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A new treefrog (Hylidae : Hyla) from southern Ecuador and a description of its antipredator behavior

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We describe a new species of Hyla from southern Ecuador that is diagnosed by the absence of an omosternum; the presence of an enlarged, curved, and pointed prepollex in males; features of coloration; and an associated suite of presumed anti-predator behaviors. Upon capture, frogs exuded a white, sticky fluid; continued perturbation elicited a defensive posture in which white patches on the posterior sides of the limbs and the vent were displayed. The phylogenetic relationships of H. tapichalaca with respect to the H. armata, H. larinopygion, and H. pulchella groups are discussed.
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Accepted: 10 February 2003
Associate Editor: Edmund Brodie III
Herpetologica, 59(3), 2003, 339–349
by The Herpetologists’ League, Inc.
A NEW TREEFROG (HYLIDAE: HYLA) FROM SOUTHERN ECUADOR
AND A DESCRIPTION OF ITS ANTIPREDATOR BEHAVIOR
DAVID KIZIRIAN
1,4
,LUIS A. COLOMA
2
,AND ARTURO PAREDES-RECALDE
2,3
1
Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, CA 90007, USA
2
Museo de Zoologı
´a, Centro de Biodiversidad y Ambiente, Departamento de Ciencias Biolo
´gicas,
Pontificia Universidad Cato
´lica del Ecuador, Av. 12 de Octubre y Roca, Aptdo. 17-01-2184, Quito, Ecuador
3
Laboratorio de Herpetologı
´a, Universidad San Francisco de Quito, Cı
´rculo de Cumbaya
´,
Aptdo. 17-12-841, Quito, Ecuador
ABSTRACT: We describe a new species of Hyla from southern Ecuador that is diagnosed by the absence of
an omosternum; the presence of an enlarged, curved, and pointed prepollex in males; features of coloration;
and an associated suite of presumed anti-predator behaviors. Upon capture, frogs exuded a white, sticky fluid;
continued perturbation elicited a defensive posture in which white patches on the posterior sides of the limbs
and the vent were displayed. The phylogenetic relationships of H. tapichalaca with respect to the H. armata,
H. larinopygion, and H. pulchella groups are discussed.
Key words: Antipredator behavior; Anura; Ecuador; Hyla armata group; Hyla larinopygion group; Hyla
pulchella group; Hyla tapichalaca; New species
ECUADOR is among the most biologically
diverse countries and hosts the richest (per
unit of area) amphibian fauna in the world,
including 425 formally described species
(Coloma and Quiguango-Ubillu
´s, 2000–
2002). In addition, Duellman’s (1999) model
of the rate of discovery and descriptions of new
species in South America predicts that this
4
CORRESPONDENCE: e-mail, kizirian@nhm.org
September 2003] HERPETOLOGICA 339
number will increase significantly. Among the
several reasons for the expected increase is
that many previously unexplored areas are now
accessible to biologists, in particular, the
Andes of extreme southern Ecuador. One
locality in this region is Cerro Tapichalaca in
the Nudo de Sabanilla, which has not been
subjected to a thorough survey of amphibians.
At this site, a large undescribed species of Hyla
was recently discovered. Herein, we describe
the new taxon and its antipredator behavior
and discuss its relationships with respect to the
H. armata group, H. larinopygion group, and
Andean members of the H. pulchella group.
