HERP

QL

668

.E257

D84

1999

bcientific Papers

Natural History Museum
The University of Kansas

30 July 1999 Number 13:1-78

Frogs of the Genus Eleutherodactylus
(Anura: Leptodactylidae) in the Andes of Northern Peru

By

William E. Duellman and Jennifer B. Pramuk

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

CONTENTS

ABSTRACT 2

RESUMEN 2

INTRODUCTION 3

Acknowledgments 3

MATERIALS AND METHODS 3

ANDES OF NORTHERN PERU 4

Geological History 4

Physiography 5

Climate 5

Vegetation 8

GENERA OF ELEUTHERODACTYLIINE FROGS 10

JT SUMMARY OF TAXONOMIC CHARACTERS 11

^ IDENTinCATION OF SPECIES 17

■ — Key to the Species 17

Clave de las especies 18

r^' ACCOUNTS OF SPECIES 20

"^ Eleutherodactylus conspicillatus Group 20

^ Eleutherod/ictyll/s N/GROvnr/iTus Group 33

■^ Eleutherodactylus orestes Group 35

S:::. Eleutherodactylus unistricatus Grovf 41

r: BIOGEOGRAPHY 69

■2> LITERATURE CITED 72

APPENDICES 75

^ Specimens Examined 75

^ Gazetteer 77

^ © Natural History Museum, The University of Kansas ISSN No. 1094-0782

a

^C/A 2 Scientific Papers, Natural History Museum, The University of Kansas

^ 7,S'*1 ABSTRACT We recognize 45 species in the leptodactylid frog genus Ekuthewdnch/his in the Andes of

northern Ecuador. Twenty-one of these species have been known previously from Peru, but three of

'T^'^0^ them are reported for the first time from the Andes. Six other species known previously from Ecuador

-^ are recorded for the first time in Peru, and 18 species are described as new. The majority (31) of the 45

! /^*7£/ species are members of the large EleiithewdiKtylus iinistrigntus group, and nine species belong to the

Eleutherodactylns conspicillatus group; one species belongs to the Eleutherodactyhis nigrovittahis group,

and four Peruvian species are in the EleutJierodnctylus orestes group, which otherwise is known from

three species in the Andes of southern Ecuador. Each species is treated in an account that includes a

diagnosis, description or reference to a description, habitat, and distribution. Keys in English and

Spanish to the species in the Andes of northern Peru are provided; 16 species are illustrated in color.

In the Andes of southern Ecuador and northern Peru, all members of the Eleutherodactyhis

conspicillatus group occur at elevations of less than 1500 m, but two species extend to elevations in

excess of 2000 m. Members of the Eleutherodactyhis nigrovittatus and orestes groups mostly are confined

St, to elevations above 3000 m, whereas species in the Eleutherodactyhis unistrigatus group range through-

o' out the elevations in the Andes to 3200 m. Among the isolated mountain ranges in northern Peru, the

o* Cordillera del Condor has the largest number of species of Eleutherodactyhis (19); of these, 11 are shared

«? N with Cordillera Oriental in Ecuador, nine with the Cordillera de Cutucu in Ecuador, and only five are

^ <D •• ' among the 16 species known from the northern part of the Cordillera Central in Peru. The Cordillera

^ ■-< '' Colan has five species of Eleutherodactyhis; one is shared with the Cordillera Oriental in Ecuador and

•Q S w another with the Cordillera Central in Peru. Of the 11 species inhabiting the Cordillera de Huancabamba,

o S --^ three are shared with the Cordillera Occidental in Ecuador, two with the Cordillera Oriental in Ecuador,

(0 § - and two with the CordiUera Occidental in Peru. The absence of collections of £/fHf/!erai(7cfi//"s from many

^ '-^ ^ mountain ranges in northern Pem suggests that the number of species far exceeds that reported here.

' ° in Key Words: Leptodactylidae, Eleutherodactyhis, Andes of northern Peru, Taxonomy, Biogeography.

S] RESUMEN Reconocemos 45 especies en el genero leptodactilido Eleutherodactyhis en los Andes del

;§ norte Peru. Vientiuna de estas especies eran conocidas previamente para Peru, pero tres de ellas se

reportano por primera vez para los Andes. Otras seis especies conocidos para Ecuador son reportadas
por primera vez para Peru, y 18 especies son descriptas como nuevas. La mayoria (31) de las 45
especies son miembros del grupo Eleutherodactyhis unistrigatus, y nueve especies pertenecen al grupo
Eleutherodactyhis conspcillatus; una especie pertenece al grupo Eleutherdactyhis nigrovittatus, y cuatro
especies peruanas estan en el grupo Eleutherodactyhis orestes, el cual tambien es conocido por tres especies
en los Andes del sur de Ecuador Para cada una de las especies se realiza un resumen que uncluye
diagnosis, descripcion o referenda a una descripcion, habitat y ciistribucion geografica. Se proveen
claves en ingles y espaiiol para la identificacion de las especies en los Andes del norte de Peru; 16
especies se ilustradan a color.

En los Andes del sur de Ecuador y el norte de Peru, todos los miembros del grupo Eleutherodactyhis
conspicillatus se encuentran en elevaciones menores a 1500 m, pero dos especies se extienden hasta
elevaciones mayores a 2000 m. Miembros de los grupos Eleutherodactyhis nigrovittatus y Eleutherodactyhis
orestes estan restringidos a elevaciones mayores de 3000 m, mientras que especies en el grupo
Eleutherodactylus unistrigatus sew extiendenpor todas las elevaciones de los Andes hasta los 3200 m.
Entre las Cordilleras aisladas en el norte del Peru, la Cordillera del Condor tiene el numero mas grande
de especies de Eleutherodactyhis (19); de estas, 11 se encuentran tambien en la Cordillera Oriental en
Ecuador, nueve en la Cordillera de Cutucu en Ecuador, y solamente cinco estan entre las 16 especies
conocidas en el parte norte de la Cordillera Central en Peru. La Cordillera Colan tiene cinco especies
de Eleutherodactyhis; una ocurre tambien en la Cordillera Oriental en Ecuador y otra en la Cordillera
Central en Peru. De las 11 especies que habitan la Cordillera de Huancabamba, tres ocurren tambien
en la Cordillera Occidental en Ecuador, dos en la Cordillera Oriental en Ecuador, y dos en la Cordillera
Occidental en Peru. La ausencia de colecciones de Eleutherodactyhis de muchas Cordilleras en el norte
de Peru sugiere que el numero de especies excede en mucho al que aqui se reporta.

Palabras claves: Leptodactylidae, Eleutherodactyhis, Andes del norte de Peru, Taxonomia, Biogeografia.

Eleutherodactylus in the Andes of Northern Peru

INTRODUCTION

Frogs of the genus Eleutherodactylus are speciose in the
cloud forests on the slopes of the Andes in Ecuador and
Colombia (Lynch et al., 1997; Lynch and Duellman, 1980,
1997). Several species of Eleuthewdacti/his have been de-
scribed from cloud forests on the Amazonian slopes of the
Andes in southern Peru (Duellman, 1978a, b), but until
recently, few species have been recorded from northern
Peru. The first were £. cajauwrceusis and £. h/uuvii, which
were collected by G. K. Noble in 1916 and described by
Barbour and Noble (1920); both species are now known to
be rather widelv distributed in northwestern Peru and
southwestern Ecuador (Lynch and Duellman, 1997). More
than half a century passed until more collections were
made in the Andes in northern Peru.

Field parties from The University of Kansas collected
amphibians and reptiles in the Andes of northern Peru in
1979, 1989, and 1991, and field parties from Louisiana State
University obtained specimens from the Cordillera de
Huancabamba in 1974 and the Cordillera Colan in 1978.
Likewise field parties from the University of Florida ob-
tained specimens in 1970 and 1972. Rainer Schulte, a resi-
dent of Tarapoto, Peru, and his field companions collected
several species of amphibians from the eastern slopes of
the Andes and outlying ranges, as well as the southern
slopes of the Cordillera del Condor. Alfonso Miranda of
the Universidad Nacional Cajamarca, Peru, obtained speci-
mens in the northern part of the Cordillera Occidental, and
Javier Icochea and Robert P. Reynolds collected several
species on the eastern slopes of the Cordillera del Condor
in 1994. Material resulting from these field expeditions led
to the descriptions of nine new species of
Eleutherodactylus — E. schultei (Duellman, 1990a) and E.
lirellus (Dwyer, 1995) from the Cordillera Central, E.
petrobardus (Duellman, 1991a) from the Cordillera Occiden-
tal, E. bearsei (Duellman, 1992a) and E. citriogaster
(Duellman, 1992b) from eastern outliers of the Andes in
Departamento San Martin, and £. ceutliospilus, rliodoplichtts,
sternothylax, and zoiensi from the Cordillera de
Huancabamba (Duellman and Wild, 1993). The latter au-
thors also provided the first Peruvian records for three
species (£. colodactylus, crxjptomelas, and phoxocephalus) pre-
viously known only from Ecuador. Reynolds and Icochea
(1997) reported the first Peruvian locality for E. condor, pre-
viously only known from Ecuador, and provided the first
locality for E. pcruvianus from the Andes of northern Peru.
Flores and Rodriguez (1997) described E. karcharias from
the northern part of the Cordillera Central. Thus, 17 spe-

cies have been reported from the Andes of northern Peru.

Among the collections of Eleutherodactylus from north-
ern Peru are specimens of 18 more unnamed species, as
well as examples of six species known from Ecuador but
previously not reported from Peru. The purposes of this
paper are to present descriptions of the new species and
to provide a review of the species of Eleutherodactylus
known to occur in the Andes and associated mountain
ranges in northern Peru — departamentos Amazonas,
Cajamarca, Piura, and San Martin. Despite the large num-
ber of species now known from the region, our knowledge
of anurans in the Andes of northern Peru is rudimentary.
More thorough collecting surely will expand the presently
known ranges of many species, and the exploration of pre-
viously uncollected, isolated mountain ranges certainly
will reveal additional species. We would not be surprised
if the present number represents no more than half of the
eleutherodactyline fauna of northern Peru.

Acknowledgments

Duellman is indebted to his field companions — Tho-
mas J. Berger, David C. Cannatella, Fernando M. Cuadros
V, Michael E. Morrison, Rainer Schulte, and John J. Wiens —
whose efforts contributed greatly to the amount of mate-
rial available for study, to many residents of northern Peru
who provided shelter, food, and assistance to the field par-
ties, and to B. Anthony Luscombe of Lima, and Rainer
Schulte of Tarapoto for logistical support. For the loan of
specimens, we are grateful to David L. Auth of the Florida
State Museum; Frank T. Burbrink, David A. Good, and
Douglas A. Rossman of Louisiana State University; and
Robert P. Reynolds of the National Museum of Natural
History. We are grateful to Rainer Schulte for providing
photographs of one species, Erik R. Wild for measuring
many of the specimens and to Analia Piigener for consid-
erable help with the Resumen and key in Spanish. The
manuscript benefited from critical review by S. Blair
Hedges and John D. Lynch and careful editing by Linda
Trueb; we thank them profusely for their efforts. Permits
for the collection and exportation of specimens were is-
sued by Luis J. Cueto Aragon, Armando Pimental
Bustamento, Gonzalo Bravo Mejia Mufioz, and Jose
Purisaca, Direccion General Forestal y de Fauna, Ministerio
de Agricultura, Lima, Peru. The research reported herein
is part of a study on patterns of speciation and biogeogra-
phy of Andean anurans supported by a grant (BSR
8805920) from the National Science Foundation (W. E.
Duellman, PL).

MATERIALS AND METHODS

Specimens in museum collections are identified by their = Academy of Natural Sciences of Philadelphia; BM = Brit-
catalogue numbers preceded by the following codes: ANSP ish Museum (Natural History); KU = Natural History

Scientific Papers, Natural History Museum, The University of Kansas

Museum, University of Kansas; LSUMZ = Museum of
Zoology, Louisiana State University; MCZ = Museum of
Comparative Zoology, Harvard University; MHNSM =
Museo de Historia Natural, Universidad Nacional Mayor
de San Marcos, Lima, Peru; MNCN = Museo Nacional de
Ciencias Naturales, Madrid, Spain; NHMG =
Naturhistoriska Museet Goteborg, Sweden; NHRM =
Naturhistoriska Riksmuseet, Stockholm, Sweden; MSNT
= Museo e Instituto di Zoologia Sistematica, Universita di
Torino, Italy; USNM = National Museum of Natural His-
tory; UF = Florida State Museum, University of Florida,
Gainesville. All specimens from northern Peru and Ecua-
dor that have been studied are listed in Appendix 1. Lo-
calities from which specimens have been examined are
listed with their geographic coordinates and elevations in
Appendix 2.

Measurements, definitions of structural characters, and
numbered characteristics in diagnoses follow the meth-
ods of Lynch and Duellman (1997). Diagnoses of E.
bromeliaceus, colodactylus, cryptomelas, proserpens, and ver-
sicolor were modified from those of Lynch (1979), and those
of E. cajamarcensis, lymani, and phoxocephahis were taken
from Lynch and Duellman (1997), whereas those of E.
incomptus, condor, galdi, ockendeni, and pentvimuis were
modified from those of Lynch and Duellman (1980) and
that of £. pecki from Duellman and Lynch (1988).

A major problem in providing adequate diagnoses of
Eleutherodactylus is the comparison of new taxa with the
plethora of existing species. In order to be able to ascertain
character states among the many species that might be
confused with the species described or discussed herein,
we tabulated character states for all diagnostic characters

used by Lynch and Duellman (1997). For the purposes of
this study, we included all described species of
Eleutherodactylus from Peru and Ecuador, as well as the
new taxa; it is extremely unlikely that species from any
other region occur in northern Peru. To the nonspecialist,
it may seem that we have elaborated unnecessarily the
diagnoses of all of the taxa and the descriptions of the new
taxa. However, the distinction between specimens of many
species of Eleutherodactylus is difficult even for the special-
ist. We present detailed diagnoses and descriptions in an
endeavor to facilitate comparisons with other taxa that
certainly will be discovered in the future.

Measurements were taken with dial calipers to the near-
est 0.1 mm. If sex and reproductive condition were not
evident externally (nuptial pads or vocal sacs in males and
eggs visible through the body wall in females), sex and
reproductive condition were determined by dissection. The
following abbreviations are used: E-N = eye-nostril dis-
tance; HL = head length; HW = head width; lOD = inter-
orbital distance; SVL = snout-vent length. Photographs
noted as (ERW) were taken by Erik R. Wild; all others were
taken by Duellman.

Except for the sites visited by Reynolds and Icochea,
who used GPS devices, geographical coordinates were
obtained from maps, principally the Mapa Fisico Politico,
1:1,000,000 (1973) but also the Carta Nacional del Peru,
1:100,000 (1986) for those regions so mapped; both sets of
maps were produced by the Instituto Geografico Militar
del Peru. Coordinates for Ecuadorian localities were ob-
tained from the 1974 edition of the Mapa de Ecuador,
1:1:000,000, produced by the Instituto Geografico Militar,
Quito. Elevations were obtained by altimeters or in some
cases from maps.

ANDES OF NORTHERN PERU

Equaled in heights and expanse only by the Himalayas,
the majestic Andean mountain chain was described by
Enock (1907) as: "Heavenward thrown, crumpled, folded,
ridged and fractured, with gleaming 'porcelain' gnomons
pointing to the sun; shattered strata and shear crevasse;
far terraces and grim escarpments, hung over with filmy
mist- veils, and robed with the white clothing of crystalised
rains and mists; the birthplace of the winds and hails; the
father of rivers whose floods are borne a thousand leagues
away — the mighty cordillera is!" This description is espe-
cially appropriate for the Andes of northern Peru
(departamentos Amazonas, Cajamarca, Piura, and San
Martin), where north-south mountain ranges are separated
by deep valleys.

Geological History

In northern Peru and southern Ecuador, a major struc-
tural and physiographic break exists in the Andes (Figs. 1,

2); this is the Huancabamba Depression, a complex sys-
tem of relatively low ridges, basins, and valleys. Therein
is the lowest pass in the Andes between Colombia and
southern Chile — Abra de Porculla at 2145 m. In the
Huancabamba Depression, there is a structural deflection
of the Andean faults that corresponds to two major tec-
tonic segments of the Andes (Sillitoe, 1974). South of the
depression the axis of the Andes is northwest to south-
east; north of the depression the axis is north-northeast-
south-southwest.

Geological evidence points to extensive marine trans-
gressions in the region of the Huancabamba Depression
during the Cretaceous (Ham and Herrera, 1963). Orogenic
events associated with plate tectonics along the west coast
of South America in the Late Cretaceous resulted in the
uplift of the Andes to elevations probably not exceeding
1000 m above sea level (Zeil, 1979). The major uplift of the

Eleutherodactylus in the Andes of Northern Peru

Andes south of the depression was in the Miocene
(Harrington, 1962; Aubodin et al., 1973; Sempere et al.,
1990); the final major uplift was in the Pliocene with some
additional orogeny in the Pleistocene (James, 1973; Gansser,
1973; Noble et al., 1990). In contrast, the uplifted terrain
resulting from the Late Cretaceous orogeny subsequently
was eroded to low hills before the major uplift of the Andes
north of the Huancabamba Depression was initiated in the
Pliocene; this uplift continued into the Quaternary, as evi-
denced by many active volcanoes in Ecuador and Colom-
bia (Herd and Naeser, 1974; Sauer, 1971; Shagam, 1975;
Simpson, 1979). Thus, the Huancabamba Depression is not
only a physiographic anomaly in the Andes and a tectonic
border, but it bridges tectonic segments that were uplifted
at different times.

The present elevations and drainage patterns in the
Andes of northern Peru and the Huancabamba Depres-
sion probably were achieved in the Pleistocene (Gansser,
1973; Harrington, 1956). During the Pleistocene, climatic
fluctuations included cooler, drier conditions during gla-
cial phases, and warmer, more moist conditions during
interglacial phases. According to Sauer (1971), during in-
terglacials, climates were depressed 1500-2000 m in the
Ecuadorian Andes, whereas in the Peruvian Andes they
were depressed 1000-1500 m on the eastern slopes and 500-
1000 m on the western slopes (Hastenrath, 1967; Dollfus,
1976). These postulated changes are in accord with the
paleoecological work in Colombia by van der Hammer
(1974), van der Hammen and Cleef (1986), and Clapperton
(1987), and in the Ecuadorian Andes by Colinvaux (1988).
These climatic fluctuations presumably resulted in alter-
nating isolation and interconnection of montane environ-
ments, thereby providing corridors for, and barriers to,
dispersal in the Andes (Simpson, 1979; Colinvaux, 1993).

Physiography

A casual glance at a physiographic map of South
America reveals the long Andean mountain chain border-
ing the west side of the continent; as such, one might ex-
pect major drainages to be east and west. Although there
are many rivers originating in the Andes, some of the most
significant drainages are north and south. This is especially
evident in Colombia where the Rio Magdalena and Rio
Cauca form broad valleys between three major north-south
Cordilleras.

Between the Nudo de Pasto (Macizo Colombiano) in
southern Colombia, the Andes are formed by two major
ranges in Ecuador — the western Cordillera Occidental and
the eastern Cordillera Oriental. Between the eastern and
western Cordilleras are 10 basins that are completely or
partially separated by transverse ridges that in most cases
connect the cordilleras. South of the dry valley of the Rio

Jubones, the Cordillera Occidental diminishes in isolated
ranges, such as the Estribuciones de Celica. On the other
hand, the Cordillera Oriental is continuous with the Cor-
dillera de Huancabamba and Cordillera de Tabacones in
northern Peru, both of which terminate north of the Rio
Chamaya, a tributary of the Rio Marahon (Fig. 2).

In Peru, the Cordillera Occidental forms the backbone
of the Andes; it is confluent with the Cordillera Oriental at
the Nudo de Pasco. To the north of the Nudo de Pasco, the
valley of the Rio Marafion separates the Cordillera Cen-
tral from the Cordillera Occidental. The northern part of
the Cordillera Oriental is separated from the Cordillera
Central by the valley of the Rio Huallaga, which, as it
curves eastward, forms the northern boundary of the Cor-
dillera Oriental. Thus the easternmost range of the Andes
in Peru north of about 7° S Lat. is the Cordillera Central.

In southern Ecuador and northern Peru, the complex
topography is associated with north and south drainages
(Fig. 2). For example, the Cordillera del Condor is bordered
on the west by the Rio Narangaritza (flowing northward)
and on the east by the Rio Cenepa (flowing southward).
The major drainage system in the region is the Rio
Maraiion, which at its confluence with the Rio Huallaga
forms the Rio Amazonas. With the exception of the Rio
Chamaya, the major tributaries of the Rio Marafion flow
southward (e.g., Rio Huancabamba, Rio Cenepa, Rio
Chinchipe) or northward (e.g., Rio Chotano, Rio
Utcubamba, Rio Chiriaco). These rivers separate distinct
north-south mountain ranges (e.g., Cordillera de
Huancabamba, Cordillera Golan; Fig. 3). The change in di-
rection (from north to east) of the Rio Marafion forms the
northern boundary of the Cordillera Central. The north-
ern part of the Cordillera Central is dissected by rivers that
flow northward (Rio Chiriaco and Rio Utcubamba) and
southward (Rio Mayo). The resulting complex topography
contains major isolated highland areas (e.g., Cordillera
Golan) and the high elevations to the east of the Rio Mayo;
many ridges more than 1000 m above sea level extend
southward almost to the Rio Huallaga. Unlike other moun-
tain ranges in the region, the highest parts of the Cordil-
lera del Condor consist of sandstone table mountains (Fos-
ter and Beltran, 1997).

Climate

Temperature is dependent primarily on elevation. In
the lower valley of the Rio Marafion, temperatures may
exceed 40 °C. At high elevations (> 3000 m), mean monthly
temperatures are relatively constant throughout the year,
but the daily fluctuation can be as much as 20 ° C with noc-
turnal lows below freezing (Schwerdtfeger, 1976).

Rainfall patterns are determined by wind patterns and
topography. The prevailing winds from the Amazon Ba-

Scientific Papers, Natural History Museum, The University of Kansas

Fig. 1 . Political map of the Andes of northern Peru and southern Ecuador showing major rivers and localities mentioned in the text.

Eleutherodactylus in the Andes of Northern Peru

Fig. 2. Physiographic map of the Andes of northern Peru and southern Ecuador

Scientific Papers. Natural History Museum. The University of Kansas

-2000

1000

Meters 81
-4000

3000

80°

79

78°

Cordillera del Condor

77°

76°
4°10'S

Meters
-4000

-3000

2000

-1000

Cordillera de
Huancabamba

Cordillera Central

5°30' S

Meters
-4000

-3000

-2000

-1000

Cordillera
Occidental

Cordillera Central

6°10'S

Fig. 3. Profiles of the Andes in southern Ecuador and northern Peru. Vertical exaggeration 35X.

sin bring moisture to the eastern slopes of the mountain
ranges throughout the region. Generally rainfall is high-
est (+ 2500 mm annually) at elevations of 500-1500 m on
the windward slopes and lower at higher elevations and
on the leeward slopes. Deep valleys, such as those of the
Rio Maraiion and its tributaries in the Huancabamba De-
pression are in rain shadows and receive only 250-500 mm
of rain annually; most of this falls in October-May.

Influenced by the cold Humboldt Current, the winds
blowing off of the Pacific Ocean have relatively little mois-
ture; some of this moisture is dissipated as fog on the dry
western flanks of the Cordillera Occidental, where annual
precipitation may be less than 150 mm. However, as noted
by Foster and Beltran (1997), the relatively low passes in
the western Cordilleras permit the westerlies to provide

rainfall on the higher mountains to the east (e.g., Cordil-
lera del Condor and Cordillera de Huancabamba). Of
course, the former receives its greatest rainfall from the
winds off the Amazon Basin. Sporadically, rainfall is in-
creased along the Pacific lowlands and on the western
slopes of the Andes by the southward movement of the
warm waters of the counter-equatorial current. El Nifio.

Vegetation

The complex topography and associated climatic pat-
terns result in a great variety of vegetation formations.
Herein we use the terminology for vegetation formations
(and associated maps) in Peru proposed by Tosi (1960,
based on Holdridge's [1967] classification) and extend
these into Ecuador, as classified and mapped by Caiiadas
(1983). As noted by Savage (1975) and Lynch and Duellman
(1997), the Holdridge system seems to be too sophisticated

Eleutherodactylus in the Andes of Northern Peru

for determining broad patterns of animal distribution.
Therefore, \ve ha\e simplified the terminology by com-
bining some of the categories.

A transect from west to east across northern Peru at
approximately 0530' S. Lat. begins in coastal desert, as-
cends dry, rocky, slopes to humid highlands, drops to an
arid \'alley and repeats this sequence until descending into
the Amazonian lowlands. In making this transect, one
passes through dry scrub forest, dry montane forest, hu-
mid montane forest, and lowland rainforest. Each of the
major vegetation formations in northern Peru is discussed
briefly below; Tosi's (1960) terminology is in parentheses.

Desert scrub. — (Desierto subtropical y tropical; Maleza
desertica subtropical y tropical). Essentially, the entire Pa-
cific lowlands of Peru is extremely dry; with annual rain-
fall less than 150 mm, only a few xeric-adapted plants are
present. In the drier regions, vegetation may be absent or
consist only of a terrestrial gray Tilla)idsin. Farther inland
and close to the base of the Andes, vegetation becomes
more diverse with legumes (e.g., Prosopis jiiUflorn and
Cnp^paris sp.) and cacti (e.g., Cerents, Leniairocereiis, and
Opimtia.

Thorn forest. — (Bosque espinoza subtropical y tropi-
cal). In regions receiving up to 500 mm of rainfall annu-
ally, this type of forest develops on the western slopes of
the Cordillera Occidental to elevations of about 1200 m. It
also is prevalent in the interior valleys of the rios Chamaya,
Chinchipe, Huancabamba, Maranon, and Utcubamba, as
well as in the Catamayo Basin in Ecuador. Trees consist
principally of legumes (Prosopis and Acacia) with other
drought-resistant trees especially near streams — Bombax,
Bursera, Jacaranda, and Pithecolobiitm. Cacti (Ceretis, Opim-
iitt, and Lemairocereus) are numerous, especially in the
middle Maranon Valley, where dense cactus forest pre-
dominates. Both terrestrial and arboreal bromeliads (e.g.,
Tillandsia) are locally abundant. In many areas, especially
near rivers, there are irrigated fields of rice or sugar cane.

Dry forest. — (Bosque seco tropical; Bosque muy seco
tropical; bosque seco subtropical). Occurring peripheral
to the thorn forests in the interior river valleys and in the
rain shadow in the lower Rio Mayo Valley, there is a forest
that develops in areas receiving 500-1000 mm of rain an-
nually. This forest has no closed canopy and is made up of
moderate-sized trees of diverse genera (e.g., Bauhinia,
Bowbax, Bursera, Cordia, Centrolobium, Ciiratella, Iiiga,
Pithecolobium, Tabebuia). The epiphytic Spanish moss
(.Tillandsia iisneoides) is prevalent locally, and both terres-
trial and arboreal bromeliads (e.g., Pitcairuia and Tilland-
sia) are abundant locally. In the lower Rio Mayo Valley and
extending to the middle part of the Rio Huallaga, the dry
forest receives more rainfall (1000-2000 mm annually). In
this region, the forest contains several genera of trees (e.g..

Brosiiniini, Ccdrcla, M\/rox\/lon, Swietenia) characteristic of
humid tropical forest.

Montane dry forest. — (Bosque seco montanobajo). Iso-
lated patches of this forest occur at elevations of 2000-2500
m in the upper Rio Maraiion Valley, in the Cordillera Occi-
dental in the vicinity of Cajamarca and southward, in the
vicinity of Chachapoyas in the Cordillera Central, and in
the mountains south and west of Loja in Ecuador. Natural
vegetation consists of trees such as Jacaranda acutifolia,
Cacsalpinia tiiictoria, and various Acacia and Mimosa. Gen-
erally, these areas are heavily cultivated and have plantings
of Agave americana and Eucalytus globulus.

Humid Montane Forest. — (Bosque hiimedo montano).
This is the dominant type of forest on the lower slopes (up
to about 2500 m) of the cordilleras Huancabamba,
Tabacones, Condor, and Colan, where it is continuous to
the Cordillera Central. Principal trees include Berberis,
Polylepis, and Eugenia. At higher elevations (>3000 m) in
the Cordillera Occidental, this vegetation is reduced to few
trees, bushes (Baccliaris) and bunch grasses. Much of this
forest has been cleared and cultivated; at higher elevations
Eucalyptus has been planted.

Very humid montane forest. — (Bosque muy hiimedo
montano). This is the "ceja de la montafia" or "cloud for-
est" characteristic mostly of windward slopes at elevations
of 2500-3000 m and receiving 1000-2000 mm of rainfall
annually. This forest exists on the higher ridges of the cor-
dilleras Huancabamba, Tabacones, Condor, and Colan, as
well as on the slopes (especially eastern) of the Cordillera
Central in Peru and the Cordillera Occidental in Ecuador.
A variety of trees, often stunted and covered with lichens
and mosses, includes Polylepis and Podocarpus. Also charac-
teristic are bushes (Bacchnris) and the spiny bamboo (Chusquea
spicata). Arboreal bromeliads are locally abundant.

Humid subtropical forest. — (Bosque hiimedo y muy
hiimedo subtropical). Covering the lower (500-1900 m)
eastern slopes of the Cordillera Central and middle Rio
Mayo Valley, and lower slopes of the Cordillera Oriental
in Ecuador, this type of forest develops in areas receiving
as little as 1000 mm of rainfall annually to others that re-
ceive rainfall in excess of 3000 mm. The forest consists of a
variety of moderate to large trees including fuglans
neotropica, Cedrela fissipes, Tabebuia, and genera common in
the Amazonian lowlands — Brositnum, Cordia, Inga, Piper,
Swietenia. In some areas, the forest has been cleared for
citrus and coffee plantations.

Subtropical pluvial forest. — (Bosque pluvial subtropi-
cal). In northern Peru, this type of forest is known only on
the highest ridges of the Cordillera Central east of the Rio
Mayo Valley, where rainfall is expected to be in excess of
4000 mm annually. This type of forest consists of stunted
trees with emergent palms, such as Euterpe.

10

Scientific Papers, Natural History Museum, The University of Kansas

Humid tropical forest. — (Bosque humedo tropical).
This is the Amazonian rainforest, which receives rainfall
in excess of 2000 mm annually and has a great richness of
tree species. The trees form a continuous, or nearly so,
canopy 30^0 m above the ground, but there are canopy
emergents, such as species of Cedrela, Ceiba, and Ficus. This
diverse forest contains many kinds of palms (e.g., Bacfris,
Iriartea, Schedea), but under certain edaphic and hydrologi-
cal conditions, the forest is clearly dominated by one spe-
cies of palm, Mauritia reflexa.

Other formations. — Local climatic and edaphic condi-
tions may result in different local vegetation formations.
For example, the Abra de Zamora at 2800 m in the Cordil-

lera Oriental of Ecuador is windswept and has vegetation
consisting of thick growths of low bushes supporting
mosses and large bromeliads. Although Tosi (1960)
mapped the highest reaches of the Cordillera Colan as very
humid montane forest, data on a field tag of an
Eleiitherodactyhis collected by Thomas S. Schulenberg at
3300 m in that cordillera noted "in a grassy bog above
treeline." In the poorly explored Cordillera del Condor,
the flat-topped sandstone mountains at 2000-2300 m are
mostly are covered by sclerophyllous shrublands — shrubs
and small trees 2-5 m high; according to Foster and Beltran
(1997), these shrublands are composed mostly of species
of Ilex, Weiniiiniiiiin, Clusia, and Persea.

GENERA OF ELEUTHERODACTYLIINE FROGS

Four genera within the tribe Eleutherodactylini, as
defined by Lynch (1971) occur in the Andes of northern
Peru. Only species in the genus Eleuthewdactyhis are treated
herein. However, we provide brief diagnoses of external
features of all four genera presently recognized in the
Andes of northern Peru, so that specimens can be allocated
to the proper genus. There is no substantial evidence that
any of the genera are monophyletic, and Eleiithewdactylus
may be paraphyletic with respect to the other genera. For
example. Lynch (1986) questioned the allocation of spe-
cies to PIn/llonnstes or Phryiiopnis, and Harvey and Keck
(1995) noted the occurrence of "diagnostic" characters of
Ischnocnema in some species of Eleuthewdactyhis.

Eleuthewdactyhis Dumeril and Bibron, 1841

Diagnosis. — Terrestrial or arboreal frogs characterized
by (1 ) skin on venter smooth or areolate; (2) relative lengths
of Fingers I and 11 variable; (3) relative lengths of Toes 111
and V variable; (4) digits bearing terminal discs and pads;
(5) tips of digits rounded, elliptical, or truncate; (6) tarsal
tubercle present or absent; (7) palmar and subarticular tu-
bercles not greatly enlarged; (8) adult size 10-120 mm SVL.

Content. — About 600 species in five subgenera, only
one of which {Eleuthewdactyhis) includes the 45 species
known from the Andes of northern Ecuador.

Distribution. — Tropical and subtropical America (ex-
clusive of arid regions), through the West Indies and into
southwestern United States.

Remarks. — The 45 species known from the Andes of
northern Peru are members of four species groups, which
are defined in the following Accounts of Species.

Ischnocnema Reinhardt and Liitken, 1862

mar and subarticular tubercles greatly enlarged; (8) adult
size 30-55 mm SVL.

Content. — Five species, of which only Ischuocuema
saxatilis and /. swimonsi are in the Andes of northern Ecua-
dor (Duellman, 1990b; Lynch, 1974a).

Distribution. — Upper Amazon Basin, Andean slopes
in southern Ecuador, northern Peni, and Bolivia.

Remarks. — Harvey and Keck (1995) questioned the
validity oi Ischnocnema.^

Phrynopus Peters, 1874

Diagnosis. — Terrestrial frogs characterized by (1) skin
on venter smooth or areolate; (2) relative lengths of Fin-
gers I and 11 variable; (3) Toe III longer than, or equal in
length to. Toe V; (4) digits not bearing terminal discs or
pads; (5) tips of digits rounded; (6) tarsal tubercle absent;

(7) palmar and subarticular tubercles not greatly enlarged;

(8) adult size 18-45 mm SVL.

Content. — More than 20 species, of which three have
been reported from the Andes of northern Peru; descrip-
tions of three others are being written (Duellman, in prep.).

Distribution. — Humid habitats at moderate to high
elevations in the Andes from Colombia to Bolivia. In the
Andes of northern Peru, Phrynopus nebulanastes and P.
parkeri are endemic to the Cordillera de Huancabamba, and
P. siiuousii is widespread in the northern part of the Cor-
dillera Occidental (Cannatella, 1984; Lynch, 1975a); one of
the undescribed species is from the northern part of the
Cordillera Central, and two are from the Cordillera Occi-
dental.

Diagnosis. — Terrestrial frogs characterized by (1) skin
on venter smooth; (2) Finger I > II; (3) Toe V > 111; (4) digits

'It is doubtful that the Amazonian Ischnoaiciim qiiixciisis and the three
Andean species are related to the type species, /. vcnucofii^, in .southeast-
, . 11 , • 1 J. 1 , 1 ,r-\ ern Brazil. If that should be shown to be true, Ori'd/iflte Jimene/ de la

bearmg small termmal discs; pads present or absent; (5) Espada, 1872 (type species by monotypy O. qmxcmi,) is an available ge-
tips of digits rounded; (6) tarsal tubercle absent; (7) pal- neric name.

Eleutherodactylus in the Andes of Northern Peru

U

Remarks. — In reviewing the osteology of Phryjwpu^,
Phi/Uoimstcs, and other genera, Lynch (1986) noted that the
Phryuopiis pcniviamis group and PIn/llonastcs may be re-
lated.

PhyUouastes Heyer, 1977

Diagnosis. — Terrestrial frogs characterized by (1) skin
on venter smooth; (2) Finger I shorter than or equal in
length to Finger 11; (3) Toe III shorter than Toe V; (4) digits
not bearing terminal discs or pads; (5) tips of at least Toes
III and IV pointed; (6) prominent tarsal tubercle present;

(7) palmar and subarticular tubercles not greatly enlarged;

(8) adult size 13-20 mm SVL.

SUMMARY OF TAXONOMIC CHARACTERS

Content. — Four species, two of which occur in the
Andes of northern Peru.

Distribution. — Pliyllonastes nn/niiecoidcs inhabits the
upper Amazon Basin, whereas three montane species have
been reported from only a few localities — P. heyeri from
Alamor, Ecuador, and the Cordillera de Huancabamba in
Peru (Lynch, 1986)-; P. locliitcs from the Cordillera del Con-
dor in Ecuador (Lynch, 1976); and P. lyiiclii from the north-
ern part of the Cordillera Central in Peru (Duellman,
1991b).

Remarks. — See Remarks under Phrynopits.

Morphological characters, size, and color pattern of
members of this genus in western Ecuador were described
in detail and illustrated by Lynch and Duellman (1997).
Consequently, we treat the characters rather briefly herein;
22 morphological characters and eight features of color
pattern for species of Eleutherodactylus in the Andes of
northern Peru are tabulated in Tables 1 and 2. As deemed
necessary, we attempt to clarify some of these characters
below.

Skin texture. — Although statements in the diagnoses
and descriptions may suggest that various states are dis-
crete, they are not in many cases. For example, the distinc-
tion between the dorsal skin being shagreen with scattered
tubercles versus finely tuberculate is not discrete, and cer-
tainly different persons may describe the conditions in
different ways. The ventral skin commonly has been de-
scribed as granular; we use the term areolate. In contrast,
fine granules on the dorsum give the skin a finely textured
appearance that we term shagreen. If the granules are large
and subcorneal or conical, we define the texture as tuber-
culate.

Dorsolateral folds on the body are evident in seven
species of the Eleutherodactylus couspicillatus Group (E.
avicuporum, citriogaster, condor, knrcliarias, lymani, and
peruvianus) and in E. araiodactylus. Of these, £. avicuporum
and E. karcharias also have interrupted, longitudinal folds
on the flanks. In the Andes of northern Peru, E. avicuporum
is unique in having a dermal interocular ridge. This fea-
ture also is present in another member of the
Eleutherodactylus couspicillatus Group, £. skydmaiiios (Flores
and Rodriguez, 1997) and in some other species of the (e.g.,
E. quinquagesimus [Lynch and Duellman, 1997]).

Tubercles. — The most noticeable and taxonomically
useful tubercles are those on the upper eyelid and heels.
We divege from Lynch and Duellman (1997) in the recog-
nition of eyelid tubercles; they used the term only for the
presence of enlarged conical or subconical tubercles on the

eyelid. No species in the Andes of northern Peru possess
such distinctive tubercles, but many species have a few
pungent tubercles on the upper eyelid. If a tubercle is
present on the heel, it usually is small, but four species
(Eleutherodactylus galdi, lanthanites, muscosus, and
quaquaversus) have large, conical tubercles on the heel.

Tuberculation on the tarsus is highly variable. Tu-
bercles are absent in several species (e.g., Eleutherodactylus
citriogaster, colodactylus,proserpens, and rufioculis). Tubercles
are present on the outer surface of the tarsus in many spe-
cies (e.g., E. auemerus, bearsei, schultei, and versicolor); the
outer tarsal tubercles are noticeably conical in E. galdi. The
inner edge of the tarsus may be unadorned (e.g., E, condor,
iufraguttatus, percnopterus, and phoxocephalus) , bear a low,
elliptical tubercle distally (e.g., E. ceuthospilus, cuneirostris,
uephophilus, and petrohardus) , or have a low fold distally
(e.g., E. atrabracus, avicuporum, melanogaster, and
rhodoplichus).

A row of tubercles may be present on the ventrolat-
eral surface of the forearm. These ulnar tubercles are ab-
sent in several species (e.g., Eleutherodactylus bromeliaceus,
incomptus, lymani, and phoxocephalus). The tubercles are
especially prominent in E. cri/ptomelas and E. iufraguttatus,
conical in £. galdi, and coalesced to form a short fold in E.
pinguis.

Two species (Eleutherodactylus anemerus and £.
proserpeus) have a noticeable tubercle (papilla) on the tip
of the snout, and E. phoxocephalus has a low vertical keel
on the snout. Eleutherodactylus karcharias is unique in hav-
ing a finlike middorsal tubercle on the body.

Tympanum. — As emphasized by Lynch and Duellman
(1997:28): "The terms 'tympanum' and 'external ear' have
been used to identify a combination of characters rather
than a single character. The 'tympanum' is a combination

=A specimen (LSUMZ 39364) from the Cordillera Colan extends the known
range of this species eastward into Departamento Amazonas, Peru.

12

Scientific Papers, Natural History Museum, The University of Kansas

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Elevtherodactyiais in the Andes of Northern Peru

13

of (1) a differentiated tympanic membrane and (2) a tym-
panic annulus." Thev recognized four states of tympanic
annuli and membranes. In most Eleiitherodactiflus in the
Andes of northern Peru, a prominent tympanic membrane
and annukis are present. The membrane is not differenti-
ated but most of the annulus is yisible beneath the skin in
£. imianos^aster, pataikof, and rufioculis, whereas only the
ventral part of the annulus is visible below the skin in E.
acmiiiimtiis, ardalonychus, qiiaquaversus, and unensi. There
is no external evidence of an ear in E. colodactyliis and £.
UreUiif.

Fingers and toes. — In all members of the
Eleiitherodactylus orestes and unistrigatus groups in the
Andes of northern Peru, the first finger (thumb) is shorter
than the second. The first finger is longer than the second
in most species in the Eleuthewdnctylus conspicillatiis Group,
but the two digits are about ecjual in length in E. ciincirostris
and E. knrcharias, as well as in E. araiodactylus in the
Eleiitherodactylus nigrovittatus Group. As shown by Lynch
and Duellman (1979), the relative lengths of Toes 111-lV is
an important taxonomic character. Except for E.
araiodactylus, in which Toes 111 and V are of equal length.
Toe V is longer than Toe III in all species in the region. In
members of the Eleutherodactylus conspicillatiis and orestes
groups. Toe V is only slightly longer than Toe III, and nei-
ther toe extends to the level of the distal subarticular tu-
bercle on Toe IV. In species in the Eleutherodactylus
unistrigatus Group, Toe V is noticeably longer than Toe 111
and extends to the middle or distal border of the distal
subarticular tubercle of Toe IV.

With the exception of species in the Eleutherodactylus
nigrovittatus and orestes groups, in which the discs on the
fingers and toes are barely expanded, the discs on Fingers
1 and 11 and all toes are expanded. Their shapes are de-
scribed arbitrarily as rounded, elliptical (much wider and
long), or truncate.

Most species have lateral fringes on the digits. We do
not distinguish between fringes (thin) and keels (thick).
Webbing between the toes is absent in all species, except
three members of the E. conspicillatiis group (E. avicupwrum,
karcharias, and metahates) in which basal webbing is present.

Coloration. — The dorsal coloration can be used readily
for identification of most species of frogs — not so for most
species of Eleutherodactylus, in which distinctive coloration
may be present only in the groin or on the anterior and /or
posterior surfaces of the thighs (Table 2). In northern Peru,
only two species are easily identified by their dorsal col-
oration — uniform green (in life) in £. acuminatus and uni-
form orange-red (in life) in E. anemerus. In preservative,
the dorsal coloration of most species is varying shades of
tan, brown, or gray, usually with darker markings. These
may consist of distinct spots (e.g., E. cuneirostris), chev-

rons (V-shaped marks with tlie apex anteriad) (e.g., E.
lauthanitcs and £. lymani), or a W-, X-, or H-shaped mark in
the scapular region (e.g., £. ardalonychus, exoristus, and
steniothi/lax). Other dorsal patterns include dark longitu-
dinal streaks or dashes (e.g., £. ceuthospilus, galdi, and
percnopterus), a large middorsal blotch (e.g., some £.
proserpens) , and pale vermiculations (E. muscosus). Identi-
fication is confused further by pattern polymorphism is
several species. For example, some individuals of £.
colodactyliis, proserpens, rhodopUclius, and sternotlnjlax have
pale dorsolateral stripes, whereas some individuals of £.
schultei and E. unensi have dark dorsolateral stripes. Some
specimens of £. bearsei and E. petrobardus have pale spots
on the dorsum.

Facial markings usually consist of dark labial bars, but
these are absent in some species (Table 2). Eleutherodactylus
cuneirostris and some specimens of E. peruvianus have a
pale labial stripe. A dark canthal stripe is evident in most
species and is especially notable in E. acuminatus and £.
galdi. The canthal stripe is incorporated into a dark face
mask encompassing the loreal region in E. peruvianus.

The coloration on the venter in most species is cream
(with or without dark flecks) to brown. However, distinc-
tive coloration is present in a few species — bright yellow
(in life) in Eleutherodactylus citriogaster, uniform black in E.
melanogaster, black on the ventral surfaces of the hind limbs
in E. atrnbracus, and brown or black marbling or reticula-
tion (e.g., E. ardalonychus, infraguttatus, muscosus, and ver-
sicolor).

Whereas the groin is colored like the flanks in most
Eleutherodactylus, distinctive markings are present in some
species. Commonly these are pale spots, which are white,
yellow, or red in life; such spots are characteristic of most
species in the Eleutherodactylus orestes Group, but also ex-
ist in E. cajamarcensis, condor, lirellus, muscosus, and rufioculis.
The groin has dark reticulation in E. phoxocephalus and is
black in £. cryptomelas.

In preserved specimens of Eleutherodactylus, the pos-
terior surfaces of the thighs usually are varying shades of
tan to brown, with or without pale flecks or spots that are
cream to red in life (Table 2). Notable exceptions are dark
reticulations (£. phoxocephalus, quaquaversus, and £.
rhodopUclius), pale mottling (£. ardalonychus, citriogaster, and
infi-aguttatiis), or black with white spots (£. lymani).

Sizes and proportions. — The sizes of adults of
Eleutherodactylus are highly variable. With the exception
of some species in the Eleutherodactylus conspicillatiis Group,
most species in the Andes of northern Peru have SVLs of
less than 35 mm (Table 3). Considerable sexual dimorphism
in size is evident in most species of Eleutherodactylus, in
which females usually are at least one third larger than
males. In the species in the Andes of northern Peru, SVLs

14

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Scientific Papers, Natural History Museum, The University of Kansas

Table 3. Body sizes (SVL in mm) of Elentherodactylus in the Andes of
northern Peru. Values include range (mean, sample size), SD = sexual
dimorphism. Unless noted otherwise, measurements are for Peruvian
specimens only.

Species

Males

Females

E. acuminatus
E. anemerus
E. araiodactylus
E. ardalonychus
E. atrabracus
E. avicuporum
E. bearsei
£. bromeliaceus'
E. cajamarcensis
. ceuthospilus
. citriognster
. colodadylus
. condor-
. cryptomelas^

ciineirostris

exoristus

galdi^

incomptus''

infraguttatus

karclwrias

lanthanites"

lirellus

lymani

melanogaster

metabates

muscosits

nephophilus

E. ockendenf

E. pataikos

E. peck?

E. percnopterus
peruvianus^
petrobnrdus
phoxocephalus
pinguis
proserpens''
quaquaversus"
rhodoplichus
rhodostichus
rufiocuUs
schultci
serendipituf
sternothytax
versicolor

E. wiensi

21.3-21.5(21.4,2)
20.4

19.7-21.9(20.8,2)
18.9
25.2

22.7-25.5 (24.0, 3)
16.7-23.2 (20.8, 14)

22.6-25.9 (24.7, 5)
19.9-25.8(21.3,35)

31.6-41.3(37.0,9)
16.8-19.6(18.3,19)

32.1-39.5 (36.5, 9)

28.2-30.2 (29.2, 4)

15.0-16.9(16.2,4)
17.1-24.8(21.0,8)
15.6-18.8(17.5,10)
15.8

21.7-26.0(23.5,20)

14.1-17.0(15.3,13)

39.7-45.3(41.9,4)

20.6-22.8(21.9,3)

29.9-32.2(31.1,2)

16.9-21.2(19.1,3)

15.3-18.7(17.3,3)
21.5-23.2(22.5,3)
26.1-30.0,28.1,8)
27.0-30.7 (28.5, 7)
22.7-27.8 (26.2, 5)

15.2-21.0(18.6,11)

19.6-22.5 (20.7, 29)

21.8-28.9 (25.8, 17)

19.3

18.1

23.5-26.5 (25.0, 10)

20.4-21.2(20.8,2)

18.3-29.1 (24.2, 39)

21.8-24.7(23.3,2)

27.8-33.0 (30.7, 6)

SD

24.3 1.14

24.5 —

27.4 1.32
22.7 1.20

31.0-34.3(32.4,6) 1.29

38.0-38.8 (38.4), 2) 1.60

22,9-28.1(26.5,5) 1.27

27.0-32.0(29.2,14) 1.18

23.5-26.7(25.2,8) 1.18

42.0-51.0(44.4,4) 1.20

19.1-23.2(21.5,9) 1.17

52.6-63.2(58.7,3) 1.61

38.6 1.32

29.1 —
21.3-23.5(22.7,6) 1.40

28.1-34.0(31,7,11) 1.51

23.7-25.9(24.5,7) 1.40

22.9-24.3(23.6,3) 1.49

30.4 —

27.5^4.8(36.2,32) 1.54

19.4-24.0(21.5,12) 1.39

48.4-66.7(58.7,4) 1.40

24.2-24.7(24.5,2) 1.12

29.6^6.1 (37.8, 4) —

24.6-34.0 (29.8, 5) —

24.6-31.5(28,0,12) 1,47

21,6 —

25.2 1.46
25.9 1.15

36,0^2,5(39.1,9) 1.39

38.9 1.48

28.4-29.8 (28.9, 3) —

20.2-23.5(22.0,4) 1.18

24.6-31.3(27.3,20) 1.32

30.1-34.2(32.5,4) 1.26

20.6 1.14

28.4-34.0(31.8,5) 1.27

28.3-36,7(32.1,6) 1,33

26,0-27,3(26,5,3) 1,14

37,0 1,21

'Provincia Morona-Santiago, Ecuador (Lynch, 1979)
-Including specimens from Provincia Morona-Santiago, Ecuador (Lynch
and Duellman, 1980)

Trovincias Loja and Santiago-Zamora, Ecuador (Lynch, 1979)
■■Cordillera Oriental, Ecuador, and Cordillera del Condor (Lynch and
Duellman, 1980)

'Provincia Napo, Ecuador (Lynch and Duellman, 1980)
'Upper Amazon Basin (Lynch, 1980)
'Amazonian Peru (Lynch, 1980)

'Including specimens from the Cordillera de Cu tucii, Ecuador ( Duellman
and Lynch, 1988)

'Cordillera Oriental, Ecuador (Lynch. 1979)
'"Cordillera Oriental, Ecuador (Lynch and Duellman, 1980)

of females are 1,12-f .61 (x = 1.25) greater than those of
males.

The three groups of Eleutherodactylus in the Andes of
northern Peru have distinctly different proportional hind-
limb lengths (Fig. 4). In members of the Eleuthewdactijlus
Orestes Group, the mean tibia/SVL ratio is 37.0-39.6 (x =
38.4, II = 4), whereas members of the Eleutherodactylus
conspicdlatus Group have much longer hind limbs; the
mean tibia/SVL ratio is 60.4-65.6 (x= 62.2, n = 9). Ratios
among species in the Eleutherodactylus unistrigatus Group
are intermediate — 43.3-56.0 (x = 49.4, n = 31). The lowest
ratios (43.3 and 44.1) are for two bromeliad-d welling spe-
cies, E. colodactylus and £. proserpens; elimination of those
two species gives a range of 47.5 (£. phoxoceplialus, also a
bromeliad-dweller), to 56.0 (£. rufiocuUs and E. zvieiisi) with
a mean of 49.9.

35

30

25

20

15

10

5
25

E 20

E

15

0!

^ 10

■ E. ctinspicilliiuts group ■

▼ E. iiigroviiuinis group

A E. orestes group

9 E. iinisrrif;tiliis grcuip

15

20

25 30 35 40 45

Snout-vent Length (mm)

Fig. 4. Correlations of tibia length (A) mmS head width (B) with
snout-vent length in Elcutlicrodnclyliis in the Andes of northern Peru.
Symbols for the species groups are the same in both graphs.

ELEUTHEKODACnLUS IN THE AnDES Ol- NORTHERN PeRU

17

IDENTIFICATION OF SPECIES

Key to the Species
This key emphasizes those external features usually
independent of maturity and sex; however, juveniles are
always difficult to identify, because the distinctness of the
tympanic membrane and annulus increases with age, and
many features of color patterns on the venter and concealed
surfaces of the limbs develop with age. This key is only a
guide; it is necessary to confirm the identifications by re-
ferring to the diagnoses and descriptions.

1. Toe V longer than Toe III 2

Toes 111 and V equal in length £, araiodactylus

2. Toe V only slightly longer than Toe III 3

Toe V much longer than Toe III 15

3. Finger I longer than, or equal in length, to Finger II;

outer fingers with greatly expanded discs 4

Finger I shorter than Finger II; outer fingers with nar-
row, rounded discs 12

4. Skin on belly areolate 5

Skin on belly smooth 6

5. Fin-shaped middorsal tubercle present; interocular

fold present E. karcharias

Fin-shaped middorsal tubercle absent; interocular fold
usually present, weak E. aviciiponiiii

6. Toes webbed basally £. metabates

Toes unwebbed 7

7. Prominent conical tubercle on heel; throat dark with

median white stripe £. lanthanites

Heel lacking tubercle; no white stripe on throat 8

8. Snout long, wedge shaped; broad, pale labial stripe

present; dorsum with many small dark spots

E. cuueirostris

Snout not wedge-shaped; labial region usually barred;
dorsal pattern not consisting of many small dark spots
9

9. Posterior surfaces of thighs dark with pale flecks .. 10
Posterior surfaces of thighs with dark and pale mot-
tling 11

10. Throat dark; belly and ventral surfaces of hind limbs

with dark mottling E. condor

Throat pale; dark flecks or reticulations on throat and
chest £. peruvianus

11. Posterior surfaces of thighs black with bold cream

mottling; dark marks on margin of lower jaw

E. lymani

Posterior surfaces of thighs brown with tan mottling;
venter pale (yellow in life) with diffuse brown spots
on throat £. citriogaster

12. Dorsum and venter black £. melanogaster

13

15

16

Dorsum brown; venter variable 13

Ventral surfaces of hind limbs black E. ntrnbracus

Ventral surfaces of hind limbs cream or tan 114

14. Large pale spot in groin; posterior surfaces of thighs

cream with brown marks £. pinguis

No pale spot in groin; posterior surfaces of thighs uni-
form brown £. pataikos

Tympanic membrane not differentiated 16

Tympanic membrane distinct with well-defined tym-
panic annulus 22

Tympanic annulus absent 17

Tympanic annulus visible beneath skin or evident only
ventrally 18

1 7. Fingers short, stubby, with round discs; labial and limb

bars absent; no pale spot in groin £. colodncfylus

Fingers long, slender, with elliptical discs; labial and
limb bars present; pale spot in groin £. lirellus

18. Tubercle present on heel 19

Tubercle absent on heel 20

19. Tubercle on heel large, conical; flanks uniform tan;

venter white £. quaqiuwcrsus

Tubercle on heel small, rounded; flanks cream with
dark mottling; venter pale with irregular brown spots
£. wiensi

20. Snout acuminate; dorsum pale (green in life); only
markings are black canthal and supratympanic stripes
£. acumi}iatiis

Snout rounded; dorsal markings consisting of at least
interorbital bar, transverse bars on limbs, and dark
markings on body 21

21. Flanks with diagonal dark bars; venter cream with

brown reticulations £. ardalonychus

Flanks dark with pale spots; venter densely flecked

with brown £. rufioculis

Tubercle present on heel 23

Tubercle absent on heel 32

Tubercle on heel large, conical 24

Tubercle on heel small, not conical 25

24. Snout acuminate in dorsal view; row of conical tu-
bercles on outer edge of tarsus; flanks uniform cream

E. galdi

Snout bluntly rounded in dorsal view; one or two small
tubercles on outer edge of tarsus; flanks brown with
white spots E. muscosus

25. Upper eyelid lacking tubercles E. pctrobardus

Upper eyelid bearing tubercles 26

26. Snout rounded in dorsal view; posterior surfaces of

22,

23

18

Scientific Papers, Natural History Museum, The University of Kansas

thighs brown with cream bars £. nephophilus

Snout subacuminate in dorsal view; posterior surfaces
of thighs not brown with cream bars 27

27. Groin and posterior surfaces of thighs black

E. cryptomelas

Groin and posterior surfaces of thighs not black.... 28

28. Flanks pale with darker markings 29

Flanks uniform tan or brown 30

29. Flanks with diagonal bars or reticulations; posterior

surfaces of thighs with dark reticulations

£. rhodoplichiis

Flanks with dark streaks; posterior surfaces of thighs
uniformly pale £. schultei

30. Posterior surfaces of thighs brown with cream spots
E. bromeliaceus

Posterior surfaces of thighs uniform brown 31

31 . Dorsal markings consisting of dark scapular W or chev-
ron; venter white with brown spots E, ockendeni

Dorsal markings consisting of dark spots and streaks;
venter densely flecked with gray E. pecki

32. Prominent tubercle on tip of snout 33

No tubercle on tip of snout 34

33. Dorsum uniformly pale (orange-red in life)

E. anemerus

Large middorsal blotch or X-shaped mark, interorbital
and limb bars present E. pivserpeiis

34. Posterior surfaces of thighs brown or black with pale

markings 35

Posterior surfaces of thighs uniform tan or brown with
or without darker markings 39

35. Posterior surfaces of thighs and groin black with white

spots E. cajamarcensis

Posterior surfaces of thighs brown with cream mark-
ings; groin unmarked 36

36. Posterior surfaces of thighs brown with cream mot-
tling; flanks tan with brown spots posteriorly

E. infrnguttattis

Posterior surfaces of thighs brown with cream spots
or flecks; flanks not so marked 37

37. Flanks uniform tan E. ceittliospiliis

Flanks tan or cream with diagonal or vertical dark bars
38

38. Dorsal markings consisting of scapular H- or W-shaped
mark and chevrons; venter tan with brown flecks

E. cxoristiis

Dorsal markings consisting of dark blotches; venter
dull white with brown or black reticulations

E. versicolor

39. Posterior surfaces of thighs tan with dark brown re-

ticulations; vertical keel on snout E. phoxocephahis

Posterior surfaces of thighs not so colored; no vertical
keel on snout 40

40. Flanks uniform tan or brown 41

Flanks with pale or dark markings 42

41. Venter tan with small black flecks; limb bars diagonal

E. percnopterus

Venter cream with brown flecks; limb bars transverse
E. screudipitus

42. Flanks brown with cream flecks E. hearsei

Flanks not brown with cream flecks 43

43. Flanks with small dark spots; eyelid tubercles present;

snout acuminate in dorsal view E. rhodosticliiis

Hanks not having small dark spots; eyelid tubercles
absent; snout not acuminate in dorsal view 44

44. Throat and belly uniform brown E. incornptus

Throat and belly cream with brown flecks

E. sternothylax

Clave de las Especies

Esta clave pone entasis en los rasgos externos
usualmente independientementes de la madurez y el sexo;
sin embargo, los juveniles son siempre dificiles de
identificar debido a que la membrana y el anillo timpanicos
se hacen mas visibles con la edad, y muchos rasgos del
patron de coloracion del vientre y superficies ocultas de
los miembros se desarrollan con la edad. Esta clave es
solamente una guia; es necesario confirmar las
identificaciones consultando las diagnosis y descripciones.

1. Dedo V del piemas largo que Dedo III 2

Dedos 111 y V del pie iquales en largo E. nraiodactylus

2. Dedo V del pie solamente un poco mas largo que Dedo

111 3

Dedo V del pie mucho mas largo que Dedo 111 15

3. Dedo 1 de la mano mas largo que, o igual de largo que
Dedo 11; dedos externos con discos muy dilitados ... 4
Dedo 1 de la mano mas corta que Dedo II; dedos
externos con discos angostos, redondeados 12

4. Piel del vientre areolada 5

Piel del vientre Hsa 6

5. Tuberculo dorso medial en forma de aleta presente;

pliegue interocular presente £. karcharias

Tuberculo dorso medial ausente; pliegue interocular
usualmente presente, incospicuo £. aviciiponiiii

6. Dedos posteriores con palmeadura basal £. metabntcs
Dedos posteriores sin palmeadura 7

7. Tuberculo conico prominente en el takin; garganta con

raya blanca mediana £. Innthaiiitca

Sin tuberculo en el talon; sin raya blanca en la garanta
8

Eleutherodactylus in the Andes of Northern Peru

19

8. Rostro largo, en forma de cuna; rnya labial ancha,
palida presente; dorso con muchas manchas oscuras

pequenas £• ciiiicimstris

Rostro no en forma de cuiia; region labial usualmente
con barras; patron dorsal no compuesto de muchas
manchas oscuras pequefias 9

9. Superficies posteriores de los muslos oscuras con

puntillos palidos 10

Superficies posteriores de los muslos con monteado
oscuro V palido 11

10. Garganta oscura; vientre del cuerpo y los miembros

posteriores con monteado oscuro E. condor

Garganta palida; puntillos o reticulaciones oscuras en
la garanta y el pecho E. peruviamis

11. Superficies posteriores de los muslos negras con
monteado crema; marcas oscuras en el margen de la

mandibula E. h/mani

Superficies posteriores de los muslos cafe oscuro con
monteado cafe palido; vientre palido (amarillo en vida)
con manchas difusas de cafe oscuro en la garganta ...
£. citriogaster

12. Dorso y vientre negro £. melanogaster

Dorso cafe; vientre variable 13

13. Superficie ventral de los miembros posteriores negra

E. atmbracus

Superficie ventral de los miembros posteriores crema
o cafe claro 14

14. Mancha palida grande en la ingle; superficies

posteriores de los muslos crema con marcas cafe

E. pinguis

Sin mancha palida en la ingle; superficies posteriores
de los muslos uniformemente cafe £. pataikos

15. Membrana timpanica no diferenciada 16

Membrana timpanica distintiva con anillo timpanico
bien defindo 22

16. Anillo timpanico ausente 17

Anillo timpanico visible debajo de la piel o evidente
solamente ventralmente 18

17. Dedos d la mano cortos y tiesos, con discos redondos;
barras de los labios y miembros ausentes; sin mancha

palida en la ingle £. colodnctybis

Dedos de la mano largos, delgados, con discos
elipticos; barras de los labios y miembros presentes;
mancha palida en la ingle £. lirellus

18. Tuberculo presente en el talon 19

Tuberculo ausente en el talon 20

19. Tuberculo en el talon grande, conico; flancos
uniformemente cafe claro; vientre bianco

£. qiiaqunversus

Tuberculo en el talon pequefio, redondeado; flancos

crema con monteado oscuro; vientre palido con
manchas cafe irregulares £. wiensi

20. Rostro acuminado; dorso palido (verde en vida); las
linicas marcas son rayas canthales y supratimpanicas

£. acumhiatiis

Rostro redondeado; marcas dorsales compuestas al
menos de una barra interorbital, barras transversales
en los miembros y marcas oscuras en el cuerpo 21

21. Flancos con barras oscuras diagonales; vientre crema

con reticulaciones cafe £. nrdnlom/chiis

Flancos oscuros con manchas palidas; vientre
densamente con puntillado cafe £. nifioculis

22. Tuberculo presente en el talon 23

Tuberculo ausente en el talon 32

23. Tuberculo en el talon grande, conico 24

Tuberculo en el talon pequefio, no conico 25

24. Rostro acuminado en vista dorsal; fila de tuberculos
conicos en el borde externo del tarso; flancos

uniformemente crema E. galdi

Rostro redondeado sin filo en vista dorsal; uno o dos
tuberculos pequefios en el border externo del tarso;
flancos cafe con manchas blancas £. muscosus

25. Parpado superior sin tuberculos E. petrobardus

Parpado superior con tuberculos 26

26. Rostro redondeado en vista dorsal; superficies

posteriores de los muslos cafe con barras crema

E. nephophilus

Rostro subacuminado en vista dorsal; superficies
posteriores de los muslos diferente de cafe con barras
crema 27

27. Ingle y superficies posteriores de los muslos color

negro £; cryptomelas

Ingle y superficies posteriores de los muslos distinto
de negro 28

28. Flancos palidos con marcas mas oscuras 29

Flancos uniformemente cafe claro o cafe oscuro .... 30

29. Flancos con barras diagonales o reticulaciones; super-
ficies posteriores de los muslos con reticulaciones

oscuras £. rhodopUchus

Flancos con rayitos oscuras; superficies posteriores de
los muslos uniformemente palido £. schidtei

30. Superficies posteriores de los muslos cafe con manchas

cremas £• bromelinceiis

Superficies posteriores de los muslos uniformemente
cafe 31

31. Marcas dorsales compuestas de una W en la region
escapular o chevron; vientre bianco con manchas cafe

E. ockendeni

Marcas dorsales compuestas de manchas y rayitas
oscuras; vientre con puntillado gris densamente

£. pecki

20

Scientific Papers, Natural History Museum, The University of Kansas

32. Tuberculo prominente en el extremo del rostro 33

Sin tuberculo en el extremo del rostro 34

33. Dorso uniformemente palido (naranja-rojo en vida) .

£. aiiemerus

Mancha grande mediodorsal o marca en forma de X;
barras interorbital y en los miembros presentes

£. proserpens

34. Superficies posteriores de los muslos cafe o negro con

marcas palidas 35

Superficies posteriores de los muslos uniformemente
cafe claro o cafe oscuro con o sin marcas mas oscuras
39

35. Superficies posteriores de los muslos y la ingle negras

con manchas blancas £. cajamarcensis

Superficies posteriores de los muslos cafe con marcas
crema; ingle sin marcas 36

36. Superficies posteriores de los muslos cafe monteado
con crema; flancos cafe claro con manchas cafe oscuras

posteriores £. iiifraguttatiis

Superficies posteriores de los muslos cafe con manchas
o puntillos crema; flancos con patron diferente 37

37. Flancos uniformemente cafe claro £. ceiithospilus

Flancos cafe claro o crema con barras diagonales o
verticales 38

38. Marcas dorsales en forma de H o W in la region
escapular o chevrones; vientre cafe claro con puntillos
cafe oscuro £. exoristus

Marcas dorsales compuestas de manchas oscuras;
vientre bianco mate con reticulaciones cafe o negroE.
versicolor

39. Superficies posteriores de los muslos cafe claro con
reticulaciones cafe oscuro; quilla vertical en el extremo

del rostro £. phoxoceplmliis

Superficies posteriores de los muslos con otro patron;
sin quilla vertical en el extremo del rostro 40

40. Flancos uniformemente cafe claro o cafe oscuro .... 41
Rancos con marcas palidas u oscuras 42

41. Vientre cafe claro con puntillos negros pecjuefios;

barras diagonales en los miembros £. pcrcnoptcrus

Vientre crema con puntillos cafe oscuro; barras
transversales en los miembros £. seredipitus

42. Flancos cafe oscuro con puntillos crema £. hcarsci

Flancos distinto de cafe oscuro con puntillos crema ..
43

43. Flancos con manchas oscuras pec[uenas; parpados
superiores con tuberculos; rostro acuminado en vista

dorsal £. rhodostichus

Flancos sin manchas oscuras; parpados superiores sin
tuberculos; rostro no acuminado en vista dorsal .... 44

44. Garganta y vientre uniformemente cafe oscuro

£. incomptiis

Garganta y vientre crema con puntillos cafe oscuro ..
£. stemothylax

SPECIES ACCOUNTS

Eleutherodactylus conspicillatus Group

The inclusion of several new species necessitates a
slight modification of the definition of this group as given
by Lynch and Duellman (1997). Most species have smooth
skin on the venter, but the skin is areolate in £. avicuporum,
caprifcr, and knrchnrias. Finger 1 is longer than Finger 11 in
most species, but the first two fingers are equal in length
in £. cuneirostris and E. karcharias. The toes are unwebbed
in most species, but basal webbing is present in £.
avicuporum, malkini, meiabates, and karcharias. Moreover,
two species, £. citriogaster and £, zeuctoiylus, apparently
are unique in Ekutherodaciylus by possessing round, in-
stead of bifid, palmar tubecrles.

Lynch and Duellman reported 29 species in the group.
This number is increased by five species in Peru — £.
karcharias and £. skydmainos (Flores and Rodriguez, 1997)
and three species described herein. Nine members of the
group are known from low to moderate elevations in the
Andes of northern Peru.

Ekutherodaciylus avicuporum new species

Holotype.— LSUMZ 39365, an adult female, from 1 2 km
[by trail] E La Peca (05°36' S, 78i9' W, 1700 m), Provincia

Bagua, Departamento Amazonas, Peru, one of a series col-
lected by L. J. Barkley on 11 June 1978.

Paratypes.— LSUMZ 39361 from 2030 m and KU
288628, LSUMZ 39367-68, 39371-72 from 1700 m, one adult
male and five adult females, all from the western slope of
the Cordillera Colan east of La Peca.

Referred specimens.— LSUMZ 39359, 39366, 39374-75,
45089, juveniles, and LSUMZ 39370, a partially smashed
adult female.

Diagnosis. — A member of the Eleutherodactylus
couspicillatus Group having (1) skin on dorsum smooth
with scattered tubercles; low interocular dermal ridge
present in females; skin on venter weakly areolate; discoi-
dal fold prominent; dorsolateral folds present; (2) tympanic
membrane smooth, and tympanic annulus prominent,
slightly higher than long, its length slightly more than V2
length of eye; (3) snout moderately long, bluntly rounded
in dorsal view, rounded in profile; (4) upper eyelid with
numerous, small, low tubercles, narrower than lOD; cra-
nial crests absent; (5) vomerine odontophores prominent,
triangular; (6) males possessing vocal slits and nonspinous
nuptial pads; (7) Finger 1 slightly longer than II; discs on

Eleutherodactylus in the Andes of Northern Peru
B C

21

Fig. 5. Dorsal and lateral views of the heads of three species in the Elculheivdactylus coiispicilbtiis group. A and D. £. aviciiporum, LSUMZ 39365.
B and E. E. aiiiciwstris, LSUMZ 39369. C and E E. metalmtes, KU 186504. Scale bars = 5 mm.

outer fingers expanded, truncate, more than twice width
of digit proximal to pad; (8) fingers bearing narrow lateral
fringes; (9) ulnar tubercles low, round; (10) heel and outer
edge of tarsus lacking tubercles; inner edge of tarsus bear-
ing low fold distally; (11) inner metatarsal tubercle ellipti-
cal, about 2-3x rounded outer metatarsal tubercle; super-
numerary plantar tubercles absent; (12) toes lacking lat-
eral fringes; webbing basal; Toe V slightly longer than III;
discs equal in size to those on outer fingers; (13) dorsum
brown with single, small, prominent, dark brown middor-
sal spot; venter tan with diffuse brown reticulations;
posterior surfaces of thighs brown with small tan spots;
(14) SVL in male 25.2 mm, in 6 females 31.0-34.3 (x =
32.4) mm.

By having an interocular dermal fold, Eleutherodactylus
avicuporwn is most similar to E. skydmainos of the Amazo-
nian lowlands and lower (<750 m) slopes of the Cordillera
Oriental in central and southern Peru; E. skydmainos dif-
fers by having the venter smooth and immaculate.
Eleutherodactylus cuueirostris is sympatric with E.
avicuporum; the former differs by having a long, wedge-
shaped snout, many round, dark spots on the dorsum, a
broad pale labial stripe, and a smooth venter.

Eleutherodactylus avicuporum is like E. caprifer of the
Chocoan Region of southern Colombia and Ecuador and
£. karcharias of the Rio Utcubamba Valley on the west slope
of the Cordillera Central in northern Peru in having ar-
eolate skin on the belly. The latter differs from E. avicuporum
by having a fin-shaped middorsal tubercle, and the former
differs by having a broad brown dorsolateral stripe and
many dark chevrons on the dorsum of the body and brown
stripes on the throat.

Description. — (n = 1 male, 6 females). Head as wide
as body; HW 38.8-42.8 (x = 40.3)% SVL; HL 40.1-46.6 (x =
42.3)% SVL; snout moderately long, bluntly rounded in
dorsal view and rounded in profile (Fig. 5A); E-N 95.1-
120.0 (x= 106.1)%. length of eye; nostrils slightly protuber-
ant, directed laterally; canthus rostralis angular, straight;
loreal region noticeably concave; lips rounded; upper eye-
lid bearing small, rounded tubercles, especially on lateral
edge; upper eyelid width 57.1-71.4 (x = 61.7)% lOD;
supratympanic fold moderately heavy; abruptly curving
downward behind tympanum, obscuring dorsal and
posterodorsal edge of tympanum; side of head nearly ver-
tical; tympanic membrane prominent; tympanic annulus
slightly higher than long; length of tympanic annulus 51 .7-

22

Scientific Papers, Natural History Museum, The University of Kansas

Fig. 6. Feet of three species in the Ek'utherodactylus couspjiciUntus
group. A. E. aviaiporum, LSUMZ 39365. B. £. cuneiivstrii, LSUMZ 39369.
C. E. meiabates, KU 186504. Scale = 5 mm.

62.9 (x - 57.2)% length of eye; two or three prominent
postrictral tubercles below tympanum. Choanae small,
ovoid, not concealed by palatal shelf of maxillary arch;
vomerine odontophores prominent, triangular in outline,
moderately separated medially, located well behind pos-
terior edges of choanae, each odontophore bearing 4-6 (x

= 5.3) teeth; tongue about twice as long as wide, barely
notched posteriorly; posterior half not adherent to floor of
mouth.

Skin on dorsum smooth to very weakly shagreen with
scattered small tubercles especially posteriorly and later-
ally; low interocular dermal ridge (absent in male); dorso-
lateral fold weakly tubercular; lateral fold nearly as pro-
nounced as dorsolateral one; skin on flanks areolate; skin
on throat and belly weakly areolate; other ventral surfaces
smooth, except coarsely areolate on proximal
posteroventral surfaces of thighs; discoidal fold prominent;
cloacal sheath short; large tubercles in cloacal region ab-
sent. Ulnar tubercles low, round (barely discernible in two
specimens; coalesced so as to form low ridge distally on
forearm in two specimens); thenar tubercle elevated, el-
liptical, slightly larger than bifurcate palmar tubercle; pal-
mar supernumerary tubercles absent; subarticular tu-
bercles prominent, subconical; fingers bearing thin lateral
fringes; Finger I slightly longer than II; discs on Fingers I
and II small, round; discs on outer fingers truncate, more
than twice width of digit proximal to pad; all fingers hav-
ing ventral pads well defined by circumferential grooves.
Upper surfaces of hind limbs smooth with scattered minute
tubercles; heel and outer edge of tarsus lacking tubercles;
inner edge of tarsus bearing low fold distally; inner meta-
tarsal tubercle elevated, elliptical, 2-3x rounded outer
metatarsal tubercle; plantar supernumerary tubercles ab-
sent; subarticular tubercles small, subconical; toes lacking

Fig. 7. Dorsal color patterns of three species in the Eleutlicwdacti/lus coiispicillnluii group. A. L. niucupormii,
LSUMZ 39369. C. E. metahates, KU 186504. Scale bars = 5 mm.

LSUMZ 39365. B. E. ciindwftris,

ELEUTHEROnACTYlVS IN THE ANDES OF NORTHERN PeRU

23

Fig. 8. Localities of known occurrence of five species in the
Eleutliewiiactylus coiifpicilhitiis group in the Andes of southern Ecuador
and northern Peru.

lateral fringes; toes webbed basally; discs on toes about
equal in size to those on outer fingers; tip of Toe V extend-
ing to distal edge of pentultimate subarticular tubercle on
Toe IV; tip of Toe 111 extending to middle of pentultimate
subarticular tubercle on Toe IV (Fig. 6A); when hind limbs
flexed perpendicular to axis of body, heels broadly over-
lapping; shank 58.0-63.9 (x= 60.7)% SVL.

Coloration in preservative: Dorsum brown with darker
brown markings consisting of small round spot
middorsally about midway between scapular region and
sacrum, canthal and supratympanic stripes, minute spots
posterolaterally from scapular region (2 specimens), row
of minute spots just ventral to dorsolateral fold (1 speci-
men); diffuse brown marking consisting of interorbital bar,
labial bars that extend onto margin of lower lip, and nar-
row transverse bars on forearms, thighs, shanks, and tarsi;
one individual with diffuse brown chevron posterior to
sacrum, and one with diffuse brown spot posterior to
sacrum followed by diffuse transverse mark posteriorly;
flanks tan. Posterior surfaces of thighs dark brown with
small tan spots, especially distally. Throat and belly tan
with diffuse brown reticulation; ventral surfaces of thighs
and shanks creamy tan with diffuse brown reticulations

and flecks; ventral surfaces of tarsi black, bordered by nar-
row creamy tan stripes.

Coloration in life: Unknown.

Measurements of holotype: SVL 33.2, tibia length 20.1,
foot length 17.7, head width 13.1, head length 13.9, lOD
4.1 , upper eyelid width 2.4, E-N 4.0, eye, 3.5, tympanum 2.0.

Distribution and habitat. — Elciitherodactylus
aviciipoyiDU is known only from elevations of 1700-2030 m
on the western slopes of the Cordillera Colan in northern
Peru (Fig. 8). All individuals were on the ground in humid
montane forest.

Etymology. — The specific name is derived from the
Latin avicupis meaning bird catcher, and the genitive plu-
ral suffix -oriDU. The name is used in recognition of the
efforts expended by ornithologists from the Museum of
Zoology, Louisiana State University, collecting amphibians
in the Cordillera Colan.

Remarks. — The five juveniles have SVLs of 14.0-19.3
(x = 17.4) mm. The interocular dermal ridge is weak in
three individuals and absent in two, both of which are
males. The coloration is like that of the adults, except that
in three of the juveniles a diffuse brown chevron in present
immediately posterior to the middorsal dark brown spot.

Eleuthewdactylus citriogaster Duellman

Eleutherodachjlus citriogaster Duellman, 1992b:24. Holotype: KU 212277,
adult female, from Cataratas Ahuashiyacu, 730 m, 14 km [by road]
NE Tarapoto, Departamento San Martin, Peru.

Diagnosis. — A member of the Eleutherodactylus
(Eleiitlicwdacti/lus) conspicillntus Group having (1) skin on
dorsum shagreen, lacking tubercles, that on venter smooth;
discoidal fold evident; dorsolateral folds indistinct; (2) tym-
panic membrane and tympanic annulus prominent, round,
its length about 307c length of eye; (3) snout subacuminate
in dorsal view, rounded in profile; canthus rostralis angu-
lar; (4) upper eyelid lacking tubercles, slightly narrower
than or equal to lOD; cranial crests absent; (5) vomerine
odontophores triangular, prominent; (6) males having vo-
cal slits and nuptial pads; (7) Finger 1 longer than 11; discs
broad, round; (8) fingers lacking lateral fringes; (9) ulnar
tubercles present distally; (10) heel and tarsus lacking tu-
bercles; (11) inner metatarsal tubercle oval, 3x oval outer
metatarsal tubercle; plantar supernumerary tubercles nu-
merous; (12) toes lacking lateral fringes; webbing basal;
Toe V longer than 111 but not extending to distal
subarticular tubercle of Toe IV; discs as large as those on
fingers; (13) dorsum grayish brown with or without nar-
row brown chevrons and interorbital bar; venter cream
with brown flecks on throat and thighs; (14) SVL in males
31.6^1.3 mm, in females 42.0-51.0 mm.

In general appearance, Eleutherodactylus citriogaster (Fig.
9) closely resembles two other species in the E. conspiciUatus

24

Scientific Papers, Natural History Museum, The University of Kansas

Eteuthcrodacn/liis atnogaster, KU 212277, temale, 42.0 mm SVL.

ElL-iilhL'ivdiict\/lus fnclanoi^iislcr, KU 212321, temale, 24.7 mm SVL.

Ek'iithcrodnctytus patatkos, KU 212320, female, 2 1 n nun s\' I

Elcuthc>vdact\/ltif pinguid, KU 181284, female, 28.5 mm SVL.

Elcuthcwdactylus ardaloni/duis, KU 212301, female, 27.4 mm SVL.

XBPTi

iZ

^^m^^^m* iH

^

B^^^^L^^^P^^^ ^ '^v ' ^j-

^^i^^'

Wf%^^^i^^M^

^^♦"inj

■

|,^,T^"5^^|

^^

1

'^^ A an»l*%<^^'fvV^^y<7^

i^H^^^^HSI

HI

'''^^JE jUytJv.^^^jpi^^ -=»g*MH

I^^HHH^^

T^

|^g^^3

i!i^

ElcuthcivdiKli/liis ccuthospiliii^, KU 219776, male, 22.2 mm SVL (ERW). Elcittltcioilikliiliif. iiili,i:^iill:ilii-. kU 212247, lemale, 22.4 mm SV L.

Fig. 9. Eight species of ElcutbcrodiR-lylu^ from the Andes of northern Peru.

Eleutherodactylus in the Andes of Northern Peru

25

Eleuthemdact\ihi> iici'lioi'liilu>. kL 2123U. , icmalc, 2>..-

ilciitJicivi1iKt\jliis petrobardus, KU 212292, male, 28.7 mm SVL.

ElctiflieivdiKtylu<. liiodoplichus, KU 219787, male, 25.7 mm SVL (ERW). Eleutherodach/lus rhodostichus, KU 212264, male, 19.3 mm SVL.

'#? ■ ' '■■I

Eleidherodactylus uifioculis, KU 212312, male, 181 mm SVL.

Eleuthcwdnctvliis achultei, KU 212222, male, 26.5 mm SVL.

Eleutherodactylus ierendipilus . KU 181279, male, 20.4 mm SVL. Ekuthcrodnctylus wiciisi. KU 219795, male, 32.1 mm SVL (ERW).

Fig. 10. Eight species of Eleutherodactylus from the Andes of northern Peru.

Scientific Papers, Natural History Museum, The University of Kansas
B ^^^ C

Fig. 11. Ventral color patterns of three large species in the
Eleiithewdactylus conspicillatus group. A. E. citriogaster, KU 212287; B. E.
condor, KU 147020; C. E. lymani, KU 181265. From Duellman (1992b).

Group in northern Peru and southern Ecuador — £. condor
and £. himnni. The three species are distinguishable on the
basis of color pattern on the posterior surfaces of the
thighs — bold black and white spots or reticulations in £.
lymani, brown with pale flecks in £. condor, and dark brown
and tan mottling in £. citriogaster (Figs. IIA, 12A). Fur-
thermore, £. citriogaster has a round, instead of bifid, pal-
mar tubercle. Other members of the £. conspicillatus Group
are smaller; of those in the upper Amazon Basin and lower
slopes of the Andes, £. conspicdiatiis, nialkini, metabates, and
peniviauus have the posterior surfaces of the thighs dark
brown with small cream to red spots, and two of these (E.
nialkini and £. metabates) have basal webbing between the
toes, whereas £. fenestratus, lanthanites, and vilarsi have
uniformly dark brown posterior surfaces of the thighs.
Furthermore, £. lanthanites has median pale stripe on a dark
throat and a prominent tubercle on the heel, and £. vilarsi
has a bifid palmar tubercle. Other small members of the E.
conspicillatus Group in the region differ in structure and
coloration. Eleutherodactylus buccinator has pink spots on
the groin on the hidden surfaces of the thigh (Rodriguez,
1994); £. karcharias has a prominent fin-shaped middorsal
tubercle, and £. avicuporum and £. skydmainos have an
interocular fold. In £. ai'icuporum and E. karcharias, the skin
on the belly is areolate.

Description. — The description by Duellman (1992b) is
adequate and contains all information known about this
species.

Distribution and habitat. — This species is known only
from elevations of 600-800 m in lower humid montane
forest on the slopes of a ridge northeast of Tarapoto,
Departamento San Martin, Peru (Fig. 8). All adults were
observed at night on rocks in and along cascading streams.

Remarks. — Duellman (1992b) overlooked the fact that
the presence of a round, instead of bifid, palmar tubercle

Fig. 12. Color patterns on the posterior surfaces
of the thighs in three large species in the Eleuthero-
dnctylus conspicillatus group. A. E. citriogaster, KU
212287. B. £. condor, KU 147020. C. E. lywani. KU
181265. From Duellman (1992b).

is shared in Eleutherodactylus only with £. zeuctotylus, a

member of the Eleutherodactylus conspicillatus Group in

northeastern South America (Lynch and Hoogmoed, 1977).

Eleutherodaclyhis cotuior Lynch and Duellman

Eleutherodactylus condor Lynch and Duellman, 1980:18. Holotype, KU
146992, a juvenile female, from the Rio Piuntza, Cordillera del Condor,
Provincia Morona-Santiago, Ecuador.

Diagnosis. — A member of the Eleutherodactylus
conspicillatus Group having (1) skin on dorsum shagreen,
that on venter smooth; discoidal fold evident; dorsolat-
eral folds present; (2) tympanic membrane smooth, and
tympanic annulus prominent, nearly round, its length
about Vi length of eye; (3) snout subacuminate in dorsal
view, rounded in profile; (4) upper eyelid lacking tubercles,
slightly narrower than or equal to lOD; cranial crests pal-
pable in large females; (5) vomerine odontophores promi-
nent, triangular; (6) males possessing vocal slits and
nonspinous nuptial pads; (7) Finger 1 longer than 11; discs
expanded, about twice width of digit proximal to pad; (8)
fingers lacking lateral fringes; (9) ulnar tubercles absent;
(10) heel and tarsus lacking tubercles; (11) inner metatar-
sal tubercle elongate, 6-8x round outer metatarsal tubercle;
supernumerary plantar tubercles few or absent; (12) toes
bearing narrow lateral fringes; webbing absent; Toe V

Eleutherodactylus in the Andes of Northern Peru

27

slightly longer than III; discs slightly smaller than those
on outer fingers; (13) dorsum brown with darker brown
markings — labial bars, interorbital bar, chevrons; venter
cream with brown blotches; throat heavily pigmented with
brown; posterior surfaces of thighs brown with cream or
white spots; (14) SVL in males 32.1-39.5 mm, in females
52.6-62.2 mm.

In general appearance, Elciitlierodnctylus condor is most
similar to E. citriogaster and E. lymaiii. The venter in E. con-
dor is creamy white with dense gray pigmentation on the
throat, belly, and ventral surfaces of the thighs (Figs. 11 B,
12B); the posterior surfaces of the thighs are brown with
pale spots. In contrast, in E. lymmu, the venter is essen-
tially unpigmented, except for dark bars on the margin of
the lower lips and posterolaterally on the throat; the pos-
terior surfaces of the thighs are black with bold cream
mottling. In E. citriogaster, the venter is yellow with brown
or gray mottling on the throat, laterally on the belly, and
on the ventral surfaces of the thighs; the posterior surfaces
of the thighs are mottled dark brown and creamy tan. Fur-
thermore, the palmar tubercle is round, instead of bifid, in
E. citriogaster. Other members of the E. cmispicillatiis Group
are smaller; of those in the upper Amazon Basin and lower
slopes of the Andes, E. conspiciUatus , malkini, metabates and
peritvianus have the posterior surfaces of the thighs dark
brown with small cream to red spots, and two of these (£.
malkini and £. metabates) have basal webbing between the
toes, whereas E. fenestratus, lanthanites, and vilarsi have
uniformly dark brown posterior surfaces of the thighs.
Furthermore, £. lanthanites has median pale stripe on a dark
throat and a prominent tubercle on the heel, and E. vilarsi
has proportionately shorter hind limbs and larger tym-
pana. Other small members of the E. conspiciUatus Group
in the region differ in structure and coloration.
Eleutherodactylus cuneirostris has a wedge-shaped snout and
many small dark spots on the dorsum, and E. buccinator
has pink spots on the groin on the hidden surfaces of the
thigh (Rodriguez, 1994); E. karcharias has a prominent fin-
shaped middorsal tubercle, and E. avicuporum and £.
skydmainos have an interocular fold. Eleutherodactylus
avicuporum and E. karcharias differ from other members of
the group in the region by having areolate skin on the belly.

Distribution and habitat. — The few records for
Eleutherodactylus condor suggest a rather limited elevational
distribution on mountain ranges to the east of the Cordil-
lera Oriental of the Andes (Fig. 13). The species is known
from 1975 m in the Cordillera de Cutucii (Duellman and
Lynch, 1980), the type locality at 1550 m- and Coangos at
1500-1600 m (Almendariz, 1997) on the western slope of
the Cordillera del Condor, and from 1750 m on the eastern

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'Lynch and Duellman (1980) erroneously gave the elevation of the type
locality as 1830 m.

Fig. 13. Localities of known occurrence of four species in the
Eleutherodactylus conspiciUatus group and one species in the Eleuthero-
dactylus nigrovittatus group (E. araiodactylus) in the Andes of southern
Ecuador and northern Peru.

slope of the Cordillera del Condor. All individuals have
been taken in humid montane forest.

Remarks. — This first Peruvian record is based on
USNM 525437, an adult female having a SVL of 60.2 mm.
The specimen was obtained at a site on the upper Rio
Comainas, at the base of Cerro Machinaza in the Cordil-
lera del Condor, by Thomas S. Schulenberg on 6 August
1994.

Eleutlierodactylus cuneirostris new species
Holotype.— LSUMZ 39369, an adult female, from 12 km
[by trail] E La Peca (05°36' S, 78°19' W, 1700 m), Provincia
Bagua, Departamento Amazonas, Peru, obtained on 11
Junel978byL. J. Barkley.

Diagnosis. — A member of the Eleutherodactylus
conspiciUatus Group having (1) skin on dorsum smooth;
that on venter smooth; discoidal fold evident; dorsolat-
eral folds absent; (2) tympanic membrane smooth, and
tympanic annulus prominent, slightly higher than long,
its length slightly more than V2 length of eye; (3) snout long,
rounded in dorsal view, protruding well beyond margin
of lower lip and acutely rounded in profile; (4) upper eye-
lid lacking tubercles, narrower than lOD; cranial crests

28

Scientific Papers. Natural History Museum, The University of Kansas

absent; (5) vomerine odontophores prominent, oblique; (6)
condition of vocal slits and nuptial pads unknown; (7)
Finger I equal in length to II; discs on outer fingers ex-
panded, tn.mcate, about twice width of digit proximal to
pad; (8) fingers lacking lateral fringes; (9) ulnar tubercles
absent; (10) heel and outer edge of tarsus lacking tubercles;
inner edge of tarsus bearing low, elliptical tubercle dis-
tally; (11) inner metatarsal tubercle elliptical, about 3x
subconical outer metatarsal tubercle; supernumerary plan-
tar tubercles absent; (12) toes lacking lateral fringes; web-
bing absent; Toe V slightly longer than III; discs slightly
smaller than those on outer fingers; (13) dorsum tan with
small dark brown spots; venter creamy tan with diffuse
brown reticulations on belly and ventral surfaces of tliighs;
posterior surfaces of thighs brown; (14) SVL in one female
29.1 mm.

Eleuthewdacti/liis cuneirostris differs from all other mem-
bers of the Elcutherodaciyhis conspicillntiis Group in the re-
gion by having a long, wedge-shaped snout and a con-
spicuous pale labial stripe. Eleuthewdnctyhis peniviainis also
has a pale labial stripe, but the dorsum usually has dark
chevrons (otherwise uniform tan, red, or green); the throat
has dense dark flecks, and there are no reticulations on
the belly. Of other small members of the £. conspicillntiis
Group, E.feiwstratiis, laiitlianites, and vilnrsi have uniformly
dark brown posterior surfaces of the thighs. Furthermore,
E. lanihanites has median pale stripe on a dark throat and
a prominent tubercle on the heel, and £. vilarsi has propor-
tionately shorter hind limbs and larger tympana. Other
small members of the £. coiispicillatus Group in the region
differ in structure and coloration. Eleutherodactylus bucci-
nator has pink spots in the groin and on the hidden sur-
faces of the thighs (Rodriguez, 1994); £. karcharias has a
prominent fin-shaped middorsal tubercle and areolate skin
on the venter, and £. avicuporum and £. sici/dinnijios have
an interocular fold. Eleutherodactylus metabates lacks a pale
labial stripe and has X- or chevron-shaped marks on the
back. In the larger species in the £. conspicillatus group in
the region, the venter in £. condor is creamy white with
dense gray pigmentation on the throat, belly, and ventral
surfaces of the thighs; the posterior surfaces of the thighs
are brown with pale spots. In £. lymani, the venter is es-
sentially unpigmented, except for dark bars on the mar-
gin of the lower lips and posterolaterally on the throat; the
posterior surfaces of the thighs are black with bold cream
mottling. In £. citriogaster, the palmar tubercle is round (in-
stead of bifid), and the venter is yellow with brown or gray
mottling on the throat, laterally on the belly, and on the
ventral surfaces of the thighs; the posterior surfaces of the
thighs are mottled dark brown and creamy tan.

Description. — (» = 1 female). Head as wide as body.
HW 40.5% SVL; HL 44.6% SVL; Snout long, rounded. in
dorsal view, protruding well beyond margin of lower lip

and acutely rounded in profile, giving wedge-shaped ap-
pearance (Fig. 5B); E-N 97.1% length of eye; nostrils nota-
bly protuberant, directed laterally; canthus rostralis angu-
lar, straight; loreal region noticeably concave; lips rounded;
upper eyelid lacking tubercles; upper eyelid width 70.6%
lOD; supratympanic fold weak; abruptly curving down-
ward behind tympanum, not obscuring edge of tympa-
num; side of head inclined ventrolaterally; tympanic mem-
brane prominent; tympanic annulus slightly higher than
long; length of tympanic annulus 55.9% length of eye; two
low, rounded postrictral tubercles posteroventral to tym-
panum. Choanae large, round, not concealed by palatal
shelf of maxillary arch; vomerine odontophores prominent,
oblic}ue, narrowly separated medially, located well behind
posterior edges of choanae, each odontophore bearing five
teeth. Tongue about twice as long as wide, barely notched
posteriorly; posterior half not adherent to floor of mouth.

Skin on dorsum smooth; dorsolateral fold absent; skin
on flanks and venter smooth, except coarsely areolate on
proximal posteroventral surfaces of thighs; discoidal fold
prominent; cloacal sheath short; large tubercles in cloacal
region absent. Ulnar tubercles absent. Thenar tubercle el-
evated, elliptical, slightly smaller than bifurcate palmar
tubercle; palmar supernumerary tubercles absent;
subarticular tubercles prominent, subconical; fingers lack-
ing lateral fringes; Finger I equal in length to 11. Discs on
Fingers I and II small, round. Discs on outer fingers trun-
cate, about twice width of digit proximal to pad; all fin-
gers having ventral pads well defined bv circumferential
grooves. Upper surfaces of hind limbs smooth; heel and
outer edge of tarsus lacking tubercles; inner edge of tarsus
bearing small, elliptical tubercle distally; inner metatarsal
tubercle elevated, elliptical, about 2x subconical outer
metatarsal tubercle; plantar supernumerary tubercles ab-
sent; subarticular tubercles small, subconical; toes lacking
lateral fringes; toes webbed basally; discs on toes about
ecjual in size to those on outer fingers; tip of Toe V extend-
ing to distal edge of penultimate subarticular tubercle on
Toe IV; tip of Toe 111 extending to base of penultimate
subarticular tubercle on Toe IV (Fig. 6B); when hind limbs
flexed perpendicular to axis of body, heels overlap by about
one-third length of shank; shank 65.6%. SVL.

Coloration in preservative: Dorsum tan with dark
brown markings consisting of many small round or ovoid
spots on dorsum of body and flanks (Fig. 7B). Canthal and
supratympanic stripes; transverse mark above vent and
small spots distally on anterior surfaces of thighs; narrow
transverse bars on limbs barely evident; broad creamv tan
labial stripe. Posterior surfaces of thighs dull brown; throat
creamy tan with minute brown flecks; belly and ventral
surfaces of thighs creamy tan with diffuse, faint brown
reticulations. Ventral surfaces of tarsi black.

Eleutherodactylus in the Andes of Northi:rn Pbru

29

Coloration in life: Unknown.

Measurements of holotype; SVL 29.1, tibia length 19.1,
foot length 15.6, head width 11.8, head length 13.0, lOD
3.4, upper evelid width 2.4, E-N 3.3, eye, 3.4, tympanum 1.9.

Distribution and habitat. — Elcuihcmdaciyliis cinicirostris
is known only from 1700 m in humid montane forest on
the western slopes of the Cordillera Colan in northern Peru
(Fig. 13).

Etymology. — The specific name is an adjective from the
Latin ciiiieiis meaning wedge and the Latin rostrum mean-
ing snout. The name refers to the wedge-shaped snout.

Remarks. — Among species of Elcntlicrodnctyliifi in Ec-
uador and Peru, the long, wedge-shaped snout of this spe-
cies is approached only by the long, subacuminate snout
in E. loii^^irostris, a member of the ElcutIu'rot1nct\/liis
(Crnugnstor) fitzhigeri group in the Chocoan region.

Eleutherodactylus karcharias Flores and Rodriguez

Eleutherodactylus knninuias Flores and Rodriguez, 1997:392. Holotype,
MCZ 89075, immature female, from Alva, 1000m, between
Chachapoyas and Bagua Grande, Departamento Amazonas, Peru.

Diagnosis. — A member of the Eleutherodactylus
couspicillntus Group having (1) skin on dorsum shagreen
with scattered tubercles, especially on flanks, prominent
fin-shaped middorsal tubercle; skin on venter areolate;
discoidal fold prominent; dorsolateral and flank folds
present; (2) tympanic membrane smooth, and tympanic
annulus prominent, slightly higher than long, its length
about M to Vi length of eye; (3) snout rounded in dorsal
view and in profile; (4) upper eyelid lacking tubercles,
broader than lOD; cranial crests present; (5) vomerine
odontophores prominent, triangular; (6) condition of vo-
cal slits and nuptial pads unknown; (7) Finger 1 equal in
length to II; discs expanded, about twice width of digit
proximal to pad; (8) fingers bearing broad lateral fringes;
(9) ulnar tubercles absent; (10) heel and tarsus lacking tu-
bercles; (11) inner metatarsal tubercle oval to elongate, 2-
6x round outer metatarsal tubercle; (12) toes bearing broad
lateral fringes; webbing basal; Toe V slightly longer than
111; discs nearly equal in size to those on outer fingers; (13)
dorsum pale grayish tan to brown with darker brown
markings — interorbital and labial bars, canthal and
supratympanic stripes, W-shaped mark in scapular region,
two dorsal chevrons (anterior one enclosing black spot with
middorsal tubercle), triangular mark enclosing vent, and
diagonal bars on limbs; venter cream with brown flecks,
most dense on chest and throat; posterior surfaces of thighs
brown with cream flecks or spots; (14) SVL in subadult
female 30.4.

Except for three species, all members of the
Eleutherodactylus conspicillatus Group have smooth venters.
By having an areolate venter, Eleutherodactylus karcharias
resembles £. capvifer of the Chocoan Region of southern

Colombia and Ecuador and £. avicuporum of the Cordil-
lera Collin. The latter differs from £. karcharias by lacking a
fin-shaped middorsal tubercle and distinct dorsal mark-
ings, and the former differs by having a broad brown dor-
solateral stripe and many dark chevrons on the dorsum of
the body and brown stripes on the throat.

Description. — Nothing can be added to the description
bv Flores and Rodriguez (1997).

Distribution and habitat. — Eleutherodactylus karcharias
is known from a single locality at an elevation of 1000 m in
the valley of the Rio Utcubamba (tributary of the Rio
Marahtin) in the northern part of the Cordillera Central in
northern Peru (Fig. 8). The specimens were "taken from
moist leaf litter in cloud forest" (Rores and Rodriguez, 1997).

Remarks. — This species is known from only seven
immature females.

Eleutherodactylus lanthanites Lynch

Eleuthcrodtictylus Inutlmnitcs Lynch, 1975b:10. Holotype, KU 146144, an
adult female, from Santa Cecilia, 340 m, Provincia Napo, Ecuador.

Diagnosis. — A member of the Eleutherodactylus
couspic Hiatus Group having (1) skin on dorsum finely tu-
berculate with scattered larger tubercles, that on venter
smooth; discoidal fold prominent; dorsolateral folds ab-
sent; (2) tympanic membrane smooth, and tympanic an-
nulus prominent, nearly round, its length about Vi length
of eye; (3) snout long, subacuminate in dorsal view,
rounded in profile; (4) upper eyelid lacking tubercles, nar-
rower than lOD; cranial crests absent; (5) vomerine
odontophores prominent, oblique; (6) males possessing
vocal slits and nonspinous nuptial pads; (7) Finger 1 longer
than 11; discs expanded, about twice width of digit proxi-
mal to pad; (8) fingers lacking lateral fringes; (9) ulnar tu-
bercles absent; (10) heel bearing conical tubercle; outer edge
of tarsus bearing row of indistinct tubercles; inner tarsal
tubercles and fold absent; (11) inner metatarsal tubercle
elongate, about 5x conical outer metatarsal tubercle; su-
pernumerary plantar tubercles few in number; (12) toes
lacking lateral fringes; webbing absent; Toe V slightly
longer than III; discs equal in size to those on outer fin-
gers; (13) dorsum brown with darker brown markings
(usually chevrons); venter cream with dark flecks; throat
heavily pigmented with brown or black, defining median
white stripe; posterior surfaces of thighs brown with cream
flecks; (14) SVL in males 21.7-27.9 mm, in females 27.5-
45.4 mm (Lynch and Duellman, 1980).

Eleutherodactylus lanthanites differs from all other mem-
bers of the Eleutherodactylus conspicillatus Group, except E.
gutturalis, by having a dark throat with a median white
stripe. The latter species, which inhabits the Guianan Re-
gion, differs from £. lanthanites by lacking a tubercle on
the heel and by having proportionately longer hind limbs
(Hoogmoed et al., 1977).

30

Scientific Papers, Natural History Museum, The University of Kansas

Distribution and habitat. — Eleutherodactyliis
Innthimites is widespread in tlie upper Amazon Basin in
southern Colombia, Ecuador, extreme western Brazil, and
northern Peru (Lynch, 1980; Duellman and Mendelson,
1995). The species ascends the eastern slopes of the Andes
to elevations of 1440 m in southern Colombia and 1490 m
in Ecuador (Lynch and Duellman, 1980). In northern Peru,
the species has been taken only at Venceremos at an eleva-
tion of 1630 m, Departamento San Martin (Fig. 13). The
single individual was on the ground by day in humid
montane forest.

Remarks. — A juvenile male has a SVL of 19.0 mm. In
life, the dorsum was tan with dull red markings. The ante-
rior, ventral, and posterior surfaces of the thighs were or-
ange, and the face mask was dark brown. The belly was
yellow with black flecks, and the throat was black with
cream spots and a median white stripe. The iris was bronze
with a median, horizontal, red streak. This juvenile is as-
signed to Eleutherodactyliis larithanites on the basis of the
throat pattern, which is unique to this species in the
Eleutherodactyliis conspic Hiatus Group.

Eleutherodactylus lymani Barbour and Noble

Eleuthewdactylus lymaiu Barbour and Noble, 1920:403. Holotype: MCZ
5422, young female, from Perico, Departamento Cajamarca, Peru.

Eleutherodactylus carrioni Parker, 1932:23. Holotype: BM 1947.2.15.99, adult
female, from Loja, Provincia Loja, Ecuador Synonymy fide Lynch,
1969:263.

Eleutherodactylus h/maiii — Lynch, 1969:263.

Diagnosis. — A member of the Eleutherodactylus
(Eleutherodactylus) conspicillatus Group having (1) skin on
dorsum shagreen with scattered tubercles, that on venter
smooth; discoidal fold prominent; dorsolateral folds
present; (2) tympanic membrane and tympanic annulus
prominent, round, its length about Vi length of eye; (3)
snout subacuminate in dorsal view, rounded in profile;
canthus rostralis angular; (4) upper eyelid lacking tu-
bercles, usually narrower than lOD; cranial crests absent;
(5) vomerine odontophores subtriangular in outline,
prominent; (6) males lacking vocal slits; nuptial pads
present; (7) Finger 1 longer than 11; discs small; (8) fingers
bearing lateral keels; (9) ulnar tubercles absent; (10) heel
and outer edge of tarsus lacking tubercles; inner surface
of tarsus bearing fold; (11) inner metatarsal tubercle elon-
gate, 4x round outer metatarsal tubercle; plantar supernu-
merary tubercles absent; (12) toes bearing lateral fringes;
webbing absent; Toe V slightly longer than 111; discs as large
as those on fingers; (13) dorsum brown with dark brown
to black markings; venter cream to white; groin and pos-
terior surfaces of thighs black with white spots or reticula-
tions; (14) SVL in males 25.7-45.3 mm, in females 48.4-
69.3 mm.

In size and general appearance, Eleutherodactylus lymani
closely resembles two other species in the E. conspicillatus

Group in northern Peru and southern Ecuador — E. condor
and E. citriogaster. The three species are distinguishable on
the basis of color pattern on the posterior surfaces of the
thighs — bold black and white spots or reticulations in E.
lymani, brown with pale flecks in E. condor, and dark brown
and tan mottling in £. citriogaster (Figs. IIC, 12C); further-
more, E. citriogaster has a round, instead of bifid, palmar
tubercle. Other members of the E. conspicillatus Group are
smaller; of those in the upper Amazon Basin and lower
slopes of the Andes, £. conspicillatus, malkini, metabates, and
peruvianus have the posterior surfaces of the thighs dark
brown with small cream to red spots, and two of these (£.
malkini and E. metabates) have basal webbing between the
toes, whereas E, fenestratus, lanthanites, and vilarsi have
uniformly dark brown posterior surfaces of the thighs.
Furthermore, E. lanthanites has median pale stripe on a dark
throat and a prominent tubercle on the heel, and E. vilarsi
has proportionately shorter hind limbs and larger tym-
pana. Other small members of the £. conspicillatus Group
in the region differ in structure and coloration.
Eleutherodactylus cuneirostris has a wedge-shaped snout and
many small dark spots on the dorsum, and £. buccinator
has pink spots in the groin and on the hidden surfaces of
the thighs (Rodriguez, 1994); £. karcharias has a prominent
fin-shaped, middorsal tubercle, and E. avicuporiim and E.
skydmainos have an interocular fold. Two members of the
E. conspicillatus Group in the region (E. avicuporum and E.
karcharias) have areolate skin on the belly.

Description. — Lynch (1969) provided a thorough rede-
scription of the species, which was augmented by Lynch
and Duellman (1997).

Distribution and habitat. — Eleutherodactylus lymanihas
a reasonably broad distribution on the drier Pacific slopes
and lowlands of southern Ecuador and northwestern Peru
(Fig. 13). Most records are at elevations of less than 1600
m, but it has been found as high as 3000 m in subparamo
in Provincia Loja, Ecuador. Three specimens (KU 196464,
196509; LSUMZ 19553) are from 28 km N of Santa Cruz,
725 m, Provincia Jaen, Departamento Cajamarca, Peru; this
village is on the road between Jaen and Bellavista in the
valley of the Rio Maraiion. The frogs were collected by
Richard Thomas on 8 July 1968; they were under rocks at
the edge of a stream in thorn forest with trees 12-18 m
high. LSUMZ 32460 is from 20 km SW of Chiriaco,
Provinica Bagua, Departamento Amazonas, Peru; this lo-
cality also is in the valley of the Rio Maranon.

Lynch (1969:266) listed MCZ 5436 as being from
Palambla in Departamento Cajamarca. We have been un-
able to find a Palambla in Departamento Cajamarca, but a
village by that name exists at 5°23' S, 79°37' W in
Departamento Piura. Although no place by that name was
cited in Noble's (1921) account of his travels in northern

Eleutherodactylus in the Andes of Northern Peru

31

Peru, the \-illage is in the immediate \-icinity of the route
described bv Noble; furthermore, Barbour and Noble
(1920:395) stated: "The towns of Huancabamba and
Palambla are on the western range of the Andes, on the
border of Piura." Duellman and Wild (1993) recorded
Eleutherodactylus lymani from Canchaque, 1120 m; Palambla
is about 2 km south of Canchaque and at the same eleva-
tion.

Remarks. — Of the specimens reported herein, one (KU
196464) is a subadult male having a SVL of 34.6 mm and
the other (LSU 19553) is a female having a SVL of 47.0 mm.

Eleutherodactylus metabates new species

Holotype.— KU 186504, an adult male, from 20 km [by
road] SW of Chiriaco (05°13' S, 78°17' W, 525 m), Provincia
Bagua, Departamento Amazonas, obtained by Richard
Thomas on 16 December 1974.

Paratypes. — LSUMZ 32460, an adult male from the type
locality.

Diagnosis. — A member of the Eleutherodactylus
couspicillatus Group having (1) skin on dorsum smooth
weakly tuberculate, that on venter smooth; discoidal fold
prominent; dorsolateral folds absent; (2) tympanic mem-
brane smooth, and tympanic annulus prominent, slightly
higher than long, its length slightly less than V2 length of
eye; (3) snout moderately long, rounded in dorsal view
and in profile; (4) upper eyelid with numerous, small, low
tubercles, as wide as lOD; cranial crests absent; (5) vomer-
ine odontophores prominent, oval; (6) males possessing
vocal slits, lacking nuptial pads; (7) Finger I slightly longer
than II; discs on outer fingers expanded, nearly truncate,
more than twice width of digit proximal to pad; (8) fingers
bearing narrow lateral fringes; (9) ulnar tubercles absent;
(10) heel and outer edge of tarsus lacking tubercles; inner
edge of tarsus bearing distinct fold distally; (11) inner meta-
tarsal tubercle elevated, elliptical, about lOx subconical
outer metatarsal tubercle; supernumerary plantar tubercles
absent; (12) toes bearing broad lateral fringes; webbing
basal; Toe V slightly longer than III; discs slightly smaller
than those on outer fingers; (13) dorsum brown with faintly
darker X- or chevron-shaped marks; venter cream with
diffuse brown flecks on throat; posterior surfaces of thighs
brown with small cream flecks; (14) SVL in two males 29.9-
32.2 (x = 31.1) mm; females unknown.

Eleutherodactylus metabates is most similar to £.
couspicillatus and E. peruvianus, both of which have small,
pale flecks on the posterior surfaces of the thighs, but both
species lack webbing and usually have dark face masks;
moreover, E. peruvianus has dark mottling on the throat
and chest. The structurally similar E. malkiui has more
webbing between the toes and black and cream mottling
on the posterior surfaces of the thighs. Of other small mem-

bers of the E. couspicillatus Group, E.feuestratus, laiithauites,
and vilarsi have uniformly dark brown posterior surfaces
of the thighs. Furthermore, E. laiithauites has median pale
stripe on a dark throat and a prominent tubercle on the
heel, and E. -oilarsi has proportionately shorter hind limbs
and larger tympana. Other small members of the £.
coiispicillatus Group in the region differ in structure and
coloration. Eleutherodactylus buccinator has pink spots in
the groin and on the hidden surfaces of the thighs
(Rodriguez, 1994); £. karcharias has a prominent middor-
sal tubercle, and E. avicuporum and E. skyduiaiuos have an
interocular fold. Eleutherodactylus avicuporum and E.
karcharias have areolate skin on the belly, and E. cuneirostris
has a wedge-shaped snout and many round, dark brown
spots on the dorsum. In the larger species in the E.
couspicillatus group in the region, the venter in E. condor is
creamy white with dense gray pigmentation on the throat,
belly, and ventral surfaces of the thighs; the posterior sur-
faces of the thighs are brown with pale spots. In E. lymani,
the venter is essentially unpigmented, except for dark bars
on the margin of the lower lips and posterolaterally on the
throat; the posterior surfaces of the thighs are black with
bold cream mottling. In £. citriogaster, the palmar tubercle
is round (instead of bifid), and the venter is yellow with
brown or gray mottling on the throat, laterally on the belly,
and on the ventral surfaces of the thighs; the posterior sur-
faces of the tliighs are mottled dark brown and creamy tan.

Description. — (n = 2 males). Head wider than body;
HW 37.1-37.3 (X = 37.1)% SVL; HL 41.8-42.2 (x = 42.0)%
SVL; snout moderately long, rounded in dorsal view and
in profile (Fig. 5C); E-N 86.0-92.7 (x = 89.4)% length of
eye; nostrils barely protuberant, directed dorsolaterally;
canthus rostralis angular, straight; loreal region nearly flat;
lips rounded; upper eyelid bearing small, rounded tu-
bercles; upper eyelid width 100.0-103.4 (x = 101.7)% lOD;
supratympanic fold moderately heavy; abruptly curving
downward behind tympanum, obscuring dorsal and
posterodorsal edge of tympanum; side of head nearly ver-
tical; tympanic membrane prominent; tympanic annulus
slightly higher than long; length of tympanic annulus 41 .3-
41.5 (x = 41.4)%i length of eye; two prominent postrictral
tubercles posteroventral to tympanum. Choanae small,
round, partly concealed by palatal shelf of maxillary arch;
vomerine odontophores prominent, oval, moderately sepa-
rated medially, located well behind posterior edges of choa-
nae, each odontophore bearing 5-6 (x = 5.5) teeth; tongue
nearly twice as long as wide, barely notched posteriorly;
posterior one third not adherent to floor of mouth.

Skin on dorsum weakly tuberculate; dorsolateral fold
absent; skin on flanks tuberculate; skin on venter smooth;
discoidal fold prominent; cloacal sheath short; large tu-
bercles in cloacal region absent. Ulnar tubercles absent;

32

Scientific Papers, Natural History Museum, The University of Kansas

thenar tubercle elevated, elliptical, slightly smaller than
bifid palmar tubercle; palmar supernumerary tubercles
absent; subarticular tubercles prominent, subconical; fin-
gers bearing lateral fringes; first finger slightly longer than
second; discs on Fingers I and 11 small, round; discs on
outer fingers nearly truncate, more than twice width of
digit proximal to pad; all fingers having ventral pads well
defined by circumferential grooves. Upper surfaces of hind
limbs smooth; heel and outer edge of tarsus lacking tu-
bercles; inner edge of tarsus bearing distinct fold distally;
inner metatarsal tubercle elevated, elliptical, about lOx
subconical outer metatarsal tubercle; plantar supernumer-
ary tubercles absent; subarticular tubercles small, rounded;
toes bearing broad lateral fringes; toes webbed to basal
subarticular tubercles; discs on toes slightly smaller than
those on outer fingers; tip of Toe V extending to middle of
penultimate subarticular tubercle on Toe IV; tip of Toe 111
extending to base of penultimate subarticular tubercle on
Toe IV (Fig. 6C); when hind limbs flexed perpendicular to
axis of body, heels overlapping by about Vi length of shank;
shank 62.2-62.4 (x = 62.3)% SVL.

Coloration in preservative: Dorsum brown with faint
darker brown markings consisting of interorbital bar, la-
bial bars, supratympanic stripe (but no canthal stripe), X-
or chevron-shaped marks on the back, and barely discern-
ible transverse bars on limbs (Fig. 7C); flanks tan with dif-
fuse brown diagonal bars; axilla cream with small brown
spots; posterior surfaces of thighs brown with cream flecks;
venter cream with diffuse brown flecks on throat.

Coloration in life: Unknown.

Measurements of holotype: SVL 32.2, tibia length 20.1,
foot length 17.0, head width 12.0, head length 13.6, lOD
3.2, upper eyelid width 3.2, E-N 3.5, eye, 4.6, tympanum 1 .9.

Distribution and habitat. — Eleiitliewdactylus nictabatcfi
is known only from the type locality at an elevation of 525
m in the Rio Marafion Valley in Departamento Amazonas,
Peru (Fig. 8). This region supports thorn forest.

Etymology. — The specific name is a Greek noun mean-
ing leaper. The name is applied to this long-legged species
that presumably is capable of lengthy jumps.

Remarks. — The paratype (adult male having a SVL of
29.9 mm) differs from the holotype only by having a more
pallid coloration.

Eleiitliewdactylus penwianus (Melin)

Hylodes penwitvms Melin, 1941:43. Holotype, NHMG 490, an adult fe-
male, from Roque, Departamento San Martin, Peru
Ekntberodactybis peniz'innus — Gorham, 1966:91.

Diagnosis. — A member of the Elcutlicrodiictyhi!^
conspicillatits Croup having (1) skin on dorsum shagreen
with scattered tubercles, that on venter smooth; discoidal
fold prominent; dorsolateral folds present; (2) tympanic

membrane smooth, and tympanic annulus prominent,
nearly round, its length about Vi length of eye; (3) snout
rounded to subacuminate in dorsal view, rounded in pro-
file; (4) upper eyelid lacking tubercles, broader than lOD;
cranial crests absent; (5) vomerine odontophores promi-
nent, triangular; (6) males possessing vocal slits and white,
nonspinous nuptial pads; (7) Finger 1 longer than 11; discs
expanded, about twice width of digit proximal to pad; (8)
fingers bearing lateral fringes; (9) ulnar tubercles absent;
(10) heel and tarsus lacking tubercles, except for small tu-
bercle on inner edge of tarsus; (11) inner metatarsal tu-
bercle oval, about 6x subconical outer metatarsal tubercle;
supernumerary plantar tubercles few in number; (12) toes
bearing lateral fringes; webbing absent; Toe V slightly
longer than III; discs equal in size to those on outer fin-
gers; (13) dorsum usually brown with darker brown chev-
rons; venter cream with brown flecks, most dense on throat;
posterior surfaces of thighs brown with cream spots; (14)
SVL in males 29.2-35.8 mm, in females 38.6-46.4 mm
(Lynch and Duellman, 1980).

Eleiitlicrodnctylns pcniviaiiiis is most similar to £.
conspicillntuf, malkiui, and nietabates, all of which have pale
spots on the posterior surfaces of the thighs. The structur-
ally similar £. conspkiUaiiis lacks dark pigmentation on the
throat and chest and lacks labial bars, whereas £. malkini
and E. mctabates have distinct webbing between the toes
(absent in E. peniviamis), and £. malkini has black and cream
mottling on the posterior surfaces of the thighs. Of other
small members of the £. conspicillatus Group, E.fencstratiis,
lanthanites, and vilnrsi have uniformly dark brown poste-
rior surfaces of the thighs. Furthermore, E. laiitluvutes has
median pale stripe on a dark throat and a prominent tu-
bercle on the heel, and E. vilarsi has proportionately shorter
hind limbs and larger tympana. Other small members of
the E. conspicillatus Group in the region differ in structure
and coloration. Elcnthcwdactylus buccinator has pink spots
in the groin and on the hidden surfaces of the thighs
(Rodriguez, 1994), and E. cuneiwstris has many small, dark
spots on the dorsum; E. karcharias has a prominent mid-
dorsal tubercle, and E. avicuporuni and E. ski/duiaiuos have
an interocular fold (Flores and Rodriguez, 1997). In the
larger species in the E. conspicillatus Group in the region,
the venter in £. condor is creamy white with dense gray
pigmentation on the throat, belly, and ventral surfaces of
the thighs; the posterior surfaces of the thighs are brown
with pale spots. In E. h/fnani, the venter is unpigmented,
except for dark bars on the margin of the lower lips and
posterolaterally on the throat; the posterior surfaces of the
thighs are black with bold cream mottling. In E. citriogaster,
the palmar tubercle is round (instead of bifid), and the
venter is yellow with brown or gray mottling on the throat,
laterally on the belly, and on the ventral surfaces of the
thighs; the posterior surfaces of the thighs are mottled dark
brown and creamy tan.

Elevtherodactylus in the Andes of Northern Peru

33

Distribution and habitat. — Elciilhcrodactylu^
pcniviaiiu> is \videspread in the upper Amazon Basin in
western Brazil, Ecuador, and Peru (Lynch, 1980; Lynch and
Duellman, 1980). Herein, we record it at elevations of 950
and 1080 m on the eastern slopes ot the Cordillera Central
in northern Pen.i — respectively, KU 217313 from 15.4 km
SW of Zapatero and KU 212291 from Abra Tangarana, 7
km [bv road] NE San Juan de Pacaysapa, both in Provincia
Lamas, Departamento San Martin (Fig. 8). The latter was
on the ground bv a stream during the day. The species as-
cends the eastern slopes of the Andes to elevations of nearly
2000 m in Ecuador and Peru (Lynch and Duellman, 1980).
It has been reported from elevations of 1700-1975 m in the
Cordillera de Cutucii (Duellman and Lynch, 1988), from
1500-1830 m on the western slopes of the Cordillera del
Condor (Almendariz, 1997; Lynch and Duellman, 1980),
and from 665-1750 m on the eastern slopes of the Cordil-
lera del Condor in Peru (Reynolds and Icochea, 1997). Two
of the latter were on vegetation at night, and one was in a
bromeliad; all were in humid montane forest.

Remarks.— Two specimens (USNM 525438-39) from
the upper Rio Comainas, 1 750 m, base of Cerro Machinaza,
Departamento Amazonas, are adult females having SVLs
of 37.8 and 37.2 mm, respectively. A third individual
(USNM 525440) from the Puesto Vigilancia, Rio Comainas,
665 m, Departamento Amazonas, is a male having a SVL
of 28.7 mm. In USNM 525438, the dorsum is uniform olive
tan, and the flanks are brown, whereas in the other speci-
mens, the dorsum is tan with brown chevrons. In the fe-
males, the throat and chest have dark gray mottling,
whereas only small gray flecks are present on the throat
and chest of the male. The coloration of living individuals
was: USNM 525438 — Dorsum reddish brown; flanks and
legs brown; dark canthal mask; chin mottled brown-yel-
low; chest and belly yellow with brown spotting; under-
side of legs yellow with brown spotting; iris bronze (R. P.
Reynolds^ field notes, 17 July 1994). USNM 5254440— Dor-
sum tan with light brown chevrons; dark canthal stripe;
legs barred; rear of thighs brown with yellow spots; chin
green; belly cream (Robert P. Reynolds field notes, 31 July
1994). The specimen from Abra Tangarana is a juvenile with
a SVL of 23.5 mm. In life, the dorsum was dull olive-tan
with brown chevrons, and the posterior surfaces of the
thighs were brown with pale red flecks; the ventral sur-
faces of the limbs were pale yellow (WED field notes, 5
February 1989).

Eleutherodactylus nigrovittatus Group

This group was defined by Lynch (1989) as having the
skin on the v^enter smooth, dorsolateral folds present or
absent. Finger I longer than Finger II, discs on digits not
expanded, and vomerine odontophores present. Lynch and
Duellman (1997) added relative length of the toes — Toe III

longer than Toe IV. The inclusion of the new species de-
scribed below necessitates a slight modification of the defi-
nition — Finger I longer than, or equal in length to. Finger
II, and Toe IV shorter than, or equal in length to Toe III.

In addition to the species described herein, the group
includes four species — E. latens and £. manipus in the Andes
of Colombia, E. elassodisctis in the Cordillera Oriental in
Ecuador, and £. iiigivvittntiis in the upper Amazon Basin
and lower slopes of the Andes in Colombia, Ecuador, and
Peru. (See map in Lynch et al., 1997.)

Eleiithewdactyhis araiodactyhis new species

Holotype.—UF 40764, subadult female, from 24 km [by
road] SW Leimebamba (06°40' S, 77 °50' W, 3370 m),
Provincia Chachapoyas, Departamento Amazonas, Peru,
obtained on 25 April 1972 by Fred G. Thompson.

Diagnosis. — A member of the Eleutherodactylus
(Eleutherodactylus) nigrovittatus Group having (1) skin on
dorsum smooth; that on venter smooth; discoidal fold evi-
dent; dorsolateral folds present; (2) tympanic membrane
smooth, and tympanic annulus prominent, nearly round,
its length about V2 length of eye; (3) snout moderately long,
rounded in dorsal view and in profile; (4) upper eyelid
lacking tubercles, narrower than lOD; cranial crests absent;
(5) vomerine odontophores prominent, elongately oval; (6)
condition of vocal slits anci nuptial pads unknown; (7)
Finger I equal in length to II; discs on outer fingers nar-
row, acutely rounded; (8) fingers bearing lateral fringes;
(9) ulnar tubercles absent; (10) heel and tarsus lacking tu-
bercles; (11) inner metatarsal tubercle elliptical, about 2x
subconical outer metatarsal tubercle; supernumerary plan-
tar tubercles few; (12) toes bearing lateral fringes; webbing
absent; Toes V and III equal in length; discs equal to those
on outer fingers; (13) dorsum tan with middorsal brown
marking; venter and posterior surfaces of thighs tan; (14)
SVL in one female 24.5 mm.

Eleutherodactylus araiodactylus is readily distinguished
from all other members of the genus in the Andes of north-
ern Peru by having a combination of smooth skin on the
venter and undilated discs on the digits. The only other
species in the region having smooth skin on the venter are
members of the Eleutherodactylus conspicillatus Group, all
of which have greatly expanded digital discs. From other
members of the Eleutherodactylus nigrovittatus Group, £.
araiodactylus differs by having Fingers I and II of equal
length (instead of Finger I > II), Toes III and IV of equal
length (instead of Toe III > V), digits acutely rounded in-
stead of nearly pointed, and a diagonal dermal ridge
posteroventral to the tympanum. Moreover, E. araiodactylus
differs from E. elassodiscus by having dorsolateral folds and
lateral fringes on the digits; the latter character also differ-
entiates E. araiodactyhis from E. nigrovittatus, a species with
proportionately shorter fingers and more robust body. The

34

Scientific Papers, Natural History Museum, The University of Kansas

Fig. 14. Dorsal and lateral views of the head of Ekutherodactylus
arniodnctyliis, UF 40764. Scale bar = 5 mm.

presence of lateral fringes on the digits also distinguishes
£. araiodactylus from E. maiiipus.

Description. — (h = 1 female). Head as wide as body.
HW 40.0% SVL; HL 44.0% SVL; Snout moderately long,
rounded in dorsal view and in profile (Fig. 14); E-N 89.7%
length of eye; nostrils slightly protuberant, directed
dorsolaterally; canthus rostralis rounded, slightly curved;
loreal region noticeably concave; lips rounded; upper eye-
lid lacking tubercles; upper eyelid width 71.0%- lOD;
supra tympanic fold moderate; abruptly curving down-
ward behind tympanum, barely obscuring upper edge of
tympanum; side of head nearly vertical; tympanic mem-
brane evident; tympanic annulus distinct; length of tym-
panic annulus 48.3% length of eye; prominent diagonal
dermal ridge (possibly coalesced postrictal tubercles)
postero ventral to tympanum. Choanae small, round, not
concealed by palatal shelf of maxillary arch; vomerine
odontophores prominent, oval, widely separated medially,
located behind posterior edges of choanae, each
odontophore bearing three teeth. Tongue about 1.5x as
long as wide, not notched posteriorly; posterior half not
adherent to floor of mouth.

Skin on dorsum smooth; dorsolateral fold low, extend-
ing from level of axilla to groin; skin on flanks pustular,
venter smooth, except coarsely areolate on proximal
posteroventral surfaces of thighs; discoidal fold prominent;
cloacal sheath short; large tubercles in cloacal region ab-
sent. Ulnar tubercles absent. Thenar tubercle elevated, el-
liptical, about twice size of inner palmar tubercle; palmar
tubercle elevated, completely divided, outer tubercle about
Vi size of inner; palmar supernumerary tubercles absent;
subarticular tubercles large, round; fingers bearing lateral
fringes; first finger equal in length to second. Discs on Fin-
gers 1 and II small, round; discs on all fingers narrow,
round, barely wider than digit proximal to pad (Fig. 15);
Fingers 11-lV having ventral pads weakly defined by cir-
cumferential grooves. Upper surfaces of hind limbs
smooth; heel tarsus lacking tubercles; inner metatarsal tu-

Fig. 15. Hand and foot of Elciitlicmdactylus nmiodnclylus , UF 40764.
Scale = 5 mm.

bercle elevated, elliptical, about 2x subcorneal outer meta-
tarsal tubercle; few plantar supernumerary tubercles on
proximal segments of Toes IV and V; subarticular tubercles
small, subconical; toes bearing lateral fringes; toes not
webbed; discs on toes about equal in size to those on fin-
gers; tips of Toes 111 and V extending to distal edge of
penultimate subarticular tubercle on Toe IV (Fig. 15); when
hind limbs flexed perpendicular to axis of body, heels over-
lap by about '4 length of shank; shank 54.3%- SVL.

Coloration in preservative: Dorsum between dorsolat-
eral folds tan with narrow, median cream stripe from snout
to vent; on body, stripe bordered by irregular brown mark-
ings 2-3x width of stripe (Fig. 16). Brown interorbital bar,
and canthal and supratympanic stripes present; short, dark
brown stripe from upper eyelid extending posteromedially
onto occiput; labial bars absent. Flanks brown; groin un-
marked; forelimbs tan with two brown spots on inner sur-
face of forearm; hind limbs brown with faint, narrow,
darker diagonal marks; venter and posterior surfaces of
thighs uniform tan.

Coloration in life: Unknown.

Measurements of holotype: SVL 24.5, tibia length 13.3,
foot length 13.5, head width 9.8, head length 9.8, lOD 3.1,
upper eyelid width 2.2, E-N 2.6, eye, 2.9, tympanum 1 .4.

Distribution and habitat. — Elcutlicrodaciylii^
araiodactylus is knt)wn only from 3370 m in very humid

Eleutherodactylus in the Andes of Northern Peru

35

montane forest in the northern part of the Cordillera Cen-
tral in Peru (Fig. 13).

Etymology. — The specific name deriveci from the Greek
rrra/c'S, meaning thin, and the Greek dnkti/lofi, meaning fin-
ger or toe. The name applies to the extremely slender dig-
its of this species.

Remarks. — This is one of the most distinctive species
of Elciitlicivdacti/lus in northern Peru; consequently, we
have no qualms about naming it on the basis of a single
specimen. The presence of a middorsal pale stripe is a color
polymorphism in many species of Elciithcwdncti/his: its
presence in the holotype probably is not indicative of the
species.

ElEUTHERODACTi'LUS ORESTES GrOUP

Inclusion in the Eleutherodactylus orestes Group of the
four species described herein necessitates a slight modifi-
cation of the definition of the group as given by Lynch
and Duellman (1997). These are that vocal slits and vomer-
ine odontophores are present (E. orestes, simoiibolivnri, and
vidua), absent (£. melanogaster and £. pataikos), vomerine
odontophores absent and vocal slits present (£. afrnbmchus),
or vomerine odontophores present and vocal slits un-
known (E. pinguis). Furthermore, the distribution of the
group needs to be changed. Members of the group inhabit
high-altitude cloud forests in the Cordillera Occidental in
western Ecuador (£. simonbolivari), paramo in the south-
ern part of the Cordillera Oriental in Ecuador (£. orestes
and £. vidua), and supra-treeline habitats in the Cordillera
Colan (£. atrabracus) and the northern parts of the Cordil-
lera Central (£. melanogaster and E. pataikos) and Cordil-
lera Occidental (£. pinguis) in Peru.

Eleutherodactylus atrabracus new species

Holotype.— LSUMZ 49144, adult female from the Cor-
dillera Colan, east of La Peca (0536' S, 78°22' W, 2963 m),
Provincia Bagua, Departamento Amazonas, Peru, obtained
on 4 September 1978 by Morris D. Williams.

Paratype.— LSUMZ 45090, adult male from the Cor-
dillera Colan, east of La Peca, 3330 m, obtained on 31 Au-
gust 1978 by Thomas S. Schulenberg.

Diagnosis. — A member of the Eleutherodactylus
(Eleutherodactylus) orestes Group having (1) skin on dorsal
surfaces finely shagreen, that on throat, belly, and ventral
surfaces of thighs coarsely areolate; discoidal fold weak
posteriorly; dorsolateral folds absent; (2) tympanic mem-
brane differentiated, tympanic annulus round; its diam-
eter slightly about Vi length of eye; (3) snout moderately
long, bluntly rounded in dorsal v^iew and in profile, in-
clined anteroventrally from level of nostrils; (4) upper eye-
lid narrower than lOD, lacking tubercles; cranial crests
absent; (5) vomerine odontophores absent; (6) vocal slits
present; nuptial excrescences absent; (7)Finger I shorter

than 11; discs on outer fingers rounded, barely wider than
digit proximal to pad; (8) fingers bearing lateral fringes;
(9) ulnar tubercles absent; (10) heel and outer edge of tar-
sus lacking tubercles; low, thick fold on distal part on in-
ner surface of tarsus; (11) inner metatarsal tubercle el-
evated, ovoid, about 5x round outer metatarsal tubercle;
plantar surfaces coarsely areolate; (12) toes bearing lateral
fringes; webbing absent; Toe V slightly longer than III; discs
as large as those on outer fingers; (13) dorsum nearly uni-
form brown; throat and belly creamy tan with fine brown
reticulations; ventral surfaces of hind limbs black; groin
and anterior surfaces of thighs dark brown with large tan
spots; (14) SVL in one adult male 18.9, in one adult female
22.7 mm.

Eleutherodactylus atrabrachus differs from all other
members of the E. orestes Group by having a pale belly
and the ventral surfaces of the hind limbs black; further-
more, it differs from £. pataikos by having large pale spots
in the groin and on the anterior surfaces of the thighs (Fig.
17). Eleutherodactylus atrabracus differs from E. melanogaster
and £. simonbolivari by having a pale, instead of black, belly,
and from E. pinguis by having the ventral surfaces of the
hind limbs black and by lacking vomerine teeth. Two other
species in the group, E. orestes and £. vidua, both of which
have vomerine odontophores, also have pale spots in the
groin; in the latter, the spots are large and lie between di-
agonal brown bars on the posterior half of the flanks, whereas
in E. vidua, the groin is black with small white spots.

Description. — (n = 1 male, 1 female). Head narrower
than body; HW 34.9-36.1 (x = 35.5)% SVL; HL 34.9-40.5 (x
= 37.7)% SVL; snout moderately long, bluntly rounded in
dorsal view; in profile, inclined anteroventrally from level
of nostrils to margin of lip; E-N 90.0-100.0 (x = 95.0)%
length of eye; nostrils slightly protuberant, directed later-
ally; canthus rostralis weakly angular, slightly curved; lo-
real region noticeably concave; lips rounded; upper eye-
lid lacking tubercles; upper eyelid width 71.4-84.0 (x =
77.7)% lOD; cranial crests absent; supratympanic fold
weak, barely obscuring upper edge of tympanum; side of
head nearly vertical; tympanic membrane differentiated;
tympanic annulus round; length of tympanic annulus 39.1-
50.0 (x = 44.6)% length of eye; postrictal tubercles low, dif-
fuse. Choanae small, round, not concealed by palatal shelf
of maxillary arch; vomerine odontophores absent; tongue
much longer than wide, shallowly notched posteriorly, pos-
terior half not adherent to floor of mouth.

Skin on dorsum finely shagreen, that on venter ar-
eolate; discoidal fold barely evident posteriorly; dorsolat-
eral folds absent; cloacal sheath short; enlarged tubercles
in cloacal region absent. Ulnar tubercles absent; thenar
tubercle ovoid, elevated, about half the size of bifid pal-
mar tubercle; supernumerary palmar tubercles round;

36

Scientific Papers, Natural History Museum, The University of Kansas

Fig. 17. Color pattern and skin texture in the groin and on the
anterior surfaces of the thighs in species in the Eleutherodactylus orestes
group. A. E. atrnbmciif, LSUMZ 49144. B. £. mchino'^iistcr, RU 218513.
C. £. pataikos, KU 212320. D. £. pinguis. KU 181283. E. £. simonbolivari,
KU 218253. F. E. uidwfl, KU 120083. G. £. oresfes, KU 141999. Not drawn
to scale.

subarticular tubercles prominent, round; fingers bearing
thick lateral fringes; Finger 1 shorter than II; discs on all
fingers small, only slightly wider than width of digits; all
fingers having ventral pads weakly defined by circumfer-
ential grooves. Upper surfaces of hind limbs coarsely
shagreen; heel and outer edge of tarsus lacking tubercles;
inner edge of tarsus bearing low, thick fold distally; inner
metatarsal tubercle elevated, ovoid, about 5x round outer
metatarsal tubercle; plantar surfaces areolate; subarticular
tubercles, round; toes bearing thick lateral fringes; web-
bing absent; discs on toes about equal in size to those on
fingers; tip of Toe V extending to proximal edge of
penultimate subarticular tubercle on Toe IV; tip of Toe III
not extending to that tubercle; when hind limbs flexed
perpendicular to axis of body, heels barely overlapping;
shank 38.6-40.5 (x = 39.6)% SVL.

Coloration in preservative: Dorsum of head, body, and
limbs nearly uniform brown; faint dark brown canthal and

Fig. 18. Ventral color pattern Eleutherodactylus atrabracus, LSUMZ
49144. SVL = 22.7 mm.

supratympanic stripes present in both specimens; in fe-
male holotype, diffuse brown dorsolateral line and dis-
tinct middorsal tan stripe from tip of snout to vent; inter-
orbital bar, labial bars and transverse bars on limbs ab-
sent. Groin and anterior surfaces of thighs dark brown with
large tan spots (Fig. 17A); throat and belly creamy tan with
fine brown reticulations and narrow transverse line in tho-
racic region; ventral surfaces of hind limbs black (Fig. 18).

Coloration in life: Unknown.

Measurements of holotype: SVL 22.7, tibia length 9.2,
foot length 9.5, head width 8.2, head length 9.2, lOD 22.5,
upper eyelid width 2.1, E-N 2.3, eye 2.3, tympanum 0.9.

Distribution and habitat. — Elciithewdactyliis atralvacus
is known only from two localities at elevations of 2963 and
3330 m on the western slopes of the Cordillera Colan in
northern Peru (Fig. 19). The paratype was in a grassy bog
above treeline.

Etymology. — The specific name is derived from the
Latin adjective, aim, meaning black and the Latin noun,
braca, meaning trousers, and refers to the black ventral
surfaces of the hind limbs in this species.

Eleutherodactylus melaiiogaster new species

Holotype.— KU 212321, adult female, from the north
slope of Abra Barro Negro (06 "41' S, 77°5T W, 3470 m), 28
km [by road] SSW Leimebamba, Provincia Chachapoyas,
Departamento Amazonas, Peru, one of a series collected
on 23 January 1989 by William E. Duellman and John J.
Wiens.

Paratypes.— KU 212322-23 and 218513, adult males,
collected with the holotype; KU 181281, an adult female.

Fl.l-l-rHHROnACTYI.US IN THE AnDES OF NORTHERN PeRU

37

77°

< HXHIm
. KKX>-:!l.>(H) lit
^_7J :(Xtl.l-31XKl ni
Pg^ *000-4l.KX) ni
^m >-)(H>Om

kil.mn'tcrx

• E.

atnilyracifs

ME.

nu'lanogasier

BE.

oresles

AE.
OE.

pataikos
pinfolds

A/:

viilua

v-

,-v '

79

Fig. 19. Localities of known occurrence of six species in tfie
Eleutherodactylus orestes group in the Andes of southern Ecuador and
northern Peru. Another member of the group, E. sinwnbclivari , occurs
farther north in Cordillera Occidental in Ecuador.

from the north slope of Abra Barro Negro, 3300 m, 25.5
km [by road] SSW Leimebamba, Provincia Chachapoyas,
Departamento Amazonas, Peru, collected on 6 March 1979
by William E. Duellman.

Diagnosis. — A member of the Eleutherodactylus
(Eleutherodactylus) orestes Group having (1) skin on most
surfaces coarsely areolate, nearly pustular dorsolaterally;
snout, anterior surfaces of thighs and inner surfaces of
shanks smooth; discoidal fold weak posteriorly, prominent
as transverse thoracic fold anteriorly; dorsolateral folds
absent; (2) tympanum indistinct, tympanic annulus round;
its diameter about 30% length of eye in males, 60% in fe-
males; (3) snout short, rounded in dorsal view and in pro-
file; (4) upper eyelid narrower than lOD, lacking tubercles;
cranial crests absent; (5) vomerine odontophores absent;
(6) vocal slits and nuptial excrescences absent; (7) Finger I
shorter than II; discs on outer fingers rounded, barely wider
than digit proximal to pad; (8) fingers lacking lateral
fringes; (9) ulnar tubercles absent; (10) heel and outer edge
of tarsus lacking tubercles; inner edge of tarsus bearing
low, indistinct fold distally; (11) inner metatarsal tubercle
elevated, ovoid, about 3x round outer metatarsal tubercle;

plantar surfaces areolate; (12) toes lacking lateral fringes;
webbing absent; Toe V slightly longer than III; discs as large
as those on outer fingers; (13) dorsum and venter black
with or without pale spots in groin and on anterior and
posterior surfaces of thighs and inner surfaces of shanks;
(14) SVL in three adult males 20.6-22.8 (x = 21.9 mm), in
two adult females, 24.2-24.7(x= 24.5 mm).

Eleutherodactylus melanogaster differs from all other
members of the E. orestes Group by having the skin on the
dorsum coarsely areolate and by being black dorsally and
ventrally With the exception of the nature of the skin on
the dorsum, £. melanogaster most resembles the smaller £.
swionbolivari, which also has pale spots in the groin and
on the hidden surfaces of the hind limbs, but £.
simonholivari has vomerine odontophores and a small,
nonconical tubercle on the heel. Eleutherodactylus orestes and
£. piiiguis also have pale spots in the groin and on the pos-
terior surfaces of the thighs, but the skin on the dorsum is
finely tuberculate, the dorsum is brown with darker brown
markings, vomerine odontophores are present, and there
is a small, nonconical tubercle on the heel in £. siiuonbolivari.
The sympatric £. pataikos is like E. melanogaster in lacking
vomerine odontophores and a tubercle on the heel, but it
differs by having a smooth dorsum that is pale brown and
by lacking markings in the groin and on the posterior sur-
faces of the thighs. Eleutherodactylus atrabracus also lacks
vomerine teeth, but it has a finely shagreen dorsum, a pale
belly, and black ventral surfaces of the hind limbs.

Description. — iu - 3 males, 2 females; proportions are
for males with followed those of by females). Head nar-
rower than body; HW 36.0-38.3 (x = 36.9), 39.2-39.4 (x =
39.3)% SVL; HL 33.8-36.4 (x = 35.1), 36.4-37.2 (x = 36.8)%
SVL; snout short, rounded in dorsal view, and in profile;
E-N 64.0-71.4 (x = 67.8), 80.0-83.0 (x = 81.5) % length of
eye; nostrils slightly protuberant, directed dorsolaterally;
canthus rostralis rounded, curved; loreal region weakly
concave; lips rounded; upper eyelid lacking tubercles;
upper eyelid width 68.0-71.4 (x = 70.3), 75.9-76.7 (x =
76.3)% lOD; cranial crests absent; supratympanic fold a
series of pustular granules, obscuring upper edge of tym-
panum; side of head nearly vertical; tympanum indistinct;
tympanic annulus thin; tympanic membrane thickened;
tympanic annulus round; length of tympanic annulus 33.3-
44.0% (37.8), 60.0-66.7 (63.3) % length of eye; postrictal
tubercles not discernible from other pustules; skin on top
of head smooth. Choanae small, round, not concealed by
palatal shelf of maxillary arch; vomerine odontophores
absent; tongue much longer than wide, not notched pos-
teriorly, posterior half not adherent to floor of mouth. Males
lacking vocal slits and nuptial excrescences.

Skin on dorsum coarsely areolate, especially
dorsolaterally; that on venter coarsely areolate; discoidal

38

SciENTinc Papers, Natural History Museum, The University of Kansas

Fig. 20. Hand and foot of Eleidherodach/lus melanogaster, KU 212321.
Scale = 5 mm.

fold barely evident posteriorly, present as distinct trans-
verse thoracic fold anteriorly; dorsolateral folds absent;
flanks coarsely areolate; cloacal sheath short; enlarged tu-
bercles in cloacal region absent. Ulnar tubercles not dis-
tinguishable; thenar tubercle ovoid, slightly elevated, about
as large as ovoid palmar tubercle; supernumerary palmar
tubercles not evident; subarticular tubercles prominent,
round; fingers lacking lateral fringes; first finger shorter
than second; discs on all fingers small, only slightly wider
than width of digits (Fig. 20); all fingers having ventral
pads weakly defined by circumferential grooves. Upper
surfaces of hind limbs coarsely areolate; heel and outer
edge of tarsus lacking tubercles; inner edge of tarsus with
low fold distally; inner metatarsal tubercle elevated, ovoid,
about 3x round outer metatarsal tubercle; plantar surfaces
areolate; subarticular tubercles large, round; toes lacking
lateral fringes; webbing absent; discs on toes about equal
in size to those on fingers; tip of Toe V extending to point
midway between penultimate and distal subarticular tu-
bercles on Toe IV; tip of Toe 111 extending to distal edge of
penultimate tubercle on Toe IV (Fig. 20); when hind limbs
flexed perpendicular to axis of body, heels not overlap-
ping; shank 36.4-39.3 x = 37.5), 37.1-37.7 (x = 37.4) % SVL.

Coloration in preservative: Female holotype and one
male (KU 212323) black above and below with small, dif-

fuse white spots in groin, on anterior and posterior sur-
faces of thighs (Fig. 17B), and on inner surfaces of shanks;
spots absent in other female (KU 181281) and in two males
(KU 212322, 218513).

Coloration in life: KU 212321 : Dorsum dull brown with
creamy tan canthal stripe bordered below by black line
(Fig. 9); black postocular stripe; belly dark gray; axilla,
groin, and ventral surfaces of limbs black with bright yel-
low spots; iris pale gold with black flecks (WED field notes,
23 January 1989).

Measurements of holotype: SVL 24.7, tibia length 9.3,
foot length 10.5, head width 9.7, head length 9.0, lOD 3.0,
upper eyelid width 2.3, E-N 2.0, eye 2.4, tympanum 1.6.

Distribution and habitat. — Eleutherodnctylus
melanogaster is known only from two localities at eleva-
tions of 3300 and 3470 m, respectively, on the road from
Balsas to Leimebamba, near the crest of the northern part
of the Cordillera Central in northern Peru (Fig. 19). All in-
dividuals were under stones by day; those from 3470 m
were in wet paramo, whereas the individual from 3300 m
was in elfin forest; both habitats fall within the very hu-
mid montane forest

Etymology. — The specific name is derived from the
Greek adjective melanos meaning black and the Green noun
gnster meaning belly; the name is used in reference to the
uniformly black ventral coloration.

Eleiitherodacti/lus pataikos new species

Holotype.— KU 212320, adult female, from the north
slope of Abra Barro Negro, (06°41' S, 77 °51' W, 3470 m), 28
km [by road] SSW Leimebamba, Provincia Chachapoyas,
Departamento Amazonas, Peru, obtained on 23 January
1989 by William E. Duellman.

Diagnosis. — A member of the Eleutherodactylus
(Eleutherodactyhis) orestes Group having (1) skin on dor-
sum smooth with few low tubercles dorsolaterally and on
hind limbs, that on venter coarsely areolate; discoidal fold
weak; dorsolateral folds absent; (2) tympanum indistinct,
tympanic annulus distinguishable anteroventrally, round;
its diameter less than Vi length of eye; (3) snout short,
rounded in dorsal view and in profile; (4) upper eyelid
narrower than lOD, lacking tubercles; cranial crests absent;
(5) vomerine odontophores absent; (6) condition of vocal
slits and nuptial excrescences unknown; (7) Finger 1 shorter
than 11; discs on outer fingers rounded, barely wider than
digit proximal to pad; (8) fingers lacking lateral fringes;
(9) ulnar tubercles absent; (10) heel and outer edge of tar-
sus lacking tubercles; inner edge of tarsus bearing single,
low tubercle; (11 ) inner metatarsal tubercle flat, ovoid about
4x round outer metatarsal tubercle; supernumerary tu-
bercles low, diffuse; (12) toes lacking lateral fringes; web-
bing absent; Toe V slightly longer than 111; discs as large as

Eleutherodactylvs in the Andes of Northern Peru

39

Fig. 21. Dorsal and lateral views of the head of Eteutherodnciyhts
piifrti'tos, KU 212320. Scale bar = 5 mm.

those on outer fingers; (13) dorsum uniform pale brown;
venter tan; posterior surfaces of thighs brown; (14) SVL in
one adult female 21.6 mm.

Eleutherodachjliis pntniko? differs from other members
of the £. orestcs Group by lacking distinctive markings.
Eleutherodactylus atrabracus, orestes, pinguis, simonboUvari,
and uiflaiiogaster have large pale spots in the groin, and
the latter, which occurs sympatrically with £. pataikos, has
coarsely areolate skin on the dorsum, which, like the ven-
ter, is black. Eleutherodactylus pinguis and E. atrabracus also
differ by having a prominent tympanum and tympanic
annulus; the former also differs by having the ulnar tu-
bercles coalesced into a fold, and the latter also differs by
having a pale belly and black ventral surfaces of the hind
limbs. Eleutherodactylus orestes and E. vidua have cream
spots at least distally on the posterior surfaces of the thighs,
which are uniform pale brown in E. pataikos; the latter (£.
pataikos) also differs by lacking vomerine odontophores, a
feature shared only with E. atrabracus and E. melanogaster
in the £. orestes Group.

Description. — (n = 1 female). Head narrower than
body; HW 39.4% SVL HL 36.1 % SVL; snout short, rounded
in dorsal view, and in profile (Fig. 21); E-N 71.4% length
of eye; nostrils slightly protuberant, directed dorsolaterally;
canthus rostrahs rounded, nearly straight; loreal region
weakly concave; lips rounded; upper eyelid lacking tu-
bercles; upper eyelid width 70.8% lOD; cranial crests ab-
sent; supratympanic fold diffuse, obscuring upper edge
of tympanum; side of head nearly vertical; tympanic an-
nulus thin, evident only anteroventrally; tympanic mem-
brane not pustular or thickened; tympanic annulus round;
length of tympanic annulus 33.3%- length of eye; two large,
round postrictal tubercles, posteroventral to tympanic an-
nulus; skin on head smooth. Choanae small, round, not
concealed by palatal shelf of maxillary arch; vomerine
odontophores absent; tongue much longer than wide, not

notched posteriorly, posterior half not adherent to floor of
mouth. Males unknown.

Skin on dorsum smooth with few low tubercles
dorsolaterally and on hind limbs, that on venter coarsely
areolate; discoidal fold weak; dorsolateral folds absent;
flanks tuberculate; cloacal sheath short and enlarged tu-
bercles in cloacal region absent; skin on throat smooth, on
belly coarsely granular; discoidal fold weak. Ulnar tu-
bercles absent; thenar tubercle ovoid, flat, about as large
as ovoid palmar tubercle; supernumerary palmar tubercles
few, large, round; subarticular tubercles prominent, round;
fingers lacking lateral fringes; Finger 1 shorter than II; discs
on all fingers small, only slightly wider than width of dig-
its; all fingers having ventral pads weakly defined by cir-
cumferential grooves. Upper surfaces of hind limbs smooth
with scattered small tubercles; heel and outer edge of tar-
sus lacking tubercles; inner edge of tarsus bearing small,
indistinct tubercle; inner metatarsal tubercle flat, ovoid
about 4x round outer metatarsal tubercle; supernumerary
tubercles low, diffuse; subarticular tubercles large, round;
toes lacking lateral fringes; webbing absent; discs on toes
about equal in size to those on fingers; tip of Toe V extend-
ing to point midway between penultimate and distal
subarticular tubercles on Toe IV; tip of Toe 111 extending to
distal edge of penultimate tubercle on Toe IV; when hind
limbs flexed perpendicular to axis of body, heels not over-
lapping; shank 37.0% SVL.

Coloration in preservative: Dorsum, flanks and hid-
den surfaces of limbs dull, pale brown; venter, dull tan.
No markings evident (Fig. 17C).

Coloration in life: Dorsum pale brown with narrow,
diffuse cream stripe on canthus and outer edge of eyelid,
and narrow, dark brown supratympanic stripe (Fig. 9);
venter creamy gray; groin dark gray; iris dull bronze with
black flecks.

Measurements of holotype: SVL 21.6, tibia length 8.0,
foot length 8.8, head width 8.5, head length 7.8, lOD 2.4,
upper eyelid width 1.7, E-N 1.5, eye 2.1, tympanum 0.7.

Distribution and habitat. — The species is known only
from one locality at 3470 m on the road from Balsas to
Leimebamba, near the crest of the northern part of the
Cordillera Central in northern Peru (Fig. 19). The only
known specimen, an adult female, was under a rock in
very humid montane forest by day.

Etymology. — The specific name pataikos is a Greek
noun in apposition for an odd-shaped dwarflike
Phoenician deity. The name refers to the small but
Rubenesque stature of this frog.

Remarks. — Loath though we are to describe a new
species of Eleutherodactylus from a single specimen, this
individual certainly is distinctive in the Eleutherodactylus
orestes Group.

40

Scientific Papers, Natural History Museum, The University of Kansas

Eleuthewdactyhis pingtiis new species

Holotype.— KU 181283, an adult female from 23 km
[by road] SW Celendin (06°58' S, 78°06' W, 3050 m),
Provincia Celendin, Departamento Cajamarca, Peru, one
of a series collected by David C. Cannatella, William E.
Duellman, and Thomas J. Berger on 8 March 1979.

Paratypes.— KU 181282 collected with the holotype,
and KU 181284 from 33 km [by road] SW Celendin, 3200
m, Provincia Celendin, Departamento Cajamarca, Peru.

Referred specimen. — UP 40766 from 57 km [by road]
N Cajamarca, 3760 m, Provincia Cajamarca, Departamento
Cajamarca, Peru.

Diagnosis. — A member of the Eleutherodacti/liis
{Ekutherodacti/lus) oresfes Group having (1) skin on dorsal
surfaces finely areolate, that on throat, belly, and ventral
surfaces of thighs coarsely areolate; discoidal fold weak
posteriorly, prominent as transverse thoracic fold anteri-
orly; dorsolateral folds absent; (2) tympanum membrane
differentiated, tympanic annulus round; its diameter
slightly more than Vi length of eye; (3) snout moderately
long, acutely rounded in dorsal view and bluntly rounded
in profile; (4) upper eyelid narrower than lOD, lacking
tubercles; cranial crests absent; (5) vomerine odontophores
prominent, oblique; (6) condition of vocal slits and nup-
tial excrescences vmknown; (7) Finger I shorter than II; discs
on outer fingers rounded, barely wider than digit proxi-
mal to pad; (8) fingers lacking lateral fringes; (9) ulnar tu-
bercles coalesced into short fold; (10) heel and outer edge
of tarsus lacking tubercles; low, thick fold on distal part on
inner surface of tarsus; (11) inner metatarsal tubercle flat,
ovoid, about 3x subconical outer metatarsal tubercle; plan-
tar surfaces coarsely areolate; (12) toes bearing lateral
fringes; webbing absent; Toe V slightly longer than 111; discs
as large as those on outer fingers; (13) dorsum brown; ven-
ter tan with brown flecks or reticulations; groin and proxi-
mal anterior surfaces of thighs dark brown with large
cream spots; (14) SVL in three adult females 28.4-29.8 (x =
28.9 mm).

Ekutherodactylns pingitis differs from other members
of the E. orestes Group by having the ulnar tubercles coa-
lesced into a low fold; it differs from E. mclanogaster and £.
simonbolivari by having a pale, as opposed to a dark, ven-
ter, and from £. atrabracus, melanogaster, and pataikos by
having vomerine teeth. Eletithcwdncti/liis piiigiiis also dif-
fers from £. pataikos by having a prominent tympanum and
tympanic annulus and distinctive markings in the groin
and pale spots on the posterior surfaces of thighs, which
are uniform pale brown in £. pataikos. Two other species
in the group, E. orestes and E. vidua, also have pale spots in
the groin; in the latter, the spots are large and lie between
diagonal brown bars on the posterior half of the flanks.

whereas in £. vidua, the groin is black with small white
spots.

Description. — (n = 3 females). Head narrower than
body; HW 36.5-38.0 (x =37.1)% SVL; HL 34.2-36.1 (x =
35.1)% SVL; snout moderately long, acutely rounded in
dorsal view, and bluntly rounded in profile; E-N 75.9-92.8
(x= 83.0)% length of eye; nostrils slightly protuberant, di-
rected dorsolaterally; canthus rostralis weakly angular,
slightly curved; loreal region noticeably concave; lips
rounded; upper eyelid lacking tubercles; upper eyelid
width 56.4-65.7 (x = 59.7)% lOD; cranial crests absent;
supratympanic fold weak, granular, obscuring upper edge
of tympanum; side of head nearly inclined ventrolaterally;
tympanic membrane weakly differentiated; tympanic an-
nulus round; length of tympanic annulus 53.6-55.2 (x =
54.6)% length of eye; postrictal tubercles not discernible
from other tubercles. Choanae small, round, not concealed
by palatal shelf of maxillary arch; vomerine odontophores
prominent, oval in outline, oblique, widely separated me-
dially, located behind posterior edges of choanae, each
odontophore bearing 3 or 4 (x = 3.8 teeth); tongue much
longer than wide, shallowly notched posteriorly, posterior
half not adherent to floor of mouth. Males unknown.

Skin on dorsum finely areolate, more coarsely areolate
laterally; that on venter coarsely areolate; discoidal fold
barely evident posteriorly, present as distinct transverse
thoracic fold anteriorly; dorsolateral folds absent; cloacal
sheath short; enlarged tubercles in cloacal region absent.
Ulnar tubercles coalesced so as to form low ridge distally
on forearm; thenar tubercle ovoid, slightly elevated, some-
what smaller than bifurcate palmar tubercle; supernumer-
ary palmar tubercles not evident; subarticular tubercles
prominent, round; fingers bearing thick lateral fringes; Fin-
ger I shorter than 11; discs on all fingers small, only slightly
wider than width of digits; all fingers having ventral pads
weakly defined by circumferential grooves. Upper surfaces
of hind limbs coarsely areolate; heel and outer edge of tar-
sus lacking tubercles; inner edge of tarsus bearing low, thick
fold distally; inner metatarsal tubercle flat, ovoid, about
3x subconical outer metatarsal tubercle; plantar surfaces
areolate; subarticular tubercles large, round; toes bearing
thick lateral fringes; webbing absent; discs on toes about
equal in size to those on Angers; tip of Toe V extending to
distal edge of penultimate subarticular tubercle on Toe IV;
tip of Toe 111 not extending to that tubercle; when liind
limbs flexed perpendicular to axis of body heels not o\'er-
lapping; shank 35.4-40.4 (x= 38.1)% SVL.

Coloration in preservative: Dorsum of head, body, and
limbs brown with faintly darker brown markings (absent
in KU 181284); markings consisting of faint interorbital
mark, numerous small irregular flecks on body and faint

Eleuthikoiimtyi.I'S in the Andes of Northern Peru

41

diagonal crossbar on shank. Dark brown markings in groin
and on proximal anterior surfaces of thighs, nearly enclos-
ing large cream spots (Fig. 17D); distal portion of poste-
rior surfaces of thighs and inner surfaces of shanks cream
with large, dark brown markings. Venter creamy tan with
minute brown flecks on throat and bellv in KU 181282,
larger clusters of flecks, especially on belly, in KU 181283,
and fine brown reticulations in KU 181284.

Coloration in life: KU 181284: Dorsum dull red becom-
ing dull green on upper flanks; venter dull yellow reticu-
lated with grayish brown; iris dull reddish bronze (Fig. 9;
WED field notes, 08 March 1989).

Measurements of holotype: SVL28.4, tibia length 10.7,
foot length 11.4, head width 10.8, head length 9.7, lOD 3.5,
upper evelid width 2.2, E-N 2.2, eye 2.9, tympanum 1.6.

Distribution and habitat. — Elctitherodactyhis pinguis is
known from three localities in the northern part of the
Corciillera Occidental in Peru (Fig. 19). Two localities are
at elevations of 3050 and 3200 m on the road between
Cajamarca and Celendfn, on the eastern slopes of Abra
Comulica. All indi\'iduals were under stones by day. Those
from 3050 m were in a wet grassy area, whereas the indi-
vidual from 3200 m was in a bunch grass-Baccharis asso-
ciation. Another specimen is from an elevation of 3760 m
north of Cajamarca; it was under a rock in a grassy region
by day.

Etymology. — The specific name is a Latin adjective
meaning fat; the name refers to the corpulent habitus of
this species.

Remarks. — A subadult female (UF 40766) having a
snout-vent length of 24.0 mm is like the type series in struc-
tural features. The color pattern is faded, so there are no
markings evident in the groin or on the thighs; no dark
flecks are present on the venter.

ElEUTHERODACTYLUS UNISTRIGATUS GrOUP

As noted by Lynch and Duellman (1997), members of
this group are characterized by areolate skin on the venter
and Toe V being much longer than Toe III. This is the larg-
est group of Eleiitherodactyhis and now contains more than
175 species, of which 31 are known from the Andes in
northern Peru.

Eleiitherodnctylus acimiiimtiis Shreve
Eleuthewdactyhis acuminatus Shreve, 1935:217. Holotype, MCZ 19951 from
Canelos, Provincia Pastaza, Ecuador.

Diagnosis. — A member of the Eleiithcrodncti/liis
(Eleuthewdacti/lus) iinistrigatus Group having (1) skin on
dorsum smooth, that on belly and proximal posteroventral
surfaces of thighs coarsely areolate; skin on other ventral
surfaces smooth; discoidal fold barely evident posteriorly;
dorsolateral folds absent; (2) tympanic membrane not dif-
ferentiated; tympanic annulus distinguishable under skin.

'-'■'-' ^ 7')" ■', 78"

77" y.(

▲

,'^

/

'A /

?" , — -

A ;
\ :
V\

^~'r

y

Y\

■■- IIMIdiU

lOIKI :i)(H.im
:iHH) lIMNlm
i llMHl 4(MH)in
Mi >4U(K)m
50 IIMl

KilumcU'ts

1

/

# E. iicumimitus
O E. anemcius
■ E. anlalonychus
D E. hciirsei
▲ E. bromeliaceus
A E. cajamarcensis

4

v-'->

(

\

r

) V,

A "l

\/ 74° 7S°

Fig. 22. Localities of known occurrence of six species in tlie
Eleiithcwdactylus unistrigntiis group in the Andes of southern Ecuador
and northern Peru.

round, its length less than Vi length of eye; (3) snout long,
acuminate in dorsal view, tmncate and posteriorly inclined
in profile; (4) upper eyelid lacking tubercles, much nar-
rower than lOD; cranial crests absent; (5) vomerine
odontophores low, ovoid; (6) males lacking vocal slits and
nuptial pads; (7) Finger I shorter than II; discs on outer
fingers elliptical; (8) fingers lacking lateral fringes; (9) ul-
nar tubercles absent; (10) heel lacking tubercles; outer edge
of tarsus bearing two or three low subconical tubercles;
inner edge of tarsus lacking tubercles or fold; (11) inner
metatarsal tubercle elevated, elliptical, about 4x conical
outer metatarsal tubercle; plantar supernumerary tubercles
absent; (12) toes lacking lateral fringes; webbing absent;
Toe V much longer than III; discs nearly as large as those
on outer fingers; (13) dorsum creamy tan (green in life)
with black canthal and supratympanic stripes and narrow
interorbital bar; limb bars absent; venter and hidden sur-
faces of thighs cream; (14) SVL in males 17.1-22.6 mm, in
females 25.6-31.3 (Lynch, 1980).

This nearly uniformly green species with an acuminate
snout, broad head, and concealed tympana is readily dis-
tinguished from all other members of the genus in the re-
gion. The only other predominately green species is £. galdi,

42

SciENTinc Papers, Natural History Museum, The University of Kansas

which has flared lips, cranial crests, distinct tympanum,
large truncate discs, conical tubercles along the tarsus, and
bars on the limbs.

Description. — The description by Shreve (1935) was
augmented by Duellman (1978c).

Distribution and habitat. — Tliis primarily Amazonian
species invades the lower reaches of the Andes in south-
ern Ecuador and northern Peru. It is known from San Jose,
830 m, Provincia Morona-Santiago, Ecuador (KU 147976)
on the western slopes of the Cordillera del Condor and
from 15.4 km [by road] SW Zapateros, 950 m, Provincia
Lamas, Departamento San Martin, Peru (KU 217308-10)
on the eastern slopes of the Cordillera Central (Fig. 22).
The latter were in arboreal bromeliads by day.

Remarks. — Two of the specimens from the Cordillera
Central are males with SVLs of 21 .3 and 21 .5 mm; the other
is a female having a SVL of 24.3 mm. Although this spe-
cies is abundant in the lowland rainforest in Amazonian
Ecuador, it has been reported only twice from Amazonian
Peru — Rio Aguayita, Departamento Loreto (Lynch, 1980),
and San Jacinto, Departamento Loreto (Duellman and
Mendelson, 1995). Additional records are from
Departamento San Martin — Rio Cainarache, 330 m, 33 km
[by road] NE Tarapoto (KU 209466), and Rio Shilcayo, 500
m, near Tarapoto (KU 209467).

Eleutherodactylus anetnerus new species

Holotype.— KU 219798, adult male, from El Tambo
(05°21' S, 79°33' W, 2770 m, 31 km [by road] ENE of
Canchaque, Provincia Huancabamba, Departamento
Piura, Peru, obtained on 9 January 1991 by Erik R. Wild.

Diagnosis. — A member of the Eleuiherodactylus
(Ekutherodactylits) unistrigatus Group having (1) skin on
dorsum finely tuberculate, that on venter areolate; discoi-
dal fold prominent; dorsolateral folds absent; (2) tympanic
membrane and tympanic annulus prominent, round, its
length slightly more than Vi length of eye; (3) snout short,
acuminate in dorsal view, truncate and posteriorly inclined
in profile; tubercle on tip of snout; canthus rostralis
rounded; (4) upper eyelid lacking tubercles, narrower than
lOD; cranial crests absent; (5) vomerine odontophores
absent; (6) males having vocal slits but lacking nuptial
pads; (7)Finger I shorter than 11; discs moderately smaU,
nearly truncate; (8) fingers bearing narrow lateral fringes;
(9) ulnar tubercles low, diffuse; (10) heel lacking tubercles;
outer edge of tarsus bearing low tubercles; inner edge of
tarsus lacking tubercles or fold; (11) inner metatarsal tu-
bercle broadly ovoid, about 5x subconical outer metatar-
sal tubercle; plantar supernumerary tubercles few, basal
only; (12) toes bearing narrow lateral fringes; webbing
absent; Toe V much longer than 111; discs about equal in
size to those on outer fingers; (13) dorsum uniform yel-

lowish tan; venter creamy white with minute black flecks;
(14) SVL in male 20.4 mm.

No other species of Eleutherodactylus in the region has
an unmarked orange-red dorsum and yellow flanks. The
dorsal coloration is reminiscent of some individuals of £.
chakeus, a member of the Eleutherodactylus diastema Group
in Chocoan Colombia and Ecuador. (See Lynch and
Duellman, 1997:plate 8.) However, £. chakeus differs from
£. anemerus in a number of features — areolate skin on the
dorsum, absence of a tympanum, papillate discs on the
fingers, and Toe V shorter than Toe 111. Five other species
of Eleutherodactylus in the Andes of northern Peru lack
vomerine odontophores; three of these (E. ntrabracus,
melanogaster, and pataikos) robust-bodied members of the
E. orestes Group that have small terminal discs on the fin-
gers, short hind limbs, and Toe V only slightly longer than
Toe 111. The other two (£. iiicomptus and £. percnopterus)
are members of the E. unistrigatus Group and have dis-
tinctive dark markings on the dorsum.

Description. — (ii - 1 male). Head as wide as body; HW
36.3% SVL; HL 35.3% SVL. Snout moderately short, acumi-
nate in dorsal view, truncate and slightly incline
posteroventrally in profile, with subconical tubercle on tip;
E-N = length of eye; nostrils noticeably protuberant, di-
rected dorsolaterally; canthus rostralis rounded, slightly
curved; loreal region weakly concave; lips rounded; up-
per eyelid width 77.0% lOD, lacking tubercle; cranial crests
absent; supratympanic fold weak, obscuring upper edge
of tympanic annulus; side of head inclined ventrolater-
ally; tympanic membrane present; tympanic annulus
round; length of tympanic annulus 55.0%' length of eye;
three small, subconical postrictal tubercles below and
posteroventral to tympanum. Choanae small, round, not
concealed by palatal shelf of maxillary arch; vomerine
odontophores absent; tongue nearly twice as long as wide,
distinctly notched posteriorly posterior half not adherent
to floor of mouth; vocal slits small, extending
posterolaterally from lateral base of tongue; vocal sac
single, median, subgular.

Skin on dorsum finely tuberculate; dorsolateral fold
absent; skin on throat, belly, and ventral surfaces of thighs
areolate; skin on other ventral surfaces smooth; discoidal
fold prominent; cloacal sheath short; large tubercles in anal
region absent. Ulnar tubercles low, diffuse; thenar tubercle
elevated, elliptical, slightly larger than bifurcate palmar
tubercle; palmar supernumerary tubercles absent;
subarticular tubercles moderately small, round; fingers
bearing narrow lateral fringes; first finger shorter than
second; discs on fingers nearly truncate, nearly IVix width
of digit proximal to pad; all fingers having ventral pads
well defined by circumferential grooves. Upper surfaces
of hind limbs smooth with small tubercles; heel lacking

ELEUTHhRonAcni.iis IN THE Andes of Northern Peru

43

tubercle; outer edge of tarsus bearing low tubercles; inner
edge of tarsus lacking tubercles or fold; inner metatarsal
tubercle elevated broadly ovoid, about 5x subconical outer
metatarsal tubercle; plantar supernumerary tubercles few,
basal; discs on toes about equal in size to those on fingers;
toes bearing narrow lateral fringes; toes unwebbed; tip of
Toe V extending to distal edge of distal subarticular tu-
bercle on Toe IV; tip of Toe 111 extending to middle of
penultimate subarticular tubercle on Toe IV; when hind
limbs flexed perpendicular to axis of body, heels broadly
overlapping; shank 52.5% SVL.

Coloration in preservative: Dorsum uniform tan; can-
thai and supratvmpanic stripes, labial and interorbital bars,
and transverse marks on limbs absent; venter creamy tan
with minute black flecks on throat and belly.

Coloration in life: Dorsum of head, body, and limbs
orange-red; flanks, ventral surfaces of limbs, upper lips,
loreal region, tympanum, and vocal sac yellow; rest of
venter cream; tubercles on dorsum, flanks, and venter
white; iris bronze with black reticulations and median,
horizontal red streak (Fig. 9; Erik R. Wild, field notes, 8
January 1991).

Measurements of holotype: SVL 20.4, tibia length 10.7,
foot length 9.9, head width 7.5, head length 7.2, lOD 2.6,
eyelid width 2.0, E-N 2.0, eye 2.0, tympanum 1.1.

Distribution and habitat. — This species is known only
from 2770 m in humid montane forest on the western
slopes of the Cordillera de Huancabamba (Fig. 22). The
male was calling from a red leaf 0.5 m above the ground
at night.

Etymology. — The specific epithet is a Greek adjective,
auenieros, meaning wild. The name is used in reference to
Erik R. Wild, collector of the only known specimen.

Remarks. — Duellman and Wild (1993) recognized this
species as being distinct from others in the Cordillera de
Huancabamba and were reluctant to name the species on
the basis of a single specimen. After our review of the
Eleutherodactyhis of the Andes of northern Peru, we are
convinced that the specimen cannot be associated with
any species described previously.

Eleiitherodacti/lus ardalonychus new species

Holotype. — KU 212301, an adult female, from the Rio
Cerranayacu (05°46' S, 77°27' W, 1200 m), 76 km [by road]
NW Rioja, Provincia Rioja, Departamento San Martin,
Peru, obtained on 2 February 1989 by William E. Duellman.

Paratypes. — KU 212299, adult male, from the west
slope of Abra Tangarana, 1080 m, 7 km NE [by road] San
Juan de Pacaysapa, Provincia Lamas, Departamento San
Martin, Peru; KU 212300, adult male, from 8 km [by road]
NE Tarapoto, 680 m, Provincia San Martin, Departamento
San Martin, Peru.

Referred specimen. — KU 212310, juvenile, from the
type locality.

Diagnosis. — A member of the Elcuthcrodactylus
(Elciiihcrodaciylus) unistrigntus Group having (1) skin on
dorsum smooth with scattered low tubercles on posterior
part of body, that on venter weakly areolate; discoidal fold
barely evident posteriorly; dorsolateral folds absent; (2)
tympanic membrane not differentiated; tympanic annu-
lus distinguishable under skin, round, its diameter about
V2 length of eye; (3) snout long, rounded in dorsal view
and in profile; (4) upper eyelid narrower than lOD, lack-
ing tubercles; cranial crests absent; (5) vomerine
odontophores low, ovoid; (6) males having vocal slits and
unpigmented nuptial pads; (7) Finger 1 shorter than II;
discs on outer fingers rounded, about twice width of digit
proximal to pad; (8) fingers bearing narrow lateral fringes;
(9) ulnar tubercles absent; (10) heel and outer edge of tar-
sus lacking tubercles; inner edge of tarsus bearing low,
elongate tubercle; (11) inner metatarsal tubercle elevated,
elliptical, about lOx low, round outer metatarsal tubercle;
supernumerary plantar tubercles numerous; (12) toes bear-
ing narrow lateral fringes; webbing absent; Toe V much
longer than 111; discs nearly as large as those on outer fin-
gers; (13) dorsum brown with darker brown markings;
venter tan with dark brown flecks or reticulations; poste-
rior surfaces of thighs brown, with or without cream flecks
or mottling; (14) SVL in two adult males 19.7 and 21.9 mm
and an adult female 27.4 mm.

The only other species in the region that can be con-
fused with Eleiitberodactylus ardalomjchus is £. versicolor,
which also has diagonal bars on the flanks and brown re-
ticulations on the venter, but the reticulations are much
coarser than those in £. ardaloin/diiis. Furthermore, E. ver-
sicolor differs by having a shagreen dorsum with scattered
fine tubercles, a prominent tympanum, and elliptical, in-
stead of rounded, discs on the outer fingers.

Description. — {n = 2 males, 1 female; proportions are
for the males with mean in parentheses, followed by those
of the female). Head noticeably wider than body; HW 37.4-
38.1 (x = 37.8), 38.0% SVL; HL 38.4-39.1 (x = 38.8), 39.1%
SVL; snout long, rounded in dorsal view and in profile;
E-N 92.3-100.0 (x = 96.2), 96.8% length of eye; nostrils
slightly protuberant, directed laterally; canthus rostralis
rounded, nearly straight; loreal region concave; lips not
flared; upper eyelid with lacking tubercles; upper eyelid
width 73.9-76.0 (x = 75.0), 70.8% lOD; cranial crests ab-
sent; supratympanic fold weak, not obscuring upper edge
of tympanic annulus; side of head nearly vertical; tym-
panic annulus thin; tympanic membrane not pustular or
thickened; tympanic annulus round; diameter of tympanic
annulus 40.0, 41.7% length of eye; single, enlarged
postrictal tubercle, posteroventral to tympanic annulus;
skin on head smooth except for single, rounded tubercle

44

SciENTinc Papers, Natural History Museum, The University of Kansas

between orbits in KU 212301; choanae large, nearly round,
not concealed by palatal shelf of maxillary arch; vomerine
odontophores, low, oblique, posteromedian to choanae,
oval in outline, each slightly smaller than choanae, sepa-
rated medially by distance greater than width of
odontophore, bearing 2-2 or 2-3 teeth in males, 3-4 in fe-
male; tongue longer than wide, its posterior border shal-
lowly notched, posterior half not adherent to floor of
mouth. Male with vocal slits extending posterolaterally
from midlateral base of tongue; vocal sac single, median,
subgular; unpigmented nuptial pads present.

Dorsum of head, body, and limbs smooth with scat-
tered low tubercles on posterior part of body, small,
subconical tubercles; dorsolateral folds absent; flanks
finely tuberculate; cloacal sheath short; skin on throat
weakly areolate, on belly coarsely areolate; discoidal fold
barely evident posteriorly. Uhiar tubercles absent; thenar
tubercle ovoid, about Vi size of bifid palmar tubercle; su-
pernumerary palmar tubercles few, minute; subarticular
tubercles prominent, round; fingers bearing distinct lat-
eral fringes; Finger I shorter than II; disc on thumb barely
expanded; disk on Finger II slightly larger; discs on Fin-
gers III-IV broadly rounded, more than twice width of
digits; all fingers having ventral pads defined by circum-
ferential grooves. Upper surfaces of hind limbs smooth;
heel and outer edge of tarsus lacking tubercles; inner edge
of tarsus bearing low, elongate tubercle; inner metatarsal
tubercle elevated, elliptical, about lOx low, round, outer
metatarsal tubercle; supernumerary plantar tubercles nu-
merous; subarticular tubercles round, subconical; toes
bearing narrow lateral fringes; webbing absent; discs on
toes equal in size to those on fingers; tip of Toe V extend-
ing to distal edge of distal subarticular tubercle on Toe IV;
tip of Toe III extending to distal edge of penultimate
subarticular tubercle on Toe IV; when hind limbs flexed
perpendicular to axis of body, heels barely overlapping;
shank 51.8-52.5 (x= 51.2), 50.0% SVL.

Coloration in preservative: Dorsum tan with dark
brown markings — KU 212264; elongate mark on snout,
interorbital bar, canthal stripe. Supratympanic stripe, la-
bial bars, W-shaped mark in scapular region (absent in KU
212229); chevron in sacral region (absent in KU 212229 and
212310). Diagonal bars on flanks and limbs, digits cream
with dark brown transverse bars, discs cream with dark
brown on distal edges. Posterior surfaces of thighs brown
with cream mottling in KU 212229-300 and cream flecks
in KU 212310. Venter creamy tan with fine dark brown
reticulations (brown flecks in KU 212301).

Coloration in life: KU 212301, female: Dorsum reddish
brown with dark brown marking; venter and anterior and
posterior surfaces of thighs gray; iris dull bronze with
median, horizontal, red streak (Fig. 9). KU 212299, male:

Dorsum tan, black dorsolaterally with greenish-tan bars
on flanks; dorsal surfaces of limbs olive-brown with yel-
lowish-tan bars; axilla, groin, and ventral surfaces of hind
limbs salmon; throat and belly dull yellow with black
flecks; salmon suffusion on belly; iris reddish copper. KU
212300, male: Dorsum mottled reddish brown and dark
brown; flanks cream with diagonal cream bars; venter
cream with gray mottling; iris bronze with median, hori-
zontal, red streak.

Measurements of holotype: SVL 27.4, tibia length 13.7,
foot length 11.6, head width 10.4, head length 10.7, lOD
2.4, upper eyelid width 1.7, E-N 3.0, eye 3.0, tympanum 1.2.

Distribution and habitat. — The species is known from
three localities at 680-1200 m on the east slope of the north-
ern part of the Cordillera Central in northern Peru, where
they were found in lower humid montane forest (Fig. 22).
The juvenile was on the ground by day, and one male was
in the axil of an elephant ear plant (Xanthosoiua) by day.
The female was on the leaf of an elephant ear plant at night,
anci a male was on the leaf of an herb at night.

Etymology. — The specific name is derived from the
Greek aninlos meaning dirty and the Greek onychos mean-
ing fingernail; the name is applied in reference to the ir-
regular, dark periphery of the discs on the fingers.

Remarks.— One juvenile male (KU 212310 ) has a SVL
of 14.4 mm. In life, the dorsum was black with a tan mid-
dorsal stripe broken by a black interorbital bar and at the
level of the forelimbs. The flanks and dorsal surfaces of
the thighs were marked with yellow bars; the throat and
chest were black with pale yellow flecks, and the belly
and groin were cream with yellow flecks. The iris was
bronze-brown.

Eleutherodactylus bearsei Duellman

Ekutherodnciylus benrsei Duellman, 1992a:2. Holotype: KU 212268, adult
female, from Cataratas Ahuashiyacu, 730 m, 14 km [by road] NE
Tarapoto, Provincia San Martin, Departamento San Martin, Peru.

Diagnosis. — A member of the Eleutherodactylus
{Elctithcwdnctylus) iiiiistrigafus Group having (1) skin on
tiorsum shagreen, bearing scattered low tubercles in males,
that on venter areolate; discoidal fold evident only poste-
riorly; dorsolateral folds absent; (2) tympanic membrane
and tympanic annulus prominent, vertically ovoid, its
length about 30% length of eye; (3) snout acutely rounded
in dorsal view, bluntly rounded in profile; canthus rostralis
angular; (4) upper eyelid lacking small tubercles, broader
than lOD; cranial crests absent; (5) vomerine odontophores
transverse, prominent; (6) males having vocal slits but lack-
ing nuptial pads; (7) Finger 1 shorter than 11; discs broad;
(8) fingers bearing lateral fringes; (9) ulnar tubercles dif-
fuse; (10) heel lacking tubercles; outer edge of tarsus bear-
ing tubercles; inner edge of lacking tubercles or fold; (11)
inner metatarsal tubercle oval, 8-1 Ox subconical outer

Eleutherodactylvs in the Andes of Northern Peru

45

metatarsal tubercle; supernumerary tubercles diffuse,
present only proximally; (12) toes bearing lateral fringes;
webbing absent; Toe V much longer than 111; discs nearly
as large as those on fingers; (13) dorsum brown with darker
brown marks on back and transverse bars on limbs; ven-
ter brown with cream flecks; (14) SVL in males 22.7-25.5
mm, in females 38.0-38.8 mm.

Only three species of Eletitliewdnctyliis in the Andes of
northern Peru have pale spots or flecks on brown flanks.
In Elcutlicwdactuliif bcnrfici, cream flecks are present on the
flanks; the posterior surfaces of the thighs are uniform
brown. The flanks have cream (E. nifiociilis) or white (£.
uuiscofiis) spots, and the posterior surfaces of the thighs
have cream spots in E. }iuiscot:iifi. The latter and £. nifiociilis
have smooth skin on the dorsum (shagreen and tubercu-
late in E. bt'iirsci); the former also differs from E. bcnrsci by
ha\'ing a conical tubercle on the heel, and the latter differs
by lacking a tympanic membrane.

Description. — The description by Duellman (1992a) is
adecjuate.

Distribution and habitat. — This species is known from
the type locahty and 30 km [by road] SW Zapatero (ca. 10
km NE San Jose de Sisa), 500 m, Provincia Lamas,
Departamento San Martin, Peru (Fig. 22). Adults were
found on mossy boulders and juveniles on herbaceous
vegetation at night in deep ravine at elevations of 500 and
730 m in lower humid montane forest.

Remarks. — As noted by Duellman (1992a), Eleiith-
erodactyhis bearsei can be grouped phenotypically with three
other species (£. dindemnttis, eiirydactyhis, and pinti/dacti/his)
in the upper Amazon Basin and on the lower Amazonian
slopes of the Andes.

Eleutherodactylus bromeliaceiis Lynch

Eleutherodnctylus hroweliaceiif Lynch, 1979:12. Holotype: USNM 199731,
adult female, from between Sapote and Suro Rancho, Provincia
Morona-Santiago, Ecuador.

Diagnosis. — A member of the Eleutherodnctylus
{Eleutherodnctylus) unistrigntus Group having (1) skin on
dorsum smooth, that on venter coarsely areolate; discoi-
dal fold prominent; dorsolateral folds absent; (2) tympanic
membrane and tympanic annulus evident, round, its
length about 30% length of eye; (3) snout subacuminate in
dorsal view, acutely rounded in profile; canthus rostralis
rounded; (4) upper eyelid bearing small tubercles, slightly
narrower than lOD; cranial crests absent; (5) vomerine
odontophores oblique, prominent; (6) males having vocal
sUts but lacking nuptial pads; (7) Finger I shorter than II;
discs elliptical; (8) fingers bearing lateral fringes; (9) ulnar
tubercles absent; (10) heel and outer edge of tarsus bear-
ing low tubercles; inner edge of tarsus bearing one tubercle;
(11) irmer metatarsal tubercle oval, 3x elongate outer meta-
tarsal tubercle; supernumerary tubercles numerous; (12)

toes bearing lateral fringes; webbing absent; Toe V much
longer than III; discs slightly smaller than those on fin-
gers; (13) dorsum tan with brown flecks or blotches on
dorsum; venter cream, with brown flecks in adult females;
(14) SVL in males 16.7-23.2 mm, in females 22.9-28.1 mm.

This species with a broad head and pointed snout has
small tubercles on the upper eyelid and on the heel. Other
species with this combination of characters are
Eleutherodactylus ceiithospilus, rhodoplichus, and schultei.
Eleutherodnctylus ceiitlwspilus differs by having the skin on
the dorsum shagreen and having a brown (as opposed to
green) dorsum in life and pale spots in the groin; E.
rhodoplichus differs by lacking vomerine odontophores and
labial bars and by having dark diagonal bars or reticula-
tions on the flanks and dark reticulations on the posterior
surfaces of the thighs, whereas E. schultei differs by hav-
ing a uniformly pale dorsum (as opposed to dark spots)
and dark streaks on the flanks, and by lacking labial bars,
pale spots on the posterior surfaces of the thighs, and dark
flecks on the venter.

Description. — The description by Lynch (1979) is ad-
equate.

Distribution and habitat. — The distribution of this spe-
cies is discontinuous. It is known from the Amazonian
slopes of the Cordillera Oriental (1707-2622 m), northern
(1500-1600 m) and western (1830 m) slopes of the Cordil-
lera del Condor, and the Cordillera de Cutucu (1700 m) in
southern Ecuador, and from Abra Pardo de Miguel (2180
m) in the northern part of the Cordillera Central,
Departamento San Martin, Peru (Fig. 22). All localities are
in humid montane forests, where individuals were found
on low vegetation at night and in bromeliads by day. At
Abra Pardo de MigueL a female was on the leaf of a herb
and a male was calling from the leaf of a bush on the night
of 31 January 1989; the call is a soft "peep."

Remarks. — Previously, this species has been recorded
only from Ecuador (Almendariz, 1997; Duellman and
Lynch, 1988; Lynch, 1980). The two Peruvian specimens
(KU 212213-14) are a subadult female having a SVL of 22.7
mm and an adult male having a SVL of 23.0 mm, respec-
tively. In life, the dorsum of the female was mottled green
and brown, and the venter was gray; the dorsum of the
male was tan with brown mottling, and the flanks, groin
and vocal sac were yellow and the belly cream. In both
individuals, the iris was red to reddish brown.

Eleutherodactylus cajamarcensis Barbour and Noble

Eleutherodactylus cajamarcensis Barbour and Noble, 1920:404. Holotype:
MCZ 5407, adult male, from ruins near Huambos, Departamento
Cajamarca, Peru.

Diagnosis. — A member of the Eleutherodnctylus
(Eleutherodnctylus) uuistrigatiis group having (1) skin on
dorsum shagreen, bearing ill-defined rows of pustules, tha t

46

Scientific Papers, Natural History Museum. The University of Kansas

on venter areolate; discoidal fold prominent; dorsolateral
folds absent; (2) tympanic membrane and tympanic an-
nulus prominent, round, its length no more than V2 length
of eye; (3) snout rounded in dorsal view and in profile;
canthus rostralis angular; (4) upper eyelid bearing small
tubercles, slightly narrower than lOD; cranial crests ab-
sent; (5) vomerine odontophores oblique, not prominent;
(6) males having vocal slits and nuptial pads; (7) Finger 1
shorter than II; discs small, round; (8) fingers lacking lat-
eral fringes; (9) ulnar tubercles indistinct or absent; (10)
heel and outer edge of tarsus lacking tubercles; inner edge
of tarsus usually bearing one tubercle; (11) inner metatar-
sal tubercle oval, 4-6x oval outer metatarsal tubercle; plan-
tar supernumerary tubercles numerous; (12) toes bearing
lateral fringes; webbing absent; Toe V much longer than
Toe III; discs as large as those on fingers; (13) dorsum gray
to pale brown with darker brown markings; venter cream
with brown or gray spots; (14) SVL in males 19.2-24.1 mm,
in females 27.1-33.8 mm.

The only other members of the Eleutherodnctylus
iiiiistrigntus Group in the Andes of northern Peru that have
pale spots in the groin are £. ceuthospihis, lirellus, imiscosiis,
and nifiociilis. In all of these, the pale spots are not set in a
black field, as in E. cnjamarceiisis. Furthermore, the larger
E. muscosus has a conical tubercle on the heel and white
(instead of brown) spots on the flanks, whereas the smaller
E. nifiociilis lacks a tympanic membrane and tubercles on
the upper eyelid, and has cream (instead of brown) spots
on the flanks. The much smaller £. lirellus lacks a tympanic
membrane and annulus. Eleiitliewdactyliis ceuthospihis dif-
fers by lacking brown spots on the flanks and having dark
flecks (instead of spots) on the venter; the spots in the groin
are bright yellow in life. In £. cajamarcensis, the large spots
in the groin are red in life.

Description. — Lynch (1969) provided a thorough rede-
scription of the species. Coloration of living individuals
from Peru was described by Duellman and Wild (1993)
and of individuals from southern Ecuador by Lynch and
Duellman (1997).

Distribution and habitat. — This species is widely dis-
tributed in the mountains of the Huancabamba Depres-
sion and around the Cuenca de Loja, where it occurs at
elevations of 1800-3100 in tropical dry forest, humid mon-
tane forest, and subparamo (Fig. 22). A few records exist
for the Pacific slopes of the Cordillera Occidental in Peru
(Huambos, Departamento Cajamarca) and in Ecuador (Luz
Maria, Provincia Azuay). Most individuals have been
found under rocks and logs or in arboreal or terrestrial
bromeliads by day; few have been observed perched on
low vegetation at night.

Remarks. — Eleiitherodactyhis cajamarcensis is the most
widespread species of the genus in western Peru and ad-
jacent Ecuador.

Elcutherodacti/liis ceiitliospitiis Duellman and Wild

Eleutlwrodactylus ceutliospiliis Duellman and Wild, 1993:8. Holotype:
KU 219775, adult male, from 15.8 km |by road] ENE Canchaque,
1800 m, Departamento Piura, Peru.

Diagnosis. — A member of the Eleiitherodactyhis
{Eleiitherodactyhis) iniistrigatiis Group having (1) skin on
dorsum shagreen with minute, low, round tubercles and
lacking folds, that on venter areolate; discoidal fold evi-
dent; dorsolateral folds absent; (2) tympanic membrane
and tympanic annulus prominent, round, its length about
'/2 length of eye; (3) snout acutely rounded in dorsal view
and in profile; canthus rostralis rounded; (4) upper eyelid
bearing small tubercles, narrower than lOD; cranial crests
absent; (5) vomerine odontophores low, round, concealed
in buccal mucosa; (6) males having vocal slits but lacking
nuptial pads; (7) Finger I shorter than II; discs large, ellip-
tical; (8) fingers bearing lateral fringes; (9) ulnar tubercles
low, diffuse; (10) heel lacking tubercles; outer edge of tar-
sus with few, low, round tubercles; inner edge of tarsus
bearing low tubercles or fold; (11) inner metatarsal tubercle
oval, 6x subconical outer metatarsal tubercle; plantar su-
pernumerary tubercles numerous; (12) toes bearing lateral
fringes; webbing absent; Toe V much longer than III; discs
nearly as large as those on fingers; (13) dorsum gray to
pale brown, usually with small brown flecks or streaks;
venter cream with minute dark flecks; (14) SVL in males
19.0-25.8 mm, in females 23.5-26.7 mm.

Eleiitherodactyhis ceiithospiliis most closely resembles E.
cajamarcensis, which differs by having dark (instead of pale)
spots on the flanks and dark spots (instead of flecks) on
the venter; moreover, the spots in the groin are dull red in
life in £. cajamarcensis and bright yellow in £. ceuthospihis
(Fig. 9). Other members of the Eleutherodactyhis unistrigatiis
group in the region that have pale spots in the groin are
the larger E. muscosus and the smaller £. lirellus and E.
rufioculis; both of the small species lack a tympanic mem-
brane (annulus also absent in E. lirellus), whereas E.
muscosus differs in color pattern (white reticulations on the
dorsum) and by having a conical tubercle on the heel.

Description. — The description given by Duellman and
Wild (1993) is adequate.

Distribution and habitat. — This species is known from
humid montane forest at elevations of 1735-2870 m on the
western slopes of the Cordillera de Huancabamba, and
from a site at 12 km [by roadjW of Lamas (1500 m) on the
Pacific slope of the Cordillera Occidental in northern Peru
(Fig. 23). Males call from low vegetation in cloud forest at
night; adults of both sexes have been found in bromeliads
by day.

Remarks. — Richard Thomas (field notes, 8 December
1974) described the call as ratchetlike, rapid sequence of
three (rarely 2 or 4) clicks repeated at intervals of 15-20 sec.

Eleutherodactylvs in the Andes of Northern Peru

47

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7M. 78'

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# £. ccutliospilus
O £^- cclodiiciylus
■ £^. cryplomt'Uis
n £. exorisnts

A £". incomptus

4

1

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'""^i^f^s;:

-6

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^

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W 78°

Fig. 23. Localities of known occurrence of six species in the
Ekiithewdactylus unistrigalus group in the Andes of southern Ecuador
and northern Peru.

Eleutherodaciylus colodactylus Lynch

Eleuthewdactylus cohdiKh/lus Lynch, 1979:15. Holotype: KU 142151, adult
female, from Abra de Zamora, 2800 m, 13.5 km [by road] E Loja,
Provincia Loja, Ecuador

Diagnosis. — A member of the Eleiitherodnctyliis
(Eleiitliewdactylus) nnistrigatiis Group having (1) skin on
dorsum and venter areolate; discoidal fold and dorsolat-
eral folds absent; (2) tympanic membrane and tympanic
annulus absent; (3) snout rounded in dorsal view, bluntly
rounded in profile; canthus rostralis rounded; (4) upper
eyelid bearing one or two small but prominent tubercles,
narrower than lOD; cranial crests absent; (5) vomerine
odontophores oval, concealed; (6) males lacking vocal slits
and nuptial pads; (7) Finger 1 shorter than II; discs small,
round; (8) fingers having lateral fringes; (9) ulnar tubercles
absent; (10) heel bearing single, conical tubercle; tarsus
lacking tubercles and fold; (11) inner metatarsal tubercle
oval, 4-5x round outer metatarsal tubercle; plantar super-
numerary tubercles numerous, continuing onto toes; (12)
toes bearing lateral fringes; webbing basal; Toe V longer
than III; discs about as large as those on fingers; (13) dor-
sum brown with pale dorsolateral stripes or interorbital
bar; venter cream with brown flecks; (14) SVL in males
14.0-20.7 mm, in females 16.5-25.8 mm.

In the region under consideration, only Elciitlwrodacti/liis
proserpens is like £. colodactylus in having short, stocky fin-
gers, but E. colodactj/lus differs from £. pwscrpciis by lack-
ing a cloacal sheath, tympanic membrane and annulus, a
papilla on the tip of the snout, and prominent vomerine
odontophores. In other Ek'uflicmdnctiiliis in the region hav-
ing digital pads that are only slightly broader than the digit
proximal to the pad (£. atrabmcus, melanogaster, patnikos,
and pi}iguis), the fingers are proportionately longer and
more slencier than those in £, colodactylus and £. proserpens;
moreover, in those robust-bodied species. Toe V is only
slightly longer than Toe III. The absence of a tympanum
and tympanic annulus distinguishes £. colodactylus from
all other species in the region, except £. lirellus; the latter
differs by having vocal slits, labial and limbs bars, a yel-
low spot in the groin, black flecks on the venter, and long
fingers bearing elliptical discs.

Description. — The original description by Lynch (1979)
was augmented with data from Peruvian specimens by
Duellman and Wild (1993).

Distribution and habitat. — This species is known from
two disjunct regions — Amazonian slopes (2195-3140 m)
of the Cordillera Oriental in Provincia Morona-Santiago,
Ecuador; the crest and Amazonian slopes (2710-2800 m)
of the Abra de Zamora in the southern Cordillera Orien-
tal; and the crest and upper eastern slopes of the Cordil-
lera de Huancabamba in northern Peru (Fig. 23). All indi-
viduals for which habitat data are available were found in
terrestrial and arboreal bromeliads by day in humid mon-
tane forest or subparamo.

Eleutherodaciylus cryptomelas Lynch

Eleutherodactylus cn/ptomelas Lynch, 1979:21 . Holotype: KU141992, imma-
ture female, from 15 km [by road] E of Loja, 2710 m, Provincia
Morona-Santiago, Ecuador.

Diagnosis. — A member of the Eleutherodactylus
(Eleutherodactylus) unistrigalus Group having (1) skin on
dorsum shagreen, bearing conical warts and ridges, that
on venter areolate; discoidal fold prominent; dorsolateral
folds absent; (2) tympanic membrane and tympanic an-
nulus prominent, its length 30-40% length of eye; (3) snout
subacuminate in dorsal view, rounded in profile; canthus
rostralis rounded; (4) upper eyelid bearing tubercles,
slightly broader than lOD; cranial crests absent; (5) vomer-
ine odontophores round to oval, prominent; (6) males lack-
ing vocal slits and nuptial pads; (7) Finger I shorter than
II; discs broad, elliptical; (8) fingers having weak lateral
fringes; (9) ulnar tubercles prominent; (10) heel and outer
edge of tarsus bearing conical tubercles; inner edge of tar-
sus bearing indistinct tubercle; (11) inner metatarsal tu-
bercle oval, 4-6x conical outer metatarsal tubercle; plan-
tar supernumerary tubercles numerous; (12) toes bearing
lateral fringes; webbing absent; Toe V longer than III; discs

48

Scientific Papers, Natural History Museum, The University of Kansas

smaller than those on fingers; (13) dorsum gray to brown
sparse darker markings; venter white to cream with brown
reticulations; (14) SVL in males 28.2-30.2 mm, in female
38.6 mm.

Eleuthewdactylus cryptoinclns is readily distinguished
from all other members of the Eleiitherodactylus unistrigatiis
Group in the region by having the groin and hidden sur-
faces of the thighs uniform black. The only other species
having extensive black coloration are short-legged mem-
bers of the Eleiitherodacti/liis orestes group — £. atmbmciis
with black on the undersides of the hind limbs and E.
melanogaster, which is nearly uniform black dorsally and
ventrally; both species have pale yellow spots in the groin.

Description. — The original description by Lynch (1979)
is adequate, but coloration in life was expanded by
Duellman and Wild (1993).

Distribution and habitat. — This species is known from
elevations of 2470-2710 m on the Amazonian slopes of the
Cordillera Oriental and the ridges (3000-3100 m) north of
the Cuenca de Loja in Ecuador, and at elevations of 2770-
2820 m on the western slope of Cordillera de Huancabamba
in Peru (Fig. 23). In Ecuador, the frogs were found in ter-
restrial bromeliads or under rocks in paramo and
subparamo by day. In Peru, the frogs were in humid mon-
tane forest; one was under a rock by day, and another was
in a tree at night

Eleuthcwdncti/lus exoristiis new species

Holotype. — KU 147051, an adult female, from the Rio
Piuntza (03°52' S, 78°15' W, 1550 m), western slope of the
Cordillera del Condor, Provincia Morona-Santiago, Ecua-
dor, one of a series collected on 3 January 1972 by John E.
Simmons, Robert Fiske, and Bruce MacBryde.

Paratypes.— KU 147048-49, 147053, 147056, adult
males, and 147047, 147052, 147054-55, 147058, adult fe-
males, from the type locality, 3-6 January 1972.

Referred specimens. — KU 147050, 147057, juveniles,
from the type locality; USNM 525442, an adult female, from
Puesto Vigilancia Comainas, 665 m, Rio Comainas,
Departamento Amazonas, Peru; USNM 525448, 525462,
525470 from Alfonso Ugarte, 1138 m, upper Rio Comainas,
Departamento Amazonas, Peru.

Diagnosis. — A member of the Elcutlwrodacti/lus
(uiiistrigatus) iiiiistrigatus Group having (1) skin on dor-
sum tuberculate with small conical tubercles in dorsolat-
eral rows, that on venter areolate; discoidal fold evident;
dorsolateral folds absent; (2) tympanic membrane differ-
entiated; tympanic annulus prominent, round, its diam-
eter slightly less than V2 length of eye; (3) snout moder-
ately long, acutely rounded in dorsal view, bluntly rounded
in profile; (4) upper eyelid slightly narrower than lOD,
bearing many conical tubercles; cranial crests absent; (5)

Fig. 24. Dorsal and lateral views of the head of Eleuthewdactylus
exoristus, KU 147051. Scale bar = 5 mm.

vomerine odontophores low, ovoid, oblique; (6) males hav-
ing vocal slits but lacking nuptial pads; (7) Finger 1 shorter
than II; discs on outer fingers elliptical, about twice width
of digit proximal to pad; (8) fingers lacking lateral fringes;
(9) ulnar tubercles absent; (10) heel and outer edge of tar-
sus lacking tubercles; inner edge of tarsus bearing low,
elongate tubercle distally; (11) inner metatarsal tubercle
elevated, elliptical, about 8x subconical outer metatarsal
tubercle; supernumerary plantar tubercles few, low; (12)
toes lacking lateral fringes; webbing absent; Toe V much
longer than III; discs nearly as large as those on outer fin-
gers; (13) dorsum brown with darker brown markings;
flanks pale tan, usually with diffuse diagonal bark bars;
venter tan with dark brown flecks; posterior surfaces of
thighs brown with small cream flecks ventrally; (14) SVL
in four adult males 15.0-16.9 (x = 16.2) mm and six adult
females 21.3-23.5 (x= 22.7) mm.

Eleutherodactylus exoristus differs from all other species
in the region by having flared lips and tubercles arranged
in sinusoidal dorsolateral rows on the body. This species
is most similar to £. percnoptenis and £. serendipitiis, both
of which lack lateral fringes on the digits and tubercles on
the heels and have tubercles on the dorsum, but the tu-
bercles are lower and not arranged in dorsolateral rows;
furthermore, these species lack conical tubercles on the
upper eyelids, are slightly larger, and vomerine
odontophores are absent in £. percnoptenis. Other species
in the region having tubercles on the dorsum are E.
anemerus, which lacks dorsal markings and has a tubercle
on the tip of the snout, the much larger E. benrsci, which
has a rounded snout, lateral fringes on the digits, and no
tubercles on the upper eyelids, and £. hiiithaiiik's, which
has a smooth venter, first finger longer than the second,
and fifth toe only slightly longer than the third.

Description. — (;/ - 4 males, 6 females). Head slightly
wider than body; HW 36.2^2.0 (x = 38.7% SVL; HL 40.0-
46.0 (x = 42.8)% SVL; snout moderately long, acutely
rounded in dorsal view, bluntly rounded in profile (Fig.
24); E-N 74.0-93.3 (x = 83.9)% length of eye; nostrils slightly

ELLi'lllLKDDACTYLUS IN THE AnDES Ol- NoRlHERN PeRU

49

protuberant, directed Interallv; canthus rostrnlis distinctly
angular, slightly cur\ed; loreal region concave; lips flared;
upper evelid bearing many conical tubercles; upper eye-
lid width 81. 5-91. 3 (v= S5.Sn lOD; cranial crests absent;
suprat\mpanic fold weak, barely obscuring upper edge
of prominent tympanic annulus; side of head nearly verti-
cal; tympanic membrane not pustular or thickened; tym-
panic annulus round, its diameter 37.9-32.3 (\ = 43.7)%
length of eye; two or three conical postrictal tubercles
postero\'entral to tympanic annulus. Choanae small, oval,
not concealed by palatal shelf of maxillary arch; vomerine
odontophores, low, oblique, medial to posterior edges of
choanae, oval in outline, narrowly separated medially, each
bearing three teeth in males (odontophore absent on one
side in each of two males), 3-6 {\- 5.8) in females; tongue
slightly more than twice as long as wide, its posterior bor-
der shallowly notched, posterior half not adherent to floor
of mouth. Males with vocal slits extending posterolaterally
from midlateral base of tongue; vocal sac single, median,
subgular; nuptial pads absent.

Dorsum of head, body, and limbs bearing conical tu-
bercles; in most individuals tubercles forming shallow, si-
nusoidal rows extending from posterior border of orbit to
point posterior to sacrum; numerous low tubercles on dor-
sum of snout; dorsolateral folds absent; flanks finely tu-
berculate; skin on throat, belly, and ventral surfaces of
thighs areolate; discoidal fold evident; cloacal sheath short;
enlarged tubercles lateral to vent absent. Ulnar tubercles
absent; thenar tubercle low, ovoid, nearly as large as low,
bifid palmar tubercle; supernumerary palmar tubercles
few, subconical; subarticular tubercles prominent,
subconical; fingers lacking lateral fringes; Finger 1 shorter
than II; disc on thumb moderately expanded; discs on other
fingers elliptical, about twice width of digits proximal to
pad; all fingers having ventral pads defined by circumfer-
ential grooves. Heel and outer edge of tarsus lacking tu-
bercles; inner edge of tarsus bearing low, elongate tubercle
distallv; inner metatarsal tvibercle elevated, elliptical, about
8x subconical outer metatarsal tubercle; supernumerary
plantar tubercles few, low; subarticular tubercles low,
round; toes lacking lateral fringes; webbing absent; discs
on toes nearly as large as those on fingers; tip of Toe III
extending to middle of distal subarticular tubercle on Toe
IV; Toe V extending to point midway between penultimate
and distal subarticular tubercles on Toe IV; when hind
limbs flexed perpendicular to axis of body, heels overlap-
ping by about 20% length of shank; shank 52.1-59.3 (x =
54.3)%'SVL.

Coloration in preservative: Dorsum of head, body, and
limbs reddish brown with darker brown to black mark-
ings (Fig. 25) consisting of distinct labial bars and trans-
verse bars on limbs and diffuse canthal stripe in all indi-
viduals, interorbital bar (8 individuals), narrow

Fig. 25. Dorsal color pattern of Elcuthewdactyhis cxoristiis, KU 147051.
SVL = 23.4 mm.

supratympanic stripe (5), W- or H-shaped mark in scapu-
lar region (4), paravertebral streaks on anterior part of body
(6), diffuse sacral chevrons (3); one individual (KU 147050)
with pale middorsal stripe, and one (KU 147058) with large
cream blotch in sacral region. Flanks cream or tan with (8)
or without (2) vertical to slightly diagonal bars; posterior
surfaces of thighs brown with intrusion of tan dorsally and
minute pale flecks ventrally; venter tan with brown flecks,
most dense on throat and chest; two individuals with pale
tan heels.

Coloration in life: Dorsum brown to reddish brown
with darker brown markings; one with white middorsal
stripe and one with yellow blotch in sacral region; flanks
yellow with brown bars; venter gray with or without
white flecks (John E. Simmons field notes, 3-6 January
1972).

Measurements of holotype: SVL 23.4, tibia length
12.5, foot length 10.5, head width 9.0, head length 9.6,
lOD 2.3, upper eyelid width. 2.1, E-N 2.8, eye 3.0,
tympanum 1.3.

Distribution and habitat. — This species is known
from one locality at 1550 m on the western slopes of the
northern part of the Cordillera del Condor and two
localities along the Rio Comainas (665 and 1138 m) on
the eastern slopes of the southern part of the Cordillera
del Condor (Fig. 23). All individuals were on low
vegetation in humid montane forest at night.

Etymology. — The specific name is a Greek adjective,
exhoristos, meaning exiled. The name is applied in the
sense of the species being restricted to the isolated
Cordillera del Condor.

50

Scientific Papers, Natural History Museum, The University of Kansas

Remarks. — The two juveniles from the type locality
have SVLs of 13.3 and 14.0 mm. They are colored like the
adults; one has bars on the flanks, and in the other the
flanks are uniform tan. Three subadult females from 1138
m on the eastern slopes of the Cordillera del Condor have
SVLs of 17.4-19.4 (x= 18.7) mm. Structurally, they are like
the topotypes, except that USNM 525462 is less tubercu-
late on the dorsum. That individual has short paraverte-
bral dark streaks and plain flanks; USNM 525470 has a
diffuse middorsal blotch and plain flanks, whereas USNM
525448 has a dark chevron in the scapular region, a trans-
verse bar in the sacral region, and barred flanks. In life,
the last two individuals had "clear green" or "clear blue
green" venters (Robert P. Reynolds field notes, 21 and 23
July 1994). The largest and most tuberculate specimen is
an adult female (USNM 525442) having a SVL of 25.6 mm.
It has a dark W-shaped mark in the scalar region, dark bars
on the flanks, and faint brown reticulations on the poste-
rior part of the belly. In life, the dorsum was brown with
darker mottling, and the flanks were tan with brown bars;
the venter was cream with brown spots, and the under-
sides of the thighs were reddish (Robert R Reynolds field
notes, 3 August 1994).

Eleutherodactylus galdi (Jimenez de la Espada)

Pristimantis galdi Yvmenez de la Espada, 1870:62. Types, MNCN (now lost)
from San Jose de Moti, Provincia Napo, Ecuador.

Hylodes festne Peracca, 1904:28. Holotype: MSNT AN413, ex. 3776, adult
male, from San Jose de Cuchipamba, Provincia Morona-Santiago,
Ecuador Synonymy fide Lynch, 1974h:16.

Hylodes nmrgarittfer Boulenger, 1912:189. Syntypes, BM 1912.11.1.54-55
(= 1947.2.16.78-79), from El Topo, Rio Pastaza, Provincia
Tungurahua, Ecuador Synonymy fide Lynch, 1969:270.

Eleutherodactylus fcstae — Peters, 1955:348.

Eleutherodactylus margaritifer — Peters, 1955:349.

Eleutherodactylus galdi— Peters, 1955:350; Lynch, 1969:270; Lynch,
1974b:16.

Diagnosis. — A member of the Eleutherodactylus
{Eleutherodactylus) uiiistrigatus Group having (1) skin on
dorsum smooth to finely shagreen with scattered small
tubercles, that on venter coarsely areolate, usually with
scattered larger warts; discoidal fold evident; dorsolateral
folds absent; (2) tympanic membrane differentiated, and
tympanic annulus prominent, nearly round, its length Vi-
Vi length of eye; (3) snout long, acuminate in dorsal view,
truncate in profile, with slightly swollen tip; (4) upper eye-
lid bearing conical tubercle, narrower than lOD; cranial
crests present, serrate in large females; (5) vomerine
odontophores prominent, oblique; (6) males possessing
vocal slits; nuptial pads absent; (7) Finger I shorter than II;
discs on outer fingers expanded, truncate, more than twice
width of digit proximal to pad; (8) fingers bearing weak
lateral fringes; (9) ulnar tubercles conical; (10) heel, outer
edge of tarsus, and outer edge of foot bearing conical tu-
bercles; inner edge of tarsus lacking tubercles; (11) inner

metatarsal tubercle elliptical, 5-6x ovoid outer metatarsal
tubercle; supernumerary plantar tubercles absent; (12) toes
bearing weak lateral fringes; webbing absent; Toe V much
longer than III; discs smaller than those on outer fingers;
(13) dorsum cream (green in life) with dark brown canthal
and supra tympanic stripes, interorbital and limb bars, and
sparse spotting on dorsum; venter cream with minute dark
flecks and usually dark streaks on throat; posterior sur-
faces of thighs cream; (14) SVL in males 17.1-24.8 mm, in
females 28.1-34.0 mm (Lynch and Duellman, 1980).

The green (in life) or cream (in preservative) coloration
in combination with an acuminate snout, flared hps, greatly
expanded, truncate discs on the outer fingers, row of coni-
cal tubercles extending from the heel to the outer edge of
the foot, and serrate cranial crests (in large females) im-
mediately distinguish Eleutherodactylus galdi from all other
species in the region. The only other predominately green
species is E. acuminatus, which has rounded lips and ellip-
tical discs and lacks tubercles along the tarsus, cranial
crests, and bars on the limbs; moreover, £. acuminatus has
an undifferentiated tympanic membrane. Eleutherodactylus
quaquaversus also has a conical tubercle on the upper eye-
lid and a conical tubercle on the heel, but it lacks a tympa-
num, cranial crests, tubercles along the outer edge of the
tarsus and foot, and large, truncate discs on the fingers.

Description. — Adequate descriptions and illustrations
were provided by Lynch (1969) and Lynch and Duellman
(1980).

Distribution and habitat. — In Ecuador, this gaudy
arboreal species is known in humid montane forests at el-
evations of 1000-1740 m on the eastern face of the Cordil-
lera Oriental (Lynch and Duellman, 1980), 1700-1975 m in
the Cordillera de Cutucu (Duellman and Lynch (1988), and
1500-1550 in the Cordillera del Condor (Almendariz, 1997;
Lynch and Duellman, 1980). The only Peruvian record is
from 12 km [by trail] east of La Peca, 1700 m on the west-
ern slope of the Cordillera Colan, Provincia Bagua,
Departamento Amazonas (Fig. 23). The specimen was
along a stream in humid montane forest.

Remarks. — The single Peruvian specimen (LSUMZ
39362) is a juvenile having a SVL of 13.9 mm. In this small
specimen, cranial crests and ventral warts are not evident.
The dorsum is yellowish tan with dark brown canthal and
supratympanic stripes, narrow interorbital bar, pair of di-
agonal (directed posteromedially) streaks in the scapular
region, chevron in sacral region, and limb bars. Several
short, dark brown streaks are present on the posterior part
of the throat.

Eleutherodactylus incoinptus Lynch and Duellman

Eleutherodactylus incoinptus Lynch and Duellman, 1980:35. Holotype; KU
143484,'adult male, from 16.5 km |by roadl NNE Santa Rosa, 1700
m, Provincia Napo, Ecuador

Eleutherodactylus in the Andes of Northern Peru

51

Diagnosis. — A member oi the Elcuthcrodncti/lus
(Elcuthcnidactyluf) unistrigatiis Group having (1) skin on
dorsum smooth with low, flat tubercles, that on venter
coarselv areolate; discoidal fold prominent; dorsolateral
folds absent; (2) tvmpanic membrane smooth, and tym-
panic annulus prominent, nearly round, its length about
30% length of eye; (3) snout moderately short, rounded in
dorsal view and in profile; (4) upper eyelid with or with-
out tubercles, as wide as lOD; cranial crests absent; (5)
vomerine odontophores not visible except in large females,
low, oblic]ue; (6) males possessing vocal slits and
nonspinous nuptial pads; (7) Finger 1 shorter than 11; discs
on outer fingers expanded, rounded, about twice width of
digit proximal to pad; (8) fingers bearing narrow lateral
fringes; (9) ulnar tubercles absent; (10) heel and outer edge
of tarsus lacking tubercles; inner edge of tarsus bearing
one or two tubercles; (11) inner metatarsal tubercle oval,
5-6x subcortical outer metatarsal tubercle; supernumerary
plantar tubercles numerous; (12) toes bearing narrow lat-
eral fringes; webbing absent; Toe V much longer than 111;
discs equal in size to those on outer fingers; (13) dorsum
brown with darker brown markings — interorbital bar, W-
shaped or diagonal shaped marks in scapular region, and
labial and limb bars; venter brown; posterior surfaces of
thighs dark brown; (14) SVL in males 15.6-18.8 mm, in fe-
males 23.7-25.9 mm (Lynch and Duellman, 1980).

Eleutherodactylus inco7nptus is most easily confused with
£. percnoptenis, which is similar in size and also lacks
vomerine odontophores; the latter differs from £. incomptus
by having the snout subacuminate in dorsal view, discs
on fingers nearly truncate, digits lacking lateral fringes,
inner edge of tarsus lacking tubercles, and males lacking
nuptial pads. Furthermore, in E. percnoptenis, the only dis-
tinct dorsal markings consist of a pair of small black spots
or streaks in the scapular region, and the venter is cream.
The only other member of the Eleutherodactylus unistrigatus
Group in the Andes of northern Peru that lacks vomerine
odontophores is £. anemerus, a frog that lacks dorsal mark-
ings and has an orange-red dorsum, yellow flanks, and a
prominent tubercles on the snout. Three other species (E.
atrabracus, melanogaster, and pataikos) are robust-bodied
members of the E. orestes Group that have small terminal
discs on the fingers, short hind limbs, and Toe V only
slightly longer than Toe III. In its absence of distinctive
features, Eleutherodactylus incomptus is like £. pecki, which
differs from the former by having vomerine odontophores,
a small tubercle on the heel and distinctly larger digital
discs. It also resembles £. exoristus, which differs by hav-
ing finely tuberculate skin on the dorsum, tubercles on the
upper eyelid, cream flecks on the posterior surfaces of the
thighs, and dark flecks on a cream venter.

Distribution and habitat. — Eleutherodactylus mcomptus
has been reported from elevations of 1370-1910 m on the

Amazonian slopes of the Cordillera Oriental in Ecuador
(Lynch and Duellman, 1980). The present specimens (KU
217319-20) represent the first records of the species from
Peru; they were in terrestrial bromeliads (Aechmea) by day
at Santa Rosa de la Yunga, 19 km north of Pongo de
Rentema on the Rio Marafion, Provincia San Ignacio,
Departamento Cajamarca, by W. Razzetto and W. A.
Alarcon on 15 July 1989. This locality is at about 1300 m on
the southern slopes of the Cordillera del Condor (Fig. 23).

Remarks. — The two specimens are a subadult female
having a SVL of 18.2 mm and a juvenile having a SVL of
13.9 mm. Both specimens have the dark brown W-shaped
mark in the scapular region.

Eleutherodactylus infragiittatus sp. nov.

Holotype.— KU 212297, adult female, from the east
slope of Abra Pardo de Miguel (05°46' S, 77 °42' W, 2180 m),
Provincia Rioja, Departamento San Martin, Peru, one of a
series of five specimens collected on 31 January 1989 by
William E. Duellman.

Paratypes.— KU 212298, 212314-16 from the type lo-
cality, and KU 217317 from 14 km [by road] west of
Venceremos, 2000 m, Provincia Rioja, Departamento San
Martin, Peru.

Diagnosis. — A member of the Eleutherodactylus
(Eleutherodactylus) unistrigatus Group having (1) skin on
dorsum smooth with scattered, small tubercles, that on
venter areolate; discoidal fold absent; dorsolateral folds
absent; (2) tympanic membrane evident, and tympanic
annulus thin, not prominent, higher than long, its length
M-'/a length of eye; (3) snout short, rounded in dorsal view
and in profile; (4) upper eyelid with small tubercles poste-
riorly, about as wide as lOD; cranial crests absent; vomer-
ine odontophores low, oval in outline; (6) males possess-
ing vocal slits but lacking nuptial pads; (7) Finger I shorter
than II; discs on outer fingers nearly truncate, more than
twice width of digit proximal to pad; (8) fingers bearing
narrow lateral fringes; (9) ulnar tubercles prominent; (10)
heel and inner edge of tarsus lacking tubercles; (11) inner
metatarsal tubercle ovoid, about 5x round outer metatar-
sal tubercle; supernumerary plantar tubercles evident; (12)
toes bearing narrow lateral fringes; webbing absent; Toe
V slighfly longer than III; discs equal in size to those on
outer fingers; (13) dorsum brown with diffuse darker mark-
ings; venter cream with dark brown mottling; throat brown
with cream mottling; groin and posterior surfaces of thighs
cream to tan with brown bars; (14) SVL in one male 15.8
mm, in three females 22.9-24.3 mm (x= 23.6).

Eleutherodactylus infraguttatus has more distinct dark
mottling on the belly than any other member of the
Eleutherodactylus unistrigatus Group in the region. Four
other species have dark reticulations on the belly, but all
have dark labial bars (absent in E. infraguttatus). Further-

52

Scientific Papers, Natural History Museum, The University of Kansas

more, of these species, £. nrdnhvn/cluis differs by lacking
tubercles of the upper eyelid and diagonal bars on the
flanks, and £. muscosiis differs by having a conical tubercle
on the heel and pale spots on the flanks; £. versicolor lacks
vocal slits and has distinct bars on the flanks, whereas in
E. cryptomelas, the groin and hidden surfaces of the thighs
are black.

Description. — (n = 4; 1 male, 3 females; proportions
are for male, followed by range and mean in parentheses
of three females). Head as wide as body in male and in
mature females, wider than body in gravid females; HW
39.2, 37.4-38.9% (x = 38.1) SVL; HL 37.9, 37.1-t0.1% (x =
38.8) SVL; snout short, rounded in dorsal view, and in pro-
file; E-N 68.3, 60.0-74.67o (x = 68.9)% length of eye; nos-
trils slightly protuberant, directed laterally; canthus
rostralis weakly angular, barely concave; loreal region
weakly concave sloping abruptly to lips; lips not flared;
upper eyelid with small tubercles posteriorly; upper eye-
lid width 63.6, 70.3-89.0% (x = 76.9)% lOD; cranial crests
absent; supratympanic fold weak, barely obscuring upper
edge of tympanic annulus; side of head nearly vertical;
tympanic annulus thin; tympanic membrane not pustular
or thickened; tympanic annulus slightly higher than long;
length of tympanic annulus 42.1, 26.9-33.3%- (x = 30.7)%
length of eye; postrictal tubercles small, subconical,
posteroventral to tympanic annulus; skin on head smooth.
Choanae small, round, not concealed by palatal shelf of
maxillary arch; vomerine odontophores posteromedian to
choanae (absent in one female), oval in outline, each about
twice size of choana, separated medially by distance less
than width of odontophore, each bearing two teeth in male
and 4-6 teeth in female in single row; tongue longer than
wide, its posterior border deeply notched, posterior half
not adherent to floor of mouth; vocal slits short, postero-
lateral to base of tongue; vocal sac single, median, subgular.

Dorsum smooth with scattered, small, subconical tu-
bercles and faint dermal ridge from posteromedian bor-
der of eyelid to scapular region (most evident in male);
dorsolateral folds; flanks smooth; cloacal sheath and en-
larged tubercles in cloacal region absent; skin on throat
and belly areolate; discoidal fold absent. Upper surfaces
of arms smooth; ulnar tubercles round; thenar tubercle
ovoid, smaller than bifid palmar tubercle; supernumerary
palmar tubercles prominent, round, smaller than round,
non conical subarticular tubercles; fingers bearing narrow
lateral fringes; Finger I shorter than II; disc on thumb not
expanded; discs on Fingers II-IV nearly truncate, twice
width of digits; all fingers having ventral pads defined by
circumferential grooves; nuptial pads absent in male. Up-
per surfaces of hind limbs smooth with scattered
subconical tubercles; heel tubercles absent; outer edge of
tarsus lacking tubercles; inner metatarsal tubercle nearly
round, flat, about 5x subconical outer metatarsal tubercle;

'^^

/■ ■ J19\ ,^ 78° 77° j:-.-^

r

-^ > A .

-^ S.-

-^ 1 IXJO m
MKHI-2(X.M1 111
^(MMI-.IIMMJni

-4°

f ) -^ ^;

KilnilKU'i^

' s J

# E. infragiittaius
O E. lireUus
■ E. miiscosus

4

-3°

D E. lU'phophilus
A E. (ickendeni
A E. pecki

-^-^H^s-

\ -^^ \

-6°

^ ^ , xo

Vj

■^ 79° 7,S' '\ 77°

Fig. 26. Localities of known occurrence of six species in the
Elcittherodactyliis unistrigntus group in southern Ecuador and northern
Peru.

supernumerary plantar tubercles small, subconical;
subarticular tubercles round, not conical; toes bearing lat-
eral fringes; webbing absent; discs on toes nearly as large
as those on fingers; tip of Toe V extending to distal edge of
distal subarticular tubercle on Toe IV; tip of Toe 111 not ex-
tenciing to that tubercle; when hind limbs flexed perpen-
dicular to axis of body, heels barely overlapping; shank
49.4%, 46.7-50.8 (x = 49.0)% SVL.

Coloration in preservative: Dorsum brown with darker
brown canthal stripe, supratympanic stripe, interorbital
bar, scapular marks, and transverse bars on limbs; flanks
creamy tan with brown suffusion or spots posteriorly; an-
terior and posterior surfaces of thighs brown, mottled with
cream; venter cream with brown marbling in females, pri-
marily brown in male.

Coloration in life: Dorsum mottled dull olive and tan;
flanks yellow and brown (Fig. 9); anterior surfaces of thighs
red; venter greenish yellow with brown mottling; iris dull
bronze with median, horizontal, brown streak (WED field
notes on holotype, 31 January 1988).

Measurements of ht)lotype: SVL 22.9, tibia length 10.7,
foot length 9.9, head width 8.9, head length 8.5, lOD 2.8,
upper eyelid width 2.0, E-N 1 .8, eye 2.5, tympanum 0.8.

ELEurHiMomcniA's in the Andks of Northern Peru

53

Distribution and habitat. — The species is known only
from two localities at 2000 and 2180 m along the road from
Abra Pardo de Miguel to Mo\obamba on the east slope of
the northern part of the Cordillera Central in northern Peru
(Fig. 26). All indixiduals were on leaves on low vegetation
(< 1 m) above the ground in Inmiid montane forest at night.

Etymology. — The specific name is derived from the
Latin prefix infra- meaning underside and the Latin adjec-
tive ^»ff(7f/(S meaning dappled; the name is applied in ref-
erence to the dark mottling on the venter.

Remarks. — Two juvenile females (KU 212314-15) have
SVLs of 17.4 and 18.2 mm.

Eleutherodactyliis UreUiis Dwyer

Ek'uthewdachihis lirfUiis Dwyer, 1995:247. Holotype: KU 212240, adult
female, from western slope of Abra Tangarana, 1080 m, 7 km [by
road] NE San Juan de Pacaysapa, Provincia Lamas, Departamento
San Martin, Peru.

Diagnosis. — A member of the Elctitherodnctyliis
{Eleutherodactyliis) iinistrigatits Group having (1) skin on
dorsum coarselv areolate with scattered small tubercles,
that on finely areolate; discoidal fold prominent; dorsolat-
eral folds absent; (2) tympanic membrane tympanic annu-
lus not visible; (3) snout subacuminate in dorsal view,
rounded in profile; (4) upper eyelid bearing many small
tubercles, narrower than lOD; cranial crests absent; (5)
vomerine odontophores prominent, oblique; (6) males
possessing vocal slits; nuptial pads absent; (7) Finger I
shorter than II; discs expanded, about twice width of digit
proximal to pad; (8) fingers bearing lateral fringes; (9) ul-
nar tubercles absent; (10) heel bearing two small tubercles;
outer edge of tarsus bearing row of tubercles; inner tarsal
tubercles and fold absent; (11) inner metatarsal tubercle
elliptical, 4-5x subconical outer metatarsal tubercle; super-
numerary plantar tubercles proximally; (12) toes bearing
lateral fringes; webbing absent; Toe V much longer than
III; discs nearly as large as those on outer fingers; (13) dor-
sum brown with darker brown markings; groin brown
with one pale spot; venter cream with dark flecks; poste-
rior surfaces of thighs dark brown with cream flecks; (14)
SVL in males 14.1-17.0 mm, in females 19.4-24.0 mm
(Dwyer, 1995).

The absence of a tympanum and tympanic annulus dis-
tinguishes Eleutherodactyliis lirelliis from all other species
in the northern Andes, except £. colodactyhis, which dif-
fers bv lacking vocal slits and having short fingers with
round discs. Furthermore, E. colodactyhis lacks labial and
limb bars and a yellow spot in the groin. The latter feature
is shared by several species in the upper Amazon Basin.
Two of these (£. toftae and E. variabilis) have a distinct tym-
panic annulus; E. carvalhoi and E. croceoingiiinis lack vocal
slits and lateral fringes on the fingers and toes, and E.
imitatrix has small yellow spots on the anterior surfaces of

the thighs. Andean species with pale spots in the groin are
E. ceiithospilus, luuscosus, and rufiociilis, all of which have
tvmpanic membranes and distinct tympanic annuli, ex-
cept E. rufiociilis, in which the tympanic membrane is ab-
sent, and the annulus is evident only ventrally. In the north-
ern Andes, other species having yellow or pale spots in
the groin are members of the Eleutherodactyliis orestes
Group; these small frogs have robust bodies, digits with
narrow terminal discs, and Toe V only sUghtly longer than
Toe III.

Description. — The description by Dwyer (1995) is
adec]uate.

Distribution and habitat. — Eleutherodactyliis lirellits is
known from three localities at elevations of 470-1200 m
on ridges of the eastern slopes of Cordillera Central in
northern Peru (Fig. 26). All individuals were found at night
on low vegetation in lower montane rainforest.

Remarks. — Dwyer (1995) provided a detailed morpho-
metric analysis of Eleutherodactylus lirellus and five Ama-
zonian species that have yellow spots in the groin. How-
ever, there is no evidence that the presence of yellow spots
in the groin is a synapomorphy.

Eleutherodactylus muscosus new species

Holotype.— KU 219482, adult female, the east slope
of Abra Pardo de Miguel (05°46' S, 77 °41' W, about 1800
m), Provincia Rioja, Departamento San Martin, Peru, one
of a series of four specimens collected on 30 July 1981 by
Rainer Schulte.

Paratypes.— KU 209479-80, subadult females, and
209481, an adult female, collected with the holotype.

Diagnosis. — A member of the Eleutherodactylus
(Eleutherodactylus) iinistrigatiis Group having (1) skin on
dorsum smooth with scattered, small tubercles, that on
venter areolate; discoidal fold not evident; dorsolateral
folds absent; (2) tympanic membrane and tympanic an-
nulus distinct, slightly higher than long, its length about
30% length of eye; (3) snout long, bluntly rounded in dor-
sal view and rounded in profile; (4) upper eyelid narrower
than lOD, with one or two round tubercles posteriorly;
cranial crests absent; (5) vomerine odontophores promi-
nent, slightly oblique; (6) males unknown; (7) Finger I
shorter than II; discs on outer fingers broadly expanded,
nearly truncate, more than twice width of digit proximal
to pad; (8) fingers bearing narrow lateral fringes; (9) ulnar
tubercles low, diffuse; (10) heel bearing conical tubercle;
outer edge of tarsus bearing one or two tubercles proxi-
mally, inner edge of tarsus lacking tubercles; (11) inner
metatarsal tubercle elevated, ovoid, about 3x subconical
roimded outer metatarsal tubercle; supernumerary plan-
tar tubercles prominent, in single row on each digit; (12)
toes bearing narrow lateral fringes; webbing basal, between

54

SciENTinc Papers, Natural History Museum, The University of Kansas

Toes III and IV, and IV and V; Toe V much longer than III;
discs slightly smaller than those on outer fingers; (13) dor-
sum with darker brown markings and with or without
narrow, irregular white venation; throat and chest tan with
brown reticulations or blotches; belly, groin, posterior sur-
faces of thighs, and ventral surfaces of hind limbs dark
brown with cream spots; (14) SVL in four females 29.6-
46.1 mm (x = 37.8).

In coloration, Eleiitherodactylus muscosus is unlike any
other species of the genus in the Andes of northern Peru.
The pale vermiculations on the dorsum are similar to those
exhibited by some specimens of £. spinosus (Lynch, 1 979:fig.
19), a species that differs in being smaller (females 28.3-
34.5 mm) and having a snout that is truncate in profile,
elongate tubercles on the upper eyelid, low cranial crests,
and subconical ulnar tubercles. Only two other species in
the region have tubercles on the upper eyelid and a coni-
cal tubercle on the heel; of these, E. galdi differs by having
an acuminate snout in dorsal view and a uniformly cream
venter, and E. quaquaversus differs by having elliptical (as
opposed to truncate) discs on the fingers and a white venter.

Description. — (n = 4 females). Head as wide as body;
HW 39.4-41.0 (x = 39.8)% SVL; HL 40.1-43.4 (x= 41.9)%
SVL; snout moderately long, bluntly rounded in dorsal
view, rounded in profile; E-N 80.7-91.0 (x= 86.1)% length
of eye; nostrils barely protuberant, directed laterally; can-
thus rostralis angular, barely concave; loreal region deeply
weakly concave, sloping abruptly to rounded lips; upper
eyelid with one or two rounded tubercles posteriorly; up-
per eyelid width 67.8-83.3 (x = 75.7)% lOD; cranial crests
absent; supratympanic fold moderately heavy, obscuring
upper part of tympanum; side of head nearly vertical; tym-
panic annulus thin; tympanic membrane not pustular or
thickened; tympanic annulus slightly higher than long;
length of tympanic annulus 31.3-38.6 (x = 35.9)% length
of eye; postrictal tubercles small, subconical, posteroventral
to tympanic annulus; skin on head smooth. Choanae small,
oval, not concealed by palatal shelf of maxillary arch;
vomerine odontophores slightly oblique, posteromedian
to choanae, elongately oval in outline, each about 4x size
of choana, separated medially by distance less than width
of odontophore, each bearing 4-6 (x = 4.9) teeth; tongue
oval, longer than wide, its posterior border not notched,
posterior third not adherent to floor of mouth.

Dorsum of head, body, and limbs smooth with scat-
tered, small, subconical tubercles; dorsolateral folds ab-
sent; flanks weakly tuberculate; cloacal sheath and en-
larged tubercles in cloacal region absent; skin on throat
and belly areolate; discoidal fold not evident. Ulnar tu-
bercles few, low, diffuse; thenar tubercle elongately ellip-
tical, about equal in size to bifid palmar tubercle; super-
numerary palmar tubercles few, minute; subarticular tu-

Fig. 27. Hand and foot of Eleiitherodactylus
KU 219482. Scale bar = 5 mm.

muscosus,

bercles prominent, round; fingers bearing narrow lateral
fringes; first finger shorter than second; disc on thumb
expanded, round; discs on Fingers II-IV broadly expanded,
nearly truncate, more than twice width of digits (Fig. 27);
all fingers having ventral pads defined by circumferential
grooves. Upper surfaces of hind limbs smooth; heel bear-
ing conical tubercle; outer edge of tarsus bearing one or
two subconical tubercles proximally; inner surfaces of tar-
sus smooth; inner metatarsal tubercle elevated, ovoid,
about 3x rounded outer metatarsal tubercle; supernumer-
ary plantar tubercles prominent, in single row on each digit;
subarticular tubercles subconical; toes bearing narrow lat-
eral fringes; webbing basal between Toes III and IV and IV
and V; discs on toes slightly smaller than those on fingers;
tip of Toe V extending beyond distal edge of distal
subarticular tubercle on Toe IV; tip of Toe III not extend-
ing to that tubercle (Fig. 27); when hind limbs flexed per-
pendicular to axis of body, heels slightly overlapping;
shank 51.0-56.3 (x = 53.3)% SVL.

Coloration in preservative: Dorsum grayish brown
with darker brown markings consisting of streaks on dor-
sum of snout, labial bars, diffuse interorbital bar,
supratympanic stripe, and transverse bars on limbs (Fig.
28). In two smallest specimens (KU 209479-80, SVLs 32.7
and 29.6 mm) dark transverse marks in scapular and sac-

Eleutherodactylus in the Andes of Northern Peru

55

Fig. 28. Dorsal and ventral color pattern of ElcuthcmiKth/us nmscosus,
KU 219462. SVL = 46.1 mm.

ral regions. In two largest specimens (KU 209481-81, SVLs
41 .9 and 46.1 mm) dorsum of body pale brown with white
vermiculations. Throat and chest of two smallest specimens
tan with brown blotches. Throat, chest, and anterior part
of belly in two larger specimens tan with fine brown mark-
ings nearly forming reticulations; flanks brown with white
spots. Posterior part of belly, ventral surfaces of hind limbs,
groin, and posterior surfaces of thighs dark brown with
cream spots.

Coloration in life: Dorsum green with many brown
and black spots and irregular white lines resembling struc-
tures made by worms or fly larvae on stones; groin and
lower surfaces of hind limbs with orange-yellow spots (R.
Schulte, photographs and field notes, 30 July 1981).

Measurements of holotype: SVL 46.1, tibia length 23.1,
foot length 24.5, head width 18.3, head length 18.5, lOD
5.9, upper eyelid width 4.0, E-N 4.2, eye 5.2, tympanum 2.0.

Distribution and habitat. — This species is known only
from the type locality on the eastern slopes of the north-
ern part of the Cordillera Oriental in northern Peru (Fig.
26). The type locality is a stream with rocky banks in hu-
mid, upper montane forest. Stones along the edge of the
stream are covered with moss. The frogs were found by
day in cavities between stones; according to R. Schulte,
the frogs are "imitating exactly a mossy stone."

Etymology. — The specific name is a Latin adjective
muscosiis, meaning mossy; the name is used in reference
to the appearance of the species, as well as its habitat.

Remarks. — The four specimens of this distinctive spe-
cies are poorly preserved; they died en route from the type
locality to the collector's home in Tarapoto. Consequently,
neither the measurements nor the descriptions of certain

morphological structures are as accurate as in our other
descriptions. Although Schulte's field notes imply white
vermiculations on all four individuals, they are evident
only in the two largest individuals; however, the distinc-
tive ventral coloration is the same in all four specimens.

Eleutherodactylus nephophihis new species

Holotype.— KU 212306, an adult female, the east slope
of Abra Pardo de Miguel (05°46' S, 77 °42' W, 2180 m,
Provincia Rioja, Departamento San Martin, Peru, one of a
series of six specimens collected on 31 January 1989 by
William E. Duellman, Michael E. Morrison, and John J.
Wiens.

Paratypes.— KU 212307-09 from the type locality; KU
217322, an adult female, from 14 km [by road] W
Venceremos, 2000 m, Provincia Rioja, Departamento San
Martin, Peru.

Referred specimens. — KU 212305 and 212317, juve-
nile females, from the type locality; KU 212311, juvenile,
from the west slope of Abra Tangarana, 7 km [by road] NE
San Juan de Pacaysapa, Provincia Lamas, Departamento
San Martin, Peru.

Diagnosis. — A member of the Eleutherodactylus
(Eleiitlwrodactylus) unistrigatus Group having (1) skin on
dorsum smooth with sinusoidal dermal ridges from orbit
to scapular region and scattered, small tubercles, that on
venter areolate; discoidal fold weak; dorsolateral folds
absent; (2) tympanic membrane evident, and tympanic
annulus distinct, slightly higher than long, its length only
20-30% length of eye; (3) snout moderately long, rounded
in dorsal view and in profile; (4) upper eyelid narrower
than lOD, with small tubercles posteriorly; cranial crests
absent; (5) vomerine odontophores prominent, oblique; (6)
males unknown; (7) Finger 1 shorter than II; discs on outer
fingers nearly truncate, more than twice width of digit
proximal to pad; (8) fingers bearing narrow lateral fringes;
(9) ulnar tubercles present; (10) heel and outer edge of tar-
sus bearing tubercles; inner edge of tarsus bearing tubercles
tending to coalesce into low ridge; (11) inner metatarsal
tubercle ovoid, about 5x subconical outer metatarsal tu-
bercle; supernumerary plantar tubercles prominent; (12)
toes bearing narrow lateral fringes; webbing absent; Toe V
slightly longer than III; discs slightly smaller than those
on outer fingers; (13) dorsum brown with or without darker
markings or paler markings; venter tan with dense dark
brown flecks; posterior surfaces of thighs brown with
cream spots; (14) SVL in five adult females 24.6-34.0 mm
(x=29.8).

Eleutherodactylus iieplwphilus is like the sympatric £.
rufioculis in having a red iris, but it differs from that smaller
species (female = 20.6 mm) in several features — tubercles
and ridges on dorsum (smooth in £. rufioculis), prominent

56

Scientific Papers, Natural History Museum. The University of Kansas

tympanum (annulus visible under skin), tubercles on up-
per eyelid (absent), oblique vomerine odontophores ob-
lique (ovoid), lateral fringes on fingers (absent), ulnar, heel,
and tarsal tubercles (absent), dark bars on limbs diagonal
(transverse), interorbital bar absent (present), labial bars
present (absent), and posterior surfaces of thighs dark with
pale spots (pale with dark spots). Eleiitherodacti/lus
iiephopliiliis is smaller than E. luiiscosus (females to 37.8
mm), which differs by having truncate discs, conical tu-
bercle on the heel, and large pale spots on the flanks and
lower surfaces of the hind limbs.

Description. — (;; = 5 females). Head slightly wider
than body; HW 37.8-40.0 (x = 39.1)% SVL; HL 37.8-39.4 (x
= 38.1)% SVL; snout moderately long, rounded in dorsal
view, and in profile; E-N 81.8-100.0 (x = 92.5)% length of
eye; nostrils slightly protuberant, directed laterally; can-
thus rostralis weakly angular, barely concave; loreal region
weakly concave sloping abruptly to lips; lips not flared;
upper eyelid with small tubercles posteriorly; upper eye-
lid width 75.8-88.9 (x = 80.4)% lOD; cranial crests absent;
supratympanic fold weak, barely obscuring upper edge
of tympanic annulus; side of head nearly vertical; tympanic
annulus thin; tympanic membrane not pustular or thick-
ened; tympanic annulus slightly higher than long; length
of tympanic annulus 21.2-37.5 (x = 28.4)% length of eye;
postrictal tubercles large, subconical, posteroventral to
tympanic annulus; skin on head smooth. Choanae small,
oval, not concealed by palatal shelf of maxillary arch;
vomerine odontophores oblique, posteromedian to choa-
nae, oval in outline, each about 4x size of choana, sepa-
rated medially by distance less than width of odontophore,
each bearing 3-5 (x = 4.2) teeth; tongue longer than wide,
its posterior border shallowly notched, posterior half not
adherent to floor of mouth.

Dorsum of head, body, and limbs smooth with scat-
tered, small, subconical tubercles; tubercles forming sinu-
soidal ridge from posteromedian edge of eyelid to scapu-
lar region; single, conical tubercle on midline anterior to
orbits in two individuals; dorsolateral folds absent; flanks
tuberculate; cloacal sheath and enlarged tubercles in cloa-
cal region absent; skin on throat and belly areolate; discoi-
dal fold evident posteriorly. Ulnar tubercles few, round to
subconical; thenar tubercle ovoid, about Vi size of bifid
palmar tubercle; supernumerary palmar tubercles few,
minute; subarticular tubercles prominent, subconical; fin-
gers bearing narrow lateral fringes; Finger 1 shorter than
II; disc on thumb barely expanded; discs on Fingers II-IV
elliptical, nearly truncate, more than twice width of digits;
all fingers having ventral pads defined by circumferential
grooves. Upper surfaces of hind limbs smooth with scat-
tered small tubercles; heel bearing subconical tubercle;
outer edge of tarsus bearing three or four subconical tu-

bercles; low fold on distal third of inner surfaces of tarsus;
inner metatarsal tubercle flat, ovoid, about 5x subconical
outer metatarsal tubercle; supernumerary plantar tubercles
prominent, round; subarticular tubercles round, not coni-
cal; toes bearing weak lateral fringes; webbing absent; discs
on toes nearly as large as those on fingers; tip of Toe V
extending to distal edge of distal subarticular tubercle on
Toe IV; tip of Toe III not extending to that tubercle; when
hind limbs flexed perpendicular to axis of body, heels
barely overlapping; shank 50.6-55.2 (x = 52.9)% SVL.

Coloration in preservative: Dorsum brown with a va-
riety of markings — KU 212306: broad, median grayish tan
blotches (snout, occipital region, scapular region, and sac-
ral region); KU 212307; broad cream middorsal stripe nar-
rowly bordered by dark brown; KU 212308 and 217322:
dark brown dermal ridges in occipital-scapular region; KU
212309: like KU 212308 but with top of head pale tan with
faint brown markings. Dorsal surfaces of limbs brown with
darker brown diagonal bars narrower than interspaces.
Flanks brown with narrow creamy tan diagonal marks or
reticulations, especially evident posteriorly; anterior and
posterior surfaces of thighs brown with pale cream verti-
cal bars dorsally and spots or reticulations ventrally; dis-
tinctive dark cloacal patch absent. Canthal stripe and in-
terorbital bar usually absent (canthal stripe present in KU
217322); three or four dark labial bars; dark postorbital
stripe encompassing upper part of tympanum or not. En-
tire venter tan heavily flecked with brown, reticulate in
KU 217322; cream midventral line on chest and belly in
KU 212307.

Coloration in life: Dorsal coloration highly variable —
KU 212306: mottled yellowish tan, olive-green, and dark
dull red; KU 212307: olive with median yellow stripe and
orange dorsolateral area (Fig. 10); KU 212308: dull brown;
KU 212309: brown with grayish-tan head. Flanks brown
(dull red in KU 212306) with cream markings; venter dull
yellow to tan with reddish, dark brown, or black flecks or
reticulations; iris red.

Measurements of holotype: SVL 29.7, tibia length 1 5.9,
foot length 13.4, head width 11.6, head length 11.4, lOD
3.3, upper evelid width 2.5, E-N 3.0, eye 3.0, tympanum 1 .0.

Distribution and habitat. — The species is known only
from three localities at 1080, 2000, and 2180 m along the
road from Abra Pardo Miguel to Moyobamba on the east
slope of the northern part of the Cordillera Central in north-
ern Peru (Fig. 26). Five adults and two juveniles were on
leaves on low vegetation (< 1 m) above the ground in hu-
mid montane forest at night; one juvenile was on the
ground by day.

Etymology. — The specific name is derived from the
Greek iicplios — meaning cloud and the Greek adjective

Elevtherodactylus in the Andes of Northern Peru

57

philia meaning fondness; the name is applied in reference
to the species inhabiting cloud forest.

Remarks.— Three juveniles (KU 212317, 212305, and
21231 1 ) have SVLs of 17.1, 16.9, and 9.1 mm, respectively.

Elciithcrodacti/lus ockeudeni (Boulenger)

Hylodes ockcudciti Boulenger, 1912:187. Syntypes, BM 1907.5.7.19-21 (=

1947.2.16.88-90), from La Union, Rio Hiuicamayo, Carabaya,

Departamento Puno, Peru.
Hi/Iodes hyliicfonnis Melin, 1941:48. Types, NHMG, from Roque,

Departamento San Martin, Peru. Synonymy fide Lynch, 1980.
Synlwphiis cntcaratus Andersson, 1945:27. Holotype, NHRM 1941, from

RioCosanga near Archidona, Provincia Napo, Ecuador. Synonymy

fide Lynch, 1974:16.
Elcuthcivdacttihii nwUiii Bokermann, 1958:95. Replacement name for

Hyhticf hylacfonnis Melin, 1941, non Pln/lhhntcs hylacformis Cope,

1875). Synonymy fide Lynch, 1974b:16.
Eleutlicwiiactylus iiinierssoni Lynch, 1968:292. Replacement name for

Synhcphus cnknrntui^ Andersson, 1945, non Hylodes cakarnliis

Boulenger, 1908. Synonymy fide Lvnch 1974b:16.
Ekutherodach/lus ockendeni— Lynch, 1974b:16; 1980:12.

Diagnosis. — A member of the Eleuthcrodnctylus
(Elciithcrodactifliis) unistrigatus Group having (1) skin on
dorsum shagreen with or without W-shaped occipital-
scapular ridges and dorsolateral folds, that on venter
coarsely areolate; discoidal fold evident; (2) tympanic
membrane and tympanic annulus usually prominent, its
length about '4-V2 length of eye; (3) snout short,
subacuminate in dorsal view, rounded in profile; (4) up-
per eyelid usually bearing tubercles, about equal in width
to lOD; cranial crests absent; (5) vomerine odontophores,
oval, oblique; (6) males having vocal slits, lacking nuptial
pads; (7) Finger 1 shorter than 11; discs moderately large,
elliptical; (8) fingers having narrow lateral fringes; (9) ul-
nar tubercles minute; (10) heel bearing rounded tubercle;
outer edge of tarsus bearing small tubercles; inner edge of
tarsus lacking tubercles or fold; (11) inner metatarsal tu-
bercle eUiptical, 4-6x round outer metatarsal tubercle; plan-
tar supernumerary tubercles few; (12) toes bearing lateral
fringes; webbing absent; Toe V much longer than 111; discs
as large as those on fingers; (13) dorsum cream to brown
with brown to black interorbital bar, subocular spots,
supratympanic stripe, dorsal chevrons, and bar posterior
to sacrum; posterior surfaces of thighs uniform brown;
venter white with or without brown flecks; (14) SVL in
males 16.9-21 .2 mm, in females 24.6-31 .5 mm (Lynch, 1980,
for Amazonian Peru).

The presence of a W-shaped scapular mark on an oth-
erwise plain pale brown dorsum is shared with
Eleiitherodactylus percuopterits, which lacks vomerine
odontophores and has a dark canthal stripe. Four other
members of the Eleidherodacti/his unistrigatus Group in the
region may have a W-shaped scapular mark; three of these
(E. exoristus, incomptus, and sternotln/lax) have diagonal bars
or streaks on the flanks, whereas E. bearsei has cream flecks

on the flanks, and E. cryptoiiiclas has black in the groin and
on the anterior surfaces of the thighs.

Description. — The description by Lynch (1974b) was
augmented by that of Lynch (1980).

Distribution and habitat. — Eleutherodactyhis ockciidcui
is distributed throughout the upper Amazon Basin from
southern Colombia to southern Peru. A few recorcis indi-
cate that the species ascends to slopes of the Andes to
moderate elevations. The species was reported from 1140-
1150 m on the eastern slopes of the Cordillera Oriental
(Lynch and Duellman, 1980) and from Miaza, 900 m, on
the western slope of the Cordillera del Condor
(Almendariz, 1997) in Ecuador, and from 1100-1280 m in
the Serrania de Sira, Departamento Huanuco, Peru
(Duellman and Toft, 1979). A single specimen from 4 km
SW of Chiriaco, 725 m, Provincia Bagua, Departamento
Amazonas, Peru, documents the occurrence of £. ockendeni
in the Rio Marafion Valley in the Andes of northern Peru
(Fig. 26).

Remarks. — The specimen (KU 196470) from the Rio
Maraiion Valley is a female having a SVL of 26.0 mm.

Eleutherodactyhis pecki Duellman and Lynch

Eleutherodactyhis pecki Duellman and Lynch, 1988:135. Holotype: KU
147040, adult male, from the Rio Piuntza, 1550 m, Cordillera del
Condor, Provincia Morona-Santiago, Ecuador.

Diagnosis. — A member of the Eleiitherodactylus
(Eleuthewdnctyliis) uiiistrigntus Group having (1) skin on
dorsum smooth with small tubercles on flanks, posterior
part of back, and upper surfaces of limbs, that on venter
areolate; discoidal fold evident; dorsolateral folds absent;
(2) tympanic membrane and tympanic annulus prominent,
its length about 32% length of eye; (3) snout short,
subacuminate in dorsal view, rounded in profile; canthus
rostralis rounded; (4) upper eyelid bearing tubercles, equal
in width to lOD; cranial crests absent; (5) vomerine
odontophores small, oval; (6) males having vocal slits and
white, nonspinous nuptial pads; (7) Finger I shorter than
II; discs moderately large, round; (8) fingers having weak
lateral fringes; (9) ulnar tubercles absent; (10) heel bearing
small tubercle; outer edge of tarsus lacking tubercles; in-
ner edge of tarsus bearing elongate tubercle; (11) inner
metatarsal tubercle oval, 3^x round outer metatarsal tu-
bercle; plantar supernumerary tubercles numerous; (12)
toes bearing lateral fringes; webbing absent; Toe V longer
than III; discs smaller than those on fingers; (13) dorsum
brown, darkest laterally; venter cream; (14) SVL in males
15.3-18.7 mm, in female 25.2 mm.

As noted by Duellman and Lynch (1988),
Eleiitherodactylus pecki is most similar to E. incomptus and
the lowland E. inartiae. It differs from E. incomptus by hav-
ing a tubercle on the heel and tubercles on the upper eye-

58

Scientific Papers, Natural History Museum, The University of Kansas

lid, and it differs from E. martiae by being larger (males of
E. pecki being as large as females of E. martiae) and in hav-
ing a prominent tympanum. Of the other species in north-
ern Peru having tubercles on the upper eyelids and small
tubercles on the heels, E. bwmeliaceus, cryptoniclas, lircUus,
nephophilus, and rhodoplichus differ from E. pecki in having
the discs on the outer fingers being elliptical (much broader
than long) instead of nearly rounded. Rounded terminal
discs also are present in E. colodactylus and E. schiiltei, both
of which have tubercles on upper eyelids and small tu-
bercles on the heels. Eleuthewdactyhis colodacti/lus differs
by lacking a tympanum, vocal slits, and nuptial pads.
Eleuthewdactyhis schidtei differs by being much larger
(males > 25 mm; females > 28 mm), by having the snout
inclined posteroventrally in profile (rounded in E. pecki),
and in coloration.

Description. — The original description by Duellman
and Lynch (1988) is expanded by the addition of a female
(USNM 525476) having a SVL of 25.2 mm. Comparison of
this specimen with the holotype and two paratypes (KU
147041-42) reveals that in the female the tubercles on the
eyelids of smaller and lower than those in the type series;
furthermore, the tubercles on the heels are barely discern-
ible. The coloration of the female resembles the males in
the type series. The dorsum is tan with three diffuse longi-
tudinal, brown streaks and diffuse dark brown dorsolat-
eral stripes. The flanks and hidden surfaces of the thighs
are dark brown, and the venter is heavily flecked with
brown. A dark brown canthal stripe and two brown labial
bars are present, but an interorbital bar is absent.

Distribution and habitat. — This species is known from
elevations of 1700 m in the Cordillera de Cutucu, 1550 m
on the western slopes of the Cordillera del Condor, and
1138 m on the eastern slopes of the Cordillera del Condor
(Fig. 26). The last locality is the only record for the species
in Peru; the specimen was on a leaf 2 m above the ground
at night in lower montane rainforest.

Remarks. — Duellman and Lynch (1988) suggested that
Eleuthewdactyhis frater, incomptus, and pecki were allopat-
ric variants of E. ockendeni in the lowlands of the upper
Amazon Basin. The latter is known from elevations as high
as 1150 m in the Cordillera del Due in Ecuador, 1280 m in
the Serrania de Sira in Peru (Lynch and Duellman, 1980),
and 900 m on the western slope of the Cordillera del
Condor in Ecuador (Almendariz, 1997). Likewise, Flores
and Vigle (1994) considered £. librarius, which is sympat-
ric with E. ockendeni in the upper Amazon Basin in Ecua-
dor, to be closely related to E. ockendeni.

Eleutlicwdactylus percnopterus new species

Holotype. — KU 21 731 8, adult female, from Santa Rosa
de la Yunga (06°05' S, 78°43' W, 1300 m), Provincia Jaen,

Departamento Cajamarca, Peru, obtained on 15 July 1989
by Weyder Razzetto and Walter A. Alarcon.

Paratypes.— KU 196505 and LSU 32462-63, adult
males, from 33 km [by road] SE Ingenio, 1830 m, Provincia
Chachapoyas, Departamento Amazonas, Peru.

Referred specimens. — KU 196506, subadult male,
from 20 km [by road] SW Chiriaco, Provincia Bagua,
Departamento Amazonas, Peru; KU 209472, subadult fe-
male, from pass at 2400 m, 5 km [by road] NW Mendoza,
Provincia Rodriguez de Mendoza, Departamento
Amazonas, Peru. USNM 525443, juvenile, from the upper
Rio Comainas at base of Cerro Machinaza, 1750 m,
Provincia Condorcanqui, Depart-amento Amazonas, Peru;
USNM 525445-46, 525449-63 from Alfonso Ugarte, 1138
m, Provincia Condorcanqui, Departamento Amazonas,
Peru.

Diagnosis. — A member of the Eleutherodactylus
(Eleutliewdactyhis) unistrigatiis Group having (1) skin on
dorsum with few small tubercles, that on venter weakly
areolate; discoidal fold prominent; dorsolateral folds ab-
sent; (2) tympanic membrane present; tympanic annulus
distinct, round, its diameter about 40% length of eye; (3)
snout moderately long, subacuminate in dorsal view,
rounded in profile; (4) upper eyelid narrower than lOD,
with or without low tubercles; cranial crests absent; (5)
vomerine odontophores absent; (6) males having vocal
slits, lacking nuptial pads; (7) Finger I shorter than II; discs
on outer fingers nearly truncate, more than twice width of
digit proximal to pad; (8) fingers lacking distinct lateral
fringes; (9) ulnar tubercles absent; (10) heel and tarsus lack-
ing tubercles; (11) inner metatarsal tubercle, elevated, el-
liptical, about 5x subconical outer metatarsal tubercle; su-
pernumerary plantar tubercles low, rounded; (12) toes lack-
ing lateral fringes; webbing absent; Toe V much longer than
III; discs nearly as large as those on outer fingers; (13) dor-
sum pale brown with dark brown marks in scapular re-
gion; venter tan with fine dark brown flecks; posterior sur-
faces of thighs tan; (14) SVL in three adult males 21.5-23.2
(x = 22.5) mm, in one adult female 25.9 mm.

Eleuthewdactyhis percnopterus is most easily confused
with E. incomptus, which is similar in size and also lacks
vomerine odontophores (except in large females); the lat-
ter differs from E. percnopterus by having the snout rounded
in dorsal view, discs on fingers rounded, digits bearing
narrow lateral fringes, inner edge of tarsus bearing one or
two tubercles, and males having nuptial pads. Further-
more, in E. incomptus, dorsal markings are more extensive
and contrasting with the ground color, and the venter is
brown. The only other member of the Eleutherodactylus
unistrigatus Group in the Andes of northern Peru that lacks
vomerine odontophores is E. aneinenis, a frog that lacks

Eleutherodactylus in the Andes of Northern Peru

59

Fig. 29. Dorsal and lateral views of the head of Eleutherodactylus
perciwpterus, KU 217318. Scale bar = 5 mm.

dorsal markings, has an orange-red dorsum, yellow flanks,
and a prominent tubercles on the snout. The presence of
scapular marks on an otherwise plain pale brown dorsum
is shared with the primarily Amazonian £. ockeiidcni, which
has vomerine odontophores and lacks a dark canthal stripe.
In its absence of distinctive features, £. perciiopteriis is like
E. pecki, which differs from the former by having vomer-
ine odontophores, a small tubercle on the heel and a dark
venter. Three other species in the region that lack vomer-
ine odontophores (£. atmhrncus, mclaiiogaster, and patnikos)
are robust-bodied members of the E. orestes group that have
small terminal discs on the fingers, short hind limbs, and
Toe V onlv slightly longer than Toe 111.

Description. — {n = 3 males, 1 female; proportions are
for males, followed by those of the female). Head as wide
as body; HW 35.8-36.3 (x= 36.1), 39.4% SVL; HL 35.8-37.6
(x = 37.0), 38.6% SVL; snout moderately long,
subacuminate in dorsal view, rounded in profile (Fig. 29);
E-N 70.1-100.0 (x = 89.6), 90.0% length of eye; nostrils
barely protuberant, directed laterally; canthus rostralis
rounded, curved; loreal region concave; lips slightly flared;
upper eyelid with low tubercles in males; upper eyelid
width 64.2-87.5 (x = 73.9), 69.4%. lOD; cranial crests ab-
sent; supratympanic fold weak, barely obscuring upper
edge of tympanic annulus; side of head slightly inclined
ventrolaterally; tympanic annulus thin; tympanic mem-
brane not pustular or thickened; tympanic annulus round;
diameter of tympanic annulus 37.0-44.0 (x = 41.0), 46.6%
length of eye; postrictal tubercles low, diffuse. Choanae
moderately large, ovoid, not concealed by palatal shelf of
maxillary arch; vomerine odontophores absent; tongue
longer than wide, its posterior border distinctly notched,
posterior half not adherent to floor of mouth. Males with
vocal slits extending posterolaterally from midlateral base
of tongue; v^ocal sac single, median, subgular; nuptial pads
absent.

Dorsum of head and body smooth with scattered,
small, subconical tubercles; dorsal surfaces of limbs with

Fig. 30. Dorsal color pattern of Eleutherodtictylus perciwpterus, KU
217318. SVL = 25.9 mm.

low tubercles; dorsolateral folds absent; flanks smooth;
cloacal sheath and enlarged tubercles in cloacal region
absent; skin on throat weakly areolate, on belly and ven-
tral surfaces of thighs coarsely areolate; discoidal fold
prominent. Ulnar tubercles absent; thenar tubercle broadly
ovoid, elevated more than Vi size of bifurcate palmar tu-
bercle; supernumerary palmar tubercles absent;
subarticular tubercles prominent, subconical; fingers lack-
ing distinct lateral fringes; Finger I shorter than II; disc on
thumb barely expanded; discs on fingers broadly ellipti-
cal, nearly twice width of digit proximal to disc; all fingers
having ventral pads defined by circumferential grooves.
Heel and tarsus lacking tubercles; inner metatarsal tubercle
elevated, elliptical, about 5x subconical outer metatarsal
tubercle; supernumerary plantar tubercles low, rounded;
subarticular tubercles subconical; toes lacking lateral
fringes; webbing absent; discs on toes nearly as large as
those on fingers; tip of Toe V extending to distal edge of
distal subarticular tubercle on Toe IV; tip of Toe 111 extend-
ing to distal edge of penultimate subarticular tubercle on
Toe IV; when hind limbs flexed perpendicular to axis of
body, heels barely overlapping; shank 55.3-56.8 (x = 55.9),
54.4% SVL.

Coloration in preservative: Dorsum pale brown with
narrow, dark brown canthal and supratympanic stripes,
faint brown labial bars and interorbital bar, faint brown
diagonal bars on limbs, and pair of small black spots or
short streaks in scapular region (Fig. 30); flanks, groin, and
hidden surfaces of thighs uniform pale brown; venter
creamy tan with minute dark flecks on throat and belly.

60

Scientific Papers. Natural History Museum, The University of Kansas

■■

-4°

/B --^^
' ( /

/

\
)

•7<'

'^ 1 3°-

\\

i:ZI < 1000 ni
:"~1 1000-:(K)0 m

■nun 4(HKIm
4n(i(| 111
II ^(t Hill

Kilpmeicr.

'^■.

• £. percnopiems
OE. petrobanius
■ E. phoxocephalus
D E. proserpens
AE. quaquaversus
A.E. rhctloplichiis

r

\

- '5"-

/

I
\ •

A

O

78°

r 'X

77°

Fig. 31. Localities of known occurrence of six species in the
Eleutherodactyhis uuistrigntiis group in southern Ecuador and northern
Peru.

Coloration in life: KU 217318: Dorsum pale grayish
brown with reddish tint on head and small black spots; ven-
ter whitish gray (Rainer Schulte field notes, 15 July 1989).

Measurements of holotype: SVL25.9, tibia length 14.1,
foot length 12.7, head width 10.2, head length 10.0, lOD
3.6, upper eyelid width 2.5, E-N 2.7, eye 3.0, tympanum 1 .9.

Distribution and habitat. — The species is known from
two localities at elevations of 1138 m and 1750 on the east-
ern slopes of the Cordillera del Condor, and one locality at
1300 m on the southern edge of the Cordillera del Condor
(Fig. 31). The species also is known at elevations of 1830
and 2400 m in the northern part of the Cordillera Central
in Peru. The latter is a pass northwest of Mendoza at the
headwaters of the Rio Chiriaco; the type locality also is in
the Rio Chiriaco Drainage. The locality 20 km SW Chiriaco
is at an elevation of less than 1000 m in the arid Rio
Marafion Valley. Thus, the known elevational distribution
is at least 1 400 m and suggests that the species may have a
continuous distribution between the Cordillera del Condor
and the Cordillera Central. The holotype was in an arbo-
real bromeliad by day in semiarid forest. Specimens from
the eastern slopes of the Cordillera del Condor were on
low vegetation at night in humid montane forest.

Etymology. — The specific name is the Greek noun
perknopteivs meaning vulture; the name is in loose refer-
ence to Vultur gryphus, the Andean condor, to which the
Cordillera del Condor refers.

Remarks. — The paratypes were collected bv Richard
Thomas on 21-22 December 1974. One (KU 196505) was
calling. The call consists of 2-5 notes ("pink") followed or
not by an ascending series of 10 or more rachetlike clicks
(Richard Thomas field notes, 21 December 1974). The sub-
adult male from 20 km SW Chiriaco (KU 196506), has a
SVL of 18.5 mm, and a subadult female from 5 km [by road]
NW of Mendoza (KU 209472) has a SVL of 21.6 mm. Both
have color patterns like those of the type series. Of the
specimens from eastern slopes of the Cordillera del Condor,
five subadult females have SVLs of 18.7-21.0 (\- 20.1 ) mm,
and 13 juveniles having SVLs of 10.7-16.0 (x = 14.6) mm.
The juveniles tend to be more boldly patterned than the
adults.

Eleutherodactyhis petrobanius Duellman

Eleutherodactylus petrobardus Duellman, I991a:6. Holotype: KU 212292,
adult male, from approximately 2 km [by road] W Huambos, 2500
m, Provincia Chota, Departamento Cajamarca, Peru.

Diagnosis. — A member of the Eleutherodactylus
(Eleutherodactyhis) uiiistrigatus Group having (1) skin on
dorsum, flanks, and limbs with small, irregularly arranged
pustules, that on venter areolate; discoidal fold evident;
dorsolateral folds absent; (2) tympanic membrane and tym-
panic annulus distinct, round, its length about 45'7( length
of eye; (3) snout rounded in dorsal view and in profile;
canthus rostralis rounded; (4) upper eyelid lacking tu-
bercles, slightly narrower than lOD; cranial crests absent;
(5) vomerine odontophores oblique, prominent; (6) males
having vocal slits but no nuptial pads; (7) Finger 1 shorter
than 11; discs broad; (8) fingers bearing lateral fringes; (9)
ulnar tubercles low, diffuse; (10) heel bearing small tu-
bercles; outer edge of tarsus lacking tubercles; inner edge
of tarsus bearing elongate tubercle; (11) inner metatarsal
tubercle ovoid, 3-4x conical outer metatarsal tubercle; plan-
tar supernumerary tubercles small; (12) toes bearing lat-
eral fringes; webbing absent; Toe V much longer than III;
discs as large as those on fingers; (13) dorsum tan with
irregular brown markings, with or without white spots;
venter cream; (14) SVL in males 27.0-30.7 mm, unknown
in females.

Of the species in the Eleutherodactylus uiiistrigatus Group
in the Andes of northern Peru, Eleutherodactylus pelrobardus
is most similar to the smaller £. versicolor, a species that
differs by lacking vocal slits and a small tubercle on the
heel, and by having brown reticulation on the venter,
whereas the belly is white with black flecks in £.
petrobardus. Also, E. petrobardus has more pustular skin on
the dorsum than do other species in the region (Fig. 10).

Elfa'thkrodactyu's in the Andes of Northern Peru

61

Description. — The description by Duellman (1991) is
adequate.

Distribution and habitat. — This species is known only
from the tvpe localit\' in dry scrub forest on the Pacific
slopes of the Cordillera Occidental in Departamento
Cajamarca, Peru (Fig. 31). Individuals were in terrestrial
bromeliads or calling from rock ledges at night.

Eleutherodactyhis pJwxocephalus Lynch

Elciillicnxlnch/his phoxoaylmlus Lynch, 1979:29. Holotype; KU 142075, adult
male, from Pilalo, 2340 m, Provincia Cotopaxi, Ecuador.

Diagnosis. — A member of the Elciitherodactyliis
(EU'iithcivdacti/lus) unistrigatus Group having (1) skin on
dorsum shagreen, that on venter areolate; discoidal fold
prominent; dorsolateral folds absent; (2) tympanic mem-
brane and tympanic annulus evident, round, its length no
more than '2 length of eye; (3) snout rounded in dorsal
view but with vertical fleshy keel, subacuminate in pro-
file; canthus rostralis rounded; (4) upper eyelid lacking
tubercles, narrower than lOD; cranial crests absent; (5)
vomerine odontophores oblique, prominent; (6) males hav-
ing vocal slits and nuptial pads; (7) Finger I shorter than
II; discs broad; (8) fingers bearing lateral fringes; (9) ulnar
tubercles absent; (10) heel and tarsus lacking tubercles and
folds; (11) inner metatarsal tubercle oval, 4-6x round outer
metatarsal tubercle; plantar supernumerary tubercles at
bases of Toes II-IV; (12) toes bearing lateral fringes; web-
bing absent; Toe V much longer than III; discs as large as
those on fingers; (13) dorsum gray to brown with few or
no darker markings; venter cream; lower flank, groin, and
concealed surfaces of limbs white with or without brown
or black reticulations; (14) SVL in males 22.3-29.9 mm, in
females 29.6-38.9 mm.

Structurally, Elciitliewdactylus phoxocephnJus is unique
in the region by having a vertical keel on the snout, which
is difficult to discern in some specimens. However, dark
reticulations in the groin and on the posterior surfaces of
the thighs distinguish E. phoxocephalus from congeners in
the region, except E. rhodoplichus, which differs by having
a small tubercle on the heel and by lacking vomerine
odontophores.

Description. — The original description by Lynch (1979)
is complete, and includes color in life of specimens from
provincias Cotopaxi and Loja, Ecuador. Coloration of liv-
ing individuals from provincias Cotopaxi and Pichincha,
Ecuador, were given by Lynch and Duellman (1997) and
for two individuals from Departamento Piura, Peru, by
Duellman and Wild (1993).

Distribution and ecology. — This species occurs in up-
per humid montane forest and subparamo at elevations
of 1800-3100 m on the Pacific slopes of the Cordillera Oc-
cidental in Ecuador and the Andes of southern Ecuador.

In Peru, it is known from only two localities — 15 km [by
road] ENE Canchaque, 1850 m (KU 181271 ) and El Tambo,
2770 m (MHNSM 15399)— on the western slopes of the
Cordillera de Huancabamba, Departamento Piura, and
from San Andres de Cutervo, about 1800 m, in the north-
ern part of the Cordillera Occidental in Departamento
Cajamarca (Fig. 31). In the Cordillera de Huancabamba,
one individual was on a mossy cliff at night and another
was calling at night from low vegetation. In Ecuador, many
individuals have been found in the axils of elephant ear
plants and arboreal bromeliads by day.

Remarks. — Four specimens (KU 221710-13) were col-
lected at San Andres de Cutervo, Provincia Cutervo,
Departamento Cajamarca, Peru, by Alfonso Miranda on
25 June 1992 and 1 April 1993. Of these, one female has a
SVL of 38.9 mm, and three males have SVLs of 26.6-27.8 (x
= 27.2) mm. All specimens have a fleshy vertical keel on
the snout; the keel is less pronounced in the female than in
the males. The dorsum is brown (female) or tan (males)
with dark brown canthal and supratympanic stripes and
irregular interorbital bar. The female and one male have
numerous small dark brown spots on the dorsum of the
head and body anterior to the sacrum; one male has a pair
of dark spots in the scapular region, and the other male
has many small dark spots in the scapular region and a
pair of spots in the sacral region. The venter is cream with
minute dark flecks, and the groin and hidden surfaces of
the thighs are uniform tan.

Structurally, these specimens compare favorably with
two individuals (KU 181271 and MHNSM 15399) from the
Cordillera de Huancabamba and with several series from
the southern part of the range in Ecuador — Provincia
Azuay (KU 131281-82), Provincia Canar (KU 142118-31),
Provincia Loja (KU 135460-62, 142113-17), and Provincia
Zamora-Chinchipe (KU 142104-12). However, there are
some minor differences in coloration between the Peru-
vian and Ecuadorian specimens. One of the specimens
from the Cordillera de Huancabamba (MHNSM 15399) and
all four from San Andres de Cutervo have minute dark
flecks over the entire venter, as opposed to small brown
spots or more dense flecking in the Ecuadorian specimens,
most all of which have dark mottling in the groin and hid-
den surfaces of the thighs.

Eleutherodactyhis proserpens Lynch

EleuthcwdaciyUis proserpens Lynch, 1979:32. Holotype: USNM 198484,
adult female, from between Sapote and Suro Rancho, 2622 m,
Provincia Morona-Santiago, Ecuador

Diagnosis. — A member of the Eleutlierodactylns
{Eleutherodactyhis) uuistrigatus Group having (1) skin on
dorsum and venter areolate; discoidal fold and dorsolat-
eral folds absent; cloacal sheath extending onto posterior
surfaces of thighs (2) tympanic membrane and tympanic

62

Scientific Papers, Natural History Museum, The University of Kansas

annulus prominent, round, its length no more than Vi
length of eye; (3) snout long, subacuminate in dorsal view,
rounded in profile; small papilla at tip of snout; (4) upper
eyelid bearing low, round tubercles, narrower than lOD;
cranial crests absent; (5) vomerine odontophores low, oval;
(6) males lacking vocal slits and nuptial pads; (7) Finger I
shorter than II; fingers short, broad, with round discs only
slightly wider than digit proximal to pad; (8) fingers bear-
ing ridgelike lateral fringes; (9) ulnar tubercles absent; (10)
heel and tarsus lacking tubercles; (11) inner metatarsal tu-
bercle elliptical, 2x round outer metatarsal tubercle; plan-
tar supernumerary tubercles numerous, some nearly as
large as subarticular tubercles; (12) toes bearing ridgelike
lateral fringes; webbing basal, coalesced with lateral
fringes; Toe V much longer than III; discs slightly smaller
than those on fingers; (13) dorsum tan to brown with darker
brown markings — interorbital bar, supratympanic stripe,
limb bars, and X-shaped mark or large, elongate blotch on
back — or pale dorsolateral stripe; posterior surfaces of
thighs cream to brown; venter pale brown; (14) SVL in
males 15.2-21.0 mm, in females 20.2-23.5 mm (Lynch,
1979).

In the region under consideration, only Elciith-
erodactylus colodaciylus is like £. proserpens in having short,
stocky fingers, but £. proserpens differs from £. colodaciylus
by having a cloacal sheath, tympanic membrane and an-
nulus, a tubercle on the tip of the snout, and prominent
vomerine odontophores. The only other species in the re-
gion with a tubercle on the tip of the snout is E. anemerus,
which has a unicolor dorsum and lacks a tubercle on the
heel and vomerine odontophores. In other Eleutherodactyhis
in the region having digital pads that are only slightly
broader than the digit proximal to the pad (£. atrahraciis,
melanogaster, pataikos, and puiguis), the fingers are propor-
tionately longer and more slender than those in E.
colodaciylus and £. proserpens; moreover, in those robust-
bodied species. Toe V is only slightly longer than Toe III.

Description. — The description by Lynch (1979), which
includes illustrations of the head and hand, is adequate.

Distribution and habitat. — In Ecuador, this species is
known from elevations of 1710-2620 m in the southern part
of the Cordillera Oriental, from 1700 m in the Cordillera
de Cutucii, and from 1550 m on the western slope of the
Cordillera del Condor (Duellman and Lynch, 1988; Lynch,
1979). The only Peruvian record is from the upper Rio
Comainas at the base of Cerro Machinaza, 1750 m, on the
eastern slopes of the Cordillera del Condor, Provincia
Cordoncanqui, Departamento Amazonas (Fig. 31). All
specimens have been found in bromeliads in humid mon-
tane forest.

Remarks. — The single Peruvian specimen (USNM
525447) is a juvenile having a SVL of 11.6 mm. Despite its

small size, it has the distinguishing characteristics of
Eleuiherodaciylus proserpens — small hands, papilla on snout,
and cloacal sheath. In preservative, the dorsum is grayish
tan with a large, dark brown, middorsal blotch extending
from the anterior edges of the orbits to the sacmm. Dark
brown canthal and supratympanic stripes and transverse
bars on the limbs are present; labial bars are absent. The
venter is heavily flecked with brown.

Eleutherodadylus quaquaversus Lynch

Eleutherodadylus quaquaversus Lynch, 1974b:9. Holotype: KU 123745, adult
female, from south slope Cordillera del Due, above Rio Coca,
Provincia Napo, Ecuador.

Diagnosis. — A member of the Eleutherodadylus
{Eleuiherodaciylus) unisirigaius Group having (1) skin on
dorsum shagreen, that on venter areolate; discoidal fold
prominent; dorsolateral folds absent; (2) tympanic mem-
brane absent; tympanic annulus absent or evident only
ventrally; (3) snout subacuminate in dorsal view, rounded
or weakly pointed in profile; (4) upper eyelid bearing coni-
cal tubercle, slightly narrower than lOD; cranial crests ab-
sent; (5) vomerine odontophores prominent, triangular; (6)
males having vocal slits but lacking nuptial pads; (7) Fin-
ger I shorter than II; discs broad, elliptical; (8) fingers bear-
ing lateral fringes; (9) ulnar tubercles absent; (10) heel bear-
ing conical tubercle; outer and inner edges of tarsus bear-
ing small tubercles; (11) inner metatarsal tubercle ellipti-
cal, 4-5x rounded outer metatarsal tubercle; plantar su-
pernumerary tubercles few; (12) toes bearing narrow lat-
eral fringes; webbing absent; Toe V much longer than III;
discs as large as those on fingers; (13) dorsum pale brown
to reddish brown with darker brown interorbital bar, chev-
rons or spots, and narrow diagonal bars on limbs; canthal
and supratympanic stripes absent; labial bars variably
present; posterior surfaces of thighs reddish brown with
brown reticulations; venter white, with or without brown
spots; (14) SVL in males 19.6-22.5 mm, in females 24.6-
31.3 mm (Lynch and Duellman, 1980).

Of the species in the Andes in northern Peru,
Eleuiherodaciylus quaquaversus is like E. galdi in having a
conical tubercle on the upper eyelid and a conical tubercle
on the heel, but the predominately green E. galdi has a
prominent tympanum, cranial crests, a row of tubercles
along the outer edge of the tarsus and foot, and larger, trun-
cate discs on the fingers. Eleuiherodaciylus quaquaversus is
superficially similar to E. ockendciii and E. percnopierus, both
of which have well-defined tympana and usually a W-
shaped mark in the scapular region (never present in £.
quaquaversus), and they lack conical tubercles on the up-
per eyelids and heels and lack brown reticulations on the
posterior surfaces of the thighs.

Description. — The original description by Lynch
(1974b) was augmented by Duellman (1978c) and Lynch
and Duellman (1980).

Elevthf.rodactylvs in the Andes of Northern Peru

63

Distribution and habitat. — Although Elciithcmdacti/lus
qiinquiivertius occurs at elevations as low as 200 m in the
Amazonian lowlands of Peru (Duellman and Mendelson,
1995), the species is distributed primarily on the eastern
slopes of the Andes, where it is known in Ecuador from
elevations of 920-1740 in the Cordillera Oriental (Lynch
and Duellman, 1980), 1700 m in the Cordillera de Cutucii
(Duellman and Lynch, 1988), and 1500-1550 m on the west-
ern slopes of the Cordillera del Condor ( Almendariz, 1997;
Lynch and Duellman, 1980). We now report the species
from the eastern slopes of the Cordillera del Condor —
upper Rio Comainas, base of Cerro Machinaza, 1750 m,
Provincia Condorcanqui, Departamento Amazonas, Peru
(Fig. 31). Nearly all individuals have been found on low
vegetation at night.

Remarks.— In life, juvenile (USNM 525444) with a SVL
of 14.8 mm from the upper Rio Comainas was described
as: "dorsum tan; brown bar between orbits; lips barred;
two spots on scapula; legs barred; chin gray; belly green."
(Robert R Reynolds field notes, 18 July 1994).

Eleuthewdachflus rJwdopUdms Duellman and Wild

Elcuthcwdncfylus rhodoplkhuf Duellman and Wild, 1993:13. Holotype: KU
21978b,"adult male, from El Tambo, 2770 m, 31 km [by road] ENE
Canchaque, Provincia Huancabamba, Departamento Piura, Peru.

Diagnosis. — A member of the Elcutherodnctylus
(Eleuthewdacti/lus) uiiistrignttis Group having (1) skin on
dorsum coarsely shagreen, bearing scattered low, round
or subcorneal tubercles, that on venter areolate; discoidal
fold prominent; dorsolateral folds absent; (2) tympanic
membrane and tympanic annulus prominent, round, its
length no more than 60% length of eye; (3) snout
subacuminate in dorsal view, rounded in profile; canthus
rostrahs rounded; (4) upper eyelid bearing low, round tu-
bercles, narrower than lOD; cranial crests absent; (5)
vomerine odontophores absent in males in most females,
small and elliptical in other females; (6) males having vo-
cal slits but lacking nuptial pads; (7) Finger I shorter than
H; discs broad, elliptical; (8) fingers bearing lateral fringes;
(9) ulnar tubercles few, low; (10) heel bearing small,
subconical tubercles; outer edge of tarsus bearing round,
diffuse tubercles; inner edge of tarsus bearing fold; (11)
inner metatarsal tubercle ellipticaL 3x conical outer meta-
tarsal tubercle; plantar supernumerary tubercles numer-
ous, low, round; (12) toes bearing lateral fringes; webbing
absent; Toe V much longer than III; discs slightly smaller
than those on fingers; (13) dorsum brown with fine, irregu-
lar darker brown markings; venter cream to tan with dark
brown flecks; (14) SVL in males 21.8-28.9 mm, in females
30.1-34.2 mm.

Of the other species in the Eleutherodactylns unistrigatus
Group in the Andes of northern Peru, only £. anemerus,
incomptiis, and percnopterus lack vomerine odontophores.

Elcutlwivdticti/lufi iiiiciiicnis differs by lacking dorsal mark-
ings and having a tubercle on the tip of the snout; both £.
iiicoiuptiis and £. pciriioptcrus lack the small tubercle on
the heel that is present in £. rlwdoplicluis (Fig. 10), and £.
incomptiis also differs by being much smaller and by lack-
ing tubercles on the upper eyelid.

Description. — The description by Duellman and Wild
(1993) is adequate.

Distribution and habitat. — This species is known only
from elevations of 2770-3050 m in humid montane forest
on the western slopes and crest of the Cordillera de
Huancabamba (Fig. 31). Individuals were found at night
on low vegetation and under rocks by day.

Eleutherodacti/liis rhodostichns new species

Holotype. — KU 212264, an adult male, from the west
slope of Abra Tangarana, 1080 m, 7 km [by road] NE San
Juan de Pacaysapa, Provincia Lamas, Departamento San
Martin, Peru, one of a series collected on 5 February 1989
by William E. Duellman and Rainer Schulte.

Paratypes.— KU 212265-67 collected with the holo-
type.

Diagnosis. — A member of the Eleiiiherodactylus
(Elcuihcrodactyhis) unisirigntus Group having (1) skin on
dorsum finely shagreen with scattered, small tubercles, that
on venter areolate; discoidal fold prominent; dorsolateral
folds absent; (2) tympanic membrane evident, and tym-
panic annulus distinct, round, its diameter about 40%
length of eye; (3) snout long, acuminate in dorsal view,
acutely rounded above and inclined posteroventrally in
profile; (4) upper eyelid narrower than lOD, with low tu-
bercles posterolaterally; cranial crests absent; (5) vomer-
ine odontophores low, ovoid; (6) males having vocal slits,
lacking nuptial pads; (7) Finger I shorter than II; discs on
outer fingers rounded, less than twice width of digit proxi-
mal to pad; (8) fingers bearing distinct lateral fringes; (9)
ulnar tubercles present; (10) heel lacking tubercles; outer
and inner edges of tarsus bearing small tubercles; (11) in-
ner metatarsal tubercle elliptical, about 4x subconical outer
metatarsal tubercle; supernumerary plantar tubercles nu-
merous; (12) toes bearing narrow lateral fringes; webbing
absent; Yoe V much longer than III; discs nearly as large as
those on outer fingers; (13) dorsum tan with brown mark-
ings; venter tan with fine dark brown flecks; posterior sur-
faces of thighs tan; (14) SVL in an adult male 19.3 mm and
a subadult female 19.7 mm.

By having a snout that is acuminate in dorsal view
and inclined posteroventrally in profile, Eleuiherodactylus
rhodostichns differs from all other members of the genus in
the region, except £. aciiminatus and £. schultei, both of
which lack a pattern on the dorsum. Furthermore, £.
acuminatus differs by lacking a tympanic membrane, and

64

Scientific Papers, Natural History Museum, The University of Kansas

£. schultei differs by having a small tubercle on the heel.
Red streaks on the dorsum of the body in living individu-
als also distinguish £. rhodostichus from other species in
the region.

Description. — (ii = 1 male, 1 female; proportions are
for the male, followed by those of the female). Head no-
ticeably wider than body; HW 38.3, 38.7% SVL ; HL 36.3,
39.6% SVL; snout long, acuminate in dorsal view, acutely
rounded above and inclined posteroventrally in profile;
E-N 76.0, 79.2% length of eye; nostrils slightly protuber-
ant, directed laterally; canthus rostralis weakly angular,
nearly straight; loreal region weakly concave sloping
abruptly to lips; lips not flared; upper eyelid with indis-
tinct tubercles posterolaterally; upper eyeHd width 85.0,
66.7% lOD; cranial crests absent; supratympanic fold weak,
barely obscuring upper edge of tympanic annulus; side of
head slightly inclined ventrolaterally; tympanic annulus
thin; tympanic membrane not pustular or thickened; tym-
panic annulus round; diameter of tympanic annulus 40.0,
41.7% length of eye; postrictal tubercles small, subconical,
posteroventral to tympanic annulus; skin on head smooth.
Choanae large, nearly round, not concealed by palatal shelf
of maxillary arch; vomerine odontophores, low, oblique,
posteromedian to choanae, oval in outline, each about size
of choana, separated medially by distance greater than
width of odontophore, bearing 1-0, 3-3 teeth; tongue longer
than wide, its posterior border shallowly notched, poste-
rior half not adherent to floor of mouth. Male with vocal
slits extending posterolaterally from midlateral base of
tongue; vocal sac single, median, subgular; nuptial pads
absent.

Dorsum of head, body, and linibs smooth with scat-
tered, small, subconical tubercles, more pronounced in fe-
male than in male; dorsolateral folds absent; flanks tuber-
culate, especially in female; cloacal sheath and enlarged
tubercles in cloacal region absent; skin on throat weakly
areolate, on belly coarsely areolate; discoidal fold evident
posteriorly. Ulnar tubercles few, low, round; thenar tubercle
ovoid, about Vi size of weakly bifid palmar tubercle; su-
pernumerary palmar tubercles few, minute; subarticular
tubercles prominent, subconical; fingers bearing distinct
lateral fringes; Finger I shorter than II; disc on thumb barely
expanded; disk on Finger II slightly larger; discs on Fin-
gers III-IV broadly rounded, nearly twice width of digits;
all fingers having ventral pads defined by circumferential
grooves. Upper surfaces of hind limbs smooth with scat-
tered small tubercles; heel lacking tubercle; outer edge of
tarsus bearing three or four subconical tubercles; inner edge
of tarsus with two or three low tubercles; inner metatarsal
tubercle flat, elliptical, about 4x subconical outer metatar-
sal tubercle; supernumerary plantar tubercles low, diffuse;
subarticular tubercles round, subconical; toes bearing nar-
row lateral fringes; webbing absent; discs on toes slightly

smaller than those on fingers; tip of Toe V extending to
middle of distal subarticular tubercle on Toe IV; tip of Toe
III not extending to that tubercle; when hind limbs flexed
perpendicular to axis of body, heels broadly overlapping;
shank 53.8, 54.3% SVL.

Coloration in preservative: Dorsum tan with sparse
brown markings — KU 212264: small spots laterally in
scapular and sacral regions, narrow interorbital bar, weakly
defined labial bars and transverse bars on hind limbs; KU
212265: short lines laterally in scapular and sacral regions,
narrow interorbital bar, diagonal marks on top of snout,
weakly defined labial bars and transverse bars on hind
limbs. Small dark spots on flanks; posterior surfaces of
thighs tan; venter pale tan with minute dark brown flecks
laterally on throat and belly.

Coloration in life: KU 212264: Dorsum green with red
and tan marks; anterior and posterior surfaces of thighs
pale green; heels and elbows tan (Fig. 10); vocal sac yel-
low; belly cream; ventrals surfaces of Hmbs pale green; iris
reddish brown.

Measurements of holotype: SVL 19.3, tibia length 10.4,
foot length 9.0, head width 7.4, head length 7.0, lOD 2.0,
upper eyelid width 1.7, E-N 1.9, eye 2.5, tympanum 1.0.

Distribution and habitat. — The species is known only
from one locality at 1080 m on the road from Abra Pardo
Miguel to Moyobamba on the east slope of the northern
part of the Cordillera Central in northern Peru (Fig. 32).
Two adults and two juveniles were in terrestrial bromeli-
ads in lower humid montane forest by day.

Etymology. — The specific name is derived from the
Greek rhodoii- meaning red and the Greek stichos meaning
line; the name is applied in reference to the red linear mark-
ings on the dorsum of the body.

Remarks. — Two juveniles (KU 212266-67 ) have SVLs
of 15.8 and 16.9 mm, respectively. Both are colored like the
female para type, except that both have a middorsal, brown,
triangular mark (apex anterior) just anterior to the level of
the sacrum.

Eleuthewdactylus mfioculis new species

Holotype.— KU 212313, a subadult female, from the
east slope of Abra Pardo de Miguel, 2180 m, Provincia
Rioja, Departamento San Martin, Peru, obtained on 31
January 1988 by Michael E. Morrison.

Paratypes.— KU 212312, collected with the holotype.

Referred specimens. — USNM 525465, subadult fe-
male, USNM 525464, 525467-68, and 525473 from the up-
per Rio Comainas, base of Cerro Machinaza, 1750 m,
Provincia Condorcanqui, Departamento Amazonas, Peru;
USNM 525474-75 from Alfonso Ugarte, 1138 m, upper Rio
Comainas, Provincia Condorcanqui, Departamento
Amazonas, Peru.

Eleutherodactylvs in the Andes of Northern Peru

65

I'i

i

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1 IIWO 2lXK)ni

tS3 2000-.iaiO ni
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O E. nifioculis
■ £. schiiUei
D £. scn'dipiliis
A. E. sU'iikiiIiyUix
▼ £. vfiwicdlor
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Fig. 32. Localities of known occurrence of seven species in the
Elcuthcwdnctylus unistrigatus group in the Andes of southern Ecuador
and northern Peru.

Diagnosis. — A member of the Eleiitherodactifliis
{Ek'uthcwdactylus) unistrigatus Group having (1) skin on
dorsum smooth, that on venter areolate; discoidal fold
present; dorsolateral folds absent; (2) tympanic membrane
smooth, and tympanic annulus visible beneath skin, its
length about 30% length of eye; (3) snout moderately long,
rounded in dorsal view and in profile; (4) upper eyelid
lacking tubercles, narrower than lOD; cranial crests absent;
(5) vomerine odontophores elongately ovoid; (6) males
lacking vocal slits and nuptial pads; (7) Finger 1 shorter
than II; discs on outer fingers expanded, about twice width
of digit proximal to pad; (8) fingers lacking lateral fringes;
(9) ulnar tubercles absent; (10) heel and tarsus lacking tu-
bercles; (11) inner metatarsal tubercle oval, 3x outer meta-
tarsal tubercle; supernumerary plantar tubercles minute,
diffuse; (12) toes bearing narrow lateral fringes; webbing
absent; Toe V longer than 111; discs slightly smaller than
those on outer fingers; (13) dorsum tan; venter cream, suf-
fused with brown; throat brown with cream flecks; groin
brown with cream spots; posterior surfaces of thighs brown
with cream flecks; (14) SVL in 1 male 18.1 mm, in 1 female
20.6 mm.

Eleutherodactyhis rufioculis is like £. colodncti/his and E.
versicolor in lacking vocal slits in males. Of these, £.

colodncti/his differs by completely lacking a tympanic mem-
brane and by having much shorter fingers with round
discs, and £. versicolor differs by having a prominent tym-
panum, diagonal bars on the flanks, and dark reticulation
on the belly. The flanks are dark brown with pale yellow
or white spots in only two species in the region — £.
rufioculis and £. inuscosus, which differs from £. rufioculis
by being larger and having tubercles on the upper eyelid
and a conical tubercle on the heel. From the sympatric £.
lu'phopliilus, which also has a red iris, £. rufioculis can be
distinguished by having a smooth dorsum and no differ-
entiated tympanum, and no tubercles on the eyelid, heel,
or tarsus.

Description. — (;/ = 1 male, 1 female; proportions are
for the male, followed by those of the female). Head as
wide as body; HW 36.5, 37.4% SVL; HL 39.8, 40.1% SVL;
snout moderately long, rounded in dorsal view and in pro-
file; E-N 135, 115% length of eye; nostrils barely protuber-
ant, directed laterally; canthus rostralis rounded, straight;
loreal region barely concave; lips rounded; upper eyelid
smooth; upper eyelid width 77.7, 70.3% lOD in female;
cranial crests absent; supratympanic fold weak, not ob-
scuring upper edge of tympanum; tympanic membrane
smooth; tympanic annulus distinctly visible beneath skin;
length of tympanic annulus 35%., 26% length of eye;
postrictal tubercles low, diffuse; side of head nearly verti-
cal. Choanae ovoid, not obscured by palatal shelf of max-
illary arch; vomerine odontophores posteromedial to choa-
nae, elongately ovoid in outline, each more than twice size
of choana, separated medially by distance about equal to
width of odontophore, each bearing 2-4 teeth in single row;
tongue more than twice as long as wide, its posterior bor-
der barely notched; posterior half not adherent to floor of
mouth; vocal slits and vocal sac absent.

Dorsum smooth; dorsolateral folds absent; flanks
smooth; cloacal sheath and enlarged tubercles in cloacal
region absent; skin on throat and belly weakly areolate;
discoidal fold weak, evident only posteriorly. Upper sur-
faces of arms smooth; ulnar tubercles absent; thenar tu-
bercle low, oval, smaller than bifid palmar tubercle, su-
pernumerary palmar tubercles minute, diffuse;
subarticular tubercles round; fingers lacking lateral fringes;
Finger 1 shorter than II; disc on thumb not enlarged; discs
on Fingers 111 and IV twice width of digits, that on Finger
II 1.5x width of digit; all fingers bearing ventral pads de-
fined by circumferential grooves; nuptial pads absent in
male. Upper surfaces of hind limbs smooth; tubercles on
heel and tarsus absent; inner metatarsal tubercle flat, oval,
3^x of outer metatarsal tubercle; supernumerary plantar
tubercles minute, diffuse; subarticular tubercles round, non
conical; toes bearing narrow lateral fringes; webbing ab-
sent; discs on toes slightly smaller than those on fingers;
tip of Toe V extending to middle of distal subarticular tu-

66

Scientific Papers, Natural History Museum, The University of Kansas

bercle on Toe IV; tip of Toe III not extending to that tu-
bercle; when hind limbs flexed perpendicular to axis of
body, heels overlapping by about V4, length of shank; shank
56.4, 55.8% SVL.

Coloration in preservative: Dorsum reddish brown
with faint darker brown interorbital bar, and flecks later-
ally in scapular region; indistinct darker brown transverse
bars on limbs; flanks pale brown with large cream spots
posteriorly; anterior surfaces of thighs creamy tan with
vertical brown marks; posterior surfaces of thighs brown
with cream flecks; venter creamy tan, heavily suffused with
brown flecks. Male with pale tan on snout anterior to in-
terorbital bar and on elbows and on heels.

Coloration in life: Dorsum olive (with two pairs of dull
red spots in female; grayish-white snout and tan elbows
and heels in male) (Fig. 10); groin and anterior surfaces
and thighs mottled yellow and dull red; venter yellow with
brown motthng; iris red (WED field notes, 3f January 1988).

Measurements of holotype: SVL 20.6, tibia length 11.5,
foot length 9.2, head width 7.7, head length 8.2, lOD 2.7,
upper eyelid width 1.9, E-N 2.2, eye 2.6, tympanum 0.7.

Distribution and habitat. — The species is known from
the type locality at 2180 m on the eastern slopes of the
northern part of the Cordillera Central and from 1138 m
and 1750 m on the eastern slopes of the Cordillera del
Condor (Fig. 32). All individuals were on low vegetation
(< 1 m) in humid montane forest at night.

Etymology. — The specific name is derived from the
Latin rufiis, meaning red, and the Latin oculis, meaning eye;
the name is used in reference to the red iris.

Remarks. — The small series from the Cordillera del
Condor consists of one subadult female having a SVL of
20.2 mm and six juveniles having SVLs of 13.4-17.3 (x =
15.1) mm. The color patterns of these specimens are like
that of the type series, except that two juveniles have dark
W-shaped marks in the scapular region and transverse bars
in the sacral region. Another juvenile has dark brown trans-
verse marks in the scapular and sacral regions and a broad,
pale bar immediately anterior to the interorbital bar.

Eletitherodactylus sclmltei Duellman

Beutherodactylus sclmltei Duellman, 1990a:348. Holotype: KU 212222, adult
male, from 5 km N Levanto, 2850 m, Departamento Amazonas, Peru.

Diagnosis. — A member of the Eleutherodactylus
(Eleiitlicrodacti/lus) iiiiistrigntiis Group having (1) skin on
dorsum shagreen, bearing low, tubercles on forelimbs and
in tympanic region, that on venter areolate; discoidal fold
absent; dorsolateral folds absent; (2) tympanic membrane
and tympanic annulus distinct, round, its length slightly
less than V2 length of eye; (3) snout subacuminate in dorsal
view, inclined posteroventrally in profile; canthus rostralis
rounded; (4) upper eyelid bearing low tubercles, narrower

than lOD; cranial crests absent; (5) vomerine odontophores
present or absent, oblique; (6) males having vocal slits but
lacking nuptial pads; (7) Finger I shorter than II; discs
broad, rounded; (8) fingers bearing lateral fringes; (9) ul-
nar tubercles present, low; (10) heel and outer edge of tar-
sus bearing many, low tubercles; inner edge of tarsus lack-
ing fold; (11) inner metatarsal tubercle ovoid, 2x subcorneal
outer metatarsal tubercle; plantar supernumerary tubercles
numerous; (12) toes bearing lateral fringes; webbing ab-
sent; Toe V much longer than III; discs equal to those on
fingers; (13) dorsum tan, reddish brown, or green, bordered
or not by narrow dark brown line from snout to point above
vent; venter white; (14) SVL in males 23.5-26.6 mm, in fe-
males 28.4-34.0 mm.

Eleutherodactylus schultei is one of three species in the
region in which the snout is acuminate or subacuminate
in dorsal view and inclined posteroventrally in profile (Fig.
10). Of these, £. nciiminatus, which like £. schultei, lacks a
pattern on the dorsum of the body, differs by lacking a
tympanic annulus; £. rhodostichus has dark markings on
the dorsum of the body and lacks a tubercle on the heel.

Description. — The description by Duellman (1990) is
adequate.

Distribution and habitat. — This species is known from
only two localities at elevations of 2400 and 2850 m in hu-
mid montane forest in the northern part of the Cordillera
Central in Departamento Amazonas, Peru (Fig. 32). The
second locality (5 km [by road] NW of Mendoza) is about
42 km [airline] east of the type locality. The frogs were
found in large terrestrial and arboreal bromeliads by day.

Eleutherodactylus serendipitus new species

Holotype.— KU 181279, adult male, from 8 km [by
road] NNE Balzapata, 1850 m, Provincia Bongara,
Departamento Amazonas, Peru, obtained on 3 March 1979
by William E. Dtiellman.

Paratype. — KU 181280, an adult male, collected with
the holotype.

Referred specimens. — LSUMZ 39360, subadult fe-
male, and LSUMZ 39363 and 39376, juveniles, from 12 km
[by trail] E La Peca, 1700 m, western slope of the Cordil-
lera Colan, Provincia Bagua, Departamento Amazonas,
Peru.

Diagnosis. — A member of the Eleutherodactylus
(Elcutlierodactylus) uiiistrigatus Group having (1) skin on
dorsum finely tuberculate, that on venter areolate; discoi-
dal fold barely evident posteriorly; dorsolateral folds ab-
sent; (2) tympanic membrane smooth, and tympanic an-
nulus distinct, its length about 40% length of eye; (3) snout
moderately long, subacuminate in dorsal view and bluntly
round in profile; (4) upper eyelid lacking tubercles, nar-
rower than lOD; cranial crests absent; (5) vomerine

ELEHTHERODACTyi.US IN THE AnDES OF NORTHERN PeRU

67

odontophores oblique; (6) males having vocal slits and
nuptial pads; (7) Finger I shorter than II; discs on outer
fingers expanded, nearlv truncate, more than twice width
of digit proximal to pad; (8) fingers lacking lateral fringes;
(9) ulnar tubercles absent; (10) heel and outer edge of tar-
sus lacking tubercles; weak inner tarsal fold on distal third
of tarsus (11) inner metatarsal tubercle elevated, elongately
elliptical, about 8x outer subconical metatarsal tubercle;
supernimierarv plantar tubercles low, subconical; (12) toes
lacking lateral fringes; webbing absent; Toe V much longer
than III; discs slightly smaller than those on outer fingers;
(13) dorsum tan with diffuse brown markings; venter
cream with brown flecks; groin cream; posterior surfaces
of thighs brown; (14) SVL in 2 males 20.4-21.2 (x= 20.8).

Eleutherodactylus sereiidipitus has uniformly brown
flanks and hidden surfaces of the thighs; three other spe-
cies in the Elciitlwrodacti/his iiiiistrigntiis group in the Andes
of northern Peru are similarly colored. Of these, £. ockendeni
and £. pt'cki differ by having tubercles on the upper eyelid
and a tubercle on the heel. Furthermore, in the former the
width of the upper eyelid equals the lOD, and in the latter
the venter is densely flecked with brown. Eleiitherodacti/liis
perciwptcriis differs by having a more rounded snout in
profile and diagonal (instead of transverse) bars on the
hind limbs, and by lacking vomerine odontophores.

Description. — (/; - 2 males). Head about as wide as
body; HW 34.9-37.3 (x = 36.1)% SVL; HL 37.7-38.7 (x =
38.2)% SVL; snout moderately long, subacuminate in dor-
sal view, bluntly rounded in profile; E-N 76.0-78.6 (x =
77.3)% length of eye; nostrils distinctly protuberant, di-
rected dorsolaterally; canthus rostralis rounded, straight;
loreal region barely concave; lips rounded; upper eyelid
smooth; upper eyelid width 70.0-71.4 (x= 70.7)% lOD; cra-
nial crests absent; supratympanic fold weak, barely obscur-
ing upper edge of tympanum; tympanic membrane
smooth; tympanic annulus prominent; length of tympanic
annulus 39.3-44.0 (x = 41.7)% length of eye; postrictal tu-
bercles low, diffuse; side of head nearly vertical. Choanae
ovoid, not obscured by palatal shelf of maxillary arch;
vomerine odontophores posteromedial to choanae, small,
elliptical, separated medially by distance more than width
of odontophore, each bearing two teeth in single row (ab-
sent in KU 181280); tongue nearly twice as long as wide,
its posterior border not notched; posterior half not adher-
ent to floor of mouth; vocal sUts and median, subgular
vocal sac absent.

Dorsum of snout smooth; other dorsal surfaces smooth
with scattered, small tubercles, especially prominent on
flanks; dorsolateral folds absent; cloacal sheath and en-
larged tubercles in cloacal region absent; skin on throat
and chest smooth, that on belly areolate; discoidal fold
barely evident posteriorly. Upper surfaces of arms smooth

with scattered, minute tubercles; ulnar tubercles absent
(indistinct ulnar fold in KU 181279); thenar tubercle ovoid,
about same size as bifid palmar tubercle, supernumerary
palmar tubercles minute; subarticular tubercles round; fin-
gers lacking lateral fringes; first finger shorter than sec-
ond; disc on thumb barely enlarged; discs on Fingers III
and IV more than twice width of digits, that on Finger 11
1.5x width of digit; all fingers bearing ventral pads de-
fined by circumferential grooves; nuptial pads absent.
Upper surfaces of hind limbs smooth; tubercles on heel
and outer edge of tarsus absent; low, indistinct inner tar-
sal fold on distal end of tarsus; inner metatarsal tubercle
elevated, elongately elliptical, 8x subconical outer meta-
tarsal tubercle; supernumerary plantar tubercles distinct,
in single row on proximal segments; subarticular tubercles
round, nonconical; toes lacking lateral fringes; webbing
absent; discs on toes slightly smaller than those on fingers;
tip of Toe V extending to middle of distal subarticular tu-
bercle on Toe IV; tip of Toe III extending to middle of
penultimate subarticular tubercle on Toe IV; when hind
limbs flexed perpendicular to axis of body, heels overlap-
ping by about V4 length of shank; shank 55.2-55.4 (55.3) %
SVL.

Coloration in preservative: Dorsum tan with diffuse
brown marks — broad interorbital bar, canthal stripe, single
labial bar, supratympanic stripe, diagonal mark on either
side in scapular region (Fig. 10); chevron (KU 181279) or
transverse mark (KU 181280) in sacral region, flanks pale
brown anteriorly, cream with minute brown flecks poste-
riorly; posterior surfaces of thigh pale brown; venter cream
with brown flecks.

Coloration in life: At night, pale gray; by day, brown
with darker brown markings; venter gray and throat dull
yellow, both heavily flecked with gray; iris dull bronze with
median, horizontal red-brown streak (WED field notes, 3
March 1979).

Measurements of holotype: SVL 20.4, tibia length 11 .3,
foot length 9.5, head width 7.6, head length 7.9, lOD 2.9,
upper eyelid width 2.0; E-N 1.9, eye 2.5, tympanum 1.1.

Distribution and habitat. — The species is known from
the type locality at 1850 m on the upper western slopes of
the northern part of the Cordillera Central, which is
drained by tributaries of the Rio Chiriaco that flows north-
ward into the Rio Marahon, and from 1700 m of the west-
ern slopes of the Cordillera Colan (Fig. 32). At the type
locality, two males were calling at night from leaves of low
herbaceous plants in highly disturbed humid upper mon-
tane forest. In the Cordillera Colan, a subadult female was
on a bush near a stream, and two juveniles were in leaf
litter in humid montane forest.

Etymology. — The specific name is the Latinized En-
glish word seredipitous, meaning characterized by

68

Scientific Papers, Natural History Museum. The University of Kansas

seredipity, which comes from the Persian fairy tale. The
Three Princes ofSerendip, who were known for finding vahi-
able things that were not sought for. The collection of am-
phibians made at the type locality was seredipitous be-
cause a stop was made there only after we were unable to
proceed on a road closed by an accident.

Remarks. — Of the specimens from the Cordillera
Colan, the subadult female has a SVL of 21.7 mm, and the
juveniles have SVLs of 13.8 and 14.6 mm. The female
(LSUMZ 39360) has narrow dorsolateral dark streaks pos-
terior to the sacrum and a short, median interorbital mark.

The type locality of this species also is the type local-
ity of Hi/hi nperoiuea Duellman, Phylloynedusa duellmani
Cannatella, and Scinax oreites Duellman and Wiens.

Eleutherodactylus steniothi/lax Duellman and Wild

Eleutherodach/lus sternothylnx Duellman and Wild, 1993:17. Holotype: KU
219793, adult male, "from 16 km |by road] ENE Canchaque, 1840 m,
Provincia Huancabamba, Departamento Piura, Peru.

Diagnosis. — A member of the Eleutherodachjlus
(Eleutherodactylus) unistrigatus Group having (1) skin on
dorsum shagreen, bearing few, low, round tubercles pos-
teriorly and laterally, that on venter areolate; discoidal fold
evident; dorsolateral folds absent; (2) tympanic membrane
and tympanic annulus distinct, round, its length about Vi
length of eye; (3) snout subacuminate in dorsal view,
acutely rounded in profile; canthus rostralis acutely
rounded; (4) upper eyelid bearing low, round tubercles,
narrower than lOD; cranial crests absent; (5) vomerine
odontophores prominent, oval; (6) males having vocal slits
but lacking nuptial pads; (7) Finger 1 shorter than II; discs
broad, tnmcate; (8) fingers bearing lateral fringes; (9) ul-
nar tubercles absent; (10) heel and tarsus lacking tubercles;
inner edge of tarsus lacking fold; (11) inner metatarsal tu-
bercle elUptical, 3-Ax round outer metatarsal tubercle; plan-
tar supernumerary tubercles diffuse; (12) toes bearing lat-
eral fringes; webbing absent; Toe V much longer than III;
discs smaller than those on fingers; (13) dorsum tan with
dark brown markings; venter cream with minute brown
flecks; (14) SVL in males 18.3-29.1 mm, in females 28.3-
36.7 mm.

The presence of an acuminate snout and absence of tu-
bercles on the upper eyelids and heels distinguishes
Eleutherodactylus sternothylax from all other members of the
Eleutherodactylus unistrigatus Group in the region, except
E. acuminatus and E. aneinerus. The former differs by lack-
ing a tympanic membrane and dark markings on the dor-
sum and flanks, whereas the latter differs by having a
prominent tubercle on the tip of the snout, as well as lack-
ing dark markings.

Description. — The description by Duellman and Wild
(1993) is adequate.

Distribution and habitat. — This species is known only
from elevations of 1735-1840 m in humid montane forest
on the western slopes of the Cordillera de Huancabamba

(Fig. 32).

Eleutherodactylus versicolor Lynch

Eleutherodactylus versicolor Lynch, 1979:45. Holotype: KU 119858, adult
male, from 13.5 km [by road] E Loja, 2800 m, Provincia Loja, Ecuador.

Diagnosis. — A member of the Eleutherodactylus
{Eleutherodactylus) unistrigatus Group having (1) skin on
dorsum shagreen with fine tubercles, that on venter ar-
eolate; discoidal fold prominent; dorsolateral folds absent;
(2) tympanic membrane and tympanic annulus prominent,
length of tympanum about 40-50% length of eye; (3) snout
subacuminate in dorsal view, rounded in profile; canthus
rostralis angular; (4) upper eyelid bearing tubercles or not,
narrower than lOD; cranial crests absent; (5) vomerine
odontophores prominent, oblique; (6) males lacking vocal
slits and nuptial pads; (7) Finger I shorter than II; discs
broad, elliptical; (8) fingers bearing weak lateral fringes or
none; (9) ulnar tubercles low, diffuse; (10) heel lacking tu-
bercles; outer edge of tarsus bearing row of small, conical
tubercles; inner edge of tarsus bearing low fold distally;
(11) inner metatarsal tubercle ovoid, 3-4x conical outer
metatarsal tubercle; plantar supernumerary tubercles nu-
merous, conical; (12) toes lacking lateral fringes; webbing
absent; Toe V much longer than III; discs slightly smaller
than those on fingers; (13) dorsum brown with darker
brown markings — chevrons, interorbital bar, canthal and
supratympanic stripes; flanks usually with vertical bars;
venter cream with brown reticulations; (14) SVL in males
19.3-25.2 mm, in females 22.7-29.8 mm (Lynch, 1979).

The only other species in the region that can be con-
fused with Eleutherodactylus versicolor is E. ardalonychus,
which also has diagonal bars on the flanks and brown re-
ticulations on the venter, but the reticulations are much
finer than those in E. versicolor. Furthermore, E. ardalonychus
differs by having a smooth dorsum, no tympanic mem-
brane and tympanic annulus visible only ventrally, and
rounded, instead of elliptical, discs on the outer fingers.

Description. — The description by Lynch (1979) requires
slight modification. Specimens herein assigned to this spe-
cies are like topotypic material except that tubercles are
present on the upper eyelids — 1/2 in one specimen, 2/3
in one, and many in four.

Distribution and habitat. — This species has been re-
ported from an elevation of 3100 m in subparamo north of
San Lucas, Provincia Loja, Ecuador and from 2500-2800
m in subparamo and cloud forest at Abra de Zamora, Ec-
uador (Lynch, 1979). The specimens reported herein are
from somewhat lower elevations on the eastern slopes of
the Cordillera del Condor and represent the first records

Eleutherodactylvs in the Andes of Northern Peru

69

of the species from Peru (Fig. 32). The following localities
are in humid montane forest in Departamento Amazonas:
upper Rio Comainas, base of Cerro Machinaza, 1750 m
(USNM 325466, 525477), Alfonso Ugarte, upper Rio
Comainas, 1138 m (USNM 525469, 525471-72); Puesto
Vigilancia Comainas, upper Rio Comainas, 665 m (USNM
525441). All of the Peau'ian specimens were collected at
night on vegetation up to 2 m above the ground.

Remarks. — Three of the specimens from the Cordillera
del Condor are females having SVLs of 26.0, 26.2, and 27.3
mm; two are males having SVLs of 21 .8 and 24.7 mm, and
one is a juvenile with a SVL of 17.5 mm. Five individuals
have dark, irregular, chevron-shaped marks on the dor-
sum; one male (USNM 525477) has pale cream dorsolat-
eral stripes extending from the posterior edge of the eye-
lid to a point above the insertion of the hind limbs.

Eleutherodactylus wiensi Duellman and Wild

Elcuthcwdactylus u'iensi Duellman and Wild, 1993:22. Holotype: KU
219795, adult male, from 12.7 km |by road] ENE Canchaque, 1600
m, Provincia Huancabamba, Departamento Piura, Peru.

Diagnosis. — A member of the Eleutherodactylus
{Eleutherodactylus) unistrigatus Group having (1) skin on
dorsum smooth with scattered, small, conical tubercles,
that on venter areolate; discoidal fold evident; dorsolat-
eral folds weak; (2) tjmipanic membrane absent and tym-
panic annulus evident only ventrally, its length about 30%
length of eye; (3) snout acutely rounded in dorsal view,
rounded in profile; canthus rostralis rounded; (4) upper
eyelid bearing tubercles on outer edge or not, narrower

BIOGEOGRAPHY

than lOD; cranial crests absent; (5) vomerine odontophores
prominent, oblique; (6) males having vocal slits but lack-
ing nuptial pads; (7) Finger I shorter than II; discs broad,
truncate; (8) fingers bearing lateral fringes; (9) ulnar tu-
bercles few, diffuse; (10) heel bearing single, conical tu-
bercle; outer edge of tarsus bearing few, low, round tu-
bercles; inner edge of tarsus bearing low fold distally; (11)
inner metatarsal tubercle ovoid, 4x subconical outer meta-
tarsal tubercle; plantar supernumerary tubercles minute;
(12) toes bearing lateral fringes; webbing basal; Toe V much
longer than III; discs equal to or slightly smaller than those
on fingers; (13) dorsum grayish tan with irregular brown
markings; venter cream with brown spots or reticulations;
(14) SVL in males 27.8-33.0 mm, in female 37.0 mm.

This is one of four species in the Eleutherodactylus
unistrigatus Group in the region that lacks a tympanic mem-
brane but has a tympanic annulus visible ventrally (Fig.
10). Of these, E. acuminatus, which lacks dorsal markings,
and £. ardalom/chus, which has dark reticulations on the
belly, differ from E. wiensi by having a tubercle on the heel;
E. rhodoplichus differs by lacking vomerine odontophores
and having tubercles on the upper eyelid.

Description. — The description by Duellman and Wild
(1993) is adequate.

Distribution and habitat. — This species is known from
only two localities at elevations of 1600 and 1735 m in hu-
mid montane forest on the western slope of the Cordillera
de Huancabamba, Departamento Piura, Peru (Fig. 32). In-
dividuals were perched on low vegetation at night.

Because of the complex topography and diverse envi-
ronments in the Andes of northern Peru and southern Ec-
uador, together with the intervening Huancabamba De-
pression, this region has many endemics, and it is the north-
ern or southern limits of distributions of many taxa, such
as toads of the Bufo spiuulosus and veraguensis groups
(Duellman and Schulte, 1992) and lizards of the genus
Pholidobolus (Montanucci, 1973; Reeder, 1996). Aside from
frogs of the genus Eleutherodactylus, many species of other
groups of anurans are known only from this region —
centrolenids (Cadle and McDiarmid, 1990; Duellman and
Schulte, 1993), Gastrotheca (Duellman, 1987; Duellman and
Trueb, 1988; Trueb and Duellman, 1978,), and
leptodactylids of the genera Ischnocnema (Duellman, 1990b;
Lynch, 1974a), Phrynopus (Cannatella, 1984, Lynch, 1975a),
and Phyllonastes (Duellman (1991b; Lynch, 1976), as well
as lizards of the genus Stcnocercus (Cadle, 1991). The dry
valleys even are barriers to the distributions of montane-
forest birds (Parker et al., 1985).

The following discussion refers to patterns of distri-
bution of Eleutherodactylus only in the Andes of northern

Peru and southern Ecuador, and in the intervening
Huancabamba Depression. Most of the species of
Eleutherodactylus in the region inhabit humid montane for-
ests, and many are restricted to that habitat or extend up-
ward into very humid montane forest (including
subparamo and paramo), where species richness is lower
(Table 4). With few exceptions, only members of the
Eleutherodactylus conspvcillatus Group occur in the dry habi-
tats in the Huancabamba Depression, where they usually
are found along streams. Species in this group are wide-
spread geographically and altitudinally in western South
America (Lynch and Duellman, 1997). Two members of
this group (E. karcharias and £. nietahates) are known only
from dry environments below 1000 m, and E. lyuiani in-
habits these same environments, as well as montane habi-
tats. In the region of the Huancabamba Depression, three
members of the group (E. avicuporum, condor, and
cuneirostris) are known only from humid montane forest.
Three other species in the group (E. citriogaster, lanthanites,
and peruvianus) primarily occupy humid tropical forest in
the Amazon Basin, but ascend to humid montane forest

70

SciENTinc Papers, Natural History Museum, The University of Kansas

Table 4. Geographic and latitudinal distribution of Ekutherodactylus in the Andes of northern Peru and southern Ecuador Numbers in columns are
elevations in meters. Species followed by an asterisk (*) also occur in the upper Amazon Basin.

Species

Cordillera Cordillera

Cordillera

Cordillera

Cordillera

Cordillera

Occidental, Oriental

de

del

de

Central,

Ecuador Ecuador

Cutucii

Condor

Colan

Peru

ylus conspicillatus Group

— —

—

—

1700-2030

—

— —

—

—

—

600-800

— —

1975

1500-1750

—

—

Cordillera Cordillera

Occidental de Huan-

Peru cabamba

£. avicuporuni
E. citriogaster
E. condor
E. cnneirostris
E. knrchnrias
E. lantlmnites*
E. b/inniiP
E. metabates-
E. peruvianus*
E. w-uigrum

E. arniodactylus

E. atrabracus
E, melanogaster
E. orestes
E. pataikos
E. pinguis
E. vidua

610-3000

1000-1490

1410-1910
1100-2220

1700-1975

800-3200
Eleutherodncti/Ius uigwvittatus Group

Eieutherodnctylus orestes Group

— 2720-3120

— 2710-3100

E. ncniiiiuatiis*

E. altanmzonictis'

E. anemerus

E. ardnlonyduts

E, atratus

E. balionotus

E. baryecuus

E. bearsei

E. broineliaceus
cajamarcensis
ceiithospilus
colodactylus
cryophilus

E. cryptomelas

E. exoristus

E. galdi

E. ganonotus

E. incomptus

E. infraguttatus

E. lirellus

E. muscosus
nepbophilus
nigrogriseus
ockcndeni'^
pecki

E. percnopterus

E. percidtus

E. percnopterus

Eleiitherodactyhis unistrigatiis Group

petrobnrdus
E. pboxocephalus
E. prolattis
E. proserpens
E. pycnodermis
E. qunquaversus
E. rhodopHchus
E. rhodostichus

665-1750

830
1500-1600

— .

2195-2850

—

—

—

2800

—

—

—

2195-1990

—

—

1710-2620

1700

1500-1600

2660-3000

—

—

—

2195-3140

3080-3200

2835-3355

—

—

—

2470-3100

—

—

—

—

—

665-1550

—

1000-1740

1700-1975

1500-1550

—

—

1700

—

—

1270-1910

—

1300

—

1180-

-2835

1700

1150

—

1000-

-1100

—

900

—

-

-

1700

1138-1550

—

-

-

—

1138-1750

—

2850

—

—

—

-

-

—

1138-1750

1800-3100

_

_

_

_

—

1140-

-1490

1700

—

—

1710-

-2620

1700

1550

—

2650-

-3000

—

—

—

920-

1740

1700

1500-1550

1700

2960-3330

1700

1000
1630

3370

3300-3470
3470

950
680-1200

2000-2180
470-1200

1800
1080-2180

1120-1850

3050-3760

2770

500-730

—

—

2180

—

—

—

2200

3050-3100

—

1500

1735-2870

—

—

2745-3110

_

_

2770-2820

1830

—

—

1830

—

2500

—

—

1800

1850-2770

2770-3050

1080

Eleutherodactylus in the Andes of Northern Peru

71

Tablt' 4 continuL'J

Cordillera

Cordillera

Cordillera

Cordillera

Cordillera

Cordillera

Cordillera

Cordillera

Occidental,

Orient.il

de

del

de

Central,

Occidental

de Huan-

Species

Ecuador

Ecuador

Cutucii

Condor

Colan

Peru

Peru

cabