Society for the Study of Amphibians and Reptiles
A New Hylid Frog from the Llanos of Colombia
Author(s): William F. Pyburn
Source: Journal of Herpetology, Vol. 7, No. 3 (Aug. 6, 1973), pp. 297-301
Published by: Society for the Study of Amphibians and Reptiles
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1973 JOURNAL OF HERPETOLOGY 7(3): 297-301
A New Hylid Frogfrom the Llanosof Colombia
William F. Pyburn
Department of Biology, The University of Texas at Arlington
Arlington, Texas 76010
ABSTRACT-Hyla kennedyi is a new, terrestrial member of the Hyla rostrata species
group from the Colombian llanos. It differs from other members of the rostrata group in
structural features, thigh pattern and breeding call.
* * *
Between 8 April and 20 April 1971, a field party from the University of Texas at
Arlington traveled over the Colombian llanos from Villavicencio, at the base of the eastern
Andes, to Mapiripan,a village in southeastern Meta on the Guaviare River. At two of our
campsites in this vast region of grassland,gallery forests and rolling hills, we collected specimens
of a long-nosed hylid frog that apparently representsan unnamedspecies. For this new form I
have chosen the name
Hyla kennedyi, sp. nov.
Holotype.-UTA A-3697, an adult male, collected by the author, 13 April 1971, ca.
110 mi ESE Puerto Gaitan, Departamentode Meta, Colombia. The frog was taken from the
base of a tall grass clump at the edge of a recently filled pond in grassland.
Paratypes.-UTA A-3694 (adult female, Fig. 1), UTA A-3695 (adult male), collected by J.
R. Glidewell, 12 April 1971, 21.5 mi E Puerto Gaitan, Departamentode Meta, Colombia;UTA
A-3696, collected by J. Taulman, other data as in holotype; UTA A-3698-3700 (3 specimens),
data as in holotype.
Diagnostic Characters.-A small member of the Hyla rostrata species group with elongate
snout projecting well beyond lower jaw; nostrils low, not elevated above snout profile; no
tubercles along edge of lower jaw; heel tubercles low, not projecting;skin relatively smooth;
anterior and posterior surfaces of thighs orangewith regulartransverseblack bars;web between
first and second toes greatly reduced;web between fingers basal or absent; finger and toe discs
small, their width about 1/2 to 2/3 diameter of tympanum; breeding call a rather long,
slow-pulsed note; habitat grassland.
Description.-Head flat, longer than wide; snout attenuate, projecting beyond lower jaw;
canthus indistinct, loreal region sloping; nostrils slightly raised, not elevated above snout profile;
eye small, length slightly less than distance from eye (anterior corner) to nostril (posterior
edge); tympanum distinct, its upper edge concealed by a prominent, arched, supratympanic
fold; tympanum diameter 2/3 eye length and separated from eye by about 2/3 tympanic
diameter; tongue cordiform, edges free; vomerine teeth in two transverse, slightly curved rows
lying between centers or posterior halves of choanae; vocal slits of male large, extending from
below mid-lateraledge of tongue to angle of jaw; female without vocal slits; lower jaw without
tubercles along edge; skin of dorsum smooth to moderately tuberculate, the tubercles most
prominent on top of head and snout; skin of belly granular,gular skin smooth to granular;
vocal pouch of male median, subgular.
No axillary membrane; no ulnar fold; wrist fold poorly developed or absent; no web
between first and second fingers, a basal web between second and third and between third and
fourth fingers; finger discs small, wider than long, about 1/2 to 2/3 diameter of tympanum;
subarticular tubercles of fingers subconical, none bifid; thenar tubercle large, oval, palmar
tubercle large, bifid; breeding males with gray nuptial pads but no pollical spines; legs slender,
297
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298 WILLIAM F. PYBURN
smooth, heel of adpressed limb ex-
?'. tending to point about midway between
eye and nostril; low rounded tubercles on
heel, a dermal heel fold present; a row of
low rounded tubercles along outer edge
of foot from heel to base of fifth toe, no
tarsal fold; web between first and second
toes vestigial, other toes 1/2 to 3/4 web-
bed; subarticular tubercles of toes
rounded to subconical, none bifid; inner
metatarsal tubercle small, oval; outer
metatarsal tubercle small, rounded; toe
discs small, wider than long, size about
equal to size of finger discs.
