Two new species of marsupial frogs (Anura: Hylidae: Gastrotheca) from the Cordillera Oriental in central Peru

Scientific Papers

Natural History Museum
The University of Kansas

17 June 2004 Number 32:1-10

Two New Species of Marsupial Frogs (Anura: Hylidae: Gastrotheca)
from the Cordillera Oriental in Central Peru

By

William E. Duellman,' Edgar Lehr,- Daniel Rodriguez/ and Rudolf von May"*

^Museum ofNatiiral History and Biodivciyity Rcscaicli Cciitci; The Llimvrsitjf of Kiiiisas, Lniorciicc, Kansas 66945, LISA

'Staatliclic Naturhistorischc Saiiniiliin^cn Dresden, h4iiscuiii fur Ticrkiindc, D-01i09 Dresden, Germain/

^Departaiiiciito de Herpetolo^ia, Museo de Historin Natural de Snn Mareos,
Apartado 14-0434, Lima, Peru

^Asociacibii vara la Conservaeibn de la Cueiu'a Aumzbniea (ACCA), Calle Cuzco 499,

' r

Puerto Maldoimdo, Madre de Dios, Peru L_ '

CONTENTS

ABSTRACT 1 ^,, ..rOs/ARD

RESUMEN 2y^iVEf^Sn 1

INTRODUCTION 2

Acknowledgments 2

DESCRIPTIONS OF NEW SPECIES 2

Gastrotheca AT\Mr.ANA new srEciEs 2

Gastrotheca zeugocystis new species 6

DISCUSSION 8

LITERATURE CITED 9

APPENDIX: Specimens Examined 9

Abstract A distinctive new species of Gastrotheea from the Rio Chanchama\'o \'alle\' in the Cordillera
Oriental in Peru is the only known member of the genus that lacks a tympanum. Another new species
from the Cordillera de Carpish is distinctive in having a pair of lateral brood pouches and only basal
webbing on the foot.

Kci/ words: Systematics; Anura; Hvlidae; Gastrotheea atympana. new species; Gastrotheca zeu;^ocystis;
Peru.

© Natural History Musoum, The University of Kansas EtTl^it fl'CVT I l''T?-' '^^"^ '^"' ^f'^'*"""*^-

Museum of Compsra.v/c Zoclosy
Harvard University

O^'

Scientific Papers

Natural History Museum
The University of Kansas

17 June 2004 Number 32:1-10

Two New Species of Marsupial Frogs (Anura: Hylidae: Gastrotheca)
from the Cordillera Oriental in Central Peru

By

William E. Duellman,' Edgar Llhr,- Daniel Rodriguez,'' and Rudolf von May"*

'Mitsciini of Niitiiral Hilton/ niui Biodiversity Rcsciiich Center, The Lliiiversiti/ of KiDisiis, Lnivreiiee, Kansas 66945, USA

-Staatliehe Nattirhistorisehe Saiiniilim^eii Dresden, MuseiDii far Tierkunde, D-iU109 Dresden, Germain/

^Departivnento de Herpetoto'.iia, Mnseo de Historia Nntnral de San Mareos,
Apartado 14-1)434, Lima, Peru

^Asoeiaeibn para la Conservaeibii de la Cueiiea Amazoniea (ACCAK Calle Cnzeo 499, "

Puerto Maldonado, Madre de Dios, Pern ^^•^^■- ■

^ 0 im

CONTENTS •'

ABSTRACT 1 ^p^RVAgD

RESUMEN 2y^lVEBSli ^

INTRODUCTION 2

Acknowledgments 2

DESCRIPTIONS OF NEW SPECIES 2

G.ASTROTHECA ,4r>MmN,4 NEW SrECIES 2

Gastrotheca zeugocystis new species 6

DISCUSSION 8

LITERATURE CITED 9

APPENDIX: Si'ecimens Examined 9

Absi KACT A distinctive new species of Gastrotheea from tlie Rio Chanchamavo Vailev in the Cordillera
Oriental in Peru is the only known member of the genus that lacks a tympanum. Another new species
from the Cordillera de Carpish is distincti\'e in having a pair of lateral brood pouches and onh' basal
webbing on the foot.

Key ipords: Svstematics; Anura; Hylidae; Gastrotheea atyinpana, new species; Gastrotlieea zen^oeystis;
Peru.

© N.itur.il Hislorv Museum, The L'ni\LTsit\' ot k.uiNcis FfP^'^ !*.'^\"f I !■'■"■,-?,'-'./ '^^'^ ^"- ' f'y-l-O''*^^

Museiim of Compara.v/e Zoclosy
Harvard University

SciHNTiiic Papi.rs, Natikai, Hi.sh)KV Ml slum. Till. Univi:rsity of- Kansas

Resumen Se describe una especie niieva de Cii>tivlliccii del valle del Rio Chancliamavi) en In Cordil-
lera Oriental del Peru; esta especie se distingue de tt)dos los otros taxa del genero en no poseer timpano.
Otra especie nue\ a de la Cordillera de Carpish se caracteriza pttr tener bolsas de incubacion pareadas
V membranas basales entre los dedos del pie.

I'alabras claves: Sistematica; Anura; Hylidae; Ciititivtlh\ii iiti/nijunia, especie nueva; Caslrothccn
zcugoci/stis, especie nueva; Peru.

INTRODUCTION

Herpetokigical exploration of the Cordillera Oriental by Lehr and his Peruvian colleagues, there are additional

in central Peru through a cooperative agreement between undescribed species in the genus; two of these are de-

the Museo de Historia Natural Universidad Mayor de San scribed herein.
Marcos and the Museum ftir Tierkunde Dresden has re-
vealed the existence of a rich montane herpetofauna con-
taining many previously unknown species. For more than Lehr thanks R. Acero for collecting and export per-

three decade.s, one of us (WED) has studied the genus mits issued by the Ministerio de Agricultura (INRENA),

Gastwtheca, a genus now containing 47 species. Among the Peru. We thank Juan Manuel Guayasamin for the resumen;

newspeciesofanurans from the eastern frt)nt of the Andes Joseph R. Mendelson, Linda Trueb, and Eric R. Wild for

in central Peru was a large species of Gastwtheca described their critical ctMiiments on tlu- manuscript; and Linda Trueb

bv Duellman et al. (2001 ). In more recent collections made for aid with the illustrations.

Ack,njowledc;mknts

MATERIALS AND METHODS

The 16 morphological measurements, 25 external de-
scripti\'e characters, and numbered diagnostic characters
are tiiose used by Duellman and Pyles (1980), Duellman
.)nd Hillis (1987), Duellman and Trueb (1988), and
Duellman et al. (2U0I); the latter also incorporated the 13
numbered diagnostic traits. All measurements are in mil-
limeters; snout-vent length is abbreviated SVL. Sex was
determineci by examination of gonads. Webbing formulae

were determined by the method proposed by Savage and
Heyer (1967), as modified by Myers and Duellman (1982).
The Museo de Historia Natural Universidad Mayor de San
Marcos, Lima, Peru, is noted as MHNSM, the Museum fur
Tierkunde Dresden, as MTD, and the University of Michi-
gan Museum of Zoology, as UMMZ. Most comparisons
were made with specimens of Gastrothcca in the Museum
of Natin-al Historv, The Universitv of Kansas (KU).

DESCRIPTIONS OF NEW SPECIES

Gastwtlicca atympaua new species

Holotype.— MHNSM 18692, an adult male, from
Pampa 1 lermosa, (10 59'33.3" S, 73"25'58.0" W), 1540 m,
Pro\incia de Tarma, Departamento de Junin, Peru, ob-
tained 28 August 2003 by Rudolf von May.

Diagnosis. — A moderate-sized species (to 46.7 mm)
having: ( I ) tibia length 51.6% SVL, much longer than foot;
(2) interorbital distance more than twice eyelid width; (3)
skin on dorsum shagreen, not co-ossified with skull, lack-
ing transverse ridges; (4) supraciliary processes absent; (5)
heel with transverse row of small tubercles; (6) tympanic
annulus absent; (7) Finger 1 slightlv shorter than Finger 11,
with discs much wider than digits; (8) fingers unwebbed;
(9) webbing extending to antepenultimale subarticuLir
tubercle on Toe IV, to penultimate subarticular tubercle on
Toe V; (10) dorsum tan with irregular trans\erse brown
markings; (11) head markings consisting of broad, dark
brown interocular bar and pale labial and canthal-postor-
bital stripes; (1 2) pale dorsolateral stripe absent; (13) flanks
cream with irregular, vertical bLu k marks; (14) \enter pale
tan; (15) position of brooel pouch unknowri.

Gastrothcca at\/iupaua differs from the other 46 mem-
bers of the genus by lacking an external tympanum. Seven
other species of Gastwtheca have a tibia length 50-55% of
SVL and Finger 1 shorter than Finger 11. Of these, G. automia
differs by having supraciliary processes, two tubercles on
the heel, and a venter that is orange or red anteriorly and
black posteriorly. Gastwtheca owphi/lax differs by being
uniformly green dorsally and ventrally, except for a cream
labial stripe. Gastwtheca argeiiteoz'iiviis and G. diiuiii differ
bv ha\'ing pale dorsolateral stripes, and G. psi/chwplilhi
ditfers by having a unitorm brow n to dull green ornearly
black dorsum and basal webbing l^etween the fingers.
Gastwtheca ochoai and G. rebeccae cliffer by having a nar-
rower interorbital regions (lOD about I.5x eyelid width);
the lormer is smaller (to 38 mm SVL) and lacks an
interocular bar, whereas the latter is about the same size
as (;. ti/nijiaiia, but lhi> markings on the head consist of a
dark canthal stripe in addition to the dark interocular bar
and pale labial stripe. Two other larger species have verti-
cal black bars on the Hanks. Of these, G. ovifera differs by
haxing tin' skin co-ossitied with the skull and a prominent

N[-:w Makslipiai. Fkocis irom Pf.ru

transverse occipital ridge; it also differs by having the
webbing extending to the distal subarticular tubercle on
Toe V, as opposed extending only to the penultimate tu-
bercle in G. ati/iiipnnn. In Gn^tivtliccn tcstudiiicn. Finger I is
longer than Finger 11, and the webbing extends to the dis-
tal subarticular tubercle on Toe V.

