New Species of Pristimantis (Anura: Terrarana: Strabomantinae) from
Lower Central America
MASON J. RYAN,1,2 KAREN R. LIPS,3 AND J. TOMASZ GIERMAKOWSKI1
1Department of Biology and Museum of Southwestern Biology, University of New Mexico,
Albuquerque, New Mexico 87131-0001 USA
3Department of Biology, University of Maryland, College Park, Maryland 20740 USA
ABSTRACT.?We describe a new species of Pristimantis in the subgenus Hypodictyon ridens Species Series
from the type locality in Parque Nacional General de Divisio?n Omar Torrijos Herrera on the Cordillera
Central in central Panama. This species is similar to Pristimantis caryophyllaceus but can be differentiated
from that form by snout?vent length (female Pristimantis educatoris , 27% larger than P. caryophyllaceus),
round, globular, and projecting subarticular tubercles on all digits (not projecting in P. caryophyllaceus);
finger disk covers even, round, disk pads ovoid, toe disk covers expanded and palmate, and toe pads even
and broadened (finger and toe disk covers and pads rounded in P. caryophyllaceus); outer metatarsal tubercle
large, elongate, and projecting (obscure and small in P. caryophyllaceus). We also report and describe female
parental care of eggs.
Pristimantis is a large genus of frogs with
three subgenera and 427 species (Heinicke et al.,
2007; Hedges et al., 2008). The genus reaches its
greatest diversity in northwestern South Amer-
ica, where its distribution ranges from sea level
to elevations close to 4,000 m in the Andes
(Hedges et al., 2008). Twelve species of Pristi-
mantis occur in lower Central America, includ-
ing five endemic forms (Pristimantis altae,
Pristimantis cerasinus, Pristimantis museosus, Pris-
timantis pardalis, Pristimantis pirrensis), and
seven species are found in both Central America
and Chocoan South America (Pristimantis acha-
tinus, Pristimantis caryophyllaceus, Pristimantis
cruentus, Pristimantis gaigeae, Pristimantis moro,
Pristimantis ridens, Pristimantis taeniatus; Hei-
nicke et al., 2007; Hedges et al., 2008). Eight of
these species belong to the P. ridens Species
Series (subgenus Hypodictyon), with P. cerasinus
in the Pristimantis rubicundus series and Pristi-
mantis gaigei in Pristimantis conspicillatus series
(Hedges et al., 2008). Wang et al. (2008) further
split the P. ridens Species Series by recognizing a
P. pardalis series comprised of P. altae, P.
pardalis, and P. pirrensis.
Frogs of the P. ridens Species Series possess
the following characteristics: small to moderate
size (16?45.2 mm SVL); a distinct tympanic
membrane and annulus; smooth, shagreened, or
tuberculate dorsum; coarsely areolate venter;
unwebbed toes; Toe V much longer than Toe III;
inner tarsal fold present; and vocal slits and
nuptial pads present or absent (Savage, 2002;
Hedges et al., 2008). One of the most recogniz-
able Central American Pristimantis is the slen-
der-bodied P. caryophyllaceus (Barbour, 1928),
which is part of the autochthonous Talamancan
herpetofauna (Savage, 1982). Barbour (1928)
collected and described P. caryophyllaceus from
La Loma, on the trail from Chiriquicito to
Boquete, on the eastern flanks of the Cordillera
de Talamanca in Chiriqu?? Province, Panama. As
currently described, P. caryophyllaceus ranges
from the Cordillera de Tilara?n of Costa Rica to
the Choco?an region of Colombia and occurs in
lowland, premontane, and montane forests
from sea level to 1,900-m elevation (Myers,
1969; Savage, 2002; IUCN, 2008).
Since 1998, KRL has been conducting long-
term population monitoring of the amphibian
fauna of Parque Nacional General de Divisio?n
Omar Torrijos Herrera, known as El Cope?
(8u409N 80u37.629W; 600?900-m elevation). Dur-
ing these surveys, we noticed that individuals
we were identifying as P. caryophyllaceus were
much larger than presumably conspecific indi-
viduals from Costa Rica and western Panama.