Color in life: dorsum of head and
w 10.5(9.9-11) body medium gray with dark gray or
'? ...... ; ~
- ':;~:~'-- ~black blotches, the dark blotches usually
FIGURE 1. Hyla kennedyi, UTA A-3694, female paratype. edged with a pale cream line; a dark inter-
ocular triangle with posteriorly directed apex usually continucus with a broad median dorsal stripe
(Fig. 1), and paired blotches over shoulders and flanks; upper surfaces of limbs medium gray, the
forearm, thigh, shank and foot barrkennedwith dark gray or black, the regular transverse bars of the
thigh extending onto the anterior and posterior surfaces and separated by dull orange spaces; a
mottled gray pattern on the proximal posterior thigh surface; skin of belly and lower sides of legs
cream gray, gula light to medium gray.
Color in preservative (alcohol after formalin): dorsum similar to color in life except pattern
less distinct because of darkening of light edges of blotches and darkening of ground color;
anterior and posterior thigh surfaces cream white with 3 or 4 black transverse bars; dark bars on
upper surfaces of forearm, shank and foot; fingers, toes and toe webbing medium gray; gula light
to medium gray; remainder of venter mostly cream white.
Measurements in millimeters of type series (mean and range of 6 males followed by measure-
ment of one female): snout-vent, 33.0 (31.1-35.3), 37.3; head length, 11.8 (10.5-12.9), 13.3; head
width 10.5 (9.9-11.2), 11.5; internares, 2.5 (2.3-2.8), 2.5; eye length (two specimens with damaged
eyes not included), 3.4 (3.3-3.5), 3.6; eye-to-nostril, 3.6 (3.1-4.2), 4.4; ear diameter, 2.2 (2.1-2.2),
2.2; tibia length, 17.3 (16.2-18.9), 20.3; width fourth toe disc, 1.2 (1.0-1.4), 1.3; width fifth toe
disc, 1.1 (0.9-1.3), 1.3; width third finger disc, 1.2 (1.0-1.3), 1.4; width fourth finger disc,
1.1 (0.9-1.2), 1.4.
Habitat-Two specimens of Hyla kennedyi (UTA A-3694, 3695) were found, oneinthe
afternoon and one at night, in tall grass
the type series were taken at night from
the periphery of an extensive temporary
pool in grassland, where they were calling
from the ground at the bases of tall grass
M,t~~
~ikne_ ,hwn eclumps.
The small digital discs and smooth
skin of Hyla kennedyi minimize body sur-
face areas and may therefore be corre-
lated with its terrestrial habit and rather
dry habitat (Fig. 2). Hyla kennedyi is the
only grassland species in the rostrata
group as that group is presently under-
FIGURE 2. Colombian llanos in vicinity of Puerto Gaitan, stood (see Duellman, 1972). With the
MtaE 2howinahabitat of Hvla kennedvi. o
JVjt;Lo, ZOIIUVViliti [Link] W- I lyl- exception of Hyla rostrata all other
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HYLID FROG FROM LLANOS 299
species are primarily rain forest inhabitants. These forest dwellers have large toe discs and various
epidermal projections which may serve to break up the body outline. H. rostrata, with
large discs but a relatively smooth skin, is ecologically intermediate, occupying the ecotone
between forest and grassland. It occurs in savanna, scrub forest and gallery forest in Colombia,
where breeding males call from tall-grass stems, shrubs and low branches of trees.
