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OCCASIONAL PAPERS

m , HARVARD

of the

MUSEUM OF NATURAL HISTORY

The University of Kansas

Lawrence, Kansas

NUMBER 147. PAGES 1-21 23 JANUARY 1992

SOME HYLID FROGS FROM THE

GUIANA HIGHLANDS, NORTHEASTERN SOUTH

AMERICA: NEW SPECIES, DISTRIBUTIONAL

RECORDS, AND A GENERIC REALLOCATION

William E. Duellman1 and Marinus S. Hoogmoed2

Abstract: Three new species of Hyla are named from the Guianan region
of northeastern South America. Hyla hadroceps, a species of unknown
relationships from southern Guyana, has a large, blunt head and tuberculate
dorsal skin. Hyla roraima, a member of the Hyla geographica group, and
Hyla warreni, a species of unknown relationships, occur on Mt. Roraima in
western Guyana. Hyla kanaima Goin and Woodley, a member of the Hyla
geographica group, is reported from Mt. Roraima and redescribed. Hyla
rodriguezi Rivero is placed in the genus Osteocephalus.

Key words: Hyla hadroceps, Hyla roraima, Hyla warreni, new species;
Osteocephalus rodriquezi; Guyana; Venezuela.

Resumen: Se denominan tres especies nuevas de Hyla de la region de las
Guyanas en el noreste de Sudamerica. Hyla hadroceps, especie de relaciones
desconocidas del sur de Guyana, tiene una cabeza grande y roma, y la piel del
dorso con tuberculos. Hyla roraima, miembro del grupo Hyla geographica,
e Hyla warreni, especie de relaciones desconocidas, occuren en el Monte
Roraima en Guyana occidental. Se reporta de Monte Roraima, Guyana, y
redescribe a Hyla kanaima Goin and Woodley, miembro del grupo Hyla
geographica. Se ubica a Hyla rodriguezi Rivero en el genero Osteocephalus.

'Curator, Division of Herpetology, Museum of Natural History, and Professor,
Department of Systematics and Ecology. The University of Kansas, Lawrence,
Kansas 66045-2454, USA.

2Head, Department of Vertebrates, and Curator, Herpetology. Nationaal
Natuurhistorisch Museum, Postbus 9517, 2300 Leiden. The Netherlands.

© Museum of Natural History. The University of Kansas, Lawrence.

2 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP., No. 147

Our independent field work in northeastern South America during the past
two decades has resulted in the collection of numerous new species of hylid
frogs. Study of our material and that in various museum collections has
resulted in arevision of the genus Stefania (Duellman and Hoogmoed, 1984),
definition of species in the Hyla granosa group (Hoogmoed, 1979a), and
descriptions of three new species of Ololygon{ Hoogmoed and Gorzula, 1979;
Duellman, 1986). During our studies of other museum collections we
discovered three unnamed species of Hyla not represented among specimens
that we collected.

Herein we report on some hylids of the genera Hyla and Osteocephalus
from the Guiana Highlands. This region encompasses the table mountains
extending from Surinam southwestward to the southern part of the Territorio
Federal de Amazonas in southern Venezuela. The area was described by
Hoogmoed (1979b); localities mentioned in the text are shown in Figure 1.

We examined material in the British Museum (Natural History) (BM),

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Fig. 1. Map ofGuianan Region showing localities mentioned in text. 1 =Auyan-
tepui; 2 = Paso del Danto; 3 = Km 144; 4 = Northeastern slope of Mt. Roraima; 5 =
Acarai Mountains; 6 = New River.

HYLID FROGS FROM THE GUIANA HIGHLANDS 3

Museum of Comparative Zoology at Harvard University (MCZ), Museum of
Natural History at The University of Kansas (KU), National Museum of
Natural History (USNM), Nationaal Natuurhistorisch Museum (formerly
Rijksmuseum van Natuurlijke Historie) (RMNH). the University of Guyana
Department of Biology (UGDB), and the University of Puerto Rico at
Mayagiiez (UPR-M ). Measurements and structural features follow Duellman
(1970), except that webbing formula is that of Savage and Heyer (1967). as
modified by Myers and Duellman ( 1982). Snout-vent length is abbreviated
SVL.

Hyla Lalrenti, 1768

Nearly 300 species, most of which are placed in one of more than 40
phenetic groups, are recognized in the paraphyletic genus Hyla. Seven of
these groups occur in the Guianan Region. These are: ( 1 ) Hyla boans group
(Duellman. 1970). (2) Hyla geographica group (Duellman. 1973). (3) Hyla
granosa group ( Hoogmoed. 1 979a) , (4) Hyla leucophyllata group ( Duellman,
1 970), (5 ) Hyla marmorata group ( Bokermann, 1 964 ), (6) Hyla microcephalia
group (Duellman. 1970). and (7) Hyla parviceps group (Duellman and
Crump. 1974).

Two of the three new species described herein cannot be relegated to any
of these recognized species groups. The other new species is a member of the
Hyla geographica group. For ease of comparison, comparable features are
numbered sequentially in the diagnoses.

Hyla hadroeeps new species

Holotype. — KU 69720, an adult male, from area north of Acarai Moun-
tains, west of New River (ca. 02°N, 58°W). Rupununi District, Guyana,
obtained in January 1962 by William A. Bently.