Subsequent morphological review uncovered
differences in toe pad shape, tubercle configu-
ration, and morphometrics between popula-
tions in Costa Rica and western Panama and
those in central and eastern Panama. Herein, we
describe the population from central and east-
ern Panama as a new species of Pristimantis.
MATERIALS AND METHODS
We used HerpNet (2008) and Global Biodi-
versity Information Facility (2008) to access data
on museum holdings of P. caryophyllaceus from
AMNH, FMNH, MCZ, MVZ, and NMNH2 Corresponding Author. E-mail: mjryan@unm.edu
Journal of Herpetology, Vol. 44, No. 2, pp. 193?200, 2010
Copyright 2010 Society for the Study of Amphibians and Reptiles
(museum abbreviations follow Leviton et al.,
1985). In addition to seven specimens deposited
at USNM, we located 19 specimens from the
LACM and seven specimens from the UMMZ
(Appendix 1). The drawings in this article of the
hands and feet were done with a camera lucida
by J. Sipiorski. We followed Savage (1987, 2002)
and Lynch and Duellman (1997) in the descrip-
tion of external morphology and the diagnosis
format of Mendelson et al. (2008). We deter-
mined sex by presence of vocal slits in males,
presence of eggs in females, and differences in
tympanum morphology (partially distinct and
visible in females, not visible to barely visible in
males). The tympanum characteristic was dis-
covered on the examination of museum mate-
rial; thus, we did not differentiate between sexes
in the field unless a female was gravid or
attending a clutch of eggs. We used digital
calipers (6 0.1 mm) to measure external
morphology of preserved specimens and scaled
them by snout?vent length. Standard measure-
ments were abbreviated as follows: snout?vent
length, SVL; tibia length, TL; leg length, LL;
head length, HL; head width, HW; eye to
nostril, EN; eye to snout, ES; and eye length,
EL (Lynch and Duellman, 1997; Savage, 2002;
Savage et al., 2004). We provide measurements
of SVL from 123 live individuals that were
released after capture. These measurements
were made with dial vernier calipers in milli-
meters during 2000?2003 at El Cope?. We
obtained mass of live animals using 5-g spring
scales (Pesola AG, Baar, Switzerland). The frogs
were placed in a plastic bag and then weighed.
We subtracted the weight of the bag to obtain
the mass of the frogs.
We analyzed the degree of morphological
difference between populations in southern
Costa Rican and western Panama (P. caryophyl-
laceus) and those in central and eastern Panama
(sp. nov.) using multivariate analysis of vari-
ance (MANOVA) and Discriminant Function
Analysis (DFA). We performed all analyses
with SPSS version 16.0 (SPSS Inc., Chicago)
with an a 5 0.05.
RESULTS
Pristimantis educatoris sp. nov.
Figures 1?4
Holotype.?USNM 572336?adult female; col-
lected from forest just below the Continental
Divide on Atlantic slope; Parque Nacional G. D.
Omar Torrijos H., Cocle?, Panama? (8u39.959N;
80u35.559W; 709-m elevation; Fig. 5), approxi-
mately 8 km north of El Cope?, Cocle? Province,
Panama, 3 July 2002, by K. R. Lips and M. J.
Ryan.
Paratypes.?All paratypes scored in our anal-
ysis are from Parque Nacional G. D. Omar
Torrijos H., Cocle?, Panama? (8u39.959N;
80u35.559W; 709-m elevation): two adult males
USNM 572328 and UMMZ 152952; three adult
females USNM 572336, UMMZ 167571, and
UMMZ 167629. Additional paratypes from
FIG. 1. Photo of a male Pristimantis educatoris from El Cope?. This animal was not collected as part of the
studied series.
194 M. J. RYAN ET AL.
Costa Rica not scored in our analysis from Cerro
Uatsi, Limo?n Province (9u37.209N; 82u56.109W;
600-m elevation): one adult female UCR 13133;
one adult male UCR 13004, all collected by G.
Chaves.
