Marsupial

A New species ot

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ZUUI. puetiman,

from the Cordillera Azul in Peru. Scientific

History Museum, The Universi

Gastrotheca)

Frog (Anura: Hylidae:

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668 NOV 05 2001
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=sScientific Papers x

UNIVERSI’ 1

Natural History Museum

: The University of Kansas
2 21 September 2001 Number 22:1-10
z A New Species of Marsupial Frog

(Anura: Hylidae: Gastrotheca)
from the Cordillera Azul in Peru
By
WIM E. DuELLMAN!, EDGAR LEHR’, AND CESAR AGUILAR?

'Natural History Museum and Biodiversity Research Center, and
Department of Ecology and Evolutionary Biology.
The University of Kansas, Lawrence, Kansas 66045-2454, USA.

*Staatliche Naturhistorische Sammlungen Dresden Museum fiir Tierkunde,
A.-B.-Meyer,-Bau Koenigsbruecker Landstrasse 159, D-01109 Dresden, Germany
3 *Departamento de Herpetologia, Museo de Historia Natural de San Marcos,
2 Apartado 14-0434, Lima 14, Peru
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ABSTRACT Anew species of Gastrotheca from the southern end of the Cordillera Azul in Peru seems
to be most closely related to G. lateonota known only from the Cordillera Huancabamba in northern

Peru. Both of these species and G. marsupiata, monticola and peruana produce tadpoles, whereas the
other species of Gastrotheca in Peruvian Andes have direct development.

Key Worps: Anura; Hylidae; Gastrotheca; new species; Peru; Andes.

© Natural History Museum, The University of Kansas

ISSN No. 1094-0782

2) SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS

RESUMEN Una nueva especies de Gastrotheca del extremo sur de la Cordillera Azul aparentemente
esta realcionada a G. lateonota que solo esta registrada para la Cordillera de Huancabamba al norte de
Peru. Ambas especies junto con G. marsupiata, monticola, y peruana producen renacuajos, mientras que
las otras especies de los Andes peruanos tienen desarrollo directo.

PaLasras Craves: Anura, Hylidae, Gastrotheca; especie nueva; Peru; los Andes.

INTRODUCTION

Marsupial frogs of the genus Gastrotheca are highly di-
verse in the Andes, especially in Colombia, Ecuador, and
Peru, where 31 species have been recorded; 14 of these
species have been reported from Peru (Duellman, 1987;
Duellman and Fritts, 1972; Duellman and Trueb, 1988;
Duellman and Wild, 1993; Trueb and Duellman, 1978). Of
these species, G. Jongipes occurs in the upper Amazon
Basin, and G. testudinea and G. weinlandii have exten-
sive distributions in cloud forests on the eastern slopes
of the Andes from Colombia to Bolivia. The other spe-
cies in Peru are restricted to the Andes, mostly at el-
evations above 2000 m.

Of these high Andean species, four have rather exten-
sive distributions. Gastrotheca marsupiata ranges from cen-
tral Peru into Bolivia and G. griswoldi occurs in central Peru;
the range of G. peruana extends from central Peru north-
ward in the Cordillera Occidental, whereas that of G.
monticola includes the northern parts of the Cordillera Oc-
cidental and Cordillera Central, the Cordillera de
Huancabamba, the Huancabamba Depression in Peru
and Ecuador, and the southern part of the Cordillera
Occidental in Ecuador. The nominal species G. lojana
was placed in the synonymy of G. monticola by
Duellman and Hillis (1987).

The other seven species have restricted distributions in
the Andes. Gastrotheca abdita is known only from the iso-
lated Cordillera Colan, and G. galeata and G. lateonota are
known only from the Cordillera de Huancabamba in north-
ern Peru. Four other species (G. excubitor, ochoai,
pacchamama, and rebeccae) have restricted allopatric distri-
butions at high elevations in the Cordillera Oriental in cen-
tral and southern Peru.

Field work in 1998 by Lehr and Aguilar in the Cordil-
lera Azul, a part of the Cordillera Oriental in central Peru,
resulted in the discovery of several new species of anurans
(Lehr, 2001; Lehr et al. 2001, 2002). Among these is a previ-
ously unknown species of Gastrotheca.

