUS OSTEOCEPHALUS 21

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 Hijla cabrerai, Cochran and Coin (1970:
217) stated: "This species, together with buckleyi and pearsoni
certainly make a closely knit group. . . . Both huckletji 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. Coin from Caiio Tui, between Mitu and Randal
de Yurupari, Comisaria de \'aupes, Colombia, was not examined.

Cochran and Coin (1970:211) based their description of HyJa
carri on one gravid female and stated: "A large Hyla with the
vomerine teeth in two A A 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 descrip-
tion.

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 propor-
tions the type specimens of the nominal taxa all fall within the
range of variation exhibited by a series of 18 males and 15 females
from Provincia Pastaza, Ecuador, except the ratio of the diameter
of the tympanum to that of the eye in the female holotype of Hyla
festae. In that specimen the ratio is 0.574, whereas the ratio in the
15 females from Provincia Pastaza is 0.587-0.905 (mean 0.736).

Ventral coloration is the most variable character among the
types. The venter in the type of Hyla festae is boldly spotted; it is

22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

distinctly spotted in cabrerai, uniform tan in carri, and tan, flecked,
or spotted in the type series of btickleyi. The ventral coloration in
series of specimens from Amazonian Ecuador encompasses that ob-
served in all of the types, except that of festae, which has more
ventral spotting than any other individual.

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

The types of the nominal taxa and series of specimens from
Guyana and Amazonian Ecuador display noticeable variation in
dorsal coloration. The variety of dorsal patterns of all of the types
is included in the variation displayed by the other specimens. All
specimens have some amount of dark spotting on the flanks; all have
vertically barred lips, on which a pale subocular spot usually is evi-
dent. 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 depres-
sions. This areolate dermal condition is present in all specimens
and does not occur in other species of Osteocephalus. The degree of
tubercularity of the skin on the dorsum is variable and sexually
dimoiphic. 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 supra-
tympanic fold.

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

The ventral coloration of the type of Hyla festae resembles that
of Osteocephalus verrucig,erus, but the type differs from verrii-
cigerus by having areolate skin on the flanks and distinct dark mark-
ings on the dorsum. In verrucigerus the skin on the flanks is smooth,

HYLID FROGS, GENUS OSTEOCEPHALUS 23

and the dorsum is uniform dark brown, except for a tan snout in
females.

Comparisons of the types of the nominal species with series of
specimens from Guyana, Colombia, Ecuador, and Peru suggest
strongly that the types are representative of one taxon, the oldest
name for which is Hyla buckleyi Boulenger, 18S2. Consequently,
we place Hijla festae Peracca, 1904, Hyla carri Cochran and Coin,
1970, and Hyla cahrerai Cochran and Coin, 1970, as junior synonyms
of Hyla buckleyi Boulenger, 1882.

Diag,nosis.—l) Size moderate, sexual dimoiphism extreme; max-
imum 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) demial roofing bones of
skull lacking exostosis; 10) demial sphenethmoid absent; 11) nasals
widely separated medially; 12) anteromedial margin of frontoparie-
tal at mid-level of orbit; 13) frontoparietal fontanelle partially ex-
posed; 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) intemiandibularis and submentalis
muscles independent; 19) supramandibular portion of inter-
hyoideus extensively developed; associated skin forming broad
loose fold.

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

Distribution.— The periphery of the Amazon Basin, in the Gui-
anas and Territorio do Amapa in northeastern Brasil; the upper
Amazon Basin from southern Colombia to east-central Bolivia; one
locality (Acevedo) in upper Rio Magdalena drainage in Colombia
(Fig. 8). All localities are at elevations of less than 700 m. Records

24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

0 500 1000

I Ill

kilometers

58°

Fig. 8. Distribution of Osteocephalus btickleyi (circles) and O. pearsoni

( triangles ) .

for Pallatanga and Santiago in Provincia ChimlDorazo, Ecuador
(high on the Pacific slopes of the Andes), are considered to be
erroneous. 78 specimens from 40 localities.

