Herpetologica, 62(1), 2006, 96–108
2006 by The Herpetologists’ League, Inc.
A NEW SPECIES OF THE HYPSIBOAS BENITEZI GROUP FROM THE
WESTERN AMAZON BASIN (AMPHIBIA: ANURA: HYLIDAE)
,DIEGO F. CISNEROS-HEREDIA
Herpetology, Division of Vertebrate Zoology, American Museum of Natural History, Central Park West at 79
York, NY 10024-5192, USA, and Department of Ecology, Evolution, and Environmental Biology, Columbia University
Department of Zoology, National Museum, 11579 Praha 1, Czech Republic
Universidad San Francisco de Quito, Ave. Interoceanica y Calle Diego de Robles, Campus Cumbaya
edif. Maxwell, Casilla Postal 17-12-841, Quito, Ecuador
Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; and Department
of Zoology, Hessisches Landesmuseum Darmstadt, Friedensplatz 1, 64283 Darmstadt, Germany
ABSTRACT: A new species of the Hypsiboas benitezi group is described from the western Amazon Basin.
The new species is characterized by its medium size (SVL 24.4–31.2 mm in adult males, 26.0–36.0 mm in adult
females), absence of nuptial excrescences, palpebral membrane lacking golden reticulation, and a mental
gland in males. The dorsal surfaces of living frogs are bright yellowish green to tan, and frequently bear an X-
shaped mark; the parietal peritoneum, pericardium and sclera are covered by guanophores. The new species
differs from other members of the Hypsiboas benitezi group (H. benitezi,H. lemai,H. microderma,H. pulidoi,
H. rhythmicus, and H. roraima), by the combination of a dorsal X-shaped marking, absence of golden
reticulation on palpebral membrane, and central portion of abdominal parietal peritoneum, pericardium, and
sclera covered by guanophores. The new species has been confused with Hyloscirtus albopunctulatus (as Hyla
albopunctulata) in the past, but the study of the type series of the latter indicates that they represent different
species. Hypsiboas hutchinsi is excluded from the H. benitezi group and transferred to the H. semilineatus
group, as it shares with H. geographicus and H. semilineatus the absence of a projecting spine in the prepollex
(the only known species of Hypsiboas with this character state), presence of pigmented nuptial excresences,
reticulated palpebral membrane, and ﬁnely tuberculated skin.
Key words: Amphibia; Anura; Hylidae; Hypsiboas nympha; New species; Western Amazon Basin
THE WESTERN Amazon Basin of Colombia,
Ecuador and Peru holds a remarkably high
biological diversity with regard to amphibians.
For example, 112 frogs have been recorded
from the Iquitos region, northeastern Peru
´guez and Duellman, 1994) and 120
amphibian species from a single site, the
Tiputini Biodiversity Station, in Amazonian
Ecuador (Cisneros-Heredia, 2003). Recent
ﬁeldwork in these regions and the study of
collections led to the discovery of a new
species of Hypsiboas.
This new species was included in the recent
analysis and taxonomic revision of the family
Hylidae by Faivovich et al. (2005), whose most
parsimonious trees showed it to be nested within
a group of western amazonian and Guayana
highland species (Hypsiboas benitezi,H.
lemai,H. microderma,H. roraima, and other
undescribed species). This clade was named
the Hypsiboas benitezi group, and is supported
mainly by molecular data, and by the presence
of what was termed a ‘‘ﬂat’’ mental gland.
Besides the already mentioned species, Faivo-
vich et al. (2005) tentatively included H.
hutchinsi,H. pulidoi, and H. rhythmicus in
Goals of this paper include (1) the de-
scription of the new species, (2) a discussion
of differences between the new species and
Hyloscirtus albopunctulatus, a species of the
H. bogotensis group (Faivovich et al., 2005)
with which it had been confused in the past,
and (3) brieﬂy discuss the composition of the
Hypsiboas benitezi group, as further study
indicates that H. hutchinsi should be excluded
MATERIALS AND METHODS
Thirty-one specimens of the new species
were examined, along additional material that
is listed in the Appendix. Snout dorsal out-
lines follow Heyer et al.’s (1990) standards.
Webbing formulae follow Savage and Heyer’s
(1967) convention as subsequently modiﬁed
by Myers and Duellman (1982). All other
terminology is that of Duellman (1970). Ab-
breviations used throughout the text are: DF3,
CORRESPONDENCE: e-mail, email@example.com
width of disc on third ﬁnger; EN, eye to nostril
distance; ED, horizontal eye diameter; FL,
foot length as the distance from the proximal
edge of the inner metatarsal tubercle to the tip
of the fourth toe; HL, head length as the
straight line distance from the posterior mouth
corner to the tip of the snout; HW, head width
at the level of mouth corners; IOD, interorbital
distance, taken between the anterior margins
of the orbits; IN, internarial distance; SVL,
snout–vent length; TL, tibia length; and, TYD,
tympanum diameter. All measurements (in
mm) were taken using a dissecting microscope
and electronic digital calipers (0.05 mm
accuracy and rounded to the nearest 0.1
mm); all measurements were taken by the
same person (DFCH), and each measurement
was taken at least three times. Sex was deter-
mined by presence or absence of secondary
sexual characters (prepollical spine and mental
gland) or by direct observation of the gonads.
Relative lengths of digits were determined by
adpressing adjacent digits to one another.
Color pattern in life was taken from ﬁeld notes
and color photographs. Internal soft anatomy
was examined by dissection of preserved
specimens. Drawings were made with the aid
of a stereomicroscope with a drawing tube
attachment and photographs were used to
document eye coloration.
Classiﬁcation of vegetation formations fol-
lows Sierra (1999). Geographic position and
elevation of collection localities were deter-
mined using collectors’ ﬁeld notes and the
2000 physical map of the Republic of Ecua-
dor 1:1,000,000 distributed by the Instituto
´ﬁco Militar, and NIMA (2003).