Breeding Call.-Calls of Hyla kennedyi were recorded at the type locality, 13 April 1971, air
temperature 24.7 C, with a Uher 4000 Report-L recorder, and analyzed in the laboratory with a
Kay Electric Sonagraph, Model 6061 B. The call (Fig. 3) is a series of single notes repeated
irregularly, the note repetition rate being about 1 to 4 notes per minute. Each note is a loud,
trilled, burst of sound with no change in pitch, and with a metallic quality. Thirteen notes of 3
individuals have a mean duration of 1.77 (0.66-2.90) seconds and a mean pulse rate of 26.2
(24.1-31.3) pulses per second. The dominant frequency is a band 400 Hz wide centered about
1200 Hz (lower and upper limits, 1050 and 1600 Hz). There is an emphasized frequency band at
about 3600 Hz (limits, 3300 and 3800 Hz), and 6 of 13 notes also have an emphasized band at
about 1700 Hz (limits, 1550 and 2100 Hz).
Comparisons.-The elongate rostrum, small eyes, reduced web between first and second toes,
truncated digital pads and bold thigh pattern of Hyla kennedyi indicate relationship with the Hyla
rostrata species group. Within this group H. kennedyi resembles H. rostrata in lacking a row of
tubercles along the edge of the lower jaw and in having a relatively smooth dorsum. However, the
two species differ in size, H. kennedyi being much smaller. Fifteen male H. kennedyi (males of the
type series and 9 specimens referable to this species from Venezuela UPR M-132-137*, 3139-3140,
3142) have a mean snout-vent length of 33.6mm (31.1-35.7 mm); 9 male H. rostrata (UTA
A-2646, A-2648, A-3701-3707) from Colombia have a mean snout-vent length of 43.2 mm
(42.0-45.2 mm).
Hyla egleri is the smallest species in the rostrata group. It differs from H. kennedyi in body
size, thigh color pattern and size of toe discs. According to Lutz (1968) adult male H. egleri range
from 24 to 30 mm in snout-vent length. A series of 34 male H. egleri (KU71783-71785,
127565-127569, 127572-127580, 127618-127623, 127625-127630, 130031-130035) from Para,
Brazil are 26.1 mm (23.3-28.3mm) in snout-vent length. In life the concealed surfaces of the
thighs of H. egleri are dull gray or violet (Lutz, 1968), in contrast to the orange of H. kennedyi. In
preservative the dark thigh markings of H. egleri are irregular bars or broken blotches, whereas the
three to four dark thigh bars of H. kennedyi are relatively regular and unbroken. Figure 4 shows
the difference between these 2 species in finger disc/tympanum ratio relative to snout-vent length.
5-
4 A* 4 w * o
e ,r- - * *
2 2-
0.2 0.4 0.6 0.8 1.0 1.2
SECONDS
FIGURE 3. Audiospectrogram of breeding call of Hyla kennedyi, type locality, 13 April 1971, air temperature
24.7 C.
*Referred to Hyla boulengeri by Rivero (1961, p. 122).
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300 WILLIAMF. PYBURN
100- 0 0
o 8
90-
8
o 0
85- o0
Z 0 00 0
80- ?
75- 0 0
5 0o
70- O o O O
u._ 0
a65-
5^0 00
60-
. 0 000
55-
50- 0
45 I I I I I I I t I *1 I I I I
22 23 24 25 26 27 28 29 30 31 32 33 34 35 36
SNOUT-VENT LENGTH
FIGURE 4. Relation between third finger disc width/tympanum ratio and snout-vent length; open circles
represent Hyla egleri males, black circles represent Hyla kennedyi males. Measurements of H. kennedyi include
the type series and 9 specimens from Puerto Ayacucho, Venezuela (UPR M-132-137, 3139-3140, 3142).