Remarks.— In life the dorsum is green with dark markings. A
male (KU 123171) from Santa Cecilia, Ecuador, was: "Dorsum
green with dark brown blotches. Anterior and posterior surfaces of
thighs dull blue. Venter brown, flecked with white. Iris greenish
bronze with brown horizontal triangles and ventromedian brown
line." (W. E. Duellman, field notes, 16 June 1968.) A female (KU
126646) from Lago Agrio, Ecuador, was: "Dorsum pale green with
darker green blotches and creamy yellow middorsal stripe. Lateral
blotches bronze-tan. Flanks tan with black blotches. Anterior sur-
faces of thighs dark brown. Dorsal and posterior surfaces of tliighs
and shanks tan with dark brown blotches. Webbing brown. Sub-
orbital 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, hor-

HYLID FROGS, GENUS OSTEOCEPHALUS 25

izontal, 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 hritti 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 hritti — Coin, 1961:13.
Osteocephalus leprieurii — Coin, 1961:13.

Justification of Synonymy— The holotype of HyJa 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 in 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
transxerse brown bands, which are more expanded and less reg-
ularly 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 hritti 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 sur-
faces 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 metalic 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

26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

the back and dorsal surfaces of the hmbs. A cream subocular spot
was evident, and the venter was creamy white.

MeHn (1941:42) stated that the holotype of Hijla Jeprieurii 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 Osteocephahis. 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 compari-
son 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—
9 ; 0.655 in britti— $ ) than in series of fresh specimens from Lago
Agrio, Ecuador (0.652-0.884, mean 0.785 in 17 males; 0.700-0.909,
mean 0.790 in 20 females ) . The smaller proportions in the types may
be due to geographic variation or to shrinkage as a result of many
years in preservative ( 130-f 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 Hijla leprieurii britti Melin,
1941, as a junior synonym.

In their account of Osteocephahis leprieurii, Cochran and Coin
(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 justifica-
tion Cochran and Coin (1970:322) included Osteocephahis plani-
ceps Cope, 1874, and Hyla vilarsi Melin, 1941, in the synonymy of
Osteocephalus leprieurii.

We do not concur with Cochran and Coin's synonymy and con-
tend 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

HYLID FROGS, GENUS OSTEOCEPHALUS 27

Coin; several of them, including CNHM (= FMNH) 69716, are
taurinus. Thus, due to Cochran and Coin's confusion of two taxa,
their comparisons of certain specimens with types has little meaning.

Cochran and Coin did not include Hyla leprieurii hritti in their
synonymy of Osteocephalus leprieurii but did discuss the name in
their account of Osteocephalus orcesi (= O. verrucigerus), as fol-
lows (1970:319): "When we first examined one of the specimens
we felt sure that we had Melin's Hyla J)ritti 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 arc orcesi and that britti is a dif-
ferent, probably valid species."

Diagnosis.— 1) Size moderate, sexual dimorphism evident; max-
imum 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 an-
terior levels of orbit; 13) frontoparietal fontanelle partially ex-
posed; 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 con-
nected; 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 Cui-
anas 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

28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

0 500 1000

■ I I I I I I l-H+H

kilometers

Fig. 9. Distribution of Osteocephalus leprieurii (circles) and O. verrucigerus

( triangles ) .

Andean slopes to elevations of 1100 m. 265 specimens from 31
localities.

Remarks.— Most adults of leprieurii have distinct transverse
markings on the back; these are variable in width, extent, and ar-
rangement. 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

HYLID FROGS, GENUS OSTEOCEPHALUS

29

Fig. 10. Ontogenetic change in color pattern in Osteocephahis leprieurii:
a. KU 126644; b. KU 126640; c. KU 126625. x2.

the back and limbs but still retain the light dorsolateral stripes,
whereas the stripes are lost in larger individuals.

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)

Htjla 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:

30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

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 Osteocephahis.