Diagnosis. — The single male has a SVL of 53.9 mm and the following
characteristics: ( 1 ) body robust; head blunt; ( 2 ) skin on dorsum bearing many
large, round tubercles: skin of head not co-ossified with underlying dermal
bones; (3) tympanum distinct; (4) fingers about two-thirds webbed; (5) toes
nearly fully webbed; (6) fringes and calcars absent on limbs; (7) axillary
membrane extending to midlength of upper arm; (8) dorsum brown with
irregular darker brown markings; venter cream with brown flecks; (9)
vomerine odontophores short, diagonal.

A subgular vocal sac immediately distinguishes Hyla hadroeeps from
species of Phrynohyas and Osteocephalus, some of which it resembles
superficially. The thick tubercular skin, large size, and absence of black and
orange or yellow flash colors distinguish it from members of the Hyla
marmorata group. The absence of dermal fringes on the limbs distinguishes
H. hadroeeps from H. tuberculosa.

4 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP.. No. 147

Description of holotype. — Adult male; body robust; head wider than
long; snout short, distance from tip of snout to eye equal to length of eye,
truncate in dorsal view and in profile, not projecting beyond margin of lip;
canthus rostralis rounded, indistinct; loreal region concave, sloping steeply to
rounded lip; nostril protuberant laterally; internarial area slightly depressed;
top of head flat; interorbital distance slightly more than width of upper eyelid;
tympanum distinct, round, its length 0.55 that of eye, separated from eye by
distance about equal to length of tympanum: supratympanic fold weak, barely
covering upper edge of tympanic annulus. Axillary membrane extending
about one half length of upper arm; forearm robust, lacking ulnar fold; fingers
short, bearing round discs; width of disc on Finger III slightly greater than
diameter of tympanum; disc on Finger I noticeably smaller than others;
dermal keel along outer edge of Finger IV; relative lengths of fingers 1 < 2 <
4 < 3; subarticular tubercles distinct, moderately small, conical; distal
tubercle on Finger IV weakly bifid; supernumerary tubercles small, round,
present only on proximal segments; two palmar tubercles, round; thenar
tubercle elongate; nuptial excrescence elliptical, unpigmented, present on
dorsal surface of proximal part of Finger I; fingers about two-thirds webbed;
webbing formula 12 — 2Vi III1/:— 2III2— l1/: IV (Fig. 2A). Hind limb
moderately short, lacking folds on tarsus; inner metatarsal tubercle elongate,
round in section, barely visible from above: outer metatarsal tubercle indis-
tinct, round, low; toes short, bearing round discs slightly smaller than those
on fingers; disc on Toe I much smaller than others; dermal keel along outer
edge of Toe V; relative lengths of toes 1 < 2 < 5 < 3 < 4; subarticular tubercles
distinct, moderately small, subcorneal; supernumerary tubercles small, round.

Fig. 2. Hand (A) and foot (B ) of Hyla hadroceps, KU 69720. Line = 5 mm.

HYLID FROGS FROM THE GUIANA HIGHLANDS 5

present only on proximal segments; toes extensively webbed: webbing
formula II— 2 III— 2 III 1 — 2 IV l1/:— 1 V (Fig. 2B). Skin on dorsal sur-
faces of head, body, shanks, and forearms with abutting, low, round tubercles;
skin on flanks, chest, belly, and proximal ventral surfaces of thighs coarsely
granular: skin on other surfaces smooth: anal sheath long; anal opening
puckered, directed posteroventrally at upper level of thighs. Vomerine
odontophores oblique, widely separated medially, diverging posteriorly just
behind posterior margins of transversely ovoid choanae, bearing 6 and 7 teeth;
tongue broadly cordiform. shallowly notched posteriorly, free posteriorly for
about one-fourth of its length; vocal slit extending from midlateral base of
tongue toward angle of jaw; vocal sac single, median, subgular with lateral
extensions.

Color in preservative: Dorsum brown with irregular darker brown mark-
ings; limbs brown with broad, transverse brown marks — one each on fore-
arm, wrist, and foot; two each on tibia, shank, and tarsus (Fig. 3). Flanks and
anterior and posterior surfaces of thighs creamy tan with brown flecks; venter
cream with brown flecks laterally on throat, over entire belly, and distally on
thishs.

Fig. 3. Holotype of Hyla hadroceps, KU 69720, male. 53.9 mm SVL.

6 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP., No. 147

Measurements (in mm): SVL53.9, tibia length 25.8, foot length 20.9, head
length 18.1, head width 1 9.6. interorbital distance 5.9, upper eyelid width 5.2,
eye-nostril distance 4.5, eye length 5.9, tympanum length 2.8.

Distribution. — The species is known only from the type locality in
extreme southern Guyana.

Etymology. — The specific name is derived from the Greek hadros,
meaning bulky, and the Greek kephale, meaning head; the name hadroceps
alludes to the short, heavy head of the frog.

Remarks. — The single specimen has been in the collection at The Univer-
sity of Kansas for many years, during which time it has been examined many
times and compared with other species, all to no avail in identifying it with a
known species. It was part of a collection containing many common and
widespread species of frogs typical of the Guianan rainforest; all were
collected at the same locality and include Adenomera andreae, Leptodactylus
mystaceus, L. pentadactylus, L. rhodomysta.x, L. wagneri, Bufo "typhonius,"
Epipedobates (- Allobates) femoral is, E. trivittatus, Allophryne ruthveni,
Hyla boans, H. geographica, H. minuta, and Osteocephalus taurinus.

Hyla roraima new species

Holotype. — BM 1979.560, an adult female, from the north slope of Mt.
Roraima (05°38'N, 60°44'W, elev. 1.480 m), Rupununi District, Guyana,
obtained on 29 August 1971 by Adrian N. Warren.

Paraty pes. — KU 1 82470 collected with the holotype, and UGDB 1 4 from
Mt. Roraima. 1,430 m, obtained by Michael Tamessar.

Diagnosis. — A member of the Hyla geographica group characterized by:
( 1 ) body slender; head distinct from body; (2) skin on dorsum of body and
limbs smooth, on head tuberculate. not co-ossified with underlying dermal
elements; (3) tympanum distinct; (4) fingers with vestigial webbing; (5) toes
about one-half webbed; (6) fringes absent on limbs: calcar conical; (7)
axillary membrane absent; (8) dorsum tan with irregular dark brown mark-
ings and middorsal dark stripe or not; flanks cream or pale gray with brown
flecks or irregular vertical lines; posterior surfaces of thighs uniform cream
or gray with brown flecks; ventral surfaces and webbing on foot cream;
palpebral membrane reticulated; (9) vomerine odontophores angular.

The presence of prepollical spines not projecting through the skin, calcars,
and vestigial webbing on the hand place H. roraima in the Hyla geographica
group, as defined by Duellman ( 1 973 ). The presence of a reticulated palpebrum
immediately distinguishes H. roraima from H. calcarata and H.fasciata,
both of which also have bold black markings on the flanks and posterior
surfaces of the thighs; furthermore, the calcar in//, calcarata is large, flat, and
triangular, and that in H.fasciata is long and tubercular. Two other members
of the Hyla geographica group have reticulated palpebral membranes; of

HYLID FROGS FROM THE GUIANA HIGHLANDS

these, H. geographica has the fingers about one-half webbed, large triangular
calcars, and many (usually paired) vertical dark bars on the flanks and
posterior surfaces of the thighs, whereas//, microderma has a low tubercle on
the heel and lacks dark marks on the flanks and posterior surfaces of the thighs.
Hyla dentei is about the same size as H. roraima but lacks calcars and has faint
transverse dorsal markings on the body and bold transverse bars on the
anterior, dorsal, and posterior surfaces of the thighs. Hyla roraima differs from
the sympatric H. kanaima by having more webbing on the feet, a reticulated
palpebral membrane, and angular instead of straight vomerine odontophores;
furthermore, H. kanaima has bold reticulations on the flanks, transverse bars
on the thighs, and dark flecks on the throat. Hyla roraima differs from the
geographically adjacent//, crepitans (Gran Sabana of Venezuela) in colora-
tion (vertical lines on flanks of//, crepitans) and by having a reticulated
palpebral membrane, calcars, and more webbing on the hand.

Description of holotype. — Adult female; body slender; head markedly
distinct from body, depressed, slightly longer than wide; snout moderately
long, truncate in dorsal view and in profile, not projecting beyond margin of
upper lip; canthus rostralis rounded: loreal region inclined ventrolaterally to
round lip; nostril protuberant laterally; internarial area slightly depressed; top
of head flat; interorbital distance 20% greater than width of upper eyelid;
length of eye slightly more than eye-nostril distance; tympanum distinct, its
length 0.43 that of eye, deflected dorsolaterally. separated from eye by
distance equal to length of tympanum; supratympanic fold weak, barely
covering dorsal edge of tympanic annulus. Axillary membrane absent;
forearm slender, having row of indistinct, low ulnar tubercles; fingers
moderately long, bearing large, round discs; width of disc on Finger III
slightly less than diameter of tympanum; relative lengths of fingers 1 < 2 < 4
< 3; subarticular tubercles distinct, large, round; supernumerary tubercles
moderately large, round, present only on proximal segments; palmar tubercle
bifid: thenar tubercle elongate with median projection distally ; basal webbing
between Fingers II— IV (Fig. 4A). Hind limb long, slender; heel bearing
conical tubercular calcar; inner tarsal fold weakly defined on distal part of
tarsus; inner metatarsal tubercle elliptical, flattened, visible from above; outer
metatarsal tubercle low, conical; toes moderately long, bearing round discs
smaller than those on fingers; relative lengths of toes 1 < 2 < 5 < 3 < 4;
subarticular tubercles large, round; supernumerary tubercles small, round,
present only on proximal segments; toes about one-half webbed; webbing
formula 12— 3112 — 3III1— 3 IV3 — 2V (Fig. 4B). Skin on head weakly
tuberculate, on other dorsal surfaces smooth, on ventral surfaces granular;
anal sheath short; anal opening directed posteriorly at upper level of thighs,
bordered below by round tubercles. Vomerine odontophores angular, abutting
medially, bearing 1 5 and 1 6 teeth; tongue broadly cordiform, slightly notched
posteriorly, barely free behind.

UNIV. KANSAS MUS. NAT. HIST. OCC. PAP.. No. 147

Fig. 4.

Hand (A) and foot (B) of Hyla roraima, BM 1979.560. Line = 5 mm.

Color in preservative: Dorsum tan with irregular dark brown markings
consisting of flecks on snout, interorbital mark, a crudely W-shaped mark in
scapular region, pair of elongate spots anterior to sacrum, two transverse
marks posterior to sacrum, and transverse bars on limbs — three on forearm,
five on thigh, five on shank, and two on tarsus; flanks and posterior surfaces
of thighs grayish cream with vertically elongate brown flecks; ventral
surfaces dull cream; palpebral membrane finely reticulated (Fig. 5).