Description of holotype. — An adult male; body mod-
erately slender; head wider than long, about as wide as
body; snout narrowly triuicate in dorsal \'iew, barely
rounded above and slightly inclined anteroventrally in
profile; canthus rostralis angular, ciuved; loreal region
barely concave; lips not flared; top of head flat; lOD 240' ,'.
EW; nostrils slightly protuberant, directed laterally just
below anterior terminus of canthus rostralis, at level just
posterior to anterior margin of lower jaw; diameter of eye
slightly greater than distance from anterior corner of orbit
to nostril; tympanum not differentiated; t\mpanic annu-

lus and columella absent; supratympanic fold elevated,
angular, detlecteci laterally, continuous with distinct, nearly
vertical fold extending ventrally to point just above inser-
tion of forelimb (Fig. I ).

Arm moderately robust; ulnar tubercles small, barely
evident; hand large; fingers long, bearing large round discs;
width of disc on Finger III nearly twice width of
penultimate segment of finger; relative lengths of fingers
1 < 2 < 4 < 3; fingers unwebbed, with narrow lateral fringes
on Fingers II-IV; subarticular tubercles moderately large,
round, none bifid; supernumerary tubercles subconical,
nearly as large as subarticular tubercles; palmar tubercle
indistinct; prepollical tubercle elongate, elliptical; nuptial
excrescences not evident (Fig. 2). Hind limb slender; tibia
length 51.6% of SVL; foot length 40.7% of SVL; heel bear-
ing transverse row of five or six small tubercles; tarsal tu-
bercles absent; inner tarsal fold low, weak on distal three-
fourths of tarsus; low, crenulated outer tarsal fold extend-
ing from heel nearly to disc on Toe V (evident in living
individual [Fig. 3] but not visible proximally and barely
visible distally in preserved specimen); outer metatarsal
tubercle absent; inner metatarsal tubercle small, elliptical,
\isible from above; toes long, bearing round terminal discs
about same size as those on fingers; relative length of toes

^ lo'O //

Fig. ]. Dorsal and lateral views of the hrad ot tlu' hcilotxpt' Fig, 2. Palmar surface ef right hand and plantar surface of

of Giiiitrothcai atyfupana. MHNSM 18692. left foot ot holotype of GiistivHiccn atyiiipnim, MHNSM 18692.

Scientific Papers, Nmural Histukv Museum, The Univeksi n ni K \\s \s

Fig. 3. A nnd B. Dors.il .ind LUimmI \'ie\vs of holotvpe oi Gaslivllimi iiti/nipiuui, MHNSM 18642, ndiilt male, 46.7 mm SVL. C
nnd D. Dorsal and lateral \'it"vvs of holohjn' ot Cdstrntlicca zcii^oci/f^tis, MHNSM 18675, adult temak', ^7.^ mm SVL. Photographs
by Edgar Lt'hr.

1<2<3 = 5<4; toes less thdii one-halt webbed, with
narrow lateral fringes distally; webbing formula I2-2II2-
3 ni2-.3IV2 -1 V; subarticular tubercles large, round; su-
pernumerarv tubercles small, diffuse (Fig. 2).

Skin on dorsum finely .shagreen, that on head not co-
ossified with underlying cranial elements; transverse der-
mal ridges absent; supraciiiary processes absent; man\
small, subconical tubercles in temporal region; skin on
flanks areolate; skin on throat, chest, belly, and
posteroventral surfaces of thighs coarsely granular; othei-
ventral surfaces smooth; transverse row of five subconical
tubercles below cloacal opening. Shape of opening of brood
pouch unknown; vocal slits and vocal sac absent. Vomer-
ine odontophores elevated, posteromedially inclined, nar-
rowly separated medially, between posti-rior margins ol
ovoid choanae, each bearing six teeth.

