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OCCASIONAL PAPERS

MUS. COMP. ioou
LrBRARY

NOV 12 1971

MUSEUM OF NATURi4tt<Mi»nibRY
The University of Kansas
Lawrence, Kansas

NUMBER 4, PAGES 1-21 JUNE 3, 1971

A TAXONOMIC REVIEW OF SOUTH AMERICAN
HYLID FROGS, GENUS PHRYNOHYAS

By

William E. Duellaian^

Few large South American hylicl frogs are so poorly known
taxonomically as the widespread Fhnjnolujas vemilosa (Laurenti,
1768) and its cogeners. Duellnian (1956) included three South
American species in the genus; four other species were recognized
in Cenhal America and Mexico. Duellman (1966) and McDiarmid
(1968) reduced all of the Middle American nominal species to
synonyms of Fhrynohijas venulosa. Duellman (1968) resurrected
Htjla coriacea Peters from the synonymy of Fhnjnohijas venulosa
and showed that coriacea was a valid species of Phnjnoliyas in the
Guianas and Amazon Basin. Bokermann (1966) listed Ilyla meso-
pliaea Hensel and Hyla imitatrix Miranda-Ribeiro in the genus
Plirynoliyas.

The purposes of the present paper are to : 1 ) redefine the hylid
genus Plirynoliyas, 2) describe the geographic variation in the wide-
spread Plirynoliyas venulosa, 3) ascertain the assignments of several
nominal species, and 4) present a synopsis of the species in the genus.

Specimens examined are designated by the following abbrevi-
ations of depositories:

AMNH American Museum of Natural History

ANSP Academy of Natural Sciences, Philadelphia

BMNH British Museum (Natural History)

CAS California Academy of Sciences

CAS-SU Stanford University collection, now in the California Academy of
Sciences

FMNH Field Museum of Natural History

'Curator, Division of Herpetology, Museum of Natmal History, University
of Kansas.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

IRSNB Institut Royal des Sciences Naturelles Belgique

KU University of Kansas Museum of Natural History

LACM Los Angeles County Museum of Natural History

MCZ Museum of Comparative Zoology, Harvard

MLS Museo de La Salle, Bogota

MJP Museo Javier Prado, Lima

MNHN Museum National d'Histoire Naturelle, Paris

MNRJ Museu Nacional, Rio de Janeiro

MZUSP Museu de Zoologia, Universidade de Sao Paulo

NHMG Naturhistoriska Museet, Goteborg

NHMW Naturhistorisches Museum, Wien

NHRM Naturhistoriska Riksnmseet, Stockholm

RMNH Riksmuseet van Natuurlijke Histoire, Leiden

SMF Senckenbergische Museum, Frankfurt

UIMNH University of Illinois Museum of Natural History

UMMZ University of Michigan Museum of Zoology

USNM United States National Museum

UZM Universitets Zoologiske Museum, Copenhagen

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

ZIUS Zoologiske Institut, Universitet Stockholm

ZMB Zoologisches Museum, Berlin

ZSM Zoologisches Staatssammlung, Munich

ACKNOWLEDGMENTS

For permission to examine specimens in their care, I am grateful
to James E. Bohlke, Werner C. A. Bokermann, F. W. Braestrup,
Nelly Carrillo de Espinoza, Gaston F. deWitte, Josef Eiselt, Alice
G. C. Grandison, Jean Guibe, Birgitta Hansson, Walter Helbnich,
M. S. Hoogmoed, Robert F. Inger, Konrad Klemmer, Alan E.
Leviton, Bertha Lutz, C. J. McCoy, Charles W. Myers, Hno.
Niceforo Maria, Giinther Peters, James A. Peters, Dorothy M. Smith,
Paulo E. Vanzolini, Greta Vestergren, Charles F. Walker, Ernest E.
Williams, John W. Wright, and Richard G. Zweifel.

I am especially indebted to Linda Trueb for her astute osteo-
logical observations and illustrations incorporated herein and for her
aid in examining specimens in many museums. Study of specimens
in European museums was made possible by a grant (No. 5063)
from the Penrose Fund of the American Philosophical Society.

PHRYNOHYAS FITZINGER, 1843

Phrtjnohtjas Fitzinger, 1843:30 [type species, Hyla zonata Spix, 1824 {= Rana
venulosa Laurenti, 1768) by monotypy and declaration of International
Commission on Zoological Nomenclature (Opinion No. 520, 1958)].

Acrodytes Fitzinger, 1843:30 [type species, Hyla venuJosa Daudin, 1803
(= Rana venidosa Laurenti, 1768) by monotypy; generic name suppressed
by International Commission on Zoological Nomenclature ( Opinion No. 520,
1958) for purposes of Law of Priority, but not for those of Law of
Homonymy] .

HYLID FROGS, GENUS PHRYNOHYAS 3

Scytotm Cope, 1862:354 [type species, Scytopis hebes Cope, 1862, by
monotypy] .

Diagnostic Definition.— 1) Skull slightly broader than long; 2)
dermal roofing bones of skull well ossified, lacking exostosis and
co-ossification; 3) nasals broad, overlapping anterior edge of sphen-
ethmoid, not articulating with maxillaries: 4) frontoparietals hav-
ing complete medial articulation, entirely covering frontoparietal
fontanelle; posterolateral margin of frontoparietal lying just medial
to epiotic eminence; 5) sphenethmoid well ossified with large portion
exposed dorsally; 6) maxillary arch complete; pars facialis of maxil-
lary moderately well developed but not ardculating with nasal;
7) squamosal weakly articulating with prootic; zygomaHc ramus
extending about one-half of distance to maxillary; 8) pterygoid
robust with complete articulations; 9) dentigerous processes of pre-
vomers curved; 10) parasphenoid large, edentate, acuminate anteri-
orly; alae oriented posterolaterally; 11) palatines moderately large,
widely separated medially, bearing dentigerous ridge; 12) neural
arches of vertebrae low, non-imbricate; 13) transverse processes of
vertebrae subequal in width; width of processes on third presacral
vertebra less than width of sacral diapophyses; 14) sacral diapophy-
ses expanded; 15) ilial shaft long, cylindrical, with low dorsal pro-
tuberance; 16) pubis cartilaginous; 17) clavicles expanded distally,
strongly arched; 18) coracoids expanded distally and proximally,
widely separated medially; 19) epicoracoids broadly overlapping;
20) suprascapula large, well ossified; cleithrum ossification incorpo-
rated into suprascapula; 21) vocal sacs paired, lateral, behind angles
of jaws; 22) submentalis muscle moderate in size, araphic; 23) inter-
mandibularis muscle undifferentiated, lacking aponeurosis; 24) in-
terhyoideus muscle forming large supramandibular lobes; no
myointegumental contact between lobes and sacs; 25) postman-
dibular septum short, attached to posterior end of intermandibular
portion of interhyoideus; 26) length of vocal slits about one-fifth of
length of mandible; 27) skin on dorsum thick, glandular, and
smooth, granular, or tuberculate; that on venter heavily granular;
28) parotoid glands extensively developed in occipital and scapular
regions; 29) diameter of tympanum 50-75 percent of diameter of
eye; upper edge covered by supratympanic fold; 30) discs large,
round; 31) fingers one-fourth to two-thirds webbed; toes about
three-fourths webbed; 32) smooth, horny nuptial excrescences in
breeding males; 33) pupil horizontal; 34) palpebrum clear; 35)
snout-vent length of adults 50-114 mm; 36) known tadpoles havmg
four upper and six lower rows of teeth.