Measurements (in mm): SVL45.5, tibia length 25.2, foot length 16.6, head
length 1 6. 1 , head width 15.7, interorbital distance 5.4. upper eyelid width 4. 1 .
eye-nostril distance 4.8, eye length 5.1. tympanum length 2.2.

Variation. — The two paratypes are females having S VLs of 38.5 and 42.2
mm and 14-14 and 13-13 vomerine teeth. They are essentially like the
holotype in proportions and structure. In coloration, there are some minor
differences. One (UGDB 14) has a narrow black middorsal line on the head
and anterior part of the body. Both have more extensive dark markings on the
body than does the holotype; each has a large dark spot on the snout, broad
interorbital bar, large X-shaped mark in the scapular region, and broad,
transverse mark in sacral region. In UGDB 14, the flanks have dark flecks, and
in KU 182470, dark vertical lines.

HYLID FROGS FROM THE GUIANA HIGHLANDS

«T 7

Fig 5. Species of Hyla. Top. Hyla kanaima, KU 182469, female. 49. 1 mm SVL.
Middle. Holotype of Hyla warreni, BM 1979.561. female. 36.2 mm SVL. Bottom.
Holotype of Hyla roraima, BM 1979.560. female. 45.5 mm SVL.

10 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP.. No. 147

Distribution. — Hyla roraima is known only from the forested slopes of
Mt. Roraima in southwestern Guyana.

Etymology. — The specific name is a noun in apposition for the type
locality, Mt. Roraima, the "legendary" tepui on the common border of
Guyana, Venezuela, and Brazil.

Remarks. — The type series of H. roraima consists of females, whereas
that of//, microderma is made up of five males having snout-vent lengths of
3 1 .0-32.7 mm (x = 3 1 .7) mm (Pybum, 1977:406). Although H. microderma
occurs far to the west (Rio Vaupes, Colombia), the only known locality is on
the western part of the Guiana Shield. Because H. microderma and H. roraima
are alike in having vestigial webbing on the hands and having reticulated
palpebral membranes, it was suspected that the specimens from Mt. Roraima
might be females of//, microderma. In addition to body size and relative sizes
of the calcars, the major differences between the two species are in coloration.
The smaller species, H. microderma lacks markings on the flanks and thighs,
whereas the larger//, roraima has dark markings on those surfaces. Duellman
(1973:523, 525) noted the ontogenetic development of dark markings on the
flanks and thighs in H calcarata and H.fasciata, but no noticeable changes
occurred after sexual maturity; also, no ontogenetic changes in shape and
proportional size of calcars were noted. Thus, the differences between adult
males of//, microderma and adult females of//, roraima are interpreted as
specific differences, not intraspecific sexual ones.

Field data accompanying Warren's specimens noted that the frogs were
collected in montane forest at night. One was on a palm frond, another on a
leaf over a slow-movinc stream, and the third on the forest floor.

'£-

Hyla warreni new species

Holotype. — BM 1979.561. an adult female, from the north slope of Mt.
Roraima (05°38'N, 60°44'W, elev. 1.480 m), Rupununi District, Guyana,
obtained on 2 September 1971 by Adrian N. Warren.

Paratopotype. — KU 182471, an adult female, obtained on 3 September
1 97 1 by Adrian N. Warren.

Diagnosis. — A moderate-sized Hyla with a maximum known snout-vent
length of 36.2 mm and having the following characters: ( 1 ) body slender; head
relatively large, blunt; (2) skin on dorsum smooth; skin on head not co-
ossified with underlying dermal elements; (3) tympanum distinct; (4) fingers
webbed basally; (5) toes about one-half webbed; (6) fringes and calcars
absent on limbs; (7) axillary membrane extending to midlength of upper arm;
(8) dorsum brown with irregular darker spots; venter cream with brown spots
on throat; (9) vomerine odontophores short, transverse.

Hyla warreni superficially resembles Hyla (= Osteocephalus) rodriguezi,
which differs from H. warreni by having angular vomerine odontophores,

HYLID FROGS FROM THE GUIANA HIGHLANDS 1 1

elongate choanae, small tubercles on the dorsum, different coloration, an
abbreviated axillary membrane, and relatively short fingers with simple
subarticular tubercles. Hyla warreni has an extensive axillary membrane and
longer fingers with bifid subarticular tubercles on Fingers III and IV. The
absence of black and yellow flash colors and extensive webbing on the hand
distinguish H. warreni from members of the Hyla marmorata group. The
absence of dermal fringes distinguishes//, warreni from H. tuberculosa, and
the smooth skin and vestigial webbing separate it from H. hadroceps.