ACKNOWLEDGMENTS

The field work was supported by the Forschungsinstitut
und Naturmuseum Senckenberg. Lehr is grateful to per-
sonnel at the Instituto Nacional de Recursos Naturales of
the Ministerio de Agricultura in Lima, Peru, for providing
collecting and export permits. Javier Icochea and Gunther
Kohler made helpful suggestions. The manuscript ben-
efited from critical reviews by Joseph R. Mendelson II] and
Erik R. Wild, and from careful editing by Linda Trueb. The
accurate renderings of the frogs are from the talented hand
of Christopher A. Sheil (Kansas).

MATERIALS AND METHODS

The 16 morphological measurements and 25 external
descriptive characters are those used by Duellman and
Pyles (1980), Duellman and Hillis (1987), and Duellman
and Trueb (1988). All measurements are in mm; snout—vent
length is abbreviated SVL. Museum codes are those of
Leviton et al. (1985) with the addition of the Museo de

Historia Natural Universidad San Marcos, Lima, Peru
(MHNSM). The distribution map is based on the Mapa
Fisico Politico (Instituto Geografico Militar del Peru, 1973).
Statistical analyses were performed with StatView 4.5 (Aba-
cus Concepts, Inc., 1992-1995). Numbered colors refer to cor-
responding colors in the color guides of Smithe (1975, 1981).

DESCRIPTION OF NEW SPECIES

Gastrotheca stictopleura new species

Holotype-—MHNSM 20319, an adult female, from
Tranca Grande at Chaglla (09°51’08" S, 75°54’37" W, eleva-
tion 3090 m), Provincia de Pachitea, Departamento de
Huanuco, Peru, obtained on 23 August 1998 by Edgar Lehr.

Referred specimens.—SMEF 8032829, juveniles, from
Cochacalla at Chaglla, Provincia de Pachitea,
Departamento de Huanuco, Peru, obtained on 23 August
1998 by Edgar Lehr.

Diagnosis.—A moderate-sized species (females to 68.3
mm) having (1) tibia length 49.9% SVL, noticeably shorter
than foot; (2) interorbital distance noticeably greater (160%)
than width of upper eyelid; (3) skin on dorsum finely
shagreen, not co-ossified with skull, lacking transverse
ridges; (4) supraciliary processes absent; (5) heel lacking
calcar or pronounced tubercles; (6) tympanic annulus
smooth; (7) Finger I shorter than Finger II; width of disks
greater than that of digits; (8) fingers unwebbed; (9) web-

NEw SPECIES OF MARSUPIAL FROG 3

Table 1. Measurements of adult females of three species of Gastrotheca.
Means and 1 SD given below ranges

Character G. stictopleura G. lateonota G. monticola
N 1 13 73
Snout-vent length* ** 68.3 54.4-63.7 46.3-73.0
a DOWER) 59.4 44.42
Tibia length* ** 34.1 26.2-30.2 24.1-38.2
— 27.6+1.11 29.82.21
Foot length 30.5 25.8-31.8 21.2-39.4
-- 29.2 + 1.34 27.6 +2.48
Head length* 20.3 16.7-20.0 14.7-22.7
— 18.5 + 0.89 18.8 + 1.18
Head width* 23.1 18.7-21.8 16.4-29.8
— 20.8 + 0.57 Pals) 17/7/
Interorbital distance* 8.0 5.6-6.7 5.0-10.0
_— 6.2 + 0.67 7.4+1.03
Eyelid width** 5.0 319-5: 3.0-5.1
— 4.3 40.32 4.2+0.35
Internarial distance* 4.1 3.3-4.1 3.24.6
_ 3.8 + 0.27 3.9+0.31
Eye diameter* 5.8 5.0-5.8 3.9-6.8
= 5.4 + 0.29 5.5 + 0.44
Eye-nostril distance 5.0 4.4-5.2 4.4-6.5
— 4.8 +0.30 5.2 = 0.51
Orbit-jaw distance* 3.0 242.9 2.1-3.7
— 2.7 £0.19 2.8 + 0.31
Nostril-jaw distance* ** ks) 3.64.3 2.6-4.8
4.0 + 0.27 3.9 +0.41
Tympanum diameter 3.0 DOOD, 2.3-3.8
— 2.8 + 0.23 3.0 + 0.36
Thumb length 12.0 QW 8.5-13.3
= 10.8 + 0.71 10.8 + 1.07
Third finger length* 21.8 17.5-21.8 15.3-24.4
— 20.3 + 0.84 19.3 + 1.76
Disc width* 3.5 D339) 2.34.2
3.00.21 3.1 £0.40