Throughout this paper we adopt the tax-
onomic changes recently introduced by Fai-
vovich et al. (2005), that resulted in the
partition of the formerly paraphyletic Hyla
into 15 genera, and the redeﬁnition of several
of its former species groups, particularly within
Hypsiboas. The literature on hylid frogs
mentioned in this paper other than Faivovich
et al. (2005) included all referred species in
Institutional abbreviations used are as fol-
low: AMNH—American Museum of Natural
History, New York; BMNH—British Museum
of Natural History, London; DFCH-USFQ—
Universidad San Francisco de Quito, Quito;
EPN—Departamento de Biologı
´cnica Nacional, Quito; FHGO-USFQ—
´gica G. Orce
deposited at Universidad San Francisco de
Quito, Quito; ICN—Instituto de Ciencias
Naturales, Universidad Nacional de Colombia,
´; KU—Natural History Museum at the
University of Kansas, Lawrence; MCZ—Mu-
seum of Comparative Zoology, Harvard Uni-
versity, Cambridge; NHMG, Naturhistoriska
¨teborg, Sweden; NMP6V—Na-
tional Museum, Prague; QCAZ—Museo de
´a, Universidad Cato
´lica del Ecuador,
Quito; ROM—Royal Ontario Museum, Tor-
onto; USNM—National Museum of Natural
History, Smithsonian Institution, Washington,
DC; ZFMK—Zoologisches Forschungsmu-
seum Alexander Koenig, Bonn.
Hypsiboas nympha sp. nov.
Hyla albopunctulata—Duellman and Mendel-
son, 1995:339; Cisneros-Heredia, 2003:17.
Hyla sp.—Moravec et al., 2002:36.
Hyla sp. 2—Faivovich et al., 2005:11.
Hypsiboas sp. 2—Faivovich et al., 2005:122.
Holotype.—DFCH-USFQ 0355, an adult
male (Fig. 1) from Ecuador: Provincia de
´os: Reserva de Produccio
Cuyabeno, (008059020S, 768129540W, 290 m
a.s.l.), ca. 3.3 km E of the Lago Agrio–Tarapoa-
Puerto El Carmen Road (remarks), collected
on 19 July 2000 by D. F. Cisneros-Heredia, M.
Brandt, A. Leo
´n, C. Ponce, and T. Sugahara.
Paratypes.—AMNH 88088, 94193, 94196,
94198, ZFMK 82884 (females): Ecuador:
Provincia de Morona-Santiago: Cusuime, Rı
FIG. 1.—Living male holotype of Hypsiboas nympha sp.
nov. (DFCH-USFQ 0355).
March 2006] HERPETOLOGICA 97
Cusuime (028409S, 778429W), 60 km airline
SE Macas, collected between May and July
1971 by B. Malkin. KU 221637 (female):
Provincia de Sucumbı
´os: San Pablo de Kant-
esiya (008159S, 768259W), collected on 9
February 1988 by J. M. Touzet. DFCH-USFQ
1012 (male) and 1122 (female): Provincia de
Orellana: Tiputini Biodiversity Station (008379
S, 768109W, 190–270 m), collected on 10
August 1999 and 22 May 2001 by D. F.
Cisneros-Heredia, A. Chiriboga, and T. Suga-
hara. QCAZ 14841 (male): Parque Nacional
´(008409S, 768249W, ca. 300 m),
collected on 21 July 1997 by S. Ron. QCAZ
18805 (male): same locality as before, collected
on 23 January 2002 by S. Padilla. QCAZ 8907
(female): Provincia de Pastaza: Lorocachi (018
379S, 758579W, 220 m), collected on 19
February 1996 by O. Torres et al. NMP6V
71202/1 (male), and 71202/2 (female): Peru:
Departamento de Loreto: ca. 5 km NW of the
village of Tarapoto, ca. 19 km SW of Iquitos
(038479120S, 738259430W, ca. 120 m),
collected on 26 March 2001 by J. Moravec and
I. Arista Tuanama. AMNH 96237 (male),
96238 (female): Yagua Indian village, head-
waters of Rı
´o Loretoyacu: ca. 100 km NW
Leticia (Colombia), collected between 22
April–2 May 1970 by B. Malkin. KU 221849
(male): San Jacinto (02818944.80S, 758519
46.00W), collected on 1 July 1993 by N.
Referred material.—AMNH 94192, 94194,
94198 (juveniles): Ecuador: Provincia de
Morona-Santiago: Cusuime, Rı
(028409S, 778429W), 60 km airline SE
Macas, collected between May and July 1971
by B. Malkin. QCAZ 14706 (male): 2 km E de
Santiago (038029S, 788029W, 220 m),
collected on 28 December 2000. QCAZ 8092
(male): Provincia de Orellana: Carretera Pom-
peya Sur-Iro km 38 (008309S, 768409W, ca.
300 m): collected on 21 January 1995. EPN
1562: Cotapino (008429S, 778289W, 600 m)
(unknown sex, dehydrated specimen), collected
by the Olalla brothers, unknown date. QCAZ
9685 (male): Parque Nacional Yasunı
m), collected on 11 May 1996; FHGO-USFQ
3674 (female): same locality as before, collected
on 04 September 2002. FHGO-USFQ 2694
(male): Bataburo Lodge, Rı
on 03 September 2001. EPN 8950: Provincia
de Pastaza: Unknown locality (unknown sex,
poorly preserved adult), collected by A.
´riz. DFCH-USFQ 0390 (juvenile):
Provincia de Sucumbios: collected at the type
locality. QCAZ 16327 (unknown sex, poorly
preserved adult): Singue (008149N, 768189W,
ca. 280 m). QCAZ without number: Puerto
Bolivar, Cuyabeno (08059190S, 768089
310W, 240 m) collected on August 2003.
ICN 47270 (male): Colombia: Departamento
de Amazonas: Vı
´a Tarapaca km. 9, Comunidad
Monilla-Amena, collected between 2 and 14
Diagnosis.—A medium-sized member of
the Hypsiboas benitezi group, (SVL 24.4–
31.2 in adult males, 26.0–36.0 in adult
females), and deﬁned by the following combi-
nation of characters: slender body, head wider
than body; snout truncate in dorsal view and
protruding in lateral view because of strongly
protuberant nostrils; large prominent eyes;
palpebral membrane lacking golden reticula-
tions; small tympanum; limbs long and slen-
der; axillary membrane absent; biﬁd distal
subarticular tubercles on ﬁngers I, III, and IV;
enlarged prepollex with slender slightly pro-
jecting bony spine in males; nuptial excres-
cences absent in males; mental gland in males;
hands one-fourth webbed, feet three-ﬁfths
webbed; no inner tarsal fold, tarsal tubercles
absent; heel tubercles and calcars absent;
cloacal sheath absent or very short; in life,
dorsal surfaces bright yellowish green to tan,
ventral surfaces translucent bluish or greenish
white, iris creamy golden with black periphery,
bones green, parietal peritoneum, pericardium
and sclera covered by guanophores.