OCCASIONAL PAPERS MUSEUM OF NATURAL IILSTORY

80

50

H ^ coriacea

^ P imitatnx

ik. R mesophaea

0 P venu/oso

-10

-20

Fig. L Map showing locality records for the species of Phrynohyas.

Content. — As defined here, die genus contains four species: P.
coriacea (Peters), P. imitatrix (Miranda-Ribeiro), P. mesophaea
(Hensel), and P. venulosa (Laurenti, 1768).

Distribution. — The lowland tropics from Sinaloa and Tamaulipas,
Mexico, southward to Chaco, Entre Rios, and Salta, Argentina, and
Rio Grande do Sul, Brasil. The genus occurs on Trinidad and
Tobago, but only two specimens are known from the Pacific low-
lands of South America ( Fig. 1 ) . Most records of occurrence are at
elevations of less than 500 m, but the frogs ascend the Amazonian
slopes of the Andes to about 800 m and are known to occur at
elevations of 1200 m in the Serra do Mar and Serra da Mantiqueira
in southeastern Brasil.

Comments. — As pointed out by Trueb and Duellman (1971:38),
only four h>'lid genera ha\e paired lateral \ocal sacs; these are
ArgentcoJu/la, Osteoceplwlus, PJirynolnjas, and Trachijcephahis.
Tyler (1971:349) provided evidence that the paired lateral vocal
sacs were derived from single subgular sacs and stated: "Conceiv-
ably the very highly specialized vocal sacs of Phrynoliyas and
Traclujcepliahis could have been derived from the simpler structures
now found in some species of Osteocephalus." Trueb and Duellman
(1971:40) concluded that Phrynohijas and Trachycephalus probably

HYLID FROGS, GENUS PHRYNOIIYAS 5

were representatives of one phyletic line. Trachycephalus differs
from Plmjnohyas principally by having a hea\dly casqued, co-ossified
skull and by lacking extensive parotoid glands.

GEOGRAPHIC VARIATION IN
PHRYNOHYAS V EN U LOS A

In 1956 I noted the presence of variation in size and coloration
in P. vemdosa in the Amazon Basin; furthermore, I recognized,
taxonomically two peripheral populations — P. hebes in the Gran
Chaco and P. ingens in the Maracaibo Basin. Examination of many
additional South American specimens reveals that the mosaic of
variation is more complex than that described previously.

Size.— In South America very large individuals of P. vemdosa
occur in the Maracaibo Basin (females to 112.5 mm in snout-vent
length) and in Amazonian Ecuador (females to 110.2 mm in snout-
vent length). Equally large individuals are known from Panama
and Costa Rica (Duellman, 1970:165). Specimens from other areas
in South America are noticeably smaller ( Table 1 ) . Measurements
and proportions of the limbs and head all are positively correlated
with snout-vent length. Duellman (1956) suggested that there were
interpopulational differences in the size of the tympanum, but data
from larger series of specimens negate that suggestion (Table 1).
Also, there seems to be no correlation between size and habitat, as
suggested by McDiarmid ( 1968 ) .

S/c/n.— Considerable variation is apparent in the thickness and
texture of the dorsal skin. Most specimens have scattered pustules
dorsally; these are especially noticeable in young individuals. In a
few specimens, such as KU 126067 from 2 km east of Leticia, Co-
lombia, and especially MNRJ 4054 from Rio Branco, Brasil, the skin
on the dorsum is highly pustulate. The parotoid glands result in
thickened skin in occipital and scapular regions in most specimens
from throughout the range. Many specimens from the upper Ama-
zon Basin in Ecuador, Peru, and Bolivia have thick, leatliery skin.
McDiarmid (1968) suggested that extensive development of the
parotoid glands is a seasonal phenomenon in frogs living in regions
having wet and dry seasons, the glands being better developed in
the dry season. Dates of collection are not available for many South
American specimens of P. vemdosa; the specimens having such data
do not provide strong evidence in support of McDiarmid's proposal.
For example, well-developed glands are evident in specimens from
the upper Amazon Basin where there is no prolonged dry season;

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

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HYLID FROGS, GENUS PIIRYNOIIYAS

A

^"^■^^^il:^

Fig. 2. Dorsal color patterns in Phrtjnolnjas vcnnlosa. A. Normal, code 4, KU
92258. B. Divided, code 3, KU 12fi()fi7. C. Spotted, code 2, NHMW 19013

(holotype of Hyla wettsteini).

specimens from Napo and Pastaza provinces in Ecuador have more
extensive glands than do individuals from the xeric Maracaibo Basin.
Coloration. — All individuals of P. venidom are brown or tan
dorsally with or without darker markings. The venter is uniform
grayish white or pale tan. The iris is golden with radiating black
lines. There is considerable variation in dorsal color pattern, but
most of the variants can be grouped into one of four categories; code
numbers indicate a decreasing amount of dark pigment dorsally
(Fig. 2):

NORMy\L: A large middorsal dark blotch extending from the
occiput to the rump; blotch divided anteriorly by a longitu-
dinal band of ground color in some specimens ( coded 4 ) .
DIVIDED: As above, but blotch divided by a broad transverse

band of ground color (coded 3).
SPOTTED: Dorsal pattern consisting of irregular small dark

spots (coded 2).
PLAIN: Dorsum uniform brown (coded 1).

The various color patterns are found throughout most of the
range of the species in South America ( Table 2 ) . The only region in
which all specimens have the normal pattern is Paraguay; the next
highest percentage of specimens having the normal pattern is Trini-
dad ( 93';'o ) . All four color pattern types were found in only four of
14 pooled samples; normal and plain patterns occurred in 12 sam-
ples, and divided and spotted patterns occurred in eight samples.
In the only large sample ( 151 specimens from Maracay, Venezuela)
the following percentages occur: normal 14.6, divided 44.3, spotted
7.9, and plain 33.2. Each of the various patterns has been found in
males, females, and juveniles. All four patterns, save the spotted
one, are found in Central America.

8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Table 2. — Variation in Dorsal Color Pattern in Phrynohyas venulosa.
See text for explanation of patterns and scoring.