Color ill prei^ervatiiv: Dorsum of head antf body gray-
ish tan with dull brown markings consisting of broad in-

terorbital bar connectecf to a triangular maik in scapular
region and irregular marks posteriorly on body; dorsal
surfaces of limbs pale gray with narrow transverse bars,
three on each forearm, thigh, and foot, six or seven on
shanks; those on thighs continuous onto anterior surfaces;
flanks pale brown anteriorly, pale grayish white posteri-
orly; interstices in areolate skin on flank black; flanks with
lour (right side) or five (left side) irregular vertical black
marks. Side of head pale grayish tan; narrow, creamy white
stripe from point just behind nostril, aUmg canthal crest,
edge of upper eyelid, and edge of supratympanic fold,
biirdered below by narrower brown stripe in loreal and
tempoial regions; margin of upper lip creamy white. Pos-
terior surfaces of thighs dark brown with triangular ex-
tensions dorsally meeting transverse bars on dorsal sur-
laces; tubercles on ventrolateral edges of forearms, heel,
below cloacal opening, anci fold on outer edge of foot wliite.
rhroat cream; chest, belh', and palmar surfaces pale brown;

Niw M\ksri>i\i Frogs ikom Pi-;ru

▲

G.

alympana

•

G.

stictopleura

■

G.

testudinea

T

G.

zeugocystis

+

G.

sp. (Yanachaga)

m

G.

griswoldi

Q

G.

marsupiata

B

G.

peruana

\

Cusco

J Madre
de Dios

jlometers

Fig. 4. Map ot clmiIimI IViu showing loctilitius oi t)ccurrence of five species of Gastivtliccii on the Andean slopes (symbols) and
generaUzed distribution patterns of three species of Gnstrothecn that occur at high elevations in the Andes (hash marks).

ventral surfaces of limbs and plantar surfaces pale gray-
ish brown.

Color ill life: Dorsum anci flanks pale gravisii tan; dor-
sal markings orange-brown with narrow dark brown
edges; labial and canthal-postorbital strip)es pale beige,
narrowly bordered below by brown; fringe on foot, ulnar
and subcloacal tubercles white; dorsal surfaces of discs and
distal parts of cligifs orange-brown; posterior flanks pale
gray to white with black markings; throat pale beige
with chocolate brown blotches; belly chocolate brown;
palmar and plantar surfaces dark gray; iris pale yel-
lowish tan above, pale gray below, with fine black re-
ticulations (Fig. 3).

Mcasinviiiciits (in iiiiii)ofliolot\/pc: SVL46.7, tibia length
24.1, foot length 19.0, head width 17.2, head length 15.1,

internarial distance, 2.6, interorbital distance 7.2, eyelid
width 3.0, eye-nostril distance 3.9, eye length 4.5, nostril-
jaw distance 2.6, orbit-jaw distance 2.4, fliird finger length
13.5, tiumib length 6.5, width of disc on third finger 1.9.

Etymology. — Tlie specific name is derived from the
Greek (7- meaning without and the Greek tyinpaniou mean-
ing ear drum; the name is used as an adjective describing
the absence of a tympanum.

Distribution and ecology. — The species is known only
from the typie locality at an eleyation of 1540 m in the val-
ley of the Rio Chanchamayo, a broad intrusion into the
Cordillera Oriental in central Peru (Fig. 4). The type local-
ity is a locally protected dense humid montane forest. The
holotype was on a branch of a tree 1 .2 above the ground at
2100 ii.

Scientific Papers, Natikai. IIisiokv Mishum, The Univi.rsh^ of Kansas

Gasiroiheca zeugocystis new species

Holotype.— MHNSM 18675, an adult female, from the
Cordillera de Carpish (09°43'58.2" S, 76 06'41.9" W), 2920
m, I'rovincia de Huanuco, Dcpartamento de Huanuco,
Peru, obtained on 24 July 2003 by Daniel Rodriguez.

Referred specimen. — MTD 45984, cleared-and-stained
male, 28.2 mm SVL, collected witli the holotype.

Diagnosis. — A small species (to 37.5 mm in females)
having ( 1 ) tibia length 46.9% SVL, as long as foot; (2) inter-
orbital distance less than twice eyelid width; (3) skin on
dorsum smooth, not co-ossified with skull, lacking trans-
verse ridges; (4) supraciliary processes absent; (5) heel lack-
ing tubercles or calcar; (6) tympanic annulus smooth; (7)
Finger 1 equal in length to Finger 11, with discs barely wider
than digits; (8) fingers unwebbed; (9) webbing absent on
toot except for basal webbing between Toes 111 and IV; (10)
dorsum brown with small, irregular black marks; (11) head
markings absent; (12) pale dorsolateral stripe absent; (13)
tlanks graxish brown; (14) venter brown; (15) brood pouch
paired, lateral.