* Differences between G. stictopleura and mean of G. lateonota
significant (Htest, P < 0.01).
** Differences between G. stictopleura and mean of G. monticola
significant (t-test, P <0.01).

bing extending to penultimate subarticular tubercle on Toe
IV, extending to distal subarticular tubercle on Toe V; (10)
dorsum essentially uniform green; (11) head markings con-
sisting of pale cream labial and canthal stripes, the latter
bordered below by narrow brown stripe; (12) pale cream
labial stripe present, bordered below by brown; (13) flanks
green with small white spots posteriorly; (14) venter
creamy tan.

Gastrotheca stictopleura most closely resembles G.
lateonota and G. monticola; all three species are similar in
size and proportions, although the single adult female of
G. stictopleura is slightly larger than females of G. lateonota.
Furthermore, there are significant morphometric differ-
ences between the two species; G. stictopleura has a sig-
nificantly larger head (Table 1). Also, the snouts in profile
are slightly different (Fig. 1), and the ventral color pattern
is different in the three species; the belly is dull grayish

Fig. 1. Heads of three species of Gastrotheca. A. G. lateonota, KU

181733. B. G. monticola, KU 181743. C. G. stictopleura, MHNSM
20319. All are adult females. Scale bar = 5 mm.

brown in G. lateonota, cream with black spots in G. monticola,
and creamy tan with a few brown flecks in G. stictopleura
(Fig. 2). Other species in the high Andes of central and
northern Peru are G. griswoldi, marsupiata, and peruana.

4 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS

Fig. 2. Ventral coloration in adult females of three species of Gastrotheca. A. G. lateonota, KU 181736, 63.3 mm SVL. B. G. monticola, KU 181743,
64.1mm SVL. C.G. stictopleura, MHNSM 20319, 68.3 mm SVL.

These are smaller species (SVL <57 mm) that have smaller
discs on the fingers and less webbing, which extends only
to the penultimate subarticular tubercle on Toe V.
Gastrotheca peruana differs further by having Fingers I and
II equal in length, a granular tympanic annulus, and pus-
tular skin on the dorsum. Furthermore, these three spe-
cies differ from G. sfictopleura in dorsal coloration.
Gastrotheca griswoldi and G. marsupiata have a dark inter-
orbital bar; in G. griswoldi, the bar usually is connected to
diagonal or paravertebral marks on the body, whereas in
G. marsupiata the bar is separate from the dark flecks or
spots on the body. Gastrotheca peruana lacks a dark interor-
bital bar and has elongate paravertebral marks on the body.

Other species of Gastrotheca inhabiting the Cordillera
Oriental in Peru (G. excubitor, ochoai, pacchamama, and
rebeccae) are smaller than G. stictopleiura (maximum SVL <
46 mm) and have direct development. Two species inhab-
iting cloud forests on the Andean slopes of the cordillera
(G. testudinea and G. weinlandii) differ from G. stictopleura
in several characters. Both are much larger, with females
attaining a SVL in excess of 80 mm, and have Finger I longer
than Finger II, discs on digits much wider than digit at
base of the disc, granular tympanic annulus, and a tan
dorsum with transverse bark brown bars (G. testudinea) or
chevrons (G. weinlandii). The latter has the skin on the head
co-ossified with the skull and has large calcars, whereas
the skin is not co-ossified in G. testudinea, which has a small
tubercle on the heel. On cursory examination, the skin on
the dorsal surface of the skull appears to be co-ossified
with the underlying cranial elements in some preserved
specimens of G. lateonota, monticola, stictopleura, and
testudinea; however, the texture of the skin reflects to sculp-

turing of the underlying bone, and the skin is moveable
on the top of the head.