Locality N Normal Divided Spotted Plain Score

Trinidad 32 30 1 1 0 SM

Venezuela: Maracaibo Basin ... 8 0 0 7 1 1.89

Venezuela: Maracay ______ ____ 151 22 67 12 50 2.38

Surinam 21 15 0 2 4 3.28

Brasil: Belem 26 23 1 0 2 3.73

Brasil: Mato Grosso 15 2 0 12 1 2.20

Brasil: Manaus _ 8 3 3 0 2 2.87

Colombia: Leticia 5 110 3 2.00

Ecuador: Napo-Pastaza 13 0 1 0 12 1.25

Peru: Pebas 10 4 0 4 2 2.60

Peru: Pucallpa 12 6 0 0 6 2.50

Bolivia: El Beni 16 1 1 2 12 1.44

Bolivia: Santa Cruz 15 4 1 3 7 2.13

Paraguay 14 14 0 0 0 4.00

Remarks. — It is evident that the various characters considered to
be of taxonomic value in this group vary independently of one
another in P. venulosa. Refined quantitative analysis must await the
acquisition of large series of specimens from throughout the range
of the species. Currently, karyological, larval, and biosonic data are

Table 3. — Alphabetical Synonymy of the Species of Phnjnohyas.
Trivial Name, Original Generic Name, Author, and Date Current Name

adenodcrma (Hyla) Lutz, 1968 P. venulosa

biifona (Hyla) Spix, 1824 P. venulosa

corastcrias (Phrynohyas) Shannon and Humphrey, 1957 P. venulosa

coriacea (Hyla) Peters, 1867 P. coriacea

hehcs (Scytopis) Cope, 1862 P. venulosa

imitatrix (Hyla) Miranda-Ribeiro, 1926 P. imitatrix

inflata (Acwdytes) Taylor, 1944 P. venulosa

inpens (Phrynohyas) Duelhnan, 1956 P. vemdosa

latifasciata (Phrynohyas) Duelhnan, 1956 P. venulosa

lichenosa (Hyla) Giinther, 1859 _ _. P. venulosa

macrotis (Hyla) Andersson, 1945 P. vemdosa

mcsophaea (Hyla) Hensel, 1867 P. mesophaea

modesta (Acrodytes) Taylor and Smith, 1945 P. venulosa

nigropunctata (Hyla) Boulenger, 1882 P. venulosa

paemdata (Hyla) Brocchi, 1879 P. venulosa

palpchrogranulata (Hyla) Andersson, 1906 P. venulosa

quadrangulum (Hyla) Boulenger, 1882 - P. coriacea

resinifictrix (Hyla) Goeldi, 1907 ____ - P- venulosa

spdomma (Hyla) Cope, 1877 P- venulosa

venulosa (Rana) Laurenti, 1768 P. venulosa

vermiculata (Hyla) Dumeril and Bibron, 1841 P. vemdosa

wettstemi (Hyla) Ahl, 1933 P- venulosa

zernyi (Hyla) Ahl, 1933 P- venulosa

zonata (Hyla) Spix, 1824 P- venulosa

HYLID FROGS, GENUS PHRYNOHVAS
B /T\ C y^r\ D

9

Fig. 3. Anterior cranial roofing bones showing positions of nasals and proportion

of dorsal exposure of the sphenethnioid. A. P. coriacea, KU 105249. B. P.

puitatrix, KU 92249. C. P. mcsophaea, KU 92257. D. P. venulosa, KU 138830.

Abbreviations: fp = frontoparietal, n = nasal; sphenethnioid is stippled.

lacking. Until such data are available for a more sophisticated
analysis, we must be content to base conclusions on the present in-
adequate series of preserved specimens, which have formed the basis
for my broad definition of Fhrijuohyas venulosa.

TAXONOMY OF PHRYNOHYAS

Data taken from series of preserved specimens, some living frogs,
skeletal preparations, and radiographs provide a basis for the recog-
nition of four species in the genus Fhnjnohijas ( Table 3) . Diagnostic
external morphological characters and features of coloration are used
in the following key. Osteological characters that are important at
the species level are discussed below.

Nasah. — The size and position (relative to one another) of the
nasals are distinctive (Fig. 3). In P. imitatrix and venulosa the
nasals are juxtaposed; in 7nesop1iaea they are narrowly separated,
and in coriacea they are widely separated.

Sp]}enetlimoid. — The sphenethnioid is well ossified in all species.
The proportion of the element exposed dorsally shows interspecific
differences ( Fig. 3 ) ; the area of exposure is dependent on the rela-
tive sizes and positions of adjacent dermal roofing bones — the nasals
anteriorly and the frontoparietals posteriorly. The sphenethnioid is
least exposed in P. imitatrix and moderately exposed in mesophaea
and venulosa. A relatively large area of the sphenethnioid is exposed
in coriacea, in which the nasals are widely separated.

Prevomers. — The dentigerous processes are curved and separated
medially in all species. The curvature is slight and the separation
moderate in P. imitatrix and venulosa, slight and widely separated in
coriacea, and strongly curved and narrowly separated in mesophaea.

Sacral Diapophyses. — The diapophyses are dilated in all species;
this dilation is symmetrical in P. mesophaea and venulosa and asym-
metrical (greater dilation posteriorly) in imitatrix and coriacea.

10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Key to the Species of Plinjnolnjas

1. Dorsal quadrangular mark outlined \\'ith white and terminating
in sacral region; thighs lacking dark transverse bars; black post-
tympanic spot present; red "flash-colors" in life P. coriacea

Dorsal cjuadrangular mark, if present, usually not outlined with
white; thighs usually having dark transverse bars (if blotches
present on dorsum); black post-tympanic spot absent; no red
"flash-colors" in life 2

2. Dorsum and flanks dark brown with broad cream dorsolateral
stripe from eyelid to groin; limbs unicolor or faintly barred
dorsally P. mesophaea

Dorsal pattern variable, but lacking dorsolateral cream strips;
limbs usually distinctly barred, if blotch present on dorsum -— 3

3. Dorsal markings consisting of large brown blotch anteriorly and
usually round spots or blotches posteriorly, darkest peripherally;
limbs with distinct dark brown, light bordered bars; dorsal skin
distinctly tuberculate .> P. imitatrix

Dorsal markings absent or consisting of large dark blotch ( trans-
versely divided in some) or small dark spots; limbs unicolor or
barred, lacking light borders; dorsal skin variable P. vemdosa

Phrynohyas coriacea (Peters)

Hyla coriacea Peters, 1867:711 [syntypes, ZMB 5807 (3 specimens) from

Surinam; H. Kappler collector].
Hyla quadmn^ulum Boulenger, 1882:367 [holotype, BMNH 1947.2.13.75 from

"western Ecuador"; Mr. Eraser collector]. New synonymy.
Phrynohyas coriacea — Duellman, 1968:205.

Distribution. — Surinam and the upper Amazon Basin in Ecuador
and Peru at known elexations of less than 500 m ( Fig. 1 ) . The record
for Machala on the Pacific coast of Ecuador probably is erroneous.
29 specimens from 11 localities.

Remarks. — The three syntypes (ZMB 5807) were reported on by
Duellman (1968). The holotype of Hyla quadrangulum (BMNH
1947.2.13.75) is an adult female having a snout- vent length of 57.5
mm and morphological characters which render it indistinguishable
from P. coriacea. The color pattern consists of a dark brown qua-
drangular dorsal blotch faintly outlined with cream on a tan dor-
sum; the limbs arc distinctly barred, and a dark post-tympanic spot
is evident.

HYLID FROGS, GENUS PHRYNOHYAS 11

The type locality of "western Ecuador" cannot be taken seriously;
several species contained in the Fraser collections supposedly from
western Ecuador have been found subsequently only in the Amazon
Basin and eastern slopes of the Andes.

Phrynohyas imitatrix (Miranda-Ribeiro)

Hyla imitatrix Miranda-Ribeiro, 1926:77 [syntypes, MNRJ 154 (2 specimens)
from Teresopolis, Rio de Janeiro, Brasil; Alipio de Miranda-Ribeiro col-
lector].
Phrynohtjafi mcsophaca — Bokermann, 1966:55.