In most species of Gnftwthecd, the brood pouch is
single, median, and dorsal. A paired of lateral pouches,
one on each side of the abdomen, is known in only three
species — G. walkeri, willianisoni, and zeiigoci/stib. Gastrothcai
xvalkcri and G. williiviisoiii, species living in northern Ven-
ezuela, differ from G. zeugocystis in many features — larger
size (70 and 54 mm SVL, respectively), tibia length more
than 50% SVL, presence of a triangular calcar on the heel
and granular interocular ridge, and fingers one-fourth (G.
ivnikcri) or one-half webbed (G. zvilliaiusoiii). Three other
species (G. abdita, litoiicdi^, and pcruniia) are like G.
zeugocystis in having the first and second fingers equal in
length and in having a tibia length less than 50% of SVL.
All three species have single dorsal brood pouches and
much more webbing on the foot, reaching the penultimate
subarticuiar tubercle of Toe V. The tympanic annulus is
granular in G. abdila and G. picruiiiui, whereas tlie snout is
rounded in dorsal view in G. litoiiedi>^ and G. pcrnaua. Vur-
thermore, G. abdita differs by having a small tubercle of
the heel; G. litoiwdi^ differs by having pale labial and dor-
solateral stripes, and G. peruana differs by having a pustu-
lar dorsum, dark canthal stripe, and pale labial stripe. Su-
perficially, G. zeugocystis and G. ochoai are similar; they re-
semble one another in SVL, absence of head markings, and
general body pattern, but the ground color in G. oclioai is
tan instead of brown. Finger 1 is shorter tiian Finger 11,
and the webbing on the foot is more e\tensi\'e (lo
penultimate subarticuiar tubercle on Toe V); turlhermore,
G. oclioai has a single, median, dt)rsal brood pouch.

Description of holotype. — Body nioderateK slender;
head about as wide as long, about as wide as body; snout
acuminate in dorsal view, rounded in profile; cantiius

5 mm

Fig. 3. Dorsal and Literal \ie\\s of the head nf the holotype
of Giibtivtiiccii zciigucystis, MHNSM 18675.

rostralis rounded, straight; loreal region shallowly concave;
lips rounded; top of head flat; interorbital distance 168%
of e\elid width; nostrils ni>t protuberant, directed later-
alK' just below anterior terminus of canthus rostralis, at
le\el of anterior margin of lower jaw; diameter of e\e much
greater than distance from anterior corner of orbit to nos-
tril; tympanum nearh- round, well differentiated, separated
from e\ i' hv distance about one-half diameter of e\e; tym-
panic annulus smooth; suprat\mpanic told rounded, cov-
ering upper edge ot txinpanum, ciu'ved postero\entrally
behind t\'mpanum to point abo\e insertion of forelimb
(Fig. 5).'

Arm robust; ulnar tubercles absent; iiand large; fin-
gers moderateh' long, bearing round discs; width of disc
on Finger III baieh wider than piMUiltiniate st'gment ot
linger; relative lengths ot lingers 1 - 2 < 4 < 3; tingers
uinvebbed, without lateral fringes; .subarticuiar tubercles

Niiw Marsiipial Frogs from Peru

Fig. 6. Polniar Miitace ot right hand and plantar surtace ot
right foot of the holotypo of Grt>trothecii zfugocysti^, MHNSM
18675.

moderately large, round, none bifid; supernumerary tu-
bercle absent; palmar tubercle absent; prepollical tubercle
massive, elliptical (Fig. b). Hind limb mocierately slender;
tibia length 46.9"n of SVL; foot length 46.1% of SVL; heels
lacking calcar or tubercles; tarsal tubercles, inner and outer
tarsal folds, outer metatarsal tubercle absent; inner meta-
tarsal tubercle small, elliptical, not visible from above; toes
moderateh' long, bearing round terminal discs smaller than
those on fingers; relative lengths of toes 1<2<3<5<4;
toes unwebbed except for basal webbing between Toes III
and IV; toes lacking lateral fringes; subarticular tubercles
diffuse, indistinct; supernumerary tubercles minute, few
on proximal segments of digits (Fig. 6).

Skin on dorsum smooth, that on head not co-ossified
with underlying cranial elements; transverse dermal ridge
absent; supraciliary processes absent; supratympanic fold
tubercular; longitudinal row of tubercles below tympanum;
skin on flanks smooth; skin of throat, chest, belh', and
posterovenfral siu'faces of thighs coarsely granular; other
surfaces smooth; pair of low, round tubercles below cloa-
cal opening. Brood pouches paired; external margin
elongateh' narrow, U-shaped; opening to pouches under
lateral dermal folds, extending broadly ventrolaterally to
pouch between body wall and skin. Tongue broadly cordi-

form, shallowlv notched behind, free posteriorly for about
one third of its length; vomerine odontophores low, trans-
verse at level slightly behind posterior margins of small,
round choanae, abutting medially, each bearing four teeth.

Color ill pivscmiUvc: Dorsum of head and body brown
with small, irregular black marks; dorsal surfaces of limbs
brown with indistinct narrow, transverse dark brown
bars — two on forearm, three on thigh, three on shank; ir-
regular dark brown marks on foot; flanks grayish brown
with few scattered dark brown marks. Side of head dark
brown; tympanum tan; canthal, labial, and dorsolateral
stripes absent. Anterior surfaces of thighs brown; poste-
rior surfaces of thighs dull yellow; tubercles below cloacal
opening cream. Thrciat, chest, and belly brown; ventral
surfaces of limbs and palmar and plantar surfaces tan; ax-
illa and adjacent venter yellowish orange.