Description of holotype.— Brooding female having a
SVL of 68.3 mm; head slightly wider than long, narrower
than body; head length 29.7% SVL; head width 33.8% SVL;
snout-moderately long, acutely rounded in dorsal view,
inclined anteroventrally in profile; nostrils slightly protu-
berant, directed anteroventrally, situated at a point 54.2%
distance from anterior level of orbit to tip of snout, at level
well behind margin of lower jaw; eye-nostril distance
86.2% length of eye. Canthus rostralis straight, elevated,
rounded in profile; loreal region noticeably concave; lips
rounded; internarial region slightly concave; top of head
depressed; interorbital distance 34.6% head width, 160%
width of upper eyelid. Tympanum vertically ovoid, sepa-
rated from eye by distance 1.4 x length of tympanum; tym-
panic annulus distinct anteriorly and ventrally, smooth;
supratympanic fold moderately weak, barely overlapping
upper edge of tympanum, extending from posterior cor-
ner of orbit to point above insertion of forearm.

Arm moderately robust; hand moderately large; fingers
unwebbed; distinct lateral fringes on Fingers II-IV; discs
moderately large with median longitudinal groove in an-
terior part of each pad; width of disc on third finger no-
ticeably greater than length of tympanum; relative length
of fingers 1 < 2 < 4 < 3; subarticular tubercles large,
rounded, none bifid; supernumerary tubercles absent; pal-
mar tubercle low, elliptical; prepollical tubercle elongately
elliptical (Fig. 3A). Hind limb moderately robust; tibia
length 49.9% SVL; foot length 44.7% SVL; heels of
adpressed limbs overlapping by about one-fourth length
of shank; calcar, heel tubercle, and outer tarsal fold absent;

New Species OF MARSUPIAL FROG 5

B

Fig. 3. Hand (A) and foot (B) of holotype of Gastrotheca stictopleura,
MHNSM 20319.

inner tarsal fold distinct, extending full length of tarsus;
outer metatarsal tubercle absent; inner metatarsal tubercle
ovoid, barely visible from above. Toes long, slender, bear-
ing distinct lateral fringes; relative length of toes a <2 <3
<5 <4; toes less than one-half webbed; webbing formula I
2—2 Il 1—3 III 2—3 IV 3—2. V; discs slightly smaller than
those on fingers; subarticular tubercles prominent, round;
supernumerary tubercles prominent, present on proximal
segments of all toes (Fig. 3B).

Skin on dorsum shagreen; skin on ventral surfaces of
forearms and shanks smooth; skin on other ventral sur-
faces granular; vertical cloacal folds present; transverse row
of four rounded tubercles on posterior surface of each thigh
lateral to cloaca, decreasing in size distally; opening of
brood pouch V-shaped, with the apex just posterior to
sacrum. Vomerine odontophores slightly oblique
posteromedially, narrowly separated medially, at posterior
margins of choanae, bearing 5 and 6 teeth; choanae mod-
erately large, nearly round; tongue bluntly ovoid, shal-
lowly notched posteriorly, free posteriorly for about one
third of its length.

Color in preservative: Dorsum of head, body, and limbs
bluish gray with narrow cream stripe originating as verti-
cal line on snout, passing along canthus rostralis, margin
of upper eyelid, and supratympanic fold, terminating on
upper flank at about midlength of body; narrow cream

stripe on margin of upper lip, followed posteriorly by two
white spots above insertion of forelimb. Axilla and me-
dian and posterodorsal surfaces of flanks bluish gray with
small white spots; groin pale grayish tan without mark-
ings. Narrow, transverse white stripe above vent; crests of
vertical folds and tubercles lateral to vent white; narrow
white stripe on ventrolateral edge of forearm; narrow white
bar on heel. Dorsal surfaces of fingers and Toes I-III tan;
other toes and all discs bluish gray. Throat pale bluish gray
with creamy white granules; other ventral surfaces and
anterior and posterior surfaces of thighs dull creamy tan;
some granules on the belly with a small amount of brown
pigmentation (Fig. 3C).