MATERIALS AND METHODS
Specimens were collected at night near
a stream at the type locality, in most cases,
by locating calling individuals. Latitude, lon-
gitude, and elevation were determined using
a Garman GPS 48 personal navigator. Muscle
and liver tissue samples were preserved in
100% EtOH and are currently stored at 70 C
at the Natural History Museum of Los Angeles
County (LACM) and at 20 C at the Museo de
Zoologı
´a of the Pontificia Universidad Cato
´lica
del Ecuador (QCAZ). Specimens were fixed in
7–10% formalin, then transferred into 70–75%
EtOH for permanent storage. Specimens of
the new species were compared to published
descriptions and museum material (Appendix
I). Figures were drawn freehand from speci-
mens or from scanned images. Cleared-and-
stained (C&S) material was prepared following
Dingerkus and Uhler (1977), with minor
modifications proposed by Linda Ford (per-
sonal communication). Osteological nomen-
clature follows Duellman and Trueb (1986)
and Trueb (1993). Institutional abbreviations
follow Leviton et al. (1985). Bilateral variation
is reported as left/right. Measurements were
made to the nearest 0.1 mm with Max-Cal
digital calipers in the manner described by
Duellman (1970). Additionally, we measured
hand length (‘‘Hand L’’—distance from the
proximal edge of the inner palmar tubercle to
the tip of Finger III) and disk diameter
(‘‘Disk’’—taken on Finger III). Webbing
formulae were determined following Savage
and Heyer (1967) with adjustments recom-
mended by Myers and Duellman (1982).
Recordings were made with a Radio Shack
CTR-101 tape recorder, a 33-3017 Optimus
Unidirectional microphone, and a Maxell IEC
Type I audio cassette. During recording, the
microphone was held 5–50 cm from the frog.
The original tape recording (QCAZ CC 36) is
deposited in the tape archive of QCAZ. Calls
were edited and analyzed using SoundEffects
0.9.1 (Alberto Ricci, 1993–1994) and Canary
1.2.4 (Cornell Laboratory of Ornithology,
2002) software packages. Calls were input in
a frequency range of 44.1 kHz. Acoustic
terminology follows Heyer et al. (1990). To
facilitate comparison, the template for the
description approximately follows Duellman
and Coloma (1993).
SYSTEMATICS
Hyla tapichalaca sp. nov.
Holotype.—QCAZ 14611 (LACM Field
Series 655), an adult male from a small stream
on the road between Yangana and Valladolid,
Reserva Tapichalaca, Provincia Zamora-Chin-
chipe, Ecuador; 048289S, 798089W; 2667 m;
obtained on 30 October 2000 by David
Kizirian.
Paratypes.—LACM 148770 and KU 291362,
adult males with same data as holotype. QCAZ
15083 (C&S male), 15084–85 (males), 16704
(male), 16705 (female), 16706 (C&S female),
17776 (male), 17777 (female) with the same
data as holotype, except that the former three
were collected on 5 February 2001 by Italo
Tapia-Caisaguano and the latter five were
collected on 17 September 2001 by Italo and
Ketty Tapia-Caisaguano.
Diagnosis.—Color pattern, associated anti-
predator behavior, and absence of an omo-
sternum distinguish Hyla tapichalaca from all
other Hylidae. The new taxon is most similar to
species in the H. armata,H. larinopygion,and
H. pulchella groups, but is diagnosed from the
two former by the presence of a curved and
pointed prepollical spine (trapezoidal in H.
larinopygion group; elongate, triangular in
H. armata) and from the latter by the presence
of a mostly black ventral color pattern
(uniformly pale in H. pulchella group) and
a small frontoparietal fontanelle (large in H.
pulchella group).
Description of holotype.—Male (Fig. 1);
SVL 59 mm; body and limbs robust; snout
nearly truncate in lateral view; head length 18
340 HERPETOLOGICA [Vol. 59, No. 3
mm; head widest at level of eyes (21 mm);
canthus rostralis with rounded ridge; loreal
region concave; lips rounded; dorsal surface of
internarial region concave; nostrils barely pro-
tuberant, directed anterolaterally, slightly pos-
terior to anterior margin of lower jaw; top of
head slightly concave; diameter of eye 6.5 mm;
distance from anterior corner of eye to nostril
4 mm; tympanic annulus indistinct; supratym-
panic fold thick, obscuring posterodorsal edge
of tympanum, extending from eye to shoulder.
Forearms robust; axillary membrane absent;
ulnar tubercles absent; fingers broad (Fig. 1A);
disks round to oval; width of disk on Finger III
3 mm; lateral fringes absent; palmar surface
with subequal tubercles, deep folds, subartic-
ular tubercles single, large, thick, rounded,
thenar tubercle thick, elliptical; prepollex
large, spine not protruding, nuptial excre-
sences absent, manus webbing formula: I3-3
II 2-3 III 3-3 IV/I3-3 II 2–3 III 3-3 IV, hind
limb robust; tibia length 31/32 mm; foot length
46 mm; heel tubercle small, pyramidal; inner
FIG. 1.—Palmar (A) and plantar (B) views of Hyla
tapichalaca (QCAZ 14611, holotype, male).
FIG. 2.—Hyla tapichalaca in life (QCAZ 15083, male).
September 2003] HERPETOLOGICA 341
tarsal fold absent; inner metatarsal tubercle
oval (Fig. 1B); outer metatarsal tubercle
absent; subarticular tubercles round; super-
numerary tubercles present; pes webbing for-
mula: I2-3 II 2-3 III 2-3 IV 3-2 V/I2-3 II 2-3
III 2-3 IV 3-2 V.
Skin on throat and anterior portion of chest
rugose, on belly and proximal portion of
ventral surface of inner thigh colliculate,
elsewhere relatively smooth; transverse fold
above vent distinct; tissue below and adjacent
to vent slightly swollen, bordered by low
bumps and folds; cloacal opening directed
posteriorly; vocal sac single, median, subgular.
Dorsum dark brown-gray with little hint of
pattern overall; dorsum and venter of Finger
disks II–IV and prepollex white; Finger I white
except for proximal and lateral portions, which
are gray; nearly chevron-shaped white blotch
below and lateral to vent, ventral portion of
which extends anteriorly nearly to abdomen as
midventral line; white blotches on posterior
and ventral surfaces of calcars and elbows;
throat and chest brown; abdomen dark gray to
TABLE 1.—Variation in Hyla tapichalaca including snout–vent length (SVL), tibia length (tibia), head length (Head L),
head width (Head W), ear diameter (Ear), internarial distance (Nares), toe disc width (Disc), and hand length (Hand L).
Museum Number Sex SVL TIbia Head L Head W Ear Nares Disc Hand L
KU 291362 M 60.0 31.0 21.0 21.0 3.5 6.0 3.5 20.5
LACM 148770 M 61.5 31.0 22.0 22.0 3.0 6.0 3.5 20.5
QCAZ 15085 M 63.8 30.4 20.2 20.1 3.6 5.5 3.6 19.7
QCAZ 15084 M 62.9 31.2 20.0 21.6 3.2 5.0 3.1 20.3
QCAZ 15083 M 62.8 30.0 17.5 20.9 3.5 5.3 2.7 19.8
QCAZ 16704 M 61.7 30.4 20.0 21.1 3.2 5.3 3.3 21.1
QCAZ 16706 M 62.4 30.8 20.5 20.7 3.4 4.8 3.8 21.6
QCAZ 17776 M 59.1 29.8 19.6 19.7 3.0 4.8 3.3 20.0
QCAZ 16705 F 66.5 32.9 21.2 21.8 3.3 5.8 3.8 20.0
QCAZ 17777 F 66.3 32.1 21.3 21.0 3.2 5.7 3.3 20.4
FIG. 3.—Presumed advertisement call of Hyla tapichalaca (QCAZ 16704, male).
342 HERPETOLOGICA [Vol. 59, No. 3
black; lips white; thin crescent-shaped white
marking on sternum; faint white reticulation
on flanks; faint white lines on anterior face of
thigh; white scars dorsally and ventrally on
right thigh.
Color in life, male (based on QCAZ 15085;
Fig. 2).—Dorsum gray with faint olive suffu-
sion; flanks gray brown, bearing diffuse pink-
orange marks that are more conspicuous in
axillary and inguinal regions; thighs and shanks
with brown transverse bands; digital disks,
lateral and lower margin of vent, knee, elbow,
and outer margin of forearm white; venter
black, except pectoral and gular regions, which
are gray; ventral portion of digital disks pink;
iris yellow-gold with fine black reticulations.