Diagnosis.?A thin, long-limbed, small species
with SVL of adult males 19.3?20.4 mm and of
adult females 23.3?37.7 mm (Table 1). Pristi-
mantis educatoris differs from the other 10
members of the P. ridens Species Series by
having the following combination of characters:
presence of an enlarged superciliary tubercle, a
pointed heel tubercle, a smooth dorsum, and an
elongated snout. Pristimantis educatoris can be
distinguished from P. caryophyllaceus by mean
SVL (males 8% smaller, females 27% smaller in
P. caryophyllaceus, Table 1); projecting subartic-
ular tubercles (nonprojecting subarticular tu-
bercles in P. caryophyllaceus; Fig. 3); expanded,
round and even finger disks and toe pads
(broadened and rounded finger disks and toe
pads in P. caryophyllaceus; Figs. 3, 4); a large,
elongate, inner metatarsal tubercle (small, elon-
gate, and obscure in P. caryphyllaceus); a large,
conical, projecting, outer metatarsal tubercle
(obscure, small, and rounded outer metatarsal
tubercle in P. caryophyllaceus); tip of Toe V not
reaching distal subarticular tubercle of Toe IV
(tip of Toe V reaches beyond distal subarticular
tubercle of Toe IV in P. caryophyllaceus; Fig. 4);
tip of Toe I much shorter than Toe II, not
reaching distal subarticular tubercle on Two II
(Toe I long, reaching beyond distal subarticular
tubercle of Toe II in P. caryophyllaceus; Fig. 4);
most individuals have wartlike white to yel-
lowish spots on dorsum, arms, and lateral
surfaces (no wartlike white spots in P. caryo-
phyllaceus, Fig 1).
Description of Holotype.?Head longer than
wide, head length 92% of HW, HL is 40% of
SVL. Snout rounded in dorsal outline and
profile, small point at tip. Eye to snout 52% of
HL, the tip of snout extends beyond nostrils;
nostril openings ovoid, opening dorsally. Can-
thus rostralis moderately sharp and slightly
FIG. 2. Dorsal (A) and lateral (B) views of the
holotype of Pristimantis educatoris sp. nov. (USNM
572336).
FIG. 3. Right hand of (A) Pristimantis caryophyllaceus and (B) Pristimantis educatoris showing differences in toe
pad and disk shapes.
NEW SPECIES OF PRISTIMANTIS FROM PANAMA 195
curved. Vomerine tooth patches triangular,
widely separated, much smaller than choanae.
Vomerine teeth are located posteriorly and
medially to choanae. The choanae are ovoid.
The tongue is pear shaped, wider in front than
behind. The upper eyelid is smooth, with a
single, enlarged pointed superciliary tubercle.
The tympanum is indistinct and the annulus
tympanicus partially visible beneath skin. The
tympanic membrane is partially visible exter-
nally. Finger II is much longer than Finger I,
relative finger length is IV 5 III . I . II; disks
on Fingers I?III expanded, disk covers are even
and palmate and disk pads are even and
broadened. Finger disks are approximately
twice as wide as digit on Fingers I?III, and
barely wider than digit on Finger IV. The
subarticular tubercles under the fingers are
ovoid, obtuse and raised at the distal margin.
The thenar tubercle is elongate and the palmar
tubercle is circular and bifid. There is no lateral
fringe on the fingers. Several small, round,
projecting tubercles form a raised ridge on
ventral trailing edge of forearm. Toe V is longer
than Toe III, the tip of Toe V does not reach the
distal subarticular tubercle on Toe IV. Toe I is
approximately half the length of Toe II. Relative
toe length is IV . V . III . II . I. The toe disk
covers are expanded and palmate with disk pads
even and broadened. The subarticular tubercles
on the toes are ovoid, projecting, and symmet-
rically globular. There is an indistinct lateral
fringe on the toes. Supernumerary or plantar
tubercles are absent. Toe webbing is absent.
Tarsal folds are absent. The inner metatarsal
tubercle is elongate and projecting, and is
approximately twice the size of outer metatarsal
tubercle. The outer metatarsal tubercle is round
and conical. There is a large pointed heel
tubercle. There are several small round tubercles
that form a raised ridge on the underside of the
ventral trailing edge of the tibia. The dorsal
surfaces are smooth, and the flanks are smooth
with small, scattered tubercles. The chin is
smooth, and the venter is finely granular.