Description of holotype. — An adult female; body moderately slender;
head relatively large, as wide as long; snout short, truncate in dorsal view and
in profile, not projecting beyond margin of lip; canthus rostralis distinct,
rounded; loreal region slightly concave, sloping gradually to rounded lip;
nostril protuberant dorsolaterally; internarial area deeply depressed; top of
head flat; length of eye greater than eye-nostril distance; interorbital distance
slightly greater than width of upper eyelid; tympanum distinct, its diameter
0.34 that of eye, separated from eye by distance slightly greater than length
of tympanum; supratympanic fold moderately heavy, covering upper part of
tympanum and curving to point above angle of jaw. Axillary membrane
extending more than one-half length of upper arm; forearm moderately
robust, with two indistinct ulnar tubercles distally; fingers long, bearing round
discs; width of disc on Finger III equal to diameter of tympanum; relative
lengths of fingers 1 < 2 < 4 < 3; subarticular tubercles moderately small,
round; distal tubercles on Fingers III and IV distinctly bifid; supernumerary
tubercles small, round, numerous on proximal segments; palmar tubercle
bifid; thenar tubercle elliptical; fingers webbed basally; webbing formula
112 — 3113" — 3 IV (Fig. 6A). Hind limb long, slender, lacking tarsal fold and
tubercles; inner metatarsal tubercle elongate, round in section, projecting,
visible from above; outer metatarsal tubercle small, subcorneal; toes moder-
ately long, bearing round discs slightly smaller than those on fingers; relative
lengths of toes 1<2<3<5<4; subarticular tubercles small, round;
supernumerary tubercles small, round, sparse on proximal segments; toes
about one-half webbed; webbing formula 12 — 2111— 2III1— 21 V2-
1+V (Fig. 6B). Skin on dorsal surfaces smooth; skin on belly and proximal
posteroventral surfaces of thighs coarsely granular; skin on other ventral
surfaces smooth; anal sheath short; anal opening directed posteriorly at upper
level of thighs, bordered below by two pairs of tubercles. Vomerine
odontophores widely separated medially, short, transverse, slightly posterior
to level of posterior margins of small, round choanae, bearing 5 and 6 teeth:
tongue circular, shallowly notched behind, free posteriorly for about one-
fourth of its length.

Color in preservative: Dorsum pale brown with numerous, irregular, small
dark brown spots; flanks creamy tan with small brown spots; limbs creamy tan
with brown transverse marks — two on forearm, five on thigh, four on shank.

12

UNIV. KANSAS MUS. NAT. HIST. OCC. PAP.. No. 147

Fig. 6. Hand (A) and foot (B ) of H via warreni, BM 1 979.56 1 . female. Line = 5 mm.

and three on tarsus; posterior surfaces of thighs tan; side of head tan with dark
brown labial bars demarcating two broad, creamy white marks — one below
eye, one posterior to eye; venter dull cream with large brown spots and
reticulations on chin (Fig. 5).

Measurements (in mm): Snout-vent length 36.2. tibia length 20.3, foot
length 14.5, head length 12.8. head width 1 2.8, interorbital distance 3.7, width
of upper eyelid 3. 1 , eye-nostril distance 3.2, eye length 4.2, tympanum length
1.8.

Variation. — The single female paratype has a snout-vent length of 32.9
mm and 7-8 vomerine teeth; the proportions and coloration are essentially the
same as in the paratype, except that the ventral spotting extends onto the chest.

Distribution. — The species is known only from the northern slopes of Mt.
Roraima in southwestern Guyana.

Etymology. — The species is named for the collector, Adrian N. Warren.

Remarks. — The specimens were collected in montane forest. According
to Warren's field notes, the holotype was on a palm frond at night, and the
paratype was sleeping by day in the fork of a fallen branch.

Two other large species of Hyla (H. kanaima and H. roraima) were found
at the same locality with H. warreni. Hyla kanaima differs by having long,
transverse vomerine odontophores, simple subarticular tubercles on the
fingers, boldly reticulated flanks, and no pale labial spots. Hyla roraima
differs by having angular vomerine odontophores, small tubercles on the
head, calcars, and unicolor lips and venter.

HYLID FROGS FROM THE GUIANA HIGHLANDS 1 3

Hyla kanaima Goin and Woodley

Goin and Woodley ( 1969) described Hyla kanaima from Mt. Kanaima,
near Amatuk Falls on Potaro River, Guyana, and suggested that the species
was related to Hyla geographica. We report here on three new specimens of
H. kanaima from a new locality for the species. These specimens provide new
information on the variation in the species. Therefore, we present a new
diagnosis of the species and a description of the new specimens.

Diagnosis. — A member of the Hyla geographica group characterized by:
( 1 ) body moderately slender; head distinct from body; (2) skin on dorsum
smooth; skin on head not co-ossified with underlying dermal elements; (3)
tympanum distinct; (4) fingers unwebbed; (5) toes about one-third webbed;
(6) fringes absent on limbs; calcar small, blunt; (7 ) axillary membrane absent;
(8) dorsum tan with irregular longitudinal markings and no middorsal dark
stripe: flanks boldly mottled brown and cream; posterior surfaces of thighs
with broad dark brown bars continuous with those on dorsal surfaces; ventral
surfaces of thighs pale gray with brown flecks: belly creamy gray with brown
flecks; webbing on feet cream with brown flecks; palpebral membrane
unpigmented; (9) vomerine odontophores long, diagonal.

The presence of calcars and prepollical spines not projecting through the
skin and the absence of webbing between the fingers place H. kanaima in the
Hyla geographica group as defined by Duellman ( 1973). The absence of a
reticulated palpebrum immediately distinguishes H. kanaima from H.
geographica, H. microderma, and H. roraima. Hyla dentei is about the same
size as H. kanaima and also has bold transverse markings on the thighs;
however, H. dentei has faint transverse markings on the dorsum. Two other
species in the Hyla geographica group. H. calcarata and H.fasciata, have
unpigmented palpebral membranes. Both of these have bold black markings
on the flanks and posterior surfaces of the thighs; furthermore, the calcar in
H. calcarata is large, flat, and triangular, and that in H.fasciata is long and
tubercular. Hyla kanaima differs from the sympatric H. roraima by having
less webbing on the feet, an unpigmented palpebral membrane, and straight
instead of angular vomerine odontophores; furthermore. H. roraima has
small brown flecks or irregular vertical lines on the flanks, uniformly cream
or pale gray (with brown flecks) posterior surfaces of the thighs, and
uniformly cream ventral surfaces and webbing.