Color in life: From color photographs and field notes
by Lehr. Dorsal surfaces of head, body, and limbs, and lo-
real and tympanic regions bright dark green (162B) with
diffuse salmon (6) on body and limbs; labial and dorsolat-
eral stripes cream (54); dorsolateral stripe bordered ven-
trally by brown (38) stripe; spots on flanks, above inser-
tion of forearm, and lateral to vent white. Throat, chest,
and ventral surfaces of arms cream (54) to yellow (56) with
diffuse green (162B) and salmon (6) spots; belly pale brown
(223D); ventral surfaces of thighs dark brown (119A);
shanks green (162B); hands and feet dark gray (79). Iris
metallic orange (17) with fine black reticulations (Fig. 4E).

Measurements of holotype: See Table 1.

Cranial osteology: By lifting the skin covering the right
posterolateral part of the skull, it was possible to deter-
mine the association of the frontoparietal and squamosal.
The squamosal lacks a medial flange and is not in contact
with the otic flange of the frontoparietal. This condition is
like that described for Gastrotheca lateonota by Duellman
and Trueb (1988) and differs from that in G. monticola, in
which a median flange of the squamosal is in broad con-
tact with the otic flange of the frontoparietal (Fig. 5).

Variation.—Iwo juveniles have SVLs of 28.5 and 32.7
mim. The coloration of the larger of these (SMF 80328) is
like that of the holotype, except that there are faint tan
markings—elongate, X-shaped mark on the dorsum be-
ginning on the upper eyelids and terminating postsacrally
with the intersection in the scapular region; a short, longi-
tudinal, middorsal mark posterior to the sacrum; irregu-
lar marks on dorsal surfaces of limbs. The posterior part
of the flanks is white with three vertical brown marks. The
dorsal coloration of the smaller specimen (SMF 80329) is
like that of the holotype, but the venter has more granules
with brown pigmentation; the flanks posterior to the axil-
lary region are white with brown spots.

Field notes on coloration in life of these two juveniles
are: SMF 80328: Dorsum green (159) with metallic salmon
(6) spots on head, body, and extremities. SMF 80329: Dor-
sum green (6), hands and feet cream (54); cloacal region

6 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS

lean fe Fou ge A
ee pores wt me = ss Scot

A. Gastrotheca monticola, female, KU 212022, 51.5 mm SVL.

eo 8 5 ae 44

. Gastrotheca lateonota, female, KU 181732, 5

ts

.6mm SVL. D. Gastrotheca lateonota, female, KU 181730, 61.2 mm SVL.

= wa

E. Gastrotheca stictopleura, female, MHNSM 20319, 68.3 mm SVL. FE. Gastrotheca stictopleura, juvenile, SMF 80329, 28.5 mm SVL.

Fig. 4. Three species of Gastrotheca from Peru and southern Ecuador.

New Species OF MARSUPIAL FROG 1

Frontoparietal

Fig. 5. Skulls of Gastrotheca monticola (left) and G. lateonota (right)
showing relationship of frontoparietal and squamosal. The condition in
G. stictopleura is like that in G. lateonota. Scale bar = 5 mm. Adapted from
Duellman and Trueb (1988).

dark brown (119B) bordered by white; canthal and dorso-
lateral stripes cream (54) bordered below by dark brown
(223B) stripe; lips cream (54); flanks white with dark brown
(119) spots; throat pale brown (92); chest and belly dark
brown (119B) with pale brown (92) spots; ventral surfaces
of extremities dark brown (119B) with horn-colored (92)
and salmon (106) spots; iris salmon (132D) (Fig. 4F).