Comparisons.—A prepollical spine is pres-
ent in all species of Bokermannohyla and most
Hypsiboas (Duellman et al., 1997; Faivovich
et al., 2004, 2005). Hypsiboas nympha is easily
distinguished from members of most species
groups of these genera by the mental gland
in males. Exceptions are other members of
the H. benitezi group (H. benitezi,H. lemai,
H. microderma, and H. roraima; character
state unknown in H. pulidoi, and H. rhythmi-
cus), H. albopunctatus group (H. heilprini),
and H. punctatus group (H. granosus and H.
punctatus). The absence of a leaf-green dorsal
coloration in life distinguishes H. nympha
from H. granosus and H. punctatus.Hypsiboas
nympha differs from H. heilprini for its smaller
size, much more slender body, less-developed
98 HERPETOLOGICA [Vol. 62, No. 1
prepollical spine, and absence of saclike folds
around the cloaca; furthermore, the mental
gland in H. heilprini is conspicuously thicker.
Hypsiboas nympha differs from all species in
the H. benitezi group for lacking a calcar
(present in H. microderma and H. roraima),
absence of golden reticulation on the palpebral
membrane (present in H. microderma and
H. roraima), absence of any pattern on the
thighs (brown markings on hidden surfaces of
thighs in H. roraima), frequent occurrence of
X-shaped dorsal marking (absent in all other
species), guanophores present on peritoneum
and pericardium (absent in H. benitezi,H.
microderma,andH. roraima; character state
unknown in H. pulidoi and H. rhythmicus),
lavender or purple coloration in preserved
specimens (brownish or grayish in all other
Juveniles of Hypsiboas nympha could be
confused with the syntopic Dendropsophus
miyatai. However, D. miyatai has red mark-
ings on a metallic yellow to pink dorsum, a pink
venter and a pinkish-tan iris, whereas juveniles
of D. nympha have green markings on a bright
yellow dorsum, a greenish venter and a greenish-
tan iris with black periphery.
Description of holotype.—Adult male; SVL
29.8; body slender; head wider than body,
slightly wider than long, HW/HL 51.02,
widest below eyes; snout truncate in dorsal
view, slightly protruding in lateral proﬁle
because of the prominent nostrils; eye-nostril
distance slightly longer than diameter of eye,
EN/ED 51.13; canthus rostralis curved,
slightly rounded in section; loreal region
concave; lips thin, slightly ﬂared; internarial
region distinctly depressed; nostrils strongly
protuberant, directed dorsolaterally. Interor-
bital area ﬂat, IOD/ED 51.90, IOD/HW 5
0.57; eyes large and protuberant, ED/HL 5
0.30, ED/HW 50.30; upper eyelid width, 3.2.
Palpebral membrane translucent, lacking golden
reticulations. Supratympanic fold poorly de-
veloped, semi-circular in outline, obscuring
uppermost part of annulus, extending up to
insertion of the arm; tympanum small, distinct,
directed dorsolaterally, separated from eye by
a distance slightly longer than tympanum
diameter; tympanum diameter slightly larger
than DF3. Arm slender, not hypertrophied,
lacking an axillary membrane; a row of barely
developed ulnar tubercles appearing as an
indistinct fold; ﬁngers moderately long, bear-
ing moderately large, round discs, diameter of
disc on Finger III slightly wider than tympa-
num; relative length of ﬁngers I ,II ,IV ,
III; subarticular tubercles small, round, distal
one of Fingers I, III, and IV biﬁd, most
prominent on Finger I; supernumerary
tubercles absent; carpal tubercles barely no-
ticeable; inner metacarpal tubercle small, ﬂat,
round; outer metacarpal tubercle ﬂat, large,
elongate; nuptial excrescences absent; pre-
pollex enlarged, slender bony spine slightly
projecting at tip; webbing basal between
ﬁngers one and two; webbing formula II
IV. Hind limb long and
slender; TL/SVL 50.51; heels overlapping
when hind limbs ﬂexed perpendicular to axis
of body; no tarsal fold; calcars and heel
tubercles absent; toes moderately long, bear-
ing round discs, slightly smaller than those on
ﬁngers; relative lengths of toes I ,II ,III ,
V,IV; subarticular tubercles moderately
large, round, elevated; supernumerary
tubercles absent; outer metatarsal tubercle
absent; inner metatarsal tubercle ovoid, ﬂat,
indistinct; webbing formula I2—2½II 2—3
III 2—3 IV 3—2 V. Skin on dorsum, head, and
dorsal surfaces of forearms and thighs ﬁnely
shagreen; skin on ﬂanks smooth; skin on belly
and ventral surfaces of thighs granular; oval
mental gland. Cloacal opening directed poste-
riorly at upper level of thighs; cloacal sheath
absent; cloacal tubercles present, scattered,
extending to midlevel of thighs. Tongue ovoid,
barely free behind; vomerine odontophores
prominent, in two fairly straight series, each
bearing 12 teeth, and with the inner margins
angled and converging medially; choanae
large, elongate; vocal slits moderately long,
extending from midlateral base of tongue,
almost reaching to angle of jaws; vocal sac
single, median, subgular.
In life, dorsal surfaces light yellowish green,
with distinct X-shaped gray marking on
dorsum, gray horizontal line in sacral region,
and gray interocular line; dark canthal stripe;
dark anterior lateral stripe between the dor-
sum and the ﬂanks; melanophores forming
weakly deﬁned, irregular transversal bars on
dorsal surfaces of limbs; throat greenish cream,
venter and ventral surfaces of limbs trans-
lucent greenish white, central portion of abdo-
minal parietal peritoneum white, visible
March 2006] HERPETOLOGICA 99
through ventral skin; webbing and discs trans-
parent white; iris creamy golden, with broad
black periphery, and sclera covered with
guanophores; bones green (from D. F. Cisneros-
Heredia and M. Brandt ﬁeld notes, 19 July
2000; and color transparencies).