Description of three adult females. — Body moderately slender: head
markedly distinct from body, depressed, slightly longer than wide; snout
moderately long, rounded in dorsal view, truncate in profile, not projecting
beyond margin of upper lip; canthus rostralis rounded; loreal region inclined
ventrolaterally to round lip; nostril slightly protuberant laterally; internarial
area noticeably depressed; top of head flat; interorbital distance equal to width
of upper eyelid; length of eye about equal to eye-nostril distance; tympanum

14

UNIV. KANSAS MUS. NAT. HIST. OCC. PAP.. No. 147

distinct, its length 0.31-0.42 (x = 0.34) that of eye, deflected dorsolaterally,
separated from eye by distance slightly less than length of tympanum;
supratympanic fold moderately heavy, covering upper third of tympanum.
Axillary membrane absent; forearm slender, lacking ulnar tubercles; fingers
moderately long, bearing large, round discs; width of disc on Finger III equal
to length of tympanum; relative lengths of fingers 1 < 2 < 4 < 3; subarticular
tubercles moderately large, round; supernumerary tubercles large, round;
palmar tubercle indistinctly trifid; thenar tubercle elongate; webbing absent
(Fig. 7A). Hind limb long, slender; heel bearing small, round protuberance;
inner tarsal fold weakly defined on distal part of tarsus; inner metatarsal
tubercle ovoid, rounded, barely visible from above; outer metatarsal tubercle
absent; toes moderately long, bearing round discs slightly smaller than those
on fingers; relative lengths of toes 1 < 2 < 5 = 3 < 4; subarticular tubercles
moderately large, subconical; supernumerary tubercles slightly smaller,
round; toes about one-third webbed; webbing formula 12 — 3II(2-2+) —
3III3— (3 -_3+)IV(3"-3+)— 2V (Fig. 7B). Skin on dorsum smooth; skin on
belly and ventral surfaces of thighs granular; skin on other ventral surfaces
smooth; anal sheath short; anal opening directed posteriorly at upper level of
thighs. Vomerine odontophores straight, abutting medially, inclined
posterolaterally. bearing 15 and 19 (x = 16.5) teeth; tongue cordiform.

Fig. 7. Hand (A) and foot (B ) of Hyla kanaima, KU 1 82469. Line = 5 mm.

HYLID FROGS FROM THE GUIANA HIGHLANDS 1 5

shallowly indented behind, barely free posteriorly.

Color in preservative: Dorsum tan with dark brown markings with
irregular edges consisting of canthal stripe, narrow interorbital bar, dorsolat-
eral stripe extending from posterior corner of eyelid medially to scapular
region and thence posterolaterally to groin, longitudinal stripe above inser-
tion of forelimb, two to four blotches in sacral region, small spots and dashes
on head and medial part of body, and transverse bars on limbs — two on upper
arm, three or four on forearm, seven or eight on thigh, four or five on shank,
and four or five on tarsus. Flanks mottled dark brown and cream; posterior
thighs marked with continuation of transverse bars on dorsal surfaces of
thighs; venter creamy gray with brown flecks, largest on throat; palpebral
membrane unpigmented (Fig. 5).

Measurements (in mm): SVL 46.0-49. I (x = 47.6), tibia length 23. 1-24.8
(x=24.2),footlength 16.3-17.3 (x = 16.8),headlength 16.8-17.7 (x = 17.2),
head width 15.3-16.7 (x = 16.1), interorbital distance 4.3-5. 1 ( X =4.7), upper
eyelid width 4.5-5.0 (x = 7.4), eye-nostril distance 4.9-5.7 (x = 5.3), eye
length 5.8-6.2 (x = 6.0). tympanum length 2.1-2.7 (x = 2.4).

Remarks.— These females (BM 1983.1428. KU 182469. UGDB 13) are
essentially like the type series, of which two females have SVLs of 46.3 and
48.0 mm, and three males have SVLs of 37.0-37.8 mm (x = 37.4) (Goin and
Woodley. 1969). The structure of the new specimens agrees with the descrip-
tion of the type series given by Goin and Woodley ( 1969). Two specimens
(BM 1983.1428 and KU 182469) have somewhat bolder color patterns but
otherwise are like the type series; a transverse dark mark connects the
dorsolateral dark stripes in the scapular region in BM 1983.1428. One
specimen (UGDB 13) has the dorsolateral stripes fragmented into a longitu-
dinal series of spots and has more distinct bars on the limbs.

The new specimens were found at an elevation of 1,430 m on the north
slope of Mt. Roraima on 29 August 1971 by Adrian N. Warren and on 26
October 1973 by Michael Tamessar. This locality is approximately 155 km
west of the type locality, Mt. Kanaima. All three females contain relative large
( 1 .8 mm diameter), pigmented oviducal eggs. The large size of these eggs and
the absence of ponds on the slopes of Mt. Roraima suggest that H. kanaima
deposits eggs in streams.