Diagnosis.— 1) Size moderate, sexual dimorphism evident; max-
imum 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) demial 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 maxillaiy 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 fonning broad loose fold.

Osteocephahis 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— Vp-pei- 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

HYLID FROGS, GENUS OSTEOCEPHALUS 31

March 1970. ) On the basis of this one subadult, it seems hkely that
reticulations on the venter develop with age.

Osteocephalus taurinus Steindachner

Osteocephalus taurimis Steindachner, 1862:77 [Holotype. — NHMW 16492 from

Barra do Rio Negro, Manaus, Territorio do Amazonas, Brasil; Johann

Natterer collector].
Osteocephalus flavolincatits 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, Pen'i; James Orton collector]. New

synonymy.
Hijla taurina — Boulenger, 1882:363 [synonymized Osteocephalus jiavolineatiis

Steindachner, 1862, with O. taurinus Steindachner, 1862].
Hyla planiceps — Bonlenger, 1882:364.
Hyla (Trachycephalus) vilarsi Melin, 1941:40 [Holotype.— NHMG 488 from

Taracua, Rio Uaupes, Territorio do Amazonas, Brasil; Donglas Melin

collector], (fide Bokermann, 1966:64.)
Hyla depressa Andersson, 1945:73 [Holotype. — NHRM 1966 from the Rio

Pastaza watershed ( ? Provincia Pastaza ) , Ecuador; William Clarke-Mac-

Intyre 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 )'ellow-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

32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

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 fomi 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 favoJineatus with
taurinus.

The holotype of OsteocephaJus 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 fronto-
parietals are distinctly elevated. The skin on the dorsum is smooth.

HYLID FROGS, GENUS OSTEOCEPHALUS 33

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, con-
tinuing as a broader light-edged one through the eye and tympanum
towards the base of the forelimb and then farther on continuim^
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 some-
what desiccated, especially the hands and feet. The coloration re-
mains 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 Coin (1970:322), who
assigned these names to the synonymy of O. leprieurii. Bokermann
(1966:64) placed Hyla vilarsi in the synonymy of Osteocephalus
taurimis 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 trans-
verse 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.S 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 frontoparie-
tals 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 frojn 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 ex-
amined the type on 3 January 1969, it was in excellent condition.

34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Andersson (1945:75) contrasted the type of Hyla depressa with
Jeprieurii and buckleyi, but he did not compare his specimen with
faurinus, from which it exhibits no distinguishing features.

Osteocephalus taurimis is a widespread and variable species,
and it has received several specific names. We are convinced that
Osteocephalus taurimis Steindachner, 1862, is the oldest available
name for this large Amazonian species. The following names are
junior synonyms: Osteocephalus favolineatus Steindachner, 1862;
Osteocephalus planiceps Cope, 1874; Hyla (Trachycephahis) 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 fre-
quently, 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) pala-
tine 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 diapophyscs; transverse processes of presacral
vertebrae 3-8 subequal in width; 18) intermandibularis and sub-
mentalis muscles connected; 19) supramandibular portion of inter-
hyoideus 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 taurimis 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 as-
cends 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

HYLID FROGS, GENUS OSTEOCEPHALUS

35

500 1000

I I I I I I I I I I I
kilometers

66° '^62°

Fig. 11. Distribution of Osteocephahis taurinus.

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 lo-
calities 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. Further-
more, 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 pre-

36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

served 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 radiat-
ing black streaks and a median, horizontal reddish brown streak."
(W. E. Duellman, field notes, 12 May 1969.) A female from Santa
Ceciha, 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 iden-
tification 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, fomiing a large median blotch, but some adults
retain the juvenile pattern. Cochran and Coin (1970:251)
erroneously identified several juveniles from Colombia as Hyla
palpebrogranulata Andersson.