Life history and development.—The holotype is a
brooding female. Opening the brood pouch revealed ova
about 5.3 mm in diameter. Eggs were not removed from
the pouch, but an estimated 82 eggs are present. At least
some of these eggs appear to be in developmental Stages
17-20 (Gosner, 1960); these stages are equivalent to Stages
16-18 described for Gastrotheca riobambae by del Pino and
Escobar (1981). The number and size of eggs indicate that
the eggs of Gastrotheca stictopleura hatch into tadpoles that
complete their development in ponds (Fig. 6). Gastrotheca
lateonota and G. monticola also produce eggs that hatch as
tadpoles. Eleven brooding females of G. lateonota contained
66-152 (x= 103.1 + 27.6) eggs in developmental Stages 31—
35 (Gosner, 1960) having diameters of 5.3-7.3 (x = 6.4 +
0.66) mm. Four brooding females of G. monticola contained
66-186 (x = 129.5 + 57.3) eggs having diameters of 3.6-4.9
(x = 4.4 + 0.57) mm. The number of eggs brooded by fe-
males of species having direct development is much
smaller (Fig. 6).

Distribution and ecology.—Gastrotheca stictopleura is
known only from the immediate vicinity of Chaglla,
Departamento Hudnuco, Peru (Fig. 7). This site is in the
southern end of the Cordillera Azul; this name is applied
to that part of the Cordillera Oriental lying to the east of

200}

O° G lateonota

O G. monticola

A G. stictopleura

@ Seven direct-developing species

Number of eggs
ra 3 dD
[o) oO (o)

[op)
[o)

35 40 45 50 55 60 65 70
Snout-vent length (mm)

Fig. 6. Regression plots of number of eggs in brood pouches and
corresponding female snout-vent length. Open symbols are for species
producing eggs that hatch as tadpoles; solid symbols are means of seven
Peruvian species producing eggs that hatch as froglets. The latter group
includes Gastrotheca abdivita (n = 8), G. excubitor (n = 7), G. galeata (n = 4),
G. griswoldi (n = 6), G. ochoai (n = 12), G. pacchamama (n = 4), and G. rebeccae
(n = 1).

the Rio Huallaga Valley. Chaglla is in the transition zone
between “matorral himedo” and “bosque humedo de
montanas” (INRENA, 1995, 1996). The former exists at el-
evations of 2600-3400 m and is characterized by evergreen
bushes. The village of Chaglla is in a region of extensive
cultivation, principally potatoes.

Cochacalla and Tranca Grande are local names for ar-
eas within Chaglla. Cochacalla is a narrow valley with
bushes and small trees along a small stream; Tranca Grande
is a marshy, grassy plain with scattered trees and bushes.
The specimens of Gastrotheca stictopleura were obtained by
local residents who claimed that the frogs were found in
trees.

Gastrotheca stictopleura is sympatric with the smaller,
terrestrial G. griswoldi at Chaglla. Other species of anurans
collected in the immediate vicinity of Chaglla include Bufo
spinulosus, anew species in the Bufo veraguensis group, and
four species of Phrynopus.

8 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS

SOO

Cordillera de

Huancabamba = q

7
© gi

\

ir
(
<
)
Jp Lambayeque A |

e
Z
an

| Cajamarca

@ G /ateonota
HG. monticola
A G. stictopleura

[ Area above 3000 m.

100 200

Kilometers

Ancash

Fig. 7. Distributions of three species of Gastrotheca in southern Ecuador and northern Peru.

Etymology.—The specific name is a Latinized feminine or spotted, and the Greek pleura, meaning side. The name
adjective derived from the Greek stiktos, meaning dappled _ refers to the spotted pattern on the flanks.

New SPEcIES OF MARSUPIAL FROG 9

DISCUSSION

In size, external morphology, nature of the frontopari-
etals and squamosals, and mode of life history, Gastrotheca
stictopleura is most like G. lateonota, known only from the
Cordillera de Huancabamba in northern Peru (Fig. 6). With
the exception of the nature of the frontoparietals and
squamosals, these two species are more like G. monticola
than any other members of the genus. Gastrotheca monticola
ranges from southern Ecuador throughout northern Peru
(Fig. 7).