All of the remaining species in the rostrata group have a row of tubercles (absent in H.
rostrata and H. kennedyi) along the edge of the lower jaw (Duellman, 1972). These species further
differ from H. kennedyi as follows: H. boulengeriis larger;the snout-ventlengthof South Ameri-
can males ranges from 35.5 mm (Leon, 1969) to 48.7 mm (Duellman, 1972). H. garbei has an
elongatetubercleon the heel; both H. proboscideaandH. epacrorhinahavea fleshy proboscisand
the latter has a mottled thigh pattern and elongate heel tubercle (Duellman, 1972) rather than
light and dark bars on the thigh and no elongate heel tubercle as in H. kennedyi.
A comparison of the breeding call characteristics of H. kennedyi with data and audiospectro-
grams presented by Duellman (1972) reveals that the call of H. kennedyi differs from the calls of
all other membersof the [Link] call of H. epacrorhinashows some resemblanceto the
call of H. kennedyi in note duration and emphasized harmonics. However, the pulse rate of H.
epacrorhina averages 68 (65-70) pulses per second, much higher than the pulse rate of H. ken-
nedyi. The pulse rate of calls of the latter species is about half that of H. rostrata (mean 51, range
50-60 pulses per second), which otherwise has the slowest pulse rate of any species in the rostrata
group.
Nelson (1972) noted that, in the call of the microhylidChiasmocleispanamensis,the pulse
rate increased by an average of about three pulses per second with a rise in temperature from 23 C
to 27.7 C. Duellman (1972) did not state temperatures at which his recordings of rostrata group
frogs were made, but it seems very unlikely that differences in temperature could account for the
great differences in pulse rate between H. kennedyi and other species in the rostrata group.
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HYLIDFROGFROMLLANOS 301
ACKNOWLEDGMENTS
This new species is named for Dr. J. P. Kennedy in recognitionof his outstandingcontri-
butions to the Society for the Study of Amphibiansand [Link] Colombiawas financed
jointly by the InternationalBiologicalProgramand the Universityof Texas at Arlington(UTA). I
am especiallyindebtedto WandaCarlPyburnfor her help in all phasesof the field work;to Evelyn
Buhl for the preparationof Figures 3 and 4; to M. J. Fouquette, Jr., Arizona State University
(ASU), for providingme with audiospectrogramsof breedingcalls; to WilliamE. Duellman,Uni-
versity of KansasMuseumof NaturalHistory (KU), for calling my attention to certainspecimens
in the Malkincollection of the AmericanMuseumof NaturalHistoryand to Dr. Jorge Hernandez
of INDERENA for advice on traveling conditions in Colombia. Drs. Duellman and Fouquette
loaned specimens in their care, as did Juan A. Rivero,Universityof PuertoRico, Mayaguez(UPR
M) and CharlesW. Myers,AmericanMuseumof NaturalHistory.J.J.R. Glidewelland J. Taulman
assistedin the field work. To all these personsI am most grateful.
LITERATURECITED
Duellman, W. E. 1972. South American Frogs of the Hyla rostrata group (Amphibia, Anura, Hylidae). Zool.
Mededel. Leiden, 47:177-192.
Leon, J. R. 1969. The systematics of the frogs of the Hyla rubra group in Middle America. Univ. Kansas Publ.
Mus. Nat. Hist. 18:505-545.
Lutz, B. 1968. New Brazilian forms of Hyla. Pearce-SellardsSer. Texas Mem. Mus. 10:1-18.
Nelson, C. E. 1972. Distribution and biology of Chiasmocleis panamensis (Amphibia: Microhylidae). Copeia
1972: 895-898.
Rivero, J. A. 1961. Salientia of Venezuela. Bull. Mus. Comp. Zool. 126:1-207.
Editor's Note: Drs. Hobart M. Smith, Department of Environmental, Population & Organismic Biology,
University of Colorado, Boulder, Colorado; Juan A. Rivero, Department of Biology, Instituteof Caribbean
Science, University of Puerto Rico, Mayaguez, Puerto Rico; and Bertha Lutz, Prof. Emeirta, Museu Nacional,
Rio de Janeiro, Brazil served as special editors for this manuscript.
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