HYLID FROGS, GENUS OSTEOCEPHALUS 37

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 Tungiirahua, Ecuador; William Clarke-Maclntyre

collector].
Hyla orccsi Funkhouser, 1956:78 [Holotype.— CAS-SU 13150 from Rio

Pacayacu, tributary of Rio Cotapino, Provincia Napo, Ecuador; collector

imknownj.
Osteocephalus orcesi — Cochran and Goin, 1970:317.
Osteocephalus verrucigerus — Triieb and Duellman, 1970:601 [Synonymized

Hyla riopastazae Andersson, 1945, and Hyla orcesi Funkhouser, 1956, with

Hyla verrucigera Werner, 1901].

Justification of Synonymy.— Trueh 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 doi^al
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.— I) Size moderate, sexual dimorphism evident; max-
imum observed snout-vent length in males 54.3 mm, in females 65.8
mm; 2) skin on dorsum in males bearing large, keratinized tuber-
cles; 3) skin on flanks smooth; 4) web extending to base of ante-
penultimate phalange on inner edge of third finger; 5) dorsum uni-
formly 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

38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

widely separated medially; 12) anteromedial margin of fronto-
parietals 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 dia-
pophyses; 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.

Distrilmtion.— 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 sub-
orbital 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 prin-
cipally 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 ) ,

HYLID FROGS, GENUS OSTEOCEPHALUS 39

namely the subgular type and the lateral type. Only four hylicl
genera, all Neotropical lowland groups, are known to possess paired
lateral vocal sacs; these are Osteocephalus, Argenteohyla, Phry-
nohyas, 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. Os-
teocephalus 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), charac-
terized 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 pre-
vomerine dentigerous processes in contrast to the large, angular
processes of Osteocephalus. The skull of Argenteohyla shows spe-
cializations, 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 Phry-
nohyas 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 contnist to Osteocephalus, the dentigerous processes of the
prevomers are curved, rather than angular in Phrynohyas. Further-
more, the latter genus is singularly distinguished from Osteocepha-
lus, Argenteohyla, and Trachycephalus by having extensively de-

40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

veloped 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 pos-
terolaterally, oriented. A dermal sphenethmoid is present, and the
parasphenoid bears odontoids. The basic structure of the skull has
many characters in common with both OsteocepJmJus and Pliry-
nohijas. 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 Trachycephahis is like that of
Phrynohyas and therefore further distinguishes it from Osteo-
cephahis.

Morphologically, Osteocephahis seems to be sufficiently diverse
and generalized so as to represent a modern derivative of an an-
cestral type which might have given rise to Phrynohyas, Trachy-
cephalus, and Argenteohyla. The specialized vocal sac structure
in Phrynohyas and T rachycephalus 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 T rachycephalus 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 forma-
tion 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 :

HYLID FROGS, GENUS OSTEOCEPHALUS 41

Osteoceplmlus buckJcyi.—No l^reeding 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. —\la\es 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 temporaiy 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
bromeiad 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 ob-
served 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 prohoscideus.

At Santa Cecilia many recently metamorphosed young and ju-
veniles 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 metamoiphosed
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 intennediate patterns
were obtained at Lago Agrio in May 1969. Probably juveniles ob-
tained in June and July are the offspring of an April or May breed-
ing. We have been unable to associate tadpoles with this species.

Osteocephalus taurinus.—A small chorus occurred at Lago Agrio

42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

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 amplec-
tant 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.

Osteocephahis 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 fol-
lows: 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 in-
cluded under one museum number, the number of specimens is
given in parentheses after the museum number. Unless noted other-
wise, all specimens are alcoholics.

Osteocephahis buckleyi

BOLIVIA: El Beivi: Ivon, BMNH 1967.2070-1. Satita Cruz: Buenavista,
CM 4333, 4339, UMMZ 66563-5.

BRASIL: Amapd: No specific locality, WCAB 13284.

COLOMBIA: Amazunas: Rio Guacaya, USNM 152759. Huila: Acevedo,

HYLID FROGS, GENUS OSTEOGEPHALUS 43

Rio Suaza, FMNH 69702. Narino: Rumiyacu, FMNH 54756. Mcta: Rio
Guejar, Gampamento La Macaiena, USNM 152199.