Like many other species of Gastrotheca in the high
Andes, G. lateonota, monticola, and stictopleura exhibit con-
siderable variation in dorsal coloration. In G. lateonota, the
dorsum is green or brown with or without darker paraver-
tebral marks, and in G. monticola, the dorsum is green or
brown, usually with a darker middorsal blotch or paraver-
tebral marks (Fig. 4A—D). The three known specimens of
G., stictopleura are green dorsally, but one specimen (SNM
80328) also has tan markings on the dorsum.

The co-occurrence of Gastrotheca griswoldi, in which
eggs hatch as froglets in the brood pouch, and G.
stictopleura, in which eggs hatch as tadpoles that complete
their development in ponds, reflects a consistent pattern
in Gastrotheca in the high Andes. All cases of sympatry in-
clude one species that produces tadpoles and another in
which the eggs undergo directed development. Thus, the
direct-developing G. griswoldi occurs sympatrically in dif-
ferent parts of its range with G. peruana and G. stictopleura,
both of which produce tadpoles. Likewise, the widespread
G. marsupiata, which produces tadpoles, occurs sympatri-
cally with slightly smaller species that exhibit direct de-
velopment (G. excubitor, ochoai, and pacchamama) through-
out their separate ranges in the Cordillera Oriental. On
the other hand, no two species that produce tadpoles
are known to occur sympatrically. The same situation
prevails in the high Andes of Bolivia, Ecuador, and Co-
lombia.

LITERATURE CITED

Del Pino, E. M., and B. Escobar. 1981. Embryonic stages of Gastrotheca
riobambae (Fowler) during maternal incubation and comparison of
development with that of other egg-brooding hylid frogs. Journal of
Morphology 167:277-295.

Duellman, W. E. 1987. Two new species of marsupial frogs (Anura:
Hylidae) from Peru. Copeia 1987:903-909.

Duellman, W. E., and T. H. Fritts. 1972. A taxonomic review of the south-
ern Andean marsupial frogs (Hylidae: Gastrotheca). Occasional Pa-
pers Museum Natural History University of Kansas 9:1—-37.

Duellman, W. E., and D. M. Hillis. 1987. Marsupial frogs (Anura: Hylidae:
Gastrotheca) of the Ecuadorian Andes: resolution of taxonomic prob-
lems and phylogenetic relationships. Herpetologica 43:141-173.

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

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

Duellman, W. E., and E. R. Wild. 1993. Anuran amphibians from the Cor-
dillera de Huancabamba, northern Peru: systematics, ecology, and
biogeography. Occasional Papers Museum Natural History Univer-
sity of Kansas 157:1-53.

Gosner, K. L. 1960. A simplified table for staging anuran embryos and
larvae with notes on identification. Herpetologica 16:183-190.

INRENA (Instituto Nacional de Recursos Naturales). 1995. Mapa Forestal
del Peru. Lima: Direccion General Forestal.

INRENA (Instituto Nacional de Recursos Naturales). 1996. Guia Explicativa
del Mapa Forestal. Lima: Direccion General Forestal.

Lehr, E. 2001. A new species of Phrynopus (Anura: Leptodactylidae) from
the eastern Andean slopes of central Peru. Salamandra 37:11-20.
Lehr, E. G. Kéhler, and E. Ponce. 2001. Anew species of Phrynopus (Anura:

Leptodactylidae) from Peru. Senckenbergiana Biologie 80:205—212.

Lehr, E., C. Aguilar, and G. Kohler. 2002. Two sympatric new species of
Phrynopus (Anura: Leptodactylidae) from a cloud forest in the Peruvian
Andes. Journal of Herpetology (in press).

Leviton, A. E., R. H. Gibbs, Jr, E. Heal, and C. E. Dawson. 1985. Stan-
dards in herpetology and ichthyology: Part I. Standard symbolic codes
for institutional resource collections in herpetology and ichthyology.
Copeia 1985:802-832.

Smithe, E. B. 1975. Naturalist’s Color Guide. Part 1. Color Guide. New York:
American Museum of Natural History.

Smithe, F. B. 1981. Naturalist’s Color Guide. Part Il. Color Guide Supple-
ment. Color Swatches. New York: American Museum of Natural His-
tory.