Measurements of Holotype.?Measurements in
millimeters. SVL 5 33.0; HW 5 12.7; HL 5 13.2;
TL5 18.4; LL5 30.3; EL5 4.3; EN5 5.4; ES5 6.9.
FIG. 4. Left feet of (A) Pristimantis caryophyllaceus and (B) Pristimantis educatoris showing differences in toe
pad and subarticular tubercle shapes, and inner and outer metatarsal tubercles.
196 M. J. RYAN ET AL.
Color in Preservative.?Dorsal ground color
medium light brown to dark brown with five
thin darker brown chevrons; fore- and hind
limbs light brown with darker lateral brown
bars and spotting. Venter immaculate off-white
including under surface of limbs; some faint
dark spotting occurs on underside of thigh and
lower forearm and occasionally on hands and
feet. Dorsum and ventral surfaces specked with
fine black dots. No contrasting pattern in the
groin or posterior surface of thigh.
Analysis of Morphological Variables.?We pro-
vide measurements and analysis of morpholog-
ical variation in preserved specimens of P.
caryophyllaceus and P. educatoris (Table 1). We
used SVL-scaled measurements in multivariate
analysis of variance (MANOVA) to determine
whether differences between the two species
exist when all characters are considered simul-
taneously and, if differences exist, which mor-
phological characters contribute most to differ-
ences. We also evaluated classification success
into separate taxa with discriminant function
analysis (DFA). Because TL was highly corre-
lated with LL and EL was highly correlated
with SVL (Pearson?s r . 0.7 in both cases), TL
and EL were excluded from multivariate anal-
yses. Kolmogorov-Smirnov tests of normality of
data for each character (calculated as percentage
of SVL) revealed that only ES differed signifi-
cantly from normality (0.05 . P . 0.01).
Levene?s tests supported homogeneity of vari-
ances for all variables (P . 0.05) except for SVL.
However, homogeneity of variance-covariance
matrices was not rejected by the Box?s M-test
(M 5 40.901, F21,3337 5 1.556, P . 0.05). The
MANOVA test of differences in morphometric
measurements between the two species was
statistically significant (Pillai?s trace 5 0.539,
F6,29 5 5.651, P , 0.001). Univariate tests using
F-statistics showed that species varied signifi-
cantly with respect to SVL (F1,34 5 20.931, P ,
0.001) and EN (F1,34 5 5.638, P , 0.024). The
single discriminant function based on these data
was able to correctly classify over 83% of
individuals.
Variation.?Adult males can be distinguished
from females by SVL and tympanum morphol-
ogy. Males possess an indistinct to barely visible
tympanic membrane, and females possess an
FIG. 5. Map of Costa Rica and Panama showing distributions of Pristimantis caryophyllaceus (closed circles)
and Pristimantis educatoris (open circles with black dots).
NEW SPECIES OF PRISTIMANTIS FROM PANAMA 197
indistinct but visible tympanic membrane.
Between May and August during the years
2000?2003, we made 123 captures of P. educatoris
at El Cope?. We identified 22 captures as females
by their larger size or the presence of eggs
(seven were gravid and two were brooding egg
masses). The average SVL of these females was
28.6 mm (range 5 16.6?37.8 mm). Two gravid
females weighed 2.2 g (31.1 mm SVL) and 2.3 g
(33 mm SVL). All other captures (juveniles,
males, and females) ranged in SVL from 11.1?
37.8 mm (average 5 20.08 6 3.685), and two
adults had masses of 0.8 g.
Dorsal ground color varied from light tan to
medium brown with 4?5 narrow medium
brown chevrons comprised of small darker
brown tubercles forming spots that continue
onto the dorsal surfaces of limbs. The ventral
surfaces are immaculate white to off-white with
occasional faint cream colored to light brown
spotting on undersides of thighs, lower fore-
arms, hands, and feet. There was no contrasting
pattern in groin or on posterior surface of thigh.
Some specimens have a broad middorsal cocoa
brown band (occasionally with darker brown
blotches) bordered by lighter brown laterally.
The dorsum is often speckled with dark brown
dots. Some individuals have numerous bright
white to creamy yellow spots superimposed on
raised tubercles laterally and on limbs (Fig. 1).