In alcohol, dorsal surfaces lavender to
purplish gray with scattered melanophores.
Faint brownish-purple lines on dorsum, ex-
tending from scapular region to midlevel of
body, forming an indistinct X-shaped marking.
Faint brownish purple horizontal line in the
interocular region, and other in sacral region.
Loreal region, lips and lateral sides of the head
pale white with scattered melanophores; dark
canthal stripe extending from eye to nostril,
larger melanophores scattered around nostrils.
Upper half of ﬂanks pale white with dark
melanophores, forming an indistinct lateral
stripe, lower half of ﬂanks pale white; venter
and throat cream; anterior, posterior, and
ventral surfaces of limbs and thighs cream;
dorsal surfaces of limbs and tights purplish
gray with densely scattered melanophores;
discs and ventral surfaces of hands and feet
creamy white; cloacal region with scattered
melanophores, cloacal tubercles enameled.
Measurements: SVL 29.8, HL 11.0, HW
11.1, ED 3.3, EN 3.8, IOD 6.3, TYD 1.5, DF3
1.2, NN 2.2, TL 15.3, FL 12.0.
Variation.—Variation of measurements and
body proportions is given in Tables 1 and 2. A
photograph of the paratype NMP6V 71202/1 is
shown in Fig. 2; dorsal and lateral aspects of
the head, and the ventral aspects of the hand
and foot of the same specimen are depicted in
Fig. 3. The vomerine odontophores occur
either in straight series anteriorly convergent
(NMP6V 71202/1), or with the inner margins
angled, converging medially (holotype, AMNH
94193, and, DFCH-USFQ 1012). Also, in the
females AMNH 88088, DFCH-USFQ 1122,
TABLE 1.—Variation of measurements (in mm) of adult Hypsioboas nympha. See text for abbreviations.
Mean 6SD Range nMean 6SD Range n
SVL 29.4 62.0 24.4–31.2 10 32.7 62.8 26.0–36.0 10
HL 10.9 61.1 8.2–11.9 10 11.5 61.3 8.7–13.0 10
HW 10.5 60.8 8.8–11.2 9 11.5 60.9 9.5–13.0 10
TYD 1.5 60.1 1.4–1.7 5 1.5 60.3 1.3–1.9 8
ED 3.2 60.4 2.5–3.7 9 3.9 60.4 3.3–4.4 9
DF3 1.3 60.2 1.2–1.5 5 1.4 60.3 1.0–1.8 8
EN 3.8 60.4 3.1–4.3 10 4.0 60.5 3.2–5.0 10
IN 2.2 60.3 1.8–2.5 10 2.3 60.3 1.8–3.0 10
IOD 5.9 60.5 4.6–6.3 9 6.2 60.5 5.2–7.0 10
TL 15.6 60.9 13.5–16.7 10 17.3 61.4 14.6.0–19.4 10
FL 10.7 60.9 9.8–12.0 9 11.5 60.9 9.4–12.5 9
TABLE 2.—Variation of proportions of adult Hypsiboas
nympha. See text for abbreviations.
Range nRange n
TL/SVL 0.49–0.55 10 0.49–0.56 10
FL/SVL 0.32–0.40 10 0.34–0.39 9
HL/SVL 0.34–0.40 10 0.33–0.39 10
HW/SVL 0.31–0.38 9 0.33–0.37 10
IOD/HW 0.52–0.63 9 0.52–0.56 10
HW/HL 0.82–1.07 9 0.90–1.14 10
EN/ED 1.01–1.46 9 0.92–1.25 9
IOD/ED 1.66–2.10 9 1.40–1.68 10
ED/HL 0.25–0.33 9 0.30–0.39 9
ED/HW 0.25–0.38 9 0.29–0.40 9
TY/ED 0.44–0.51 4 0.33–0.51 7 FIG. 2.—Living male paratype of Hypsiboas nympha
sp. nov. (NMP6V 71202/1).
100 HERPETOLOGICA [Vol. 62, No. 1
KU 221637, and ZFMK 82884, the vomerine
odontophores are slightly S-shaped. A some-
what intermediate condition is exhibited by
AMNH 94195. Number of vomerine teeth
varies in some paratypes as follows: males (n5
4) 7–11 (
X58.75) / 9–10 (
(n58) 9–15 (
X511.25) / 10–17 (
The development of the supratympanic fold is
indistinct or diffuse in some specimens
(AMNH 96237–238), probably as a result of
ﬁxation artifacts. The toe-webbing formulae
vary as follow: I(2–2
–3) IV (3
V. The size of the mental gland is variable. In
the paratype AMNH 96237 the glandular
tissue covers approximately the distal third of
the gular area, whereas in NMP6V 71202/1 the
glandular tissue extends posteriorly to the
anterior margin of the distended vocal sac,
covering approximately two thirds of the gular
region (Fig. 4A); the size of the gland in KU
221849 is intermediate between the latter (Fig.
The partially dissected male paratype KU
221849 has ovoid, unpigmented testes, the
lengths (1.8 mm) of which are about 30% of
kidney length; wolfﬁan ducts are notconvoluted.
Overall dorsal coloration in preservative
varies from lavender/purple (DFCH-USFQ
0335, 1012, NMP6V 71202/1–2, AMNH
94193, 14841, QCAZ 18805), pinkish purple
(AMNH 88088, 94192, 94194–195, 94198,
DFCH-USFQ 1122, KU 221637, 221849,
QCAZ 8907, ZFMK 82884) to cream (AMNH
96237–238). Dorsal pattern varies mostly
regarding presence and distinctiveness of the
dark gray interocular band, dorsal X-shaped
marking, and the lateral stripe. The X-shaped
marking is present in most specimens, with the
exception of AMNH 94193 and KU 221849.
On NMP6V 71202/1 and DFCH-USFQ 1122
the marking is barely visible, but was very
distinct in life (photo in Moravec et al., 2002:
Figs. 1, 2). The canthal stripe is less marked in
AMNH 88088 and DFCH-USFQ 1122. The
lateral stripe is absent in NMP6V 71202/1
(color photographs of this specimen published
by Moravec et al., 2002) and AMNH 96238. In
juveniles, the lines that make up the marking
are broader than in adults which exhibit less
prominent narrow lines.