Color ill life: Dorsum reddish brown with dark brown
markings; groin and anterior surfaces of thighs lavender;
posterior surfaces of thighs orange; dorsal surfaces of digits
bluish gray; dorsal surfaces of discs on digits orange (Fig. 3);
venter brown; iris reddish bronze with black reticulations.

Mensurciiients (in nun) ofholotypc: SVL 37.5, tibia length
17.6, foot length 17.3, head width 11.8, head length 11.7,
internarial distance 2.3, interorbital distance 4.2, eyelid
width 2.5, eye-nostril distance 2.8, eye length 3.7, tympa-
num length 2.4, nostril-jaw distance 20, orbit-jaw distance
1.3, third finger lengtii 11.0, thiunb length 6.1, width of
disc on third finger 1.8.

Etymology. — The specific name is derived from die Greek
zci{'^ofi, meaning pair, and the Grcvk ki/stis, meaning sac; tlie name
refers to the paired brood pouches in this species.

Distribution and ecology. — Gastwtlicca zeugocystis is
known only from the type locality in the Cordillera de
Carpish, a north-south outlier of the Cordillera Oriental
between Huanuco and Tingo Maria in central Peru (Fig.
4). The type locality is reached by a narrow "road" that
ascends the mountain immediately to the north of the
Carpish Tunnel on the Huanuco-Tingo Maria road. The
frogs were found cluring the day under leaves and under
a piece of wood in cloud forest. The forest on the upper
slopes of the Cordillera de Carpish is made up of low trees
with moss-covered branches. The two adults were main-
tained for a short while in a terrarium, where they actively
climbed about on the vegetation at night and remained
quiet in bromeliads or under leaves and moss by day.

Remarks. — Although investigations on anurans in the
Cordillera de Carpish are in their infancy, an interesting
array of anurans has been discovered there, including new
species of Pliri/iiopiis (Lehr et al., 2002) and of several taxa
yet to be described. High degrees of species diversity have
been reported for birds (Parker and O'Neill, 1976) and
mammals (Pacheco, 2002) in the Cordillera de Carpish.

8

SciENTii'ic Papers, Nau ral Hisiorv IVIii.si;iiM, Thi-; UNivKRsin oi Kansas

DISCUSSION

Oneot us (WED) hcis examined more than 41)00 speci- Three species (Gastivtiicca luarsiipiata, pcrnann, and

mens of Gastwtlieca, of which the holotype of G. af\/mpivia stictopleura) that occur at high elevations (> 2300 m) in the

is unique in having no external tympanum; thus, the nam- central and southern parts of the Cordillera Oriental of the

ingof a new species based on a one specimen is warranted. Peruvian Andes produce many (63-112) relatively small

Moreover, the color pattern and relatively large and promi- eggs that hatch as tadpoles and complete their develop-

nent supernumerary tubercles on the fingers also are ment in ponds, drainage ditches, and other lentic waters

unique in the genus. The frog does not fit readily into the (Duellman and Fritts, 1972; Duellman et al., 2001). On the

phenetic groups recognized by Duellman et al. (1988). other hand, species of Gastwtlieca that inhabit montane

However, tissues from the holotype will be used in a forth- forests in central and southern Peru, usually at elevations

coming analysis of mitochondrial and nuclear genes that of less than 3000 m produce far fewer (14-22) eggs that

will help to elucidate the phylogenetic relationships. undergo direct development and hatch as froglets. Based

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Fig. 7. Elevatioiial eiistributions of species of Gn^-trotlhra that omn iii I't'iu. I>l.u k bars are spi'eies ih.il piMduee eggs that
undergo direct development; grav bars are species that produce eggs that hatch as Irei'-sw Imniing tadpoles. The lower part
of the range of G. nwnticoln is in the i luancabamba Depression.

Nfw Marsupial Frogs from Peru

solely on elevation, we presume that G. atynipaun has di-
rect development (Fig. 7). Three other species of Gnstivtlura
are known from montane forests in the central part of tlie
Cordillera Oriental (departamentos of Ayacucho, Huanuco,
and junin). These include the larger G. tcstiidinca, which
occurs at elevations of 1560-2590 m and has direct devel-
opment, and G. zciigocystis, which is known only from an
elevation of 2920 m and presumably also has direct devel-
opment. The low number (6) and large size (3.5 mm) of
ovarian eggs in G. zcugoa/stis suggest that direct develop-
ment occurs in this species.

The third species, Gnstwtheca stictopleurn, which has a
large number of small eggs that presumably hatch as tad-
poles (Duellman et al., 2001), has been found at an eleva-
tion of 2625 m on the Cordillera de Carpish. CiKtrotlwca
zcugocystifi is known from 2920 m in the same mountain
range; thus, the two species may occur sympatrically.
Gaftwtlicca stictoplciirn formerly was known only from the
type locality in Departamento de Huanuco (Duellman et
al., 2001), but we now report it from elevations of 2500-
3090 m over a distance of about 200 km on the eastern face
of the Cordillera Oriental and associated ranges (Fig. 4).

LITERATURE CITED

Duellnicin, VV. E., and T. H. Fritts. 1972. A taxonomic review of
the southern Andean marsupial frogs (Hylidae Ga^tivtltcca).
Occasional Papers Museum of Natural History University
of Kansas 9:1-37.

Duellman, W. E., and D. M. Hillis. 1487. Marsupial frogs (Anura:
Hylidae: Gastivtiwca) of the Ecuadorian Andes: resolution
of taxonomic problems and phylogenetic relationships.
Herpetologica 43:141-173.

Duellman, W. E., E. Lehr, and C. Aguilar. 2001. A new species of
marsupial frog (Anura: Hylidae: Gnstiothcca) from the Cor-
dillera Azul in Peru. Scientific Papers Natural History Mu-
seum University of Kansas 22:1-10.

Duellman, W. E., L. R. Maxson, and C. A. Jesiolowski. 1988. Evo-
lution of marsupial frogs (Hylidae: Hemiphractinae): immu-
nological evidence. Copeia 1988:527-543.

Duellman, VV. E., and R. A. Pyles. 1980. A new marsupial frog
(Hylidae: Gastrotlieca) from the Andes of Ecuador Occasional
Papers Museum of Natural History University of Kansas
84:1-13.

Duellman, W. E., and L. Trueb. 1988. Cryptic species of h\lid
marsupial frogs in Peru. Journal of Herpetology 22:159-179.

Lehr E., D. Rodriguez, and Jesiis H. Cordova. 2002. A new spe-
cies of Phn/iiopus (Amphibia, Anura, Leptodactylidae) from
the Cordillera de Carpish (Departamento de Huanuco, Peru).
Zoologische Abhandlungen Dresden 52:65-70.

Myers, C. W., and W. E. Duellman. 1982. A new species of H\/la
from Cerro Colorado, and other tree frog records and geo-
graphical notes from western Panama. American Museum
Novitates 2752:1-32.

Pacheco, V. 2002. Proteccion de la biodiversidadd en bosques
montanos fragmentados y propuesta para bosque de
Carpish, Huanuco. Report I, II, CONCYTEC, Lima.

Parker, T. A., and J R O'Neil. 1976. Introduction to bird-finding
in Peru: Part II. The Carpish Pass region of the eastern Andes
along the central highway. Birding 20 (3):205-216.

Savage, J. M., and W. R. Hever 1967. Variation and distribution
in the tree-frog genus Plu/llojiicdiisd in Costa Rica, Central
America. Beitrage zur Neotropische Fauna 5:111-131.

APPENDIX

Specimens Examined

Gnstrothccn nhdita: PERU: AiiinzLvnif: Cordillera Colan, 2970-
3330 m, E La Peca, KU 196833-35.

GnstivHiccn antomur. COLOMBIA: Ghoc'o: San Jose del Palmar,
KU 289245. Vallc: Paso de Galapagos, KU 289244.

Gnffrotlicai ni'^ciitcovircuf: COLOMBIA: Cniicn: Coconuco,
2800 m, KU 145062, 145066-69; 2 km E Silvia, 2550, KU 181168-
73; 6 km E. Silvia, 2690 m, KU 181174-84. Vallc del Cmica: 7 km
NE Tenerife, 2850 m, KU 169403-12.

Gastrothccn atympnnn: PERU: ]unni: Pampa Hermosa, 2540
m, MHNSM 18692 (holotype).

Gnsfwthccn dmini: COLOMBIA: Autwqiiur. San Pedro, KU
204114-5.

Gastwthccn Ittoucdis: ECUADOR: Azuay: Cuenca, 2600 m, KU
120709-10, 120712, 120718-22, 203442; 2.1 km S Cutchil, 2720 m,
KU 141572; 3.5 km S Cutchil, 2785 m, KU 141579; 10 km NW
Ciron, 2750 m, KU 202690; Laguna de Zurucuchu, 16 km NW
Cuenca, 3200 m, KU 203441; Rio Matadero, 12 km E Cuenca, KU
129779-96; Rio Mazan, near Casablanca, 3150 m, KU 207521.
Canar: Biblian, 2620 m, KU 141571, 141573, 142620-24, 147113.

Gastrothcca ochoai: PERU: Chsco; Chilcn, 10 km N
Ollantaytamho. 2760 mKU 138628-65, 139202-09, 148572, 173499-
500 (C&S); Rio Marcapata, below Marcapata, 2745 m, KU 196951-
52. Pinw: Ollachea, 53 km N Macusani, 2800 m, KU 138666-718.