Trueb, L., and W. E. Duellman. 1978. An extraordinary new casque-headed
marsupial frog (Hylidae: Gastrotheca). Copeia 1978:498—-503.

APPENDIX
SPECIMENS EXAMINED

Gastrotheca lateonota: PERU: Piura: El Tambo, 31.5 km E Canchaque,
2180 m, KU 181729, 181730 (holotype), 18173139, 181836-37 (skeleton),
MHNSM 1635.

Gastrotheca monticola: ECUADOR: Azuay: Giron, 2240-2500 m, KU
13840103. Loja: Celica, 2130 m, BMNH 1931.11.3.3-4; Loja, 2150 m, BMNH
1931.2.12.10-13, 1933.6.3.18-44, 1935.11.3.26-32, 1947.2.31.6-12,
1931.2.31.13 (holotype of G. lojana), 1931.2.31.14-18, KU 120673-74, USNM
258851-58; 2 km N Loja, 2100 m, KU 142846 (tadpoles); 5 km N Loja,
2150 m, KU 138235-36, 138237 (skeleton); 2 km E Loja, 2200 m, KU 120675,
USNM 258849-50; 6.8 km E Loja, 2640 m, KU 217511-12; 9 km E Loja,
2660 m, KU 121387 (tadpoles); 10 km E Loja, 2600 m, KU 142855 (tads),
178470-76; 2 km S Loja, CAS 93898; 3 km W Loja, 2150 m, KU 138233; 5.2
km W Loja, 2310 m, KU 202688, 203547 (tadpoles); 5.5 km W Loja, 2330,
KU 142603-08, 148549-51; 7.9 km W Loja, 2440 m, KU 203548 (tadpoles);
10 km W Loja, 2500 m, KU 138234; Saraguro, 2500 m, KU 138404-09,

138410 (skeleton), 148568. Zamora-Chinchipe: Zamora (?), BMNH
1933.6.24.45.

PERU: Amazonas: N slope Abra Barro Negro, 27 km WSW
Leimebamba, 3440 m, KU 212078; Chachapoyas, 2340 m, KU 13823841,
215627 (tadpoles), MCZ 88897-901, MHNSM 6277 (tadpoles); 20.5 km
WSW Leimebamba, 3130 m, KU 181741; 22 KM WSW Leimebamba, 3220
m, KU 212495 (tadpoles), MHNSM 6294 (tadpoles); 24 km WSW
Leimebamba, 3370 m, FSM 30080; 5 km N Levanto, 2850 m, KU 212021; 6
km NW Mendoza, 2200 m, KU 209421; Molinopampa, 2400 m, KU 212022—
31, 212493 (young), 212494 (tadpoles), MHNSM 6116-21, 6287 (tadpoles),
6292 (young); Pomacochas (Florida), 2180 m, KU 181742-70, 181838-39
(skeleton), 212032-36, MHNSM 1040 (5), 6122-31. Cajamarca: No specific

Cutervo, 2620 m, KU 212055-66, NMW 6483; 8 km NW Cutervo, 2560 m,
KU 212067, 212492 (tads); Querocotillo, MCZ 5328-30. Piura: Ayabaca,

10 ScrentTiFic PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS

2700 m, MHNSM 702 (2); W slope Cerro Chinguela on Huancabamba— __m, KU 209769-70, MHNSM 15420-21; 18.5 km WSW Huancabamba, 2740
San Ignacio trail, 2620 m, KU 196819; Huancabamba, 1960 m, AMNH m, KU 181874 (tadpoles).

7551, KU 219771, MCZ 5290 (holotype), 5291-93, 5296-97, 5299-300, 5302, Gastrotheca stictopleura: PERU: Huanuco: Chaglla, MHNSM 20319 (ho-
5304-07, 5309, 5312-15, 5317, 5319, 5328-30; 1.8 km N Huancabamba, _ lotype), SMF 80328-29.

1980 m, KU 219767-68, MHNSM 15418-19; 4 km N Huancabamba, 1900

3 2044 062 534 334

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