Duellman and Mendelson (1995) described
the coloration in life of male paratype KU
221849 (using the name Hyla albopunctulata)
as ‘‘At night, pale purple; by day, pale dull
green changing to dull tan on dorsum; pale
purple on venter and hidden surfaces of limbs;
peritoneum white; iris silvery bronze.’’ The
color in life of a juvenile (DFCH-USFQ 0390,
SVL 15.5) was ‘‘bright yellow dorsum with
green stripes, greenish tan iris with black
periphery, translucent lemon green thighs
with yellow marks, translucent lemon yellow
FIG. 3.—Male paratype of Hypsiboas nympha sp. nov.
(NMP6V 71202/1). (A) Dorsal, and (B) lateral views of the
head, (C) palmar, and (D) plantar view of right hand and
foot. Scale bar equals 5 mm.
March 2006] HERPETOLOGICA 101
venter, some internal organs visible [liver and
intestines]’’ (from D. F. Cisneros-Heredia and
M. Brandt ﬁeld notes, 20 July 2000). Gross
morphological features are fairly invariant,
with the exception of NMP6V 71202/2 and
DFCH-USFQ 0390, which have a subacumi-
nate snout in dorsal view instead of rounded.
The density of the guanophores covering the
abdominal portion of the parietal peritoneum
is fairly variable, and they could be partially or
totally lost in badly preserved or dehydrated
Distribution and ecology.—Hypsiboas nym-
pha is distributed in the western Amazon Basin
(Fig. 5). Specimens recorded herein originate
from the northern and southern regions of
eastern lowland Ecuador and from northeast-
ern Peru at elevations below 600 m, and from
FIG. 4.—Gular region showing the structure of the mental gland and its variation in males of (A) Hypsiboas nympha sp.
nov. (paratype KU 221849), (B) Hypsiboas nympha sp. nov. (paratype NMP6V 71202/1), (C) Hypsiboas lemai (KU
166831), (D) Hyloscirtus colymba (AMNH 98365), (E) Hyloscirtus palmeri (ICN 20087), (F) Hyloscirtus albopunctulatus
(syntype BMNH 18220.127.116.11). Scale bar equals 5 mm.
102 HERPETOLOGICA [Vol. 62, No. 1
lowlands of Colombia around Leticia. The
northernmost locality of H. nympha is
‘‘Singue,’’ province of Sucumbı
(QCAZ 16327), whereas the southernmost
record is Leticia, Colombia (K.-H. Jungfer,
personal communication May 2004; and
referred specimen ICN 47270). The species
probably also occurs in adjacent western Brazil.
Hypsiboas nympha seems to prefer the
primary Amazonian Lowland Evergreen for-
ests ﬂooded by white waters (varzea) and black
waters (igapo), and is found only occasionally
in Lowland Evergreen nonﬂooded forests
(terra-ﬁrme forest). The type locality is a
swampy area (water depth 5–20 cm), approx-
imately 50 m distance from a stream. The
arboreal vegetation at the type locality includes
species of the families Verbenaceae, Laura-
ceae, Arecaceae, Bombacaceae, Actinidiaceae,
and Rubiaceae, with stem diameters up to 450
mm; the herbaceous coverage was poor, the
average canopy height was between 12–23 m
(emergent trees up to 40 m). The general
topography of the type locality consists of low
hills with ﬂat or slightly rounded tops and
slightly convex slopes; the lowermost areas
(where the holotype was collected) were
ﬂooded. The soils at the type locality are
lixiviated, acid, brownish-red typical distro-
pepts. The holotype was collected at night
(2200 h) sitting on a leaf ca. 40 cm above water.
Moravec et al. (2002) reported two specimens
of H. nympha (as Hyla sp.) collected ‘‘in
a swampy area of primary forest with abundant
epiphytes and vines and dense growth of
herbaceous plants, low palms...perched on
the same plant ca. 10 cm beside each other, ca.
130 cm above ground.’’ One female paratype
(DFCH-USFQ 1122) contains approximately
70 unpigmented oviductal eggs of ca. 2 mm.
The advertisement call and larvae are un-
Other hylids collected at the type locality
of Hypsiboas nympha are Dendropsophus
leucophyllatus,D. marmoratus,D. miyatai,
Hypsiboas boans,H. calcaratus,H. fasciatus.
H. geographicus,H. granosus,H. lanciformis,
Osteocephalus cabrerai,O. planiceps,O.
taurinus,O. yasuni,Scinax ruber, and Sphae-
FIG. 5.—Schematic map showing the known distribution of Hypsiboas nympha sp. nov. Square indicates the type
locality. Shaded areas indicate elevations above 500 m a.s.l.
March 2006] HERPETOLOGICA 103
Etymology.—The speciﬁc name is derived
from the Greek ny
eand in allusion to the
beautiful goddesses in Greek mythology that
lived in woods and marshes. The name is used
here as an invariable noun in apposition.
Remarks.—The type locality was reached by
way of a 3320 m transect cut into the primary
rainforest of the Cuyabeno Reserve; this
transect was made during the environmental
survey of the N
˜anpaz foundation (a NGO
partner in 2000 of City Investing Company,
now part of EnCana Corp.). The transect had
its initial point at the southeastern border of
the ‘‘Cielito Lindo’’ farm, on the Lago Agrio-
Tarapoa-Puerto El Carmen road (ca. km 85),
at 008059040S, 768149460W, and runs
eastwards. Two paratypes (NMP6V 71202/1–
2) come from ‘‘ca. 5 km NW of the village of
Tarapoto’’ in the department of Loreto, Peru.
This locality must not be confused with the
much better known town of Tarapoto in the
department of San Martin. The locality
´o Cusuime’’ where Boris Malkin
collected some paratypes of Hypsiboas nym-
pha (AMNH 88088, 94192–195, 94198,
ZFMK 82884) is called ‘‘Cusuimi, Rı
uimi’’ in the physical map of the Republic of
Ecuador 1:1,000,000. Also the locality San
Pablo de Kantesiya where Jean-Marc Touzet
collected one paratype (KU 221637) is called
‘‘San Pablo de Kantesyia’’ in the physical map
of the Republic of Ecuador and in Touzet
Throughout this paper we employ the term
‘‘mental gland’’ to describe the glandular tissue
present in the mental area of males of several
species of the Hypsiboas benitezi group.