The iris color is variable and may be bicolored,
and a white eye-ring may be present or absent.
In some individuals, the color of top portion the
eye may range from blood red to yellow-orange,
whereas the bottom half varies between dark
purple to dark grey. Other individuals may
have a completely red, yellow, to grey eye, with
the color strongest on the outer edges of the eye
and fading in strength nearer the pupil. The
pupil is horizontal.
Etymology.?The species name honors Jay
Mathers Savage and refers to his tutoring and
fostering of multiple generations of Neotropical
herpetologists, including the two senior au-
thors. The root educator refers to one who
nurtures and tutors. This name has a double
meaning because the females of this species also
nurture and care for their direct-developing
eggs.
Distribution.?Pristimantis educatoris occurs
from Santa Fe?, Veraguas Province in central
Panama eastward into Colombia, at eleva-
tions of 450?1,450 m, with a disjunct population
in extreme southeast Costa Rica near the
Panama border on the Caribbean versant of
Cerro Uatsi, Bratsi, Limo?n Province (Fig. 5).
Specimens are needed from the area between
Fortuna and Santa Fe? to more precisely deter-
mine the limits of the ranges of these two
species.
Natural History.?Myers (1969) described two
female P. caryophyllaceus brooding clutches of 19
and 25 eggs from Cerro Sapo in Darie?n
Province, Panama, that we reassign to P.
educatoris. We observed two females brooding
eggs at El Cope?, one in May 2000 and another in
June 2001. On 18 May 2000, we encountered an
adult female P. educatoris (SVL 38.70 mm)
sleeping on top of an egg clutch with approx-
imately 15?20 eggs. The adult and clutch were
positioned along the midvein on the distal third
of the top surface of a palm leaf approximately
3 cm wide and 98 cm above the ground. We
estimated the eggs to be at developmental
stages 1?5 (Townsend and Stewart 1985) but
were unable to ascertain an exact stage or egg
number without disturbing the attending frog.
Eyes were first noted in embryos 11 days after
discovery (stage 6 for Eleutherodactylus coqui,
Townsend and Stewart, 1985). On 14 June 2000,
28 days after the initial discovery of the
brooding frog, the adult was missing. We
turned the leaf over to count and stage the
eggs, which resulted in the immediate hatching
of the eggs which produced 15?20 froglets. We
estimated developmental period at 24?28 days
TABLE 1. Average and standard deviation of key morphological traits of Pristimantis caryophyllaceous from
Costa Rica and western Panama and Pristimantis educators from central and eastern Panama. Ranges in
parenthesis; all measurements in millimeters.
P. caryophyllaceus P. educatoris
= (N 5 12) R (N 5 9) = (N 5 4) R (N 5 11)
Snout?vent 17.7 6 2.7 (12.0?21.2) 21.5 6 2.8 (16.7?26.2) 20 6 0.5 (19.3?20.4) 28.5 6 4.9 (21.1?37.7)
Tibia length 10.7 6 1.7 (7.2?13.0) 12.5 6 1.5 (10.3?15.1) 12.4 6 0.4 (12.1?12.8) 16.3 6 2.4 (12.0?18.8)
Leg length 18.3 6 3.2 (12.8?22.4) 21.9 6 3.2 (16.8?26.6) 21.4 6 1.8 (18.8?22.7) 27.8 6 3.7 (21.2?32.7)
Head length 7.5 6 1.2 (5.4?10.0) 9.2 6 1.5 (6.9?10.7) 8.4 6 0.4 (8.0?8.9) 12.0 6 1.6 (9.0?13.9)
Head width 6.7 6 0.9 (4.9?7.8) 8.2 6 0.9 (6.6?9.7) 7.3 6 0.4 (7.0?7.7) 10.8 6 1.8 (7.8?12.7)
Eye?snout 3.6 6 0.5 (2.5?4.2) 4.2 6 0.6 (3.0?4.9) 4.2 6 0.2 (4.1?4.4) 5.7 6 0.8 (4.3?6.9)
Eye?nostril 2.7 6 0.5 (1.9?3.5) 3.2 6 0.7 (2.1?4.2) 3.3 6 0.1 (3.2?3.4) 4.6 6 0.7 (3.4?5.6)
Eye length 2.8 6 0.3 (2.2?3.3) 3.3 6 0.3 (2.9?3.9) 3.2 6 0.3 (2.9?3.4) 3.9 6 0.5 (3.1?4.9)
198 M. J. RYAN ET AL.
and estimated SVL of hatchlings at 5.5 mm
using a scaled photograph.
Habitat.?At El Cope?, P. educatoris is fairly
common and found in 20?30-year-old second-
ary forest (A. Jaslow, pers. comm.) with a well-
developed understory of palms and herbaceous
plants. In three three-month field seasons
between 2000 and 2003, we made 123 captures
of P. educatoris. All individuals were found at
night, and 95% of captures occurred along
terrestrial trail transects, and only 5% of
captures occurred along streams. Average perch
height for terrestrial nocturnal captures was
87 cm above the ground, and most individuals
were on leaves.
DISCUSSION
Costa Rica and Panama are home to 195
species of frogs, and 13 of these species (6%)
belong to the genus Pristimantis (formerly
Eleutherodactylus) (Hedges et al., 2008; IUCN,
2009). Of the 427 species of Pristimantis, 13
species occur in Central America including P.
educatoris (IUCN, 2009).
Our analysis of morphological features sup-
ports the hypothesis that P. educatoris popula-
tions in central and eastern Panama are distinct
from P. caryophyllaceus from western Panama
and Costa Rica. This inference is consistent with
previously recognized biogeographic patterns
of differentiation between species in the Cordil-
lera de Talamanca to the northeast and those in
the Cordillera Central more toward the east
(e.g., Myers, 1969; Savage, 1982, 2002; Wang et
al., 2008). The high valley of the R??o Chiriqu??
effectively separates the two mountain chains
and is a natural biogeographic break for many
montane amphibians (Myers and Duellman,
1982). For example, this region is the eastern-
most extent for three species of Craugastor:
Craugastor gulosus (Savage and Myers, 2002),
Craugastor rayo (Savage et al., 2004), and
Craugastor obesus (Campbell and Savage, 2000).
This report brings the total number of lower
Central American (Costa Rica and Panama) frog
species to 196, with 11 (14%) having been
described in the past seven years (Iba?n?ez et
al., 2001; IUCN, 2009). In the last 25 years, the
western and central highlands of Panama alone
have produced seven new species of strabo-
mantines and Craugastor (Frost, 2008), with
many more likely to be described with addi-
tional exploration (Savage, 2002). The use of
molecular techniques, such as DNA bar coding,
is likely to facilitate the identification of cryptic
species and is especially important for the large
number of small brown leaf litter frogs of the
strabomantine and Craugastor lineages that can
be difficult to differentiate by morphology alone
(Ko?hler et al., 2005; Fouquet et al., 2007). The
amphibian fauna of El Cope? experienced dra-
matic declines that resulted in the loss of many
species and a reduction of species abundance of
more than 60% in 2004 (Lips et al., 2006).
Therefore, the discovery and recognition of
undescribed, cryptic species is particularly
important at the time of global extinction of
amphibians (Stuart et al., 2004; Wake and
Vredenburg, 2008).
Acknowledgments.?We would like to thank J.
Robertson, L. Witters, and R. Brenes for field
assistance. Thanks to R. Brenes for the photo-
graph (Fig. 1) and K. Miller and D. Lightfoot for
use of camera equipment for Figure 2. Thanks to
J. Sipiorski for drawing Figures 3 and 4. We
thank R. Nussbaum from UMMZ, J. A. Seigel
from LACM, and K. de Quieroz, J. Poindexter,
and R. Wilson from USNM for providing
museum specimens used for comparison in this
study. We thank ANAM for issuing research and
collecting permits and the Smithsonian Tropical
Research Institute (STRI) Visitor?s Service Office
for assistance in obtaining research and collecting
permits. We thank the personnel of Parque
Nacional G. D. Omar Torrijos H (El Cope?).