Gaftmthcca owphylax: ECUADOR: Cnrchi: El Carmelo, 2710
m, KU 178569; 5.7 km NW El Carmelo, 2910 m, KU 178570-84,

209578, 209579 (C&S), 209580; Santa Barbara, 2650 m, KU 190022-
29; 1 km E Santa Barbara, 2650 m, KU 202693-94. Napw: 3 km E.
Papallacta, 2900 m, KU 117981; 11 km ESE Papallacta, 2660 m,
KU 164243-44, 178568; 12 km ESE Papallacta, 2630 m, KU 155469-
70, 164242.

Gdi^tivthcca ovifcra: VENEZUELA: ,4r(7y;((j; Paso Portochuelo,
1170 m, KU 185733; Rancho Grande, 1100 m, KU 133388-89,
166760-64, 185732, 185785.

Giiftivthccii peruana: PERU: Ancafli: Chavin de Huantar, 3230
m, KU 138514-26; Chiquian, 3200-3400 m, KU 138495-512;
Huaraz, 3250 m, KU 138513; 5 km N Recuay 3450 m, KU 138527-
44. Cajamarca: E slope Abra Gelic, 20 km E Celendin, 2740 m, KU
212071; S slope Abra Quilsh, 26 km NNW Cajamarca, 3500 m,
KU 212068; S slope Abra Quilsh, 28 km NNW Cajamarca, 3520
m, KU 212069; Cajamarca, 2800 m, KU 138494; 55 km N
Cajamarca, 3600 m, KU 212072-75; 8 km S Cajamarca, 3050 ni,
KU 212070; 23 km SW Celendin, 3050 m, KU 181740; Cutervo,
2620 m, KU 212055-57, 212060-66. Hiiniiuco: 5 km NE La
Union, 3100 m, KU 138411-52. ///;/;);: between Casa Pato
and Anascancha, ca. 10 km S Carhuamayo, 4050 m, KU
139189-90; Odonores, 4115 m, KU 207815. La Libcrtnd:
Huamachuco, 3350 m, KU 138453-59; Laguna Sacsacocha,
12 km E Huamachuco, 3200 m, KU 138460-91; Obizco, 2730
m, KU 138545-46.

10

SciENTiru r^M'i Ks, Nvn KM Hisi()K-> Ml SI I \i. Iiii l'\i\hKsiiv oi Kansas

Gaftrotlwaipsi/chivphiln: liCUADOI^: l.oja-Zaiiuna-Oundupc:
13-15 km ELoja, 2770-2850 m, KU 12076-62, 141586, 142631-37,
148599, 164233-34.

Castrotheca rebeccae: PERU: Ayacucho: 7.5 km SW Cant), 2970
m, KU 163302-03; Ccarpa, below Tambo on Vallo tlo Apurimac
Trail, 2470 m, KU 196806-11; Yuraccvacu on rambo-Valle l\v
Apurimac Trail, 2680 m, KU 196800-05.

Cii^twthccn ftictoplcura: I'oru: llunnuco: Chaglla, 3090 m,
MHNSM 20319; Cordiik>ra Av Carpish, 2625 m, MTD 45615. Paf.co:
Auqiiimarca, 2650 m, MHNSM 19815, MTD 44755-56, 4569);
above Auquimarca, MHNSM 17821, 17832, 17834, 17838, 17846,
17856, 18106; Paugmaray, Hiiachon, 2600 m, MHNSM 19166,
19168-79, 19172, 19186; MTD 45230; Uchuhuerta, 2500 m,
MHNSM 19873, MTD 45909.

Gnstrotluru tc>linii}icn: PERU: Ayncucho: Rio I'ione, lutumbaro,
1840 m, KU 163271-74, 163275 (skcl), 163276-78. Huanuco: base
of Bosque Zapata-Cocha, above Acamayo, 2545 m, KU 196815;
Huaylaspampa, 2590 m, KU 196817, Rio Huaylaspampa, 2530
m, KU 196816. Pasco: Yaupi, MTD 4.5907.

Giistwthcca walkeri: VENEZUELA: Aragiia: Km 26, Maracay-
Ocumare de la Costa Road, 770 m, KU 166766; Km 29, Maracay-
Ocumare de la Costa Road, 650 m, KU 166767-69, 185737-8;
Rancho Grande, 1100 m, KU 133390-91, 166765, 185734-36.

CiKtroihcca williaiiisoni: VENEZUELA: Cnrabobo: San Esteban,
UMMZ 55559.

GiKtivthccn zcuiioc\/stis: PERU: Hmitinco: Cordillera de
Carpish, 2920 m, MHNSM 18675 (holotype), MTD 45984 (C&S).

Gnstrothcra sp.: PERU: Prtsai; San Alberto, Parquc Nacional
Yanachaga, 2200, MHNSM 19901, MTD 45943.

3

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