Unlike the morphology commonly seen in
the Hyloscirtus bogotensis group, in which the
tissue forms a conspicuous, thick, glandular
mass of tightly packed glandular units
(Fig. 4D–F), the tissue described for species
of the Hypsiboas benitezi group is a thinner
layer composed of looser glandular units (Fig.
4A–C) (for this reason it was termed ‘‘ﬂat’’
mental gland by Faivovich et al., 2005). In the
context of the phylogenetic hypothesis of
Hylinae advanced by Faivovich et al. (2005),
the mental glands present in the Hypsiboas
benitezi group, in the H. granosus þH.
punctatus clade, in H. heilprini, and in the
Hyloscirtus bogotensis group arose indepen-
dently. Detailed histological work is necessary
to more appropriately characterize the mental
gland present in the aforementioned species of
Hypsiboas and to compare it with that present
in the Hyloscirtus bogotensis group (Romero
´rez and Ruiz-Carranza, 1996). Additional
research is also needed to better understand
morphological and perhaps physiological var-
iation of the gland within the H. bogotensis
group, as Ruiz-Carranza and Ardila-Robayo
(1991) noticed that the mental gland of H.
lynchi is not apparent externally, but visible
through dissections, a different condition from
that noticed in the species of Hyloscirtus
examined in the present study.
Duellman and Mendelson (1995) referred
two specimens (KU 221637, 221849), here
designated paratypes of Hypsiboas nympha,
to Hyloscirtus albopunctulatus, and used one
of them for a detailed redescription of the
species. The allocation of that specimen to
H. albopunctulatus was based on the original
description by Boulenger (1882) as well as
notes and pictures of the type series taken in
1969 (Duellman and Mendelson, 1995). How-
ever, our recent examination of the syntypes of
Hyloscirtus albopunctulatus revealed several
differences between the latter and Hypsiboas
nympha. As correctly mentioned by Duellman
and Mendelson (1995), only one syntype (for-
merly untagged, now BMNH 18.104.22.168) has
white spots on the dorsum (Fig. 6A), whereas
in the others the dorsum is uniform pale brown
with small, scattered melanophores (Fig. 6B).
However, all syntypes have wide fringes on
ﬁngers and toes (a putative synapomorphy of
FIG. 6.—Dorsal view of two syntypes of Hyloscirtus
albopunctulatus. (A) BMNH 1822.214.171.124, (B) BMNH
104 HERPETOLOGICA [Vol. 62, No. 1
Hyloscirtus; Faivovich et al., 2005), a thicker
mental gland (putative synapomorphy of the
Hyloscirtus bogotensis group; Duellman, 1972),
a relatively robust body, short and robust
limbs, a well-developed supratympanic fold,
a white supracloacal ridge, a white ridge on
the arm, and a white ridge on the heel extend-
ing along the tarsus, and the outer edge of
the ﬁfth toe. These distinctive characters are
absent in Hypsiboas nympha, suggesting an
incorrect allocation of these specimens to
Hyloscirtus albopunctulatus by Duellman
and Mendelson (1995).
Specimens of Hypsiboas nympha commonly
have been confused in collections, having been
identiﬁed either as Hyloscirtus albopunctula-
tus or centrolenids. The unique lavender or
pinkish dorsal coloration of some preserved
specimens of Hypsiboas nympha is similar to
the coloration found in some species of
Centrolene and Cochranella (Centrolenidae),
and is very different from that of most other
hylids; the only other case we are aware of is the
Dendropsophus rubicundulus group (Napoli
and Caramaschi, 1998). Both H. nympha and
species of Centrolene (e.g., C. prosoblepon,C.
grandisonae)orCochranella (e.g., C. cochra-
nae,C. megacheira) have a greenish dorsal
coloration in life, which, after a short time in
preservative, turns to lavender, pink, or even
cream (variation probably related with the
preservation methods or time in preservative).
Starrett and Savage (1973) discussed the dorsal
color patterns in Costa Rican centrolenids and
termed the pigment that turns purple after
preservation ‘‘pigment A,’’ tentatively identify-
ing it as guanine. Possibly, the pigment in the
skin of H. nympha that changes from green
to lavender is the same found in centro-
lenids. In addition, H. nympha has a parietal
peritoneum, pericardium, and sclera covered
by guanophores, a feature also present in
centrolenids and other Hylinae (e.g., Hyloscir-
tus bogotensis group; Ruiz-Carranza and
In the analysis of Faivovich et al. (2005),
Hypsiboas nympha (as Hypsiboas sp. 2) is
recovered as the sister taxon of H. microderma.
These two taxa are the only species of the H.
benitezi group that have a western Amazonian
distribution, while the other species are re-
stricted to the Guayana Highlands. Current
knowledge of the biology of species of the
H. benitezi group is quite poor. Males are
reported to call along fast ﬂowing streams in
H. benitezi (Donnelly and Myers, 1991; Heyer,
1994; Myers and Donnelly, 1997), H. lemai
(Duellman, 1997), and H. rhythmicus
˜aris and Ayarzagu¨ ena, 2002). Males of
H. roraima call from bromeliads, very close
to small streams (D. B. Means, personal
communication). In H. microderma, however,
males are reported to call in swampy areas
(Pyburn, 1977; Moravec et al., 2002). As
reported earlier, males of H. nympha were
observed in swampy areas, in one case close to
a stream. J. D. Lynch (personal communica-
tion) reports to have seen this species in
Leticia, Colombia, calling along streams.
Relatively large (diameter »2.0–2.5 mm),
unpigmented eggs have been reported for
Hypsiboas lemai (Duellman, 1997) and
H. nympha (this paper); eggs of H. roraima
are also large (diameter »2.0–2.2 mm) and
mostly unpigmented, with the exception of
a small light brown area in the animal pole
(J. Faivovich, personal observation on ROM
35189). Duellman (1997) reported that an
amplectant pair of H. lemai kept in a plastic
bag deposited eggs on a leaf. Based on this,
and occurrence of calling males on vegetation
above a stream, he suggested that eggs are
normally deposited on vegetation overhanging
streams in wich the tadpoles develop. The
occurrence of similar eggs in the other species
could be taken to suggest a similar reproduc-
Hypsiboas hutchinsi was tentatively placed
in the H. benitezi group by Faivovich et al.