Distribution. — Mountains of southeastern Brasil — Serra do Mar,
Serra da Bocaina, and Serra da Mantiqueira — in the states of Rio de
Janeiro, Sao Paulo, and Santa Catarina ( Fig. 1 ) ; the known localities
are at elevations between 800 and 1200 m. Eight specimens from
three localities.

Phrynohyas mesophaea (Hensel)

Hyla mesophaea Hensel, 1867:154 [holotype, ZMB 6810 from Porto Alegre, Rio

Grande do Sul, Brasil; Reinhold Hensel collector].
Phrynohyas mesophaea — Bokerman, 1966:55.

Distribution. — Coastal lowlands of southeastern Brasil from
Bahia to Rio Grande do Sul (Fig. 1). 259 specimens from 13
localities.

Phrynohyas venulosa (Laurenti)

Rana vemdosa Laurenti, 1768:31 [based on a plate in Seba ( 1734, vol. 1, pi. 72,

fig. 4)].
Hyla vemilosa — Daudin, 1803:71.
Hyla zonata Spix, 1824-41 [type formerly in ZSM, now lost, from Lago Tefe,

Amazonas, Brasil; collector rmknown].
Hijla hufonia Spix, 1824:42 [type formerly in ZSM, now lost, from Eca (= Tefe),

Amazonas, Brasil; collector unknown].
Hyla vermiculata Dumeril and Bibron, 1841:563 [holotype, MNHN 4797 from

"Amerique Sept."; presented by Mr. Harlan].
Phrynohyas zonata — Fitzinger, 1843:30.
Acrodijtes venulosa — Fitzinger, 1843:30.
Hyla lichenosa Giinther, 1859 [lectotype, BMNH 1936.12.3.119 from Amazonas,

Brasil; collector unknown].
Scytopis hehes Cope, 1862:354 [holotype, USNM 5837 from "Paraguay" (re-
stricted to Asuncion by Duellman, 1956:16); T. J. Page collector]. New

synonymy.
Scytopis veniilosus — Cope, 1866:85.
Hyla spilomma Cope, 1877:86 [holotype, unknown, apparently lost, from

Cosamaloapam, Veracruz, Mexico; Francis Sumichrast collector].
Hyla paenulata Brocchi, 1879:21 [holotype, formerly in MNHN, now lost, from

western {=. southern ?) Guatemala; collector unknown].
Hyla nigroptinctata Boulenger, 1882:366 [syntypes, BMNH 59.9.20.2 and

81.10.31.20 from Jalapa, Veracruz, Mexico; Alphonse Hoege collector].
Hyla palpehrogranulata Andersson, 1906:14 [holotype, ZIUS A133 fnmi

Tatarenda, Santa Cruz, Bolivia; Erland Nordenskiold collector]. New

synonymy.

12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Hijla resinifictrix Goeldi, 1907:135 [holotype, BMNH 1947.2.23.24 from San

Antonio do Prata, Para, Brasil; Emil Goeldi collector].
Htjla zerntji Ahl, 1933:27 [holotype, NHMW 16503 from Taperinha, near

Santarem, Para, Brasil; H. Zeniy collector]. New synonymy.
Hijla zerijni Alil, 1933:27 [holotype, NHMW 16503 from Taperinha, near

Santarem, Para, Brasil; H. Zerny collector]. New Synonymy.
Acrodytes infkita Taylor, 1944:63 [holotype, FMNH 100046 from La Venta,

Guerrero, Mexico; Edward H. Taylor collector].
Acrodytes spilomma — Taylor, 1944:64.
Acrodytes modesta Taylor and Smith, 1945:594 [holotype, USNM 115013 from

Cruz de Piedra, near Acacoyaqua, Chiapas, Mexico; Hobart M. Smith

collector].
HyJa macrotis yVndersson, 1945:70 [holotype, NHRM 1958 from the Rio Pastaza

watershed, Ecuador; William Clarke-Maclntyre collector]. New synonymy.
Phrynohyas hches — Duellman, 1956: 16.
Plirynohyas iuflata — Duellman, 1956:19.
Phrynohyas mgens Duellman, 1956:22 [holotype, UMMZ 55570 from La Fria,

Pueblo Nuevo, Zulia, Venezuela; H. B. Baker collector]. New synonymy.
Phrynohyas latifasciata Duellman, 1956:24 [holotype, BMNH 83.2.7.1 from

Presidio, Sinaloa, Mexico; Alfonso Forrer collector].
Phrynohyas modesta— DueWman, 1956:25.
Phrynohyas spilomma — Duellman, 1956:28.
Phrynohyas corastcrias Shannon and Humphrey, 1957:15 [holotype, UIMNH

67060 from . 4.8 miles east of San Bias, Nayarit, Mexico; Francis L.

Humphrey and Frederick A. Shannon collectors].
PJirynohyas vcmdosa — Hemming, 1958:172.
HyJa tihiatrix tibiatrix — Rivero, 1961:127.
Hyhi tihiatrix in^^ens — Ri\ero, 1961:131.
Hyla ventdosa vcmdosa — Ri\ero, 1964:493.
HyJa adenoderma Lutz, 1968:3 [holotype, MNRJ 4054 from Rio Branco, a

tributary of the Rio Madeira, Rondonia, Brasil; F. Casper collector]. New

syiion^iny.

Distribution. — In South America — the Caribbean lowlands, in-
cluding Trinidad and Tobago, the Amazon Basin, the Chaco, and
northeastern Brasil ( Fig. 1 ) . Only one specimen is known from the
Pacific lowlands; Duellman (1956:42) reported that specimen as
being from Hacienda San Miguel, Milagro, Guayas Province,
Ecuador, but James A. Peters, the collector, informed me that it
actually came from 7 kilometers southeast of Buenavista, El Oro
Province, Ecuador. All specimens are from elevations of less than
800 m. 475 specimens from 107 localities.

Remarks. — This widespread, variable species has received 19
names in 200 years. The earliest name, Rana ventdosa Laurenti,
1768, was based on a plate in Seba's "Thesaurus" (1734). Type
specimens of Hyla zonata Spix, //. hufonia Spix, H. spilomma Cope,
and H. paenulata Brocchi are no longer extant. I have examined the
types of the other 14 nominal species. Duellman ( 1961 ) provided
evidence that P. corastcrias Shannon and Humphrey, 1957, was not
specifically distinct from P. infata ( Taylor, 1944 ) . Duellman ( 1966)
showed that P. modesta (Taylor and Smith, 1945) was a color morph

HYLID FROGS, GENUS PHRYNOHYAS 13

of P. spilomma (Cope, 1(S77). McDiarmid (1968) analyzed variation
in Middle American PJnynolnjas and concluded that P. spilomma
(Cope, 1877), P. infiata (Taylor, 1944), and P. latifasciaia Ducllman,
1956, were synonymous with P. vemilosa (Laurenti, 1768). My re-
cent studies (Duellman, 1970) support McDiarmid's conclusions;
all Middle American populations seem to be correctly allocated to
Plirynolujas vemilosa.

Of the names in the foregoing synonymy, Duellman ( 1956) listed
Hijla vermictdata Dumeril and Bibron, H. licJwnosa Giinther, and
//. resinifctrix Goeldi in the synonymy of Phrynohyas zonata (Spix)
[= P. vemilosa (Laurenti)]. Duellman (1956) included Hyla nigro-
piinctata Boulenger in the synonymy of Phrynohyas spilomma
(Cope), now considered to be a synonym of P. vemilosa (Laurenti).

Duellman (1956) distinguished Phrynohyas hehes from vemilosa
by the wider dark limb bands in the former. Examination of addi-
tional material from throughout the range of vemilosa and especially
from Argentina, Bolivia, Paraguay, and southern Brasil reveals that
the supposed differences between vemilosa and hehes are not con-
stant. Thus, I now consider Scytopis hehes Cope, 1862, to be a
junior synonym of Phrynohyas vemilosa (Laurenti, 1768).

Duellman ( 1956) distinguished Phrynohyas ingens from vemilosa
chiefly on the basis of the larger size of the former and by its dorsal
coloration consisting of small brown spots on a tan ground color, as
opposed to a large dark blotch on the dorsum in vemilosa. It is now
apparent that the coloration considered to be diagnostic of ingens
appears in populations throughout most of the range of vemilosa
(Table 2). Furthermore, the size of ingens is equaled or exceeded
by some individuals from Ecuador and Costa Rica. Consequently,
I now consider Phrynohyas ingens Duellman, 1956, to be a junior
synonym of Phrynohyas vemilosa (Laurenti, 1768).

The holotype of Ilyla vermiculafa Dumeril and Bibron (MNHN
4797) is an adult male having a snout- vent length of 65.2 mm. The
specimen is soft and has been abraded. The dorsum is medium
brown with distinct reddish brown marks; dark marbling is evident
on the flanks and posterior surfaces of the thighs. The remnants of
the color pattern are reminiscent of one of the color morphs of
Phrynohyas vemilosa common in northeastern South America.

Hyla lichenosa Giinther was based on several juveniles; Smith
and Taylor (1948:75) designated BMNH 1936. 12..3. 119 as the lecto-
type. This specimen has the usual tuberculate skin of young
Phrynohyas, and it has a color pattern resembling that of young P.
venulosa from Belem, Para, Brasil.

14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

The holotype of Htjhi resinifictrix Goeldi (BMNH 1947.2.23.24)
is a male having a snout-vent length of 75.3 mm. The color pattern
is well preserved. The ground color is golden tan; anteriorly, on the
dorsum, there is a reddish brown blotch, which is narrowly sepa-
rated from a large, posterior blotch. The limbs bear distinct cross
bars; the skin on the dorsum is distinctly pustulate. In features of
sti-ucture and coloration the holotype is like most individuals of P.
venulosa from the lower Amazon Basin.

The holotype of Hijla palpebro^ronukita Andersson (ZIUS A133)
is a juvenile having a snout-vent length of 32.3 mm. The skin on the
dorsum is tubercular. The dorsum is yellowish tan with a large
brown blotch beginning on the head and extending nearly to the
vent. Broad brown cross bars are present on the limbs. A pair of
pigment clusters above the anal opening suggest that the specimen
is a recently metamorphosed young. The iris has the markings
characteristic of P. venulosa; broad black streaks radiate from the
pupil. This juvenile specimen certainly is a Vhnjnohyas venulosa.
Cochran and Coin (1970) recognized a monotypie "Hyla palpebro-
granuliita group." I have examined all of the Colombian specimens
that they referred to //. palpehrogranulata and find that all are
young Osteocephalus taurinus.

The holotype of Hyla icettstemi Ahl (NHMW 19013) is a female
having a snout-vent length of 61.3 mm and a moderately tuberculate
dorsum. The dorsal surfaces of the head, body, and limbs are pale
brown with scattered small brown spots (Fig. 2c). The pattern is
identical to that of many individuals of P. venulosa from the middle
and upper Amazon Basin and from Venezuela and Guyana.

The holotype of Hyla zernyi Ahl (NHMW 16503) is a juvenile
having a snout-vent length of 20.4 mm and only slightly tubercular
skin on the dorsum. The dorsum is pale tan with a narrow, elongate
brown blotch beginning on the eyelids and extending to the sacrum.
The flanks are brown, and the limbs are marked with irregular dark
brown cross bars. The upper lips are boldly marked by three vertical
bars, the first two of which fuse with a broad, diffuse canthal stripe,
whereas the third enters the orbit. A broad, postorbital brown stripe
encompasses the tympanum and is confluent with the dark flanks.
The edge of the lower jaw is dark brown interrupted by creamy
white spots. In structure, this specimen is like juvenile P. venulosa;
the color pattern is bold but falls within the range of \ariation of
P. venulosa

The holotype of /////(/ macrotis Andersson (NHRM 1958) is an
adult female having a snout- vent length of 110.2 mm. The skin on

HYLID FROGS, GENUS PHRYNOHYAS 15

the dorsum is thick and glandular; the parotoid glands are extensive,
diffuse, and form a moderately heavy supratympanic fold. The
dorsum is uniform reddish brown; the loreal region is slightly paler
brown, and the tympanum and groin are pale cream. The dentiger-
ous processes of the prevomers are nearly transverse elevations be-
tween the choanae and bear 10-10 teeth. Specimens resembling the
t\'pe ha\e been examined from several localities in Amazonian
Ecuador; some of these have small dark spots on the dorsum and
thus resemble the smaller spotted color morph known from Ama-
zonian Brasil, Peru, and Bolivia. One of the large mocrotls-Wkc frogs
was prepared as a skeleton; no osteological differences between that
specimen and several of P. venulosa were apparent. Consequently,
Hyla macrotis Andersson, 1945, is considered to be a junior synonym
of Plnyuoliyas venulosa (Laurenti, 1768).

The holotype of Hyla adenodenna Lutz, 196S (MNRJ 4054) is
a female having a snout- vent length of 66.0 mm. The skin on the
dorsal surfaces of the head and body is greatly thickened and
glandular, more so than in any other specimen of Phrynohyas that I
ha\'e seen. The dorsal color pattern consists of a network of bold
dark brown reticulations on a tan ground color ( Lutz, 1968, fig. 1 ) ;
dark brown cross bars are evident on the limbs. The following data
supplement Lutz's description: The discs on the hands are large and
equal in diameter to the tympanum, the upper edge of which is
covered by a heavy supratympanic fold. There are 7-9 pre vomerine
teeth on narrowly separated transverse elevations between the
choanae. Although the iris is faded, there is a faint indication of
radiating black streaks. This specimen seems to represent an exti'eme
in glandular development of the skin and of a bold reticulated pat-
tern. Otherwise, it is unquestionably a P. vemdosa. Individuals
having a fainter reticulated pattern are known from various localities
in the Amazon Basin and Guianas. One specimen from Leticia,
Colombia ( KU 126067), has dorsal skin nearly as thick and glandular
as that of the type of Hyh adenodenna, but the Colombian specimen
has a large divided blotch on the dorsum. I consider HyJa adeno-
derma Lutz, 1968, to be a junior synonym of Phrynohyas vemdosa
(Laurenti, 1768).

SUMiMARY

PJirynohyas is one of four genera in the Hylidae having paired
lateral vocal sacs behind the angles of the jaws; the other genera
are Argenieohyla, Osteocephalus, and Traehycephahis. The four
species of PJirynohyas exhibit differences in size, external stiuctiu-al

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

features, coloration, and cranial osteology. Phnjnohyas venulosa is
widespread in the tropical lowlands from northern Mexico to north-
ern Argentina; the other three species have much more restricted
distributions — P. coriocea in Surinam and the upper Amazon Basin,
P. mesophaea on the coastal lowlands of southeastern Brasil, and
P. imitatrix in the uplands of the coastal ranges of southeastern
Brasil. Only P. coriacea and venulosa are known to occur sym-
patrically.

The variability in size, coloration, and texture of the skin on the
dorsum in P. venulosa has resulted in 19 specific names being pro-
posed for this species. Seven nominal species are herein relegated
to the synonymy of Phnjnohyas venulosa (Laurenti, 1768); these
are: Scytopis hebes Cope, 1862; Hyla palpebwgranulata Andersson,
1906; Hyla icettsteini Ahl, 1933; Hyla zernyi Ahl, 1933; Hyh macrotis
Andersson, 1945; Phrynohyas ingens Duellman, 1956; and Hyla
adenoderma Lutz, 1968. Hyla quadrangulum Boulenger, 1882, is
placed in the synonymy of Phrynohyas coriacea (Peters, 1867).

RESUMEN

Phrynohyas cs uno de cuatro generos en la familia Hylidae que
poseen sacos vocales laterales apareados, localizados detras de los
angulos de las articulaciones mandibulares; los otros generos son:
Argenteohyla, Osfeocephalus, y Trochycephalus. Las cuatro especies
de Phrynohyas exiben diferencias en tamano, rasgos estructurales
externos, coloracion, y osteologia cranial. Phrynohyas venulosa esta
ampliamente distribuida en las tierras bajas tropicales, desde el
norte de Mexico al norte de Argentina; las otras tres especies tienen
distribuciones mas restringidas — P. coriacea en Surinam y la parte
alta de la cuenca amazonica, P. mesophaea en las tierras bajas de la
costa del sureste del Brasil, y P. imitatrix en las tierras altas que
costean el sureste brasileiio. Se sabe, que solo P. coriacea y venulosa
ocurren simpatricamente.

Diez y nueve nombres especificados han sido propuestos para
P. venulosa, debido a la variabilidad del tamaiio, coloracion, y
textura de la piel dorsal de esta especie. Siete especies nominales
son relegadas aqui a la sinonimia de Phrynohyas venulosa (Lau-
renti, 1768); estas son: Scytopis hebes Cope, 1862; Hyla palpebro-
granulafa Andersson, 1906; Hyla icettsteini Ahl, 1933; Hijla zernyi
Ahl, 1933; Hyla macrotis Andersson, 1945; Phrynohyas ingens Duell-
man, 1956; y Hyla adenoderma Lutz, 1968. Hyla quadrangulum
Boulenger, 1882, es pucsta en la sinonimia de Phrynohyas coriacea
(Peters, 1867).

HYLID FROGS, GENUS PHRYNOHYAS 17

SPECIMENS EXAMINED

The localities for each of the specimens examined are given
below. The arrangement of the data is as follows: alphabetically
by country, state (department or province), and locality; alpha-
betically by the first letter in the abbreviations for the museum, and
numericallv 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 which
are too vague to be located precisely are given in quotation marks.
Where more than one specimen is included under one museum
number, the number of specimens is given in parentheses after the
museum number. Unless noted otherwise, all specimens are pre-
served in alcohol.

Phrynohijas coriacea

ECUADOR: "western Ecuador," BMNH 1947.2.13.75. El Oro: Machala,
WCAB 40100. Napo: Lago Agrio, KU 126656; Limon Cocha, KU 107025,
UIMNH 87799, 90315-6; Santa Cecilia, KU 105249, UMMZ 129.323. Pastaza:
"Rio Pastaza watershed," NHRM 1950 (7).

PERU: Loreto: Pebas, CAS-SU 12622; Puerto Oriente, 5 km. above
Contamana, Rio Ucavali, UMMZ 123915.

SURINAM: No specific locality, BMNH 70.3.10.69, ZMB 5807 (3); "Spoor-
l)an," RMNH 9606. Mawwijne: Albino, Rio Maroni, ZMB 7606 (3); Maroni,
ZMB 7605 (2); Nassaugebergte, RMNH 7296.

Ph njnoh yas im itatrix
BRASIL: Rio de Janeiro: Teresopolis, MNHN 31/45-46, NHMW 18433.5,
SMF 58269. Santa Catarina: Rio Vernielho, KU 92249 (skeleton), 92250-1.
Sclo Paulo: Sao Jose do Barreiro, Serra da Bocaina, SMF 62562.

Ph rynoh yas m esophaca
BRASIL: No specific locality, UZM 14223, ZMB 3122. Bahia: Santa Cruz,
ZMB 30444. Guanahara: Rio de Janeiro, KU 92252-6, 92257 (skeleton), SMF
30024-9. Rio de Janeiro: Angra dos Reis, ZSM 60/1947; "Guapy," SMF 39251-
3; 36 km from Rio PetropoHs road, BMNH 1940.4.14.13; Teresopolis, SMF
2515, ZMB 26.322. Rio Grande do Sid: Porto Alegre, ZMB 6256, 6810. Santa
Catarina: No specific locality, UZM 14224; Blumenau, NHMW 6279-80; Hansa
(= Corupa), BMNH 1928.11.5.89-106, SMF 2.500-14, 30998, 31001-27; Jaragua,
NHMW 6293 (4); Joinville, MNHN 6310 (5), NHMW 6282-3, 6287 (4),
6288 (21), 6289 (2), 6290 (6), 6291 (5), 6294 (5), 6295 (7), 6296 (6),
6297 (5), 6298 (6), 6299 (9), 6.300 (6), 6301 (6), 6302 (7), 6,303 (4),
6.304 (3), 6,305 (3), 6306 (7), 6307 (9), 6309 (3), SMF 2516-,34, 5120, 5131;
"Rio Humboldt," BMNH 1910.7.26.3-6, 1923.6.1.84-91; "Rio Novo," NHMW
6286 (5), 6295 (5).

Phrynohyas vemdosa

AMERICA: No specific locality, MNHN 4797.

ARGENTINA: No specific locality, USNM 7,3.523. Chaco: Cien-o Petizo,
KU 128948-50; Colonia Eliza, NHRM 1503. Entre Rios: Puerto Marquez, 2
km N La Paz, SMF 43634. Salta: Rio Pescado, KU 128946-7.

BOLIVIA: No specific locality, NHMW 6177 (3). Cochabamha: Rio
Chapare, ZSM 9/1948. El Beni: E.speranza, BMNH 1920.11.29.15-16; Rer-

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

renabaque, UMMZ 57528, 57531 (2). La Paz: Ixiamas, MCZ 10087, UMMZ
57529 (4), 57530 (3); Santa Ana de Movimas, BMNH 98.6.9.38. Santa Cruz:
Buenavi.sta, BMNH 1927.8.185-6, CM 2582, 3828, 3892-3, 3971, .3974, 3986,
4467-8, UMMZ 60526, 63960 (2), 66559-60; Rio de la Cal, ZMB 32107; Santa
Cruz, BMNH 1904.10.29.177, 1907.10.31.72, 1927.8.1.87, CM 2514; Tatarenda,
ZIUS A1.33.

BRASIL: "lower Amazonia," USNM 28926-7. Amazonas: No specific lo-
cality, BMNH 1936.12.3.119; Aiapua, ZMB 30303; Fonte Boa, NHMW 6104;
Manacapinii, Rio Solim5es, ZMB 37080, ZSM 23/1924; Manaus, AMNH 51755,
IRSNB 2681 (14), KU 92258-61, UMMZ 66647; Rio Caicara, NHMW 6111
(2), 6119 (2), 13262 (2); Rio Juma, BMNH 1904.11.4.12; Rio Purus, ZMB
.30307; Tefe, NHMW 6100. Bahia: No specific locality, BMNH 64.1.19.34,
NHMW 6281. Sao Goncalo, 30 km SW Feira de Santana, KU 29438, MCZ
1531. Mato Gwsso: Barro do Tapirape, CAS-SU 12304-10, 12312, KU 40219;
Chapada dos Guimaraes, BMNH 1903.3.26.54-6; Posto Jacaru, Rio Xingi'i, KU
92262-3; Villa Murtinho, UMMZ 56770. Minas Gerais: La Filomena, NHMW
6102 (2), 6112 (2), 6114 (3), 6115; Manga, Rio Sao Francisco, UMMZ
108894; Piraporinha, USNM 98536-7; Sao Luis dos Caceres, CAS-SU 12298-9.
Para: Belem (= Para), BMNH 45.8.25.143, 96.1.7.2, CAS-SU 12390-3, NHMW
6098, ZMB 33189, ZSM 119/1911 (2); Cachoeira de Arari, Ilha de Marajo,
ZMB 29999, ZSM 120/1911; Cameta, Rio Tocantins, NHMW 6099, 6107, 6113;
"Ilha de Marajo," BMNH 1923.11.9.26; IPEAN, 3 km E Belem, KU 127963-4,
128557-61, 130121-6; Livramento, CAS-SU 12383-7; "San Antonio do Prata,"
BMNH 1947.2.23.24; Santarem, BMNH 56.3.25.14 (2); Taperinha, near San-
tarem, NHMW 16.503, 19013. Permnnhuco: No specific locality, BMNH
58.11.28.48; Bonita, MCZ 4998. Piaui: Bosqus, BMNH 1926.5.26.4. Rondonia:
Rio Branco, MNRJ 4054. Roraima: Serra Grande, below Boa Vista, Rio Branco,
NHMW 15893.

COLOMBIA: Amazonas: Leticia, KU 124897-900, 124955-6 (tadpoles); 2
km E Leticia, KU 126067. Caqtietd: Morelia, ANSP 25313. Cundinamarca:
Honda, MLS 1-2. Magdalcna: Barranquilla, MLS 3; Fundacion, UMMZ 48263.
Norte de Santander: Rio Zulia, N of Cucuta, MLS 4.

ECUADOR: No specific locality, USNM 14054. El Oro: 7 km SE Buena-
vista, USNM 192507. Napo: Avila, USNM 165979-80; Lago Agrio, KU 126657;
Rio Cotapino, KU 138736, USNM 165977-8. Pastaza: Montalvo, CAS-SU
10318:9, USNM 165981 (skeleton), 165982-3; Rio Bobonaza, USNM 165984;
upper Rio Oglan, USNM 165986; "Rio Pastaza watershed," NHRM 1958; Rio
Villano, USNM 165988.

FRENCH GUIANA: Caijenne: Cayenne, BMNH 53.2.89-90, NHMW
6101, 6103; Mt. Cabasson, LACM 446.38.

GUYANA: F.ast Demerara: Georgetown, MCZ 2618, UMMZ 80495. Es-
sequiho: Better Hope, BMNH 80.11.22.5-6. Mazartini-Potaro: Kaieteur Falls,
AMNH 23128. North West: Santa Rosa Island, Momka River, UMMZ 55834.
Ru))itnttni: north of Acaray Mts., west of New River, KU 69778-82.

PARAGUAY: No specific locality, AMNH 19916-7, 19919, USNM 5837;
"Alto Paraguay," ZMB 26181. Boqueron: Loma Plata, KU 73432. Central:
Asuncion, BMNH 94.3.14.168; Patifio, FMNH 10799. Concepcion: Apa-Berg-
land, ZSM 1.37/1933. La Cordillera: San Berbardino, NHMW 6106 (2), ZMB
26054, 26115. Olimpo: Puerto 14 de Mayo, Bahia Negra, BMNH 98.6.3.19.

PERU: Loreto: "Caseirio Libertad, Rio Amazonas," MJP 514; Orellana,
USNM 127168-9, 127180; Pebas, CAS-SU 3159, 6376, 12622; Pucallpa, MJP
84, 86 (3), 90, 92, 93 (2), 95, 97 (2), 99; Rio Ucayali, FMNH .3562, UMMZ
48210; Roaboya, AMNH 4,3316, 4.3534; "Transval, near Pebas," CAS-SU 3162.

SURINAM: No specific locality, BMNH 70.3.10.68-9, NHMW 6097, 6109
(2), USNM 14618. Brokopondo: Brokopondo, RMNH 16692 (2). Coronie:
Coronieweg, RMNH 16693. Marowiine: Camp 3, RMNH 16694. Suriname:

HYLID FROGS, GENUS PHRYNOHYAS 19

Berg en Dal, ZMB 7267; Coropina Creek, RMNH 16695; Kwattaweg, RMNH
16696; "Lelydorp," RMNH 16697 (2); Paraniaraibo, AMNH 4009, ANSP 2169-
70, BMNH 1946.4.2.24-25, MCZ 7662, RMNH 16698 (9), USNM 1.3820-1,
ZMB 25966 (2), 26076.

TRINIDAD AND TOBAGO: Tobago: Milford Bay, MCZ 4087; "Wliim,"
BMNH 1926.1.19.1. Tritiidad: No specific locality, FMNH 41683, MCZ .3517,
5971; "Churchill-Roosevelt Hwy," AMNH 55712-15 (16), 55717-19; "Cumpia,"
SMF 2646-47; Fyzabad, BMNH 1929.11.25.6; "Imperial College of Agriculture,"
BMNH 1932.9.16.41; "Mt. Harris," FMNH 49701; "Mt. Thabor," CM 5257;
"Oarino River," BMNH 1929.11.25.4; "Piarco," KU 84714; "St. Augu.stine, '
AMNH 55716.

VENEZUELA: "Los Canales," USNM 128792. Anzoatequi: Carapa, USNM
80612. Aragtia: Maracay, ZSM 304/1929 (20), ,305/1929 (46), 61/19.30 (22),
120/1930 (4), 292/1933 (6), 43/19.34 (10), 105/1935 (11), 196/1965 (5).
Distrito Federal: Caracas, SMF 2645; La Guaira, USNM 22545, 27797. Mo-
nagas: Caicara, AMNH 16904-6, 16911-2, USNM 36.377; Caripito, USNM
117098-9. Trujillo: Savana de Mendoza, UMMZ 57397. Zulia: La Fria, Pueblo
Nuevo, UMMZ 55567-70; Oropc, FMNH 2604.