Thanks to G. Chaves, T. Kennedy, S. Poe, and
two anonymous reviewers for helpful comments
on this manuscript. Thanks to G. Chaves for
access to recently collected material from Costa
Rica. This work was funded by Sigma Xi, ASIH
Gaige Fund, and IdeaWild (to MJR) and by the
National Science Foundation grants (DEB
0213851, 0234386, 0130273, 9996355) and the Bay
and Paul Foundation (to KRL).
LITERATURE CITED
BARBOUR, T. 1928. New Central American frogs.
Proceedings of the New England Zoological Club
10:25?31.
CAMPBELL, J. A. 1999. Distribution patterns of amphib-
ians in Middle America. In W. E. Duellman (ed.),
Patterns of Distribution of Amphibians: A Global
Perspective, pp. 111?210. Johns Hopkins Universi-
ty Press, Baltimore, MD.
CAMPBELL, J. A., AND J. M. SAVAGE. 2000. Taxonomic
consideration of Middle American frogs of the
Eleutherodactylus rugulosus group (Anura: Leptodac-
tylidae): a reconnaissance of subtle nuances among
frogs. Herpetological Monographs 14:186?292.
FOUQUET, A., A. GILLES, M. VENCES, C. MARTY, M. BLANC,
AND N. J. GEMMELL. 2007. Underestimation of
species richness in Neotropical frogs revealed by
mtDNA analyses. PLoSONE 2:e1109.
FROST, D. R. 2008. Amphibian Species of the World:
An Online Reference. Vers. 5.2. Electronic data-
base accessible at http://research.amnh.org/
herpetology/amphibia/index.php. American Mu-
seum of Natural History, New York (Accessed: 15
July 2008).
NEW SPECIES OF PRISTIMANTIS FROM PANAMA 199
GLOBAL BIODIVERSITY INFORMATION FACILITY. 2008. Elec-
tronic database accessible at: http://www.gbif.net
(Accessed: 22 July 2008).
HEDGES, B. S., W. E. DUELLMAN, AND M. P. HEINICKE.
2008. New World direct-developing frogs (Anura:
Terrarana): molecular phylogeny, classification,
biogeography, and conservation. Zootaxa
1737:1?182.
HEINICKE, M. P., W. E. DUELLMAN, AND S. B. HEDGES.
2007. Major Caribbean and Central American frog
faunas originated by ancient oceanic dispersal.
Proceedings of the National Academy of Science
104:10092?10097.
HERPNET. 2008. Electronic database accessible at:
http://www.herpnet.org/ (Accessed: 22 July 2008).
IBA?N?EZ, D. R., F. A. SOLIS, C. A. JARAMILLO, AND A. S.
RAND. 2001. An overview of the herpetology of
Panama. In J. D. Johnson, R. G. Webb, and O.
Flores-Villela (eds.), Mesoamerican Herpetology:
Systematics, Zoogeography, and Conservation,
pp. 159?170. University of Texas at El Paso, Texas.
IUCN (CONSERVATION INTERNATIONAL, AND NATURE-
SERVE). 2008. Global Amphibian Assessment.
Available: http://www.globalamphibians.org
(Accessed: 15 June 2008).
KO?HLER, J., D. R. VIEITES, R. M. BONETT, F. H. GARCIA, AND
F. GLAW. 2005. New amphibians and global con-
servation: a boost in species discoveries in a
highly endangered vertebrate group. BioScience
55:693.
LEVITON, A. E., R. H. GIBBS JR., E. HEAL, AND C. E.
DAWSON. 1985. Standards in herpetology and
ichthyology. Part I. Standard symbolic codes for
institutional resource collections in herpetology
and ichthyology. Copeia 1985:802?832.
LIPS, K. R., F. BREM, R. BRENES, J. D. REEVE, R. A. ALFORD,
J. VOYLES, C. CAREY, L. LIVO, A. P. PESSIER, AND J. P.
COLLINS. 2006. Emerging infectious disease and the
loss of biodiversity in a Neotropical amphibian
community. Proceedings of the National Academy
of Science 103:3165?3170.
LYNCH, J. D., AND W. E. DUELLMAN. 1997. Frogs of the
genus Eleutherodactylus in western Ecuador. Uni-
versity of Kansas Natural History Museum Special
Publication 23:1?236.