(2005) based on its overall similarity, as
perceived from the description of Pyburn
and Hall (1984). Examination of the holotype
suggests that this allocation was incorrect, as
the male holotype lacks a mental gland, so far
the only putative morphological synapomor-
phy of the group. Also, unlike species with
known eggs in the H. benitezi group (see
above), H. hutchinsi has small (diameter ,1
mm) and pigmented eggs (Pyburn and Hall,
There is some evidence to associate Hyp-
siboas hutchinsi with the H. semilineatus
group, as it shares with H. geographicus and
H. semilineatus the prepollex not modiﬁed into
a projecting spine (as far as we are aware, the
only species of Hypsiboas where this character
March 2006] HERPETOLOGICA 105
state occurs), the presence of pigmented
nuptial excresences (otherwise apparently
present only in H. melanopleura of the H.
pulchellus group; Duellman et al., 1997, but
see Lehr and May, 2004), reticulated palpebral
membrane (shared with H. boans,andH.
wavrini, two other species of the H. semi-
lineatus group; otherwise present in H. micro-
derma, and H. roraima; Duellman and
Hoogmoed, 1992; Pyburn, 1977), and the
ﬁnely granular dorsal skin texture of H.
geographicus and H. semilineatus (present as
well in H. microderma according with Pyburn,
1977; not visible in preserved holotype).
Other evidence in potential conﬂict with
a relationship of Hypsiboas hutchinsi with the
H. semilineatus group is that larvae of
H. hutchinsi have the oral disc completely
sourrounded with marginal papillae and a 4/7
labial tooth row formula (Pyburn and Hall,
1984). Larvae of species currently included in
the H. semilineatus group have an anterior gap
in the marginal papillae and a 2/3, 2/4, 2/5, or
3/5 labial tooth row formula (Duellman, 1978;
D’Heursel and de Sa
´, 1999; Martins and
Moreira, 1991). Within Hypsiboas, the larvae
known to have the oral disc completely
sourrounded with marginal papillae are the
ones tentatively assigned to H. benitezi (Myers
and Donnelly, 1997), H. heilprini (Noble,
1927), and those of a still undescribed species
of Hypsiboas that was called H. semiguttatus
by Faivovich (1996). Labial tooth row formu-
lae approaching the reported 4/7 of H.
hutchinsi are known to occur in Hypsiboas in
the larvae tentatively assigned to H. benitezi,
with 4/8 (Myers and Donnelly, 1997), and in
H. heilprini, with 6/9 (Noble, 1927). Note that
in the context of the results of Faivovich et al.
(2005), the oral disc completely sourrounded
with marginal papillae in the tadpole of H.
benitezi is more parsimoniously interpreted as
a plesiomorphy, as this character state occurs
in Aplastodiscus, most Bokermannohyla,Hylo-
Based on the previous discussion, we ﬁnd
that the four putative synapomorphies shared
by Hypsiboas hutchinsi,H. geographicus,and
H. semilineatus outweight the alternative
hypotheses of relationships of H. hutchinsi
with other Hypsiboas. For this reason, we
advance the hypothesis that H. hutchinsi is
a member of the H. semilineatus group.
Acknowledgments.—For the loan of specimens, pro-
vision of working space or kind hospitality, we thank W.
¨hme (ZFMK), B. T. Clarke (BMNH), L. Ford and T.
Grant (AMNH), J. D. Lynch (ICN), L. Trueb and W. E.
Duellman (KU), R. W. McDiarmid (USNM), J. Hanken
(MCZ), G. Nilson (NHMG); L. Coloma (QCAZ), R.
Murphy (ROM), J. A. Campbell (UTA), A. Almenda
(EPN), I. Arista Tuanama (IIAP, Iquitos), and A. Ma
Burgos (UNAP, Iquitos). T. Grant and three anonymous
reviewers read the manuscript and provided useful
criticisms. K.-H. Jungfer, J. D. Lynch, and D. B. Means
kindly provided unpublished data. DFCH is grateful to M.
Brandt, K. Swing, H. Valdebenito, V. Zak, T. Sugahara, A.
´n, L. Zurita and S. Ca
´rdenas for ﬁeld company and
assistance. Work of JM was supported by the project
MK0CEZ99F0201 and conducted in cooperation with the
Museo de Zoologia – UNAP, Iquitos (Andre
Burgos, research authorization N8452-2000-INRENA-
Loreto) under the auspices of the Universidad de la
´a Peruana, Iquitos (the agreement signed by J. T.
´squez and M. Stloukal). Field and laboratory work of
DFCH was supported by the Tiputini Biodiversity Station
and College of Biological Sciences, Universidad San
Francisco de Quito, 2002 Research Training Program,
National Museum of Natural History, Smithsonian In-
stitution, Smithsonian’s Women Committee, and M. E.
and L. Heredia. The Ministry of Environment of Ecuador
provided the scientiﬁc research authorization N819-IC-
FAU-DFN. JF acknowledges the American Museum of
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Environmental Biology, Columbia University, AMNH
Roosevelt Grant, and National Science Foundation Grant
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Associate Editor: Maureen Kearney
Bokermannohyla alvarengai.BRAZIL: Minas Gerais:
Serra do Cipo
´: AMNH 88644, 89373.
˜o Paulo: Bor-
aceia: AMNH 54522–54524.
March 2006] HERPETOLOGICA 107
Bokermannohyla pseudopseudis.BRAZIL: Goias: Cha-
pada dos Veadeiros: AMNH 90194
Dendropsophus miyatai.ECUADOR: Orellana: Tiputini
Biodiversity Station: DFCH-USFQ 231–235; PERU: Lor-
eto: Anguilla: NMP6V 71259.
Hyloscirtus albopunctulatus.ECUADOR: BMNH
126.96.36.199-162 (syntypes); Sarayacu: BMNH
Hyloscirtus alytolylax.ECUADOR: Cotopaxi: Las Pampas:
Hyloscirtus colymba.PANAMA: Chiriqui: Valle de la
Sierpe: ZFMK 38445–448. Cocle
´: Cont. Div. N. El Cope
600 m: AMNH 98364–365.