LITERATURE CITED

Ahl, E.

1933. Liber einige neue Frosche aus Brasilien. Zool. Anz., 104:25-32.
Andersson, L. G.

1906. On batrachians from Bolivia, Argentina, and Peru collected by Erland
Nordenskiold and Nils Holmgren 1901-1902 aild 1904-1905. Arkiv
Zool., 3 (12): 1-19.
1945. Batrachians from east Ecuador collected 1937, 1938 by Wm. Clarke-
Maclntyre and Rolf Blomberg. Ibid, ,37A (2): 1-88.

BOKEHMANN, W. C. A.

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

BOULENGER, G. A.

1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of
the British Museum, ed. 2, London, .\vi-|-503 pp.
Brocchi, p.

1879. Sur divers batrachiens anoures de I'Amerique Centrals. Bull. Soc.
Philoni. Paris, ser. 7, 3:19-24.
Cochran, D. M., and C. J. Goin

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

1862. Catalogue of the reptiles obtained during the explorations of the
Parana, Paraguay, Vermejo, and Uruguay rivers, by Capt. Thos. J.
Page, U.S.N., and of those procured by Lieut. N. Michler, U.S. Top.
Eng., Commander of the expedition conducting the sur\ey of the
Atrato River. Proc. Acad. Nat. Sci. Philadelphia, 14:346-359.
1866. On the structures and distribution of the genera of arciferous Anura.

Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 6:67-112.
1877. Tenth contribution to the herpetology of tropical America. Proc.
Amer. Philos. Soc, 17:85-98.
Daudin, F. M.

1803. Histoire naturelle et particuliere des reptiles. Paris, 8: 439 pp.
Duellmax, W. E.

1956. The frogs of the hylid genus PlmjnoJiyas Fitzinger, 1843. Misc. Publ.
Mus. Zool. Univ. Michigan, 96:1-47.

20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

DUELLMAN, W. E.

1961. The amphibians and reptiles of Michoacan, Mexico. Univ. Kansas

Puhl. Mus. Nat. Hist., 15:1-148.
1966. Ta.xonomic notes on some Mexican and Central American hylid frogs.

Ihid, 17:263-379.
1968. The taxonomic status of some American hvlid frogs. Herpetologica,

24:194-207.
1970. The hylid frogs of Middle America. Monog. Mus. Nat. Hist., Univ.

Kansas, 1: xi+753 pp.
DuMERiL, A. M. C, and G. Bibrox

1841. Erpetologie generale ou histoire naturelle complete des reptiles. Paris,

8:792 pp.

FiTZINCER, L.

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

1907. Description of Hijla resinifictrix Goeldi, a new Amazonian tree frog
peculiar for its breeding habits. Proc. Zool. Soc. London (1907): 135-
140.

GiJNTHER, A. L. C.

1859. Catalogue of the Batrachia Salientia in the collection of the British
Museum. London, x\i+160 pp.
Hemming, F. (editor)

1958. Opinion 520. Suppression under the plenary powers of the specific
name tihiatrix Laurenti, 1768, as published in the combination Hyla
tibiatrix, and of the generic name Aciodtjtes Fitzinger, 1843, and in-
terpretation of Rana vcnulosa Laurenti, 1768 (Class Amphibia).
Opin. Declar. Intl. Comm. Zool. Nomencl., 19:169-200.
Hensel, R. F.

1867. Beitrage zur Kenntniss der Wirbelthiere Siidbrasiliens. Wiegmann's
Arch. Naturg., .33: 120-162.
Laurenti, J. N.

1768. Specimen medicum, exhibens synopsin Reptilium emendatum cum
experimentis circa venena et antidota reptilium Austriacorum. Vienna,
214 pp.
LuTZ, B.

1968. New Brazilian forms of //(//fl. Pearce-Sellards Ser., 10:1-18.

McDiARMID, R. W.

1968. Populational variation in the frog genus Phnjnohyas in Middle Ameri-
ca. C(mtr. Sci. Los Angeles Co. Mus., 134:1-25.

Miranda-Ribeiro, A. de

1926. Notas para servirem ao estudio dos Gymnobatrachios (Anura) bra-
sileiros. Arch. Mus. Nac. Rio de Janeiro, 27: 227 pp.

Peters, W.

1867. tJber Flederthiere und Amphibien. Monats. Akad. Wissen. Berlin
(1867):703-712.

RiVERO, J. A.

1961. Salientia of Venezuela. Bull. Mus. Comp. Zool., 126:1-207.
1964. The distriljution of \>nezuelan frogs VI. The llanos and the delta
region. Caribbean Jour. Sci., 4:491-495.

Seba, a.

1734. Locupletissimi rerum naturalium thesauri accurata descriptio, et in-
conibus artificissimis expressio, per universam physices historiam.
Amsterdam, 1: xxxiv+178pp, 111 pis.

Shannon, F. A., and F. L. Humphrey

1957. A new species of Phrynohijas from Nayarit. Herpetologica, 13:15-18.

HYLID FROGS, GENUS PHHYNOIIYAS 21

Smith, H. M., and E. H. Taylor

1948. An annotated checklist and key to the Amphibia of Mexico. Bulk
U. S. Natl. Mus., 194:iv+118 pp.
Spix, J. B. vox

1824. Aninialia no\a, si\e species noxae testndinuni et lanaiuni, ciuas in
itinere per Biasiliani, annis 1817-1820. Monaco, 53 pp.
Taylor, E. H.

1944. The hyhd genus Aciodyfcs with comments on tlie Mexican forms.
Univ. Kansas Sci. Bulk, 30:63-68.

Taylor, E. H., and H. M. Smith

1945. Sunmiary of the collection of amphibians made in Mexico under the
Walter Rathbone Bacon Traxeling Scholarship. Proc. U. S. Natl. Mus.,
95:521-613.

Thueh, L., and W. E. Duellman

1971. A synopsis of Neotropical In lid frogs, genus Osteocephalus. Occas.
Papers Mus. Nat. Hist. Univ. Kansas, 1:1-47.
Tyler, M. J.

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

University of Kansas Publications
MUSEUM OF NATURAL HISTORY

The University of Kansas Publications, Museum of Natural
History, beginning with volume 1 in 1946, was discontinued
with volume 20 in 1971. Shorter research papers formerly pub-
lished in the above series are now published as Occasional
Papers, Museum of Natural History. The Miscellaneous Pub-
lications, Museum of Natinal History, began with number 1 in
1946. Longer research papers are published in that series.
Monographs of the Museimi of Natural History were initiated
in 1970.

Institutional libraries interested in exchanging publications
may obtain the Occasional Papers and Miscellaneous Publica-
tions by addressing the Exchange Librarian, University of Kan-
sas Library, Lawrence, Kansas 66044. Individuals may pur-
chase separate numbers of all series. Prices may be obtained
upon request addressed to Pul:)lications Secretary, Museum of
Natural History, University of Kansas, Lawrence, Kansas 66044.