MENDELSON, J. R., III., J. M. SAVAGE, E. GRIFFITH, H. ROSS,
B. KUBICKI, AND R. GAGLIARDO. 2008. Spectacular
new gliding species of Ecnomiohyla (Anura: Hyli-
dae) from central Panama. Journal of Herpetology
42:750?759.
MYERS, C. W. 1969. The ecological geography of cloud
forest in Panama. American Museum Novitates
2396:1?52.
MYERS, C. W., AND W. E. DUELLMAN. 1982. A new species
of Hyla from Cerro Colorado, and other tree frog
records and geographical notes from western
Panama. American Museum Novitates 2752:1?32.
SAVAGE, J. M. 1982. The enigma of the Central
American herpetofauna: dispersal or vicariance?
Annals of the Missouri Botanical Garden
69:464?547.
???. 1987. Systematics and distribution of the
Mexican rainfrogs of the Eleutherodactylus gollmeri
group (Amphibia: Leptodactylidae). Fieldiana Zo-
ology 33:1?57.
???. 2002. The Amphibians and Reptiles of Costa
Rica. University of Chicago Press, Chicago.
SAVAGE, J. M., AND C. W. MYERS. 2002. Frogs of the
Eleutherodactylus biporcatus group (Leptodactyli-
dae) of Central America and northern South
America, including rediscovered, resurrected,
and new taxa. American Museum Novitates
3357:1?48.
SAVAGE, J. M., B. D. HOLLINGSWORTH, K. R. LIPS, AND A.
P. JASLOW. 2004. A new species of rainfrog (genus
Eleutherodactylus) from the Serran??a de Tabasara?,
west?central Panama, and reanalysis of the fitzin-
geri species group. Herpetologica 60:519?529.
STUART, S. N., J. S. CHANSON, N. A. COX, B. E. YOUNG, A.
S. L. RODRIGUES, D. L. FISCHMAN, AND R. W. WALLER.
2004. Status and trends of amphibian declines and
extinctions worldwide. Science 306:1783?1786.
TOWNSEND, D. S., AND M. M. STEWART. 1985. Direct
development in Eleutherodactylus coqui (Anura:
Leptodactylidae): a staging table. Copeia
1985:423?436.
WAKE, D. B., AND V. T. VREDENBURG. 2008. Are we in the
midst of the sixth mass extinction? A view from
the world of amphibians. Proceedings of the
National Academy of Sciences 105:11466?11473.
WANG, I. J., A. J. CRAWFORD, AND E. BERMINGHAM. 2008.
Phylogeography of the Pygmy Rain Frog (Pristi-
mantis ridens) across lowland wet forests of
isthmian Central America. Molecular Phyloge-
netics and Evolution 47:992?1004.
Accepted: 18 June 2009.
APPENDIX 1
Material Examined
Pristimantis (formerly Eleutherodactylus) caryophylla-
ceus.?Costa Rica: Provincia de San Jose?: Bajo la
Hondura: LACM 154009 (female); LACM 154034,
LACM 154053, LACM 154071, LACM 154075 (males);
Provincia de Heredia: La Selva: LACM 154006, LACM
154081 (females), LACM 154007, LACM 154011
(males); Provincia de Cartago: Tapant??: LACM
154062, LACM 154090, LACM 154092 (females),
LACM 154033, LACM 154056 (males); Panama:
Provincia de Chiriqu??: Fortuna: LACM 154000?001,
LACM 153997 (males); Provincia de Chiriqu??: Los
Planes: LACM 153993, LACM 153998 (females),
LACM 153999 (males).
Pristimantis educatoris.?Panama: Provincia de Ver-
aguas: Santa Fe?, Cerro Arizona UMMZ 167574,
UMMZ 155686 (females); Provincia de Cocle?, Parque
Nacional General de Divisio?n Omar Torrijos Herrera,
approximately 8 km north of El Cope? USNM 572238,
USNM 572336, USNM 572831, SIUC H-07323, UMMZ
167571, UMMZ 167629, UMMZ 168290 (females);
UMMZ 152951?953, UMMZ 155682 (males).
200 M. J. RYAN ET AL.