Hyloscirtus palmeri.COLOMBIA: Valle del Cauca:
Jimenez: BMNH 19188.8.131.52–33 (syntypes); Antioquia:
Frontino: Vereda Venados, Parque Nacional Natural Las
Orquideas, quebrada El Retiro, 850 m: ICN 20087.
Hyloscirtus phyllognathus.PERU: Roque: NHMG 474
Hyloscirtus torrenticola.ECUADOR: Napo: Cascada San
Rafael, on Rio Quijos, at km 102, INECEL station, 1350
m: USNM 286399.
Hypsiboas albomarginatus.BRAZIL: 20 km N Rio de
Janeiro: ZFMK 38801–802, 39495–496.
Hypsiboas albopunctatus.BOLIVIA: Santa Cruz: P.N.
Noel Kempff Mercado: ZFMK 67049, 72698.
Hypsiboas benitezi.VENEZUELA: Bolivar: North side of
Cerro Guaiquinima: AMNH 133843–133848; Amazonas:
Mt. Duida, 2300 ft.: USNM 291067.
Hypsiboas boans.BOLIVIA: Santa Cruz: P.N. Noel
Kempff Mercado: ZFMK 67046; GUYANA: Tableau Pond,
4 mi. (by Rd.) NW Dubulay Ranch house: AMNH
141014–14017; PERU: Loreto: Anguilla: NMP6V 71249;
Madre de Dios: Puerto Maldonado: ZFMK 35909.
Hypsiboas calcaratus.ECUADOR: Orellana: Tiputini
Biodiversity Station; DFCH-USFQ 206, 208–10; PERU:
Loreto: km 31 on road Iquitos–Nauta: NMP6V 71250.
Hypsiboas dentei.BRAZIL: Amapa
´: Vila Nova: AMNH
140755; Serra do Navio: AMNH 140756.
Hypsiboas fasciatus.BOLIVIA: Pando: Cobija: ZFMK
67143–144; ECUADOR: Orellana: Tiputini Biodiversity
Station: DFCH-USFQ 207; PERU: Loreto: 20–30 km SW
of Iquitos: NMP6V 71149, 71251/1–2; Aguajito: ZFMK
Hypsiboas geographicus.BOLIVIA: Beni: Rı
cure: ZFMK 62829; Santa Cruz: P.N. Noel Kempff
Mercado: ZFMK 67047, 72690; ECUADOR: Orellana:
Tiputini Biodiversity Station: DFCH-USFQ 109, 147–8;
´os: Reserva de Produccio
DFCH-USFQ 377; PERU: Loreto: Anguilla: NMP6V
71252/1–2; GUYANA: Berbice River Camp, at ca. 18 mi.
(linear) SW Kwakwa (ca. 2 mi. downriver from Kurundi
River conﬂuence): AMNH 166146–157.
Hypsiboas granosus.BOLIVIA: Pando: Cobija: ZFMK
66820; ECUADOR: Napo: near Tena: DFCH–USFQ 712–
13; GUYANA: Roraima area: ZFMK 47694–695; PERU:
Loreto: Iquitos region: NMP6V 71185–86, 71255.
Hypsiboas heilprini.DOMINICAN REPUBLIC: Provincia
Duarte: Los Bracitos: AMNH 11401 (holotype); 31297–
300; 31305–07; 37477–78 (paratypes).
Hypsiboas hutchinsi.COLOMBIA: Vaupe
´s: ca. 2 km SW
´to: UTA 24819 (holotype).
Hypsiboas lanciformis.BOLIVIA: Pando: Cobija: ZFMK
66776–777; BRAZIL: Acre: km 29 Rio Branco-Puerto Acre
Rd.: AMNH 139307–308.
Hypsiboas lemai.GUYANA: Mount Ayangana: ROM
39569, 39570. VENEZUELA: km 112 road El Dorado–Santa
Elena de Uaire
´n: KU 166831.
Hypsiboas lundii.BRAZIL: Minas Gerais: Serra do Cipo
km 110: AMNH 89374.
Hypsiboas microderma.COLOMBIA: Vaupe
UTA 5012 (holotype); PERU: Loreto: Puerto Almendras,
17 km SW Iquitos: NMP6V 71258/1; Anguilla, 50 km
W Iquitos: NMP6V 71258/2.
Hypsiboas multifasciatus.BOLIVIA: Santa Cruz: P.N.
Noel Kempff Mercado: ZFMK 67048.
Hypsiboas ornatissimus.SURINAM: Marowijne: Loe
Creek: 54 km (air) S. Oelemari: AMNH 90790.
Hypsiboas pellucens.ECUADOR: Pichincha: Santo Dom-
ingo de los Colorados (650 m): AMNH 89690–89695.
Hypsiboas picturatus.ECUADOR: Pichincha: San Vicente
de Andoas: DFCH-USFQ 0BM5; COLOMBIA: Narin
Guayacana (500 m): AMNH 87921.
Hypsiboas polytaenius.BRAZIL: Petropolis: ZFMK 38807.
66813; Santa Cruz: La Florida: ZFMK 60367–373; San
Ramon: ZFMK 60393-394; PARAGUAY: vicinity of Asuncio
ZFMK 55249–250; PERU: Loreto: Puerto Almendras:
NMP6V 71188/1–3; Andoas: ZFMK 39319–322.
Hypsiboas pulidoi.VENEZUELA: Territorio Amazonas:
Monte Duida, 2000 pies: MCZ 72499 (holotype).
Hypsiboas raniceps.PARAGUAY: Canendiyu: AMNH
Hypsiboas roraima.GUYANA: Mount Ayangana: ROM
39611, 39616, 39623–624.
Hypsiboas ruﬁtelus.COSTA RICA: between Limon and
Cahuita: ZFMK 42112; La Selva: ZFMK 56550; PANAMA:
Bocas del Toro: Rambala: ZFMK 47788–791.
Hypsiboas semilineatus.BRAZIL: Espiritio Santo: Soor-
etama, Linhares: USNM 164162–163.
Hypsiboas sibleszi.GUYANA: Mount Ayangana: ROM
39553, 39554, 39556, 39558.
108 HERPETOLOGICA [Vol. 62, No. 1