ECUADOR: No specific locality, NHMW 6209, WGAB 35499. Chimbo-
razo: Pallatanga, BMNH 1947.2.13.46; Santiago, FMNH 42529. Monma-
Santiaf^o: "Rio Santiago" {= Rio Zaniora), MIZS 2950. Napo: Lago Agrio,
KU 126646; Puerto Libre, Rio Agnarico, KU 123172; Santa Gecilia, AUM
8138, KU 105208-9, 109506, 123171. Pastaza: Alpayacu, BMNH 1912.11.1.64;
Ganelos, BMNH 1947.2.13.40-1, 1947.2.13.43-5; Colonia Mena, Rio Gonambo,
ZSM 33/1962; Don Tonias, USNM 166014; Guache, Rio Pa.staza, AMNH
79986; Rio Bobonaza, USNM 166005; Rio Gapahuari, USNM 166554; Rio
Gonambo at Rio Shiona-yacu, USNM 166018; Rio Gopataza, 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, MGZ 26090, ZMB 10166.

GUYANA: Mazariini-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 Dememra: Dunoon,
UMMZ 52449, 52508.

PERU: Jiinm: Ghanchamayo, BMNH 1911.12.13.79-80. Loreto: Andoas,
AMNH 79984-5; Gashilioya, AMNH 43454; San Antonio, Rio Itaya, AMNH
43218. Puno: Yahuaramayo, BMNH 1913.2.25.7.

SURINAM: Suriname: Powakka, GM 44217.

SOUTH AMERIGA: No specific locality, NHMW 6208.

Osteocephaliis leprieurii

BRASIL: Acre: Tarauaca, FMNH 83247. Amazonas: Rio Javari, Benja-
min Constant, CAS-SU 12620; Rio Uaupes, north of Rio Japu, NHMG 489.

COLOMBIA: Amazonas: Gino-goje, lower Rio Apopoiis, MGZ 28038,
28040-2, 28044, USNM 152136-8.

ECUADOR: No specific locality, WGAB 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, WGAB 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: Ganelos, BMNH 1947.2.13.42,
KU 120915; Rio Alpayacu, UMMZ 92079; Rio Arajuno, USNM 166560-2,
WGAB 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. Iniui: 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.

44 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

SURINAM: No specific locality, MCZ 2036, RMNH 1 1468. Marotvijne:
Camp 3, RMNH 13045-6; Wane Creek North, RMNH 11469-70. Saranmcca:
Right Coppenanie 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.
PERtJ: 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. Sauta Cruz: Buena-
vista, 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 locaUty: "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 Mannelo, 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; Randal de la Playa, lower Rio Apoporis, MCZ
28050; Rio Apoporis, MCZ 28060. Boijacd: 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. Esmcraldas: Carondelet,
UIMNH 53560-9; Lagartera, Rio Caoni, UIMNH 53441, 53458-79. Morona-
SantiaRo: Macuma, UIMNH 63142-3, 63145, 63147, 63151, 63154, 63157,
USNM 166060. }^apo: 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;

HYLID FROGS, GENUS OSTEOCEPHALUS 45

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 Ollagiianga, 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 SoHs, 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. Mazamni-Potaro: Chinapora River,
upper Potaro River, BMNH 1905.11.1.20-1; Kamakusa, AMNH 21416, 21418-9,
21422; Kartalio, 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 Di.strict, BMNH 1905.11.1.40; Rockstone, FMNH 26591. North
West: Amakura River, Haulover, UMMZ 83735. Ruptinuni: 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. Huaniico: Monte Alegre,
Rio Pachitea, AMNH 43014, 43019. Loreto: Achinamisa, Rio Huallaga,
AMNH 42178, 42502; Andoas, Rio Pastaza, AMNH 79991; Cashiboya, AMNH
43388, 4.3453; Estiron, Rio Ampiyacu, CAS 93264-74, 9.3276, 93278-9, 93281,
93283-6, 93289, 93311, 93327; Igarape Champuia, upper Rio Curiuja, MZUSP
10339; Iquitos, AMNH 42204, 42442, 4.3468, 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 Brasiiian frontier,
AMNH 43137; Sobral, Rio Tamaya, AMNH 43242; Yurimaguas, BMNH
84.2.18.50. San Martin: Cainarachi, AMNH 42763; Moyobamlia, ZSM
19/1914.