Color in life, female (based on QCAZ
16705–06, 17777).—Dorsum pale brown;
flanks, dorsum of manus and pes gray; digital
disks, margins of lips, lateral and ventral
margin of vent, knee, elbow, and outer margin
of forearm white; yellow-cream stippling on
rostral keel, canthus rostralis, supraorbital and
supratympanic margins; inguinal region and
anterior surface of thighs with pink suffusion
in QCAZ 16706; interdigital membranes gray;
iris yellow-gold.
Intraspecific variation.—Midventral portion
of white vent marking absent in KU 291362,
LACM 148770, QCAZ 15083, 15085, 17776;
faint in QCAZ 16705. Marking on sternum
bow-tie shaped in LACM 148770, QCAZ
16704–05; nearly sinusoid and extends across
chest in QCAZ 15084, 16706, 17776–77;
indistinct in QCAZ 15083, 15085, 16705.
Banding on hind limbs (anterior and posterior
surface of thighs, dorsal inner tarsus) distinct
in QCAZ 15084–85, 16706. White vent
marking subrectangular in LACM 148770.
White elbow and knee markings restricted to
that region or extend as stripe of various
lengths along outer and ventral margins of
adjacent forearm and tarsus, respectively;
extends along nearly entire forearm and left
tarsus in QCAZ 17777. White color pattern
on dorsum of Finger I varies from distal one-
third of length (KU 291362, QCAZ 15083,
17776) to nearly entire length of digit, at least
medially (LACM 148770, QCAZ 14611,
15084–85, 16706). QCAZ 15084 has much
paler color pattern overall, being gray dorsally
with pale orange reticulations on flanks and
hind limbs.
FIG. 4.—Dorsal (A) and ventral (B) views of skull and
(C) mandible and hyobranchium of Hyla tapichalaca
(QCAZ 16706, female). Stippling represents cartilage.
Drawings of posteromedial processes may be distorted
because they are not in a horizontal plane. Scale 55 mm.
September 2003] HERPETOLOGICA 343
Sexual dimorphism.—Females exhibit great-
er SVL than males (Table 1). Females have less
robust forelimbs than males. Females possess
a rostral keel, which is bordered by white
(yellow-cream in life) stipples that form a line
that continues along the canthus rostralis,
supraorbital, and supratympanic dorsolateral
margins; stipples diminish in number posteri-
orly; few stipples on supratympanic fold in
QCAZ 17777.
Call.—A recording of QCAZ 16704 was
made in the field at 2100–2200 h on 17
September 2001 at an air temperature of
10.1 C and 99% relative humidity. The
behavioral significance of this call is unknown,
although a female was found near the calling
male (I. Tapia-Caisaguano, personal observa-
tion). The calling frog was located approxi-
mately 2–3 m from a stream margin, near
a small waterfall (2 m in height), on a branch
approximately 2.5 m from the ground. A
recording of 31.5 s contained 565 notes, which
were emitted singly (n521) or in bouts (n5
144) of 2–7 notes. A 10-min segment of the
recording contained 62 bouts and one note.
Within this segment, bout duration was 1.38 s
(two notes)–8.2 s (seven notes) and intervals
between bouts and the note varied from 4.39–
14.01 s (x58.12, n562, SD 52.11). A bout
of 8.2 s included seven notes, which had
a duration of 129.5–161.6 ms (x5147.2 ms,
SD 510.202) (Fig. 3). Six intervals between
notes varied from 877.4–1757 ms (x51185
ms, SD 5375.771). The dominant frequency
coincides with the fundamental frequency,
which begins at 1.082 and ends at 1.943 kHz
with a mean frequency peak at 1.378 kHz. Four
harmonics are visible, the first two having peak
frequencies at about 3.27 kHz and 4.82 kHz.
Cranium and hyoid (Fig. 4).—Sutures
between prootic and frontoparietal present;
frontoparietals fused; crista parotica directed
laterally; supraorbital process large, flat, elon-
gate, with irregular margins; posterior process
of tympanic ring present in male, small process
present on left side of female; tympanic ring
incomplete dorsally; pars externa plectri of
stapes with small cartilaginous process; maxil-
lary teeth extend posteriorly nearly to anterior
edge of tympanic ring; vomers sutured to
overlying sphenethmoid; five/five teeth on
vomerine ondontophore; anterior flange of
vomer with two distinctly bilaterally asymmet-
rical cusps, anteriormost cusps oriented to-
wards medial end of maxilla; lateral branch of
premaxilla distinctly shorter than palatine;
dorsal alary process of premaxilla oriented
obliquely posterodorsal to pars dentalis in
male, nearly perpendicular in female; septum
nasi cartilaginous; dorsal alary process longer
than pars dentalis; otic branch of squamosal
does not extend posteriorly beyond stapes;
extreme anterior end of zygomatic branch of
squamosal cartilaginous in male, bony in
female; hyoglossal sinus of hyoid ‘‘U-’’ to ‘‘V’’-
shaped.
Pectoral girdle (Fig. 5).—Suprascapula os-
sified, except posterior margin in female;
cleithrum ossified; omosternum absent; epi-
coracoid mineralized anteriorly, more than
FIG. 5.—Ventral view of pectoral girdle of Hyla tapichalaca (QCAZ 16706, female). Stippling represents cartilage.
Calcified areas are indicated by irregular stippling on the overlay. Scale 55 mm.
344 HERPETOLOGICA [Vol. 59, No. 3
75% in male, about 50% in female; mesoster-
num entirely mineralized in male, approxi-
mately 75% mineralized in female; internal
edge of bifurcation of xiphisternum ‘‘U-’’ to
‘‘V’’-shaped; xiphisternum more than 50%
ossified in male, entirely cartilaginous in
female.
Vertebral column (Fig. 6).—Transverse pro-
cesses of third presacral vertebra posteroven-
trally directed with respect to sagittal plane;
extreme distal portion of transverse processes
of sacrum ossified; sacrum-coccyx fusion
absent in male, present in female; postero-
laterally directed processes on anterolateral
region of coccyx absent.
Limb osteology (Figs. 7–8).—Extreme an-
terior end of ilium cartilaginous; prepollex
ossified; prepollex of female composed of two
segments, proximal segment forms a nearly
isosceles triangle, distal segment (left) forms
a sharp sigmoid spine; prehallux completely
ossified in male, first and second of three
segments partially cartilaginous in female.
Distribution and ecology.—Hyla tapichala-
ca is known only from the type locality,
a small cascading stream in Montane Cloud
Forest (Valencia et al., 1999) in the Nudo de
Sabanilla (Fig. 9). Annual mean precipitation
in this region is 1000–2000 mm and annual
mean temperature is 12–18 C (Can
˜adas-
Cruz, 1983).
The geographic distribution of H. tapicha-
laca is between that the H. larinopygion group
(Colombia and Ecuador), H. armata group
(central Peru to central Bolivia), and Andean
members of the H. pulchella group (central
Peru to central Argentina) (Duellman et al.,
1997). Despite their relatively large size and
striking color patterns, species in these groups
are not commonly encountered, in part,
because their calls are barely audible over
stream noise (Duellman and Hillis, 1990).
Indeed, all species in the H. larinopygion
group (sensu Duellman and Hillis, 1990;
Duellman et al., 1997) have become known
in the last 30 yr (Duellman, 1973). Given that
H. armata is phylogenetically nested within the
FIG. 6.—Dorsal view of vertebral column and pelvic
girdle of Hyla tapichalaca (QCAZ 16706, female). Scale 5
5 mm.
September 2003] HERPETOLOGICA 345
H. larinopygion group (Duellman et al., 1997;
Paredes-Recalde, 1999), it seems likely that
focused efforts might reveal additional species
of related frogs in the Andes of northern and
central Peru.
Reproduction.—Three females collected in
September 2001 contain eggs, QCAZ 17777
has small ovarian eggs, while QCAZ 16705 and
16706 contain large ovarian eggs. The latter
has 428 eggs that are approximately 2.96 mm
in diameter (range 52.76–3.4, n515, SD 5
0.189).
Behavior.—Upon being captured, H. tapi-
chalaca exuded a sticky white fluid with
a strong odor. During attempts to photograph
one specimen (LACM 148770; LACM Photo-
graphic Collection 1387–88 [www.nhm.org/
research]; Fig. 10), it assumed a posture in
which the white markings on the posterior
surfaces of the elbow, heel, and vent were
more obviously displayed. In addition, the
hind limbs were outstretched laterally and the
forelimbs were positioned under the chin. The
posture was similar to that described for
Xenohyla truncata (Napoli, 2001; H. da Silva,
personal communication), except that the
lungs were not inflated in H. tapichalaca.
When disturbed, H. pantosticta (www.puce.
edu.ec/Zoologia) and H. staufferorum (A. Glue-
senkamp, personal communication) position
their forelimbs in a fashion similar to H.
tapichalaca but have not been observed to
raise the posterior end of the body and do not
have distinctive isolated markings on the vent,
elbow, or knee as does H. tapichalaca.
Mostly superficial scars are present on the
hind limbs, especially anteroventrally, of
QCAZ 14611–12, 15085, and LACM 148770.
The injuries leading to scars could have been
incurred if a frog’s outstretched limbs snagged
on marginal spines of narrow leaves (present
on plants on which some H. tapichalaca were
found). Similar scars are not present on two
specimens of H. truncata examined, but are
present on some H. lindae,H. pacha,H.
pantosticta,H. ptychodactyla,H. psarolaima,
FIG. 7.—Dorsal views of (A) left forelimb and (B) left
hind limb osteology of Hyla tapichalaca (QCAZ 16706,
female). Scale 55 mm.
346 HERPETOLOGICA [Vol. 59, No. 3
and H.staufferorum. If scars are correlated
with the antipredator posture, then the behav-
ior described for H. tapichalaca might also be
exhibited by other species of the H. larinopy-
gion and H. pulchella groups.
One male specimen (QCAZ 16704) has
scars on its head, and another male (QCAZ
17776) has scars on the dorsum in the supra-
scapular region. The position of these scars
suggests that they may have been incurred as
a result of combat between males, as has been
hypothesized for Plectrohyla (Duellman and
Campbell, 1992).
Tadpole.—Duellman et al. (1997:13) de-
scribed tadpoles from cascading streams at
3300 and 3450 m in northern Peru (KU
181882–83) that may represent H. tapichalaca.
A recently metamorphosed individual (KU
uncataloged material), collected in southern
Ecuador in the same month and year as some
type material, may also represent H. tapicha-
laca.
Etymology.—The specific epithet is an
indeclinable word based on the name of the
protected forest where all known specimens
were collected. Reserva Biolo
´gica Tapichalaca
is owned by Fundacio
´n Jocotoco and was
created to protect the habitat of rare birds,
including the Jocotoco Antpitta, Grallaria
FIG. 8.—Dorsal view of left forelimb osteology of Hyla
tapichalaca (QCAZ 15083, male). Scale 55 mm.
FIG. 9.—All known localities for the Colombian and
Ecuadorian members of the Hyla larinopygion group and
H. tapichalaca:1.H. caucana,2.H. sarampiona,3.H.
larinopygion,4.H. psarolaima,5.H. pantosticta,6.H.
lindae,7.H. psarolaima,8.H. ptychodactyla,9.H. lindae,
H. pacha,H. psarolaima, 10. H. tapichalaca. Andean
members of the H. pulchella group occur in southern Peru,
Bolivia, and northern Argentina. Dotted line represents
600-m contour line and stippling represents areas over
3000 m.
September 2003] HERPETOLOGICA 347
ridgelyi (Krabbe et al., 1999), which is known
only from this region.
REMARKS
The assignment of H. tapichalaca to a spe-
cies group in Hylidae is problematical. Hyla
tapichalaca possesses an enlarged, curved, and
pointed prepollex, which is diagnostic of the H.
pulchella group, but also has a black venter
(with white markings) and a small frontopari-
etal fontanelle as do members of the H.
larinopygion group (Duellman et al., 1997).
In addition, the humerus in H. tapichalaca is
intermediate between that of the H. larinopy-
gion and H. pulchella groups in that it is more
robust than that in the species of the H.
larinopygion group, but lacks the distal flange
present in the H. pulchella group (Fig. 8;
Duellman et al., 1997, their figure 10).
Speculation on phylogenetic relationships of
the new species is further complicated by the
fact that, despite comments to contrary (Duell-
man et al., 1997:36, 38), the H. larinopygion
group cannot be diagnosed exclusive of H.
armata (Duellman et al., 1997, their figure 19;
Paredes-Recalde, 1999).
Hyla tapichalaca may be unique among
hylids in lacking an omosternum; however, the
condition is unknown for many species in the
family. Furthermore, this structure exhibits
homoplastic evolution across Anura (e.g.,
Lynch, 1973; Trueb, 1973) and the relation-
ships and monophyly of Hylidae, with respect
to other taxa lacking an omosternum, such as
Centrolenidae and Allophryne ruthveni, are
ambiguous (e.g., Austin et al., 2002; Duellman
and Trueb, 1986; Ford and Cannatella, 1993).
RESUMEN
Se describe una especie nueva de Hyla del
sur de Ecuador. El taxo
´n nuevo se diagnostica
por carecer de omosternum, tener un pre-
pollex grande, curvo y puntiagudo, por aspec-
tos de la coloracio
´n y un set asociado de
comportamientos antipredatorios. Cuando son
capturadas, las ranas secretan un fluido blanco,
pegajoso. A mayor perturbacio
´n asumen una
postura que enfatiza las manchas blancas en las
partes posteriores de las extremidades y regio
´n
cloacal. Se discuten las relaciones de H.
tapichalaca con respecto a los grupos H.
larinopygion,H. armata eH. pulchella.
Acknowledgments.—We thank N. Acosta for assisting
with the acoustic analysis; M. Bustamante for preparing
Figure 2; H. da Silva for assisting with identification of
cranial elements, offering an opinion on hylid phylogeny,
and providing unpublished data on the behavior of H.
truncata; W. Duellman for loaning comparative material
and admonishing D. Kizirian prior to his foray into hylid
systematics; K. Flum and T. Howard for rendering Figures
8 and 10, respectively; I. Tapia and K. Tapia for the call
recording; and E. Wild for clarifying matters regarding
Duellman et al. (1997). The W. M. Keck Foundation and
Center of Biodiversity and Environment of the Pontificia
Universidad Cato
´lica del Ecuador provided funds for
fieldwork.
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Accepted: 6 December 2002
Associate Editor: Joseph Mendelson III
APPENDIX I
Specimens Examined
Locality data are given in Duellman et al. (1997) and
Duellman and Hillis (1990) or are provided below.
Hyla alytolylax: QCAZ 496: Ecuador: Provincia Coto-
paxi: San Francisco de Las Pampas.
H. armata: KU 163342, 173219; 173220 (male, C&S);
164085 (female, dry skeleton).
H. psarolaima: KU 164314–15 (paratypes).
H. lindae: KU 202729–30; QCAZ 10483-84 (C&S):
Provincia Napo: Oyacachi.
H. larinopygion: QCAZ 10486 (C&S), 10490 (C&S):
Provincia Imbabura: Cuellaje.
H. pantosticta: KU 202732–33; QCAZ 4506, 10488:
Provincia Sucumbı
´os: Santa Ba
´rbara.
H. pacha: KU 202760 (paratype), KU 202763.
H. ptychodactyla: KU 209781 (paratype); QCAZ 2275
(C&S): Provincia Cotopaxi: Environs of San Francisco de
las Pampas.
H. pulchella: LACM 51175–76: Argentina: Buenos
Aires: Esteban Echeverria, 2 km W Monte Grande;
132507: Argentina: Co
´rdoba: Achiras.
H. staufferorum: KU 217694, 217696 (paratypes);
QCAZ 11146 (C&S): Provincia Napo: 27 km N Jondachi,
Cordillera de Guacamayos.
H. truncata: KU 92190–91: Brasil: Rio de Janeiro: Barra
de Sa
˜o Joa
˜o.
September 2003] HERPETOLOGICA 349
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