Osteocephalus Steindachner, 1862

Trueb and Duellman (1971) recognized five species in the genus
Osteocephalus: Duellman (1974) placed Hyla langsdorffii Dumeril and
Bibron in the genus, and Martins and Cardoso ( 1987) named O. subtilis from
southwestern Brazil. Herein we transfer Hyla rodriguezi Rivero to the genus,
thereby bringing the number of recognized species of Osteocephalus to eight.
We are aware of an undescribed species from Auyan-tepuf in southern

16 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP.. No. 147

Venezuela (specimens in RMNH and USNM) and at least one undescribed
species in the middle Amazon Basin and the Guianan lowlands that is
structurally similar to the larger sympatric Osteocephalus taurinus. This new
species and O. taurinus differ in breeding habits and advertisement calls
(Walter Hodl and Barbara Zimmerman, pers. comm.; MSH, pers. observa-
tion).

Because O. rodriguezi has less webbing on the hands and feet than do any
of the other species, the webbing characters listed in the diagnosis of the genus
by Trueb and Duellman ( 197 1:7) should be modified to read: fingers no more
than one-third webbed, and toes more than one-half webbed. Our material of
O. rodriguezi necessitates a new description of the species using the charac-
ters and numerical diagnosis equivalent to that of Trueb and Duellman ( 1 97 1 ).

Osteocephalus rodriguezi (Rivero), new combination

Diagnosis. — ( 1 ) Size small, sexual dimorphism slight: maximum ob-
served snout-vent length in males 34.7 mm, in females 38.1 mm; (2) skin on
dorsum in males bearing numerous minute, spinous tubercles; (3) skin on
flanks smooth: (4) web extending nearly to base of antepenultimate phalanx
of Finger III; (5) dorsum dark brown with faint darker middorsal blotch
extending from eyelids to sacral region; (6) venter dull cream with brown
flecks or reticulations on margin of chin and in some individuals extending
onto chest; (7) narrow pale labial stripe expanded below orbit; (8) flanks gray
with small, dark brown spots; (9) dermal roofing bones of skull slightly
exostosed; (10) dermal sphenethmoid absent; (11) nasals widely separated
medially; (12) anteromedial margin of frontoparietal between mid- and
anterior levels of orbit: (13) frontoparietal fontanelle partially exposed; ( 14)
palatine not serrate; (15) parasphenoid lacking odontoids; (16) zygomatic
ramus of squamosal extending about one-third distance to maxillary arch;
(17) transverse processes of Vertebra III wider than sacral diapophyses;
transverse processes of Vertebrae IV-VII narrower, subequal in length; (18)
intennandibularis and submentalis muscles connected; ( 19) supramandibular
portion of interhyoideus muscle forming simple tubular posterolateral exten-
sion; associated skin unmodified.

Osteocephalus rodriguezi differs from all other species in the genus by its
small size; the maximum snout-vent length in males of O. rodriguezi is 34.7
mm, whereas the smallest adult male of any other species is 37.9 mm in O.
buckleyi, a species that differs further from O. rodriguezi by having extreme
sexual dimorphism in size, the skin on the dorsum in males bearing a mixture
of large and small, nonspinous tubercles, and the skin on the flanks areolate.
Structurally, O. rodriguezi is most like O. leprieurii, which differs in being
larger (smallest male 41 .2 mm ) and in having a pattern of transverse dark bars
on the dorsum and the venter and flanks immaculate.

HYLID FROGS FROM THE GUIANA HIGHLANDS 17

Description of species. — ( 1 7 males, 2 females, 2 juveniles, including type
series). Body moderately slender: head wider than body, nearly as wide as
long, depressed: snout moderately long, round in dorsal view and in profile,
projecting slightly beyond margin of lip: canthus rostralis slightly elevated,
angular: loreal region concave: lip rounded; nostril slightly protuberant
dorsolaterally; internarial area depressed: top of head flat; skin not co-ossified
with underlying dermal bones: tympanum distinct, separated from eye by
distance equal to about one-half length of tympanum; supratympanic fold
moderately heavy, covering upper one-fourth of tympanum. Axillary mem-
brane extending one-fourth length of upper arm; forearm moderately slender,
bearing row of low tubercles ventrolaterally: fingers moderately long, bearing
round discs; width of disc on Finger III about two-thirds length of tympanum;
relative lengths of fingers 1 < 2 < 4 < 3; subarticular tubercles moderately
large, round, simple: supernumerary tubercles small, subconical, present only
on proximal segments; palmar tubercle bifid; thenar tubercle elliptical; brown
nuptial excrescences present; fingers barely webbed basally. Hind limb
moderately short, robust: small tubercle present or not on heel; tarsal folds and
tubercles absent; inner metatarsal tubercle elliptical, flat, visible from above;
outer metatarsal tubercle indistinct or absent; toes moderately short, bearing
round discs slightly smaller than those on fingers; subarticular tubercles large,
round; supernumerary tubercles absent; toes about one-half webbed; web-
bing formula I( 1 '/: -2)— (2-2'/:)IIl '/: —(2'/: -3)1111 '/: — (21/: -3)IV2— ( 1-
1 '/:)V. Skin on dorsum smooth, bearing numerous small, pointed tubercles in
males and scattered spicules in females; skin on flanks smooth; skin in post-
tympanic region and axilla weakly areolate: skin on belly and proximal
posteroventral surfaces of thighs coarsely granular; other ventral surfaces
smooth; anal sheath long; anal opening directed ventrally at lower level of
thighs, opening bordered laterally by small tubercles. Vomerine odontophores
angular, diverging posteriorly, narrowly separated medially, between diago-
nally elliptical choanae. each bearing 6-10 teeth; total number of vomerine
teeth 12-18 (x= 14.9) in males, 18-19 (x = 18.5) in females; tongue broadly
cordiform, shallowly notched behind, barely free posteriorly; vocal slit small,
at posterolateral margin of tongue; vocal sac bifid, posterior on throat with
lateral extension behind angle of jaw.

Color in preservative: Dorsum brown with faint darker brown middorsal
blotch (only evident peripherally in some individuals) extending from eyelids
and/or interorbital region to sacrum or postsacral region; dark brown, irregu-
lar or transverse marks on dorsal surfaces of limbs — two on forearm and three
each on thigh, shank, and tarsus; dark brown canthal and supratympanic
stripes; area above latter usually paler than rest of dorsum; posterior surfaces
of thighs brown; labial stripe creamy tan, expanded below orbit; flanks gray
with small dark brown spots; anal and ulnar tubercles cream; venter cream
with brown flecks on margin of chin; brown flecks or reticulations on middle

18 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP.. No. 147

of throat and chest in some specimens.

Color in life: Dorsum dark brown with pinkish tan supratympanic mark;
side of head dark brown; labial stripe cream; flanks grayish brown with dark
brown spots; anal tubercles creamy white; throat creamy yellow with gray
flecks; belly cream; hidden surfaces of limbs dull reddish brown; iris dull
grayish bronze with fine black reticulations (Fig. 8).

Measurements (in mm) and proportions ( 17 males followed by 2 females):
snout-vent length (SVL) 27.6-34.7 (x = 31.8), 35.9-38.1 (x = 37.0); tibia
length/SVL 0.450-0.569 (x = 0.509), 0.501-0.5 15 (x = 0.508); foot length/
SVL 0.350-0.396 (x = 0.379), 0.383-0.384 (x = 0.384); head length/SVL
0.330-0.374 (x = 0.348), 0.339-0.340 (x = 0.340); head width/SVL 0.318-
0.360 ( x = 0.340), 0.325-0.33 1 ( x = 0.328); tympanum/eye 0.7 1 1-0.875 ( X
= 0.781),0.766-0.771 (x = 0.769).

Distribution. — Osteocephalus rodriguezi is known from elevations of
1 , 100-1 ,2 10 m on the north face of the Sierra de Lema and the northern part
of the Gran Sabana in Estado de Bolivar, southeastern Venezuela. Locality
records are: Km 144 on El Dorado-Santa Elena de Uairen road, 1210m [KU
166998-7013, 167767 (C&S)]: Paso del Danto (MCZ 64740, UPR-M 2207,
2209-11).

Remarks. — The specimens examined include the type series. The above
description differs from that of Rivero ( 1 968 ) in that he stated erroneously that
the skin on the dorsum was granular and that the feet were three-fourths
webbed. Rivero ( 1968) described the species from "Paso del Danto ... ca.
1400 m above San Isidro." Paso del Danto is a narrow, steep incline adjacent
to the Salto del Danto encompassing elevations of 1 ,000-1 , 1 50 m above sea
level. Rivero found the frogs in terrestrial bromeliads in March, a relatively

i

Fig. 8. Osteocephalus rodriguezi, KU 167007, female, 35.9 mm SVL.

HYLID FROGS FROM THE GUIANA HIGHLANDS

o.o

0.2

0.4

0.6

0.8

1.0

Time in Seconds

Fig. 9. Advertisement call of Osteocephalus wdriguezi. KU Tape 1 298; effective
band width 45 Htz; recorded at 16°C on 22 July 1974 at Km 144 on El Dorado-Santa
Elena de Uairen road.

dry time of year in the Sierra de Lema. On 1 7 and 22 July, 1 974, the frogs were
found breeding in shallow pools on the Gran Sabana at Km 144 on the El
Dorado-Santa Elena de Uairen road, 1 .2 10 m (26 km by road S of Paso del
Danto). No frogs of this species were found in terrestrial bromeliads at Paso
del Danto in July 1 974, May and June 1 978, or January 1 979. These limited
data on habitat and behavior, together with the short hind limbs, suggest that
this species may be terrestrial, as opposed to the arboreal habits of the other
species in the genus.

At Km 144 males were calling from low vegetation and from shallow
water; the latter were not floating in the water but were partially submerged
and grasping vegetation with their hands. The mating call consists of 5-8
short, biphasic notes (Fig. 9). Analysis of recordings (KU Tapes 1296-1298)
of three individuals recorded at 16°C reveals that the mean number of notes
per call is 6.6. The repetition rate is 1 4-20 ( x = 1 6 ) calls per min; the dominant
frequency is at 1,700-2,000 Htz.

ACKNOWLEDGMENTS

We thank Barry Clarke, Alice G. C. Grandison, Juan A. Rivero, and Ernest
E. Williams for the loan of specimens. Anne M. Musser executed Figures 1
and 2; Debra Bennett executed the other drawings. Duellman is grateful to
his field companions — Juan R. Leon, John E. Simmons, Linda Trueb, and

20 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP.. No. 147

Dana Trueb Duellman in 1974, and Stephen J. Gorzula and Glenda Medina
in 1979. His field work was supported by grants (DEB 74-01998 and 76-
09986) from the National Science Foundation. Hoogmoed's field work was
supported by grants from the Treub Foundation, the Treub Society, and the
Netherlands Foundation for Tropical Research (WOTRO grants W84-191,
W87-78, WR87-131). He thanks his field companion, Peter Gibbs, who
accompanied him in 1978.

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