SURINAM: No specific locality, BMNH 70.3.10.67, NHMW 18433.3.
Brokopondo: Afobaka, RMNH 16.536; Brownsweg, RMNH 16537; Railway km.
121, RMNH 16534. Maroivijne: Djai Creek, RMNH 16513-4; Maroni River,
ZMB 8240, 8.531; Nassaugebergte, RMNH 16517-33; Paloemeu, USNM
159025; Swamp Camp, RMNH 16515. Nickerie: Sipaliwini, RMNH 16538.
Sararnacca: Left Coppename River, RMNH 165.35; Tibiti, RMNH 16516.

46 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Suriname: Berlijn, RMNH 15064; Powakka, CM 44226; Zanderij, CM 50568.
VENEZUELA: Amazonas: Cerro Diiida, UPR-M 2875; Cerro Marahuaca,
UPR-M 114-5; Esmeralda, AMNH 23174; Iniridi, SMF 2640; La Culebra,
MCZ 28572, UPR-M 117; Lagiina, liet\\'een Tama Tama and Esmeralda,
UPR-M 2760; Rio Pescado, AMNH 23177; Tapara, UPR-M 113. Distrito
Federal: Caracas, BMNH 51.7.17.182.

Osteocephalus vernicigerus
COLOMBIA: Htiila: Acevedo, Rio Suaza, FMNH 69709-10.
ECUADOR: No specific locality, ZMB 16589. Napo: Axila, 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. Pasiaza: Abitagua, CAS-SU 5067, FMNH 25791,
27619, UMMZ 90414, 92092; AJpayaca, Rio Pastaza, BMNH 1912.11.1.64;
Mera, UMMZ 90412(4). Tungurahua: Banos, NHRM 1960.

PERC: Aijacucho: La Mar, Sivia, Rio Apurimac, FMNH 39853. Huannco:
Rio Pachitea, midway between Puerto \'ictoria and Puerto Inca, CAS-SU
17745. Junin: Satipo, MJP 38.

LITERATURE CITED

Andersson, L. G.

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

BOKERMAXX, 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 Briti.sh Museum, ed. 2, London, xvi+503 pp.
CocHRAX, D. M. and C. J. Coix

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.
Duellmax, W. E.

1970a. Identity of the South American hylid frog Garheana garhei. Copeia,

(3):534-538.
1970b. The hylid frogs of Middle America. Monog. Mus. Nat. Hist., LTniv.

Kansas, l:.\i+753 pp.

Dumeril, a. M. C. and G. Bibrox

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

FiTZIXGER, L.

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

HYLID FROGS, GENUS OSTEOCEPHALUS 47

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.

Goix, G. J.

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

1941. Contribution to the knowledge of Amphibia of South America.
Goteborgs 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. Gomp., Univ. Torino, 19:1-41.

Steixdachxer, F.

1862. Uber zwei noch unbeschriebene Batrachier. x\rch. Zool. Anat.

Fisiol., 2:77-82.
1867. Amphibien. Novara Expedition. Zool. Theil, 1, Vienna, 70 pp.
Truer, L.

1970a. The evolutionary relationships of casque-headed treefrogs with
co-ossified skulls (family Hylidae). Univ. Kansas Publ. Mus. Nat.
Hist., 18:547-716.
19701). The generic status of Hyla siemersi Mertens. Herpetologica,
26:254-267.
Truer, L. and W. E. Duellman

1970. The systematic status and life history of Hijla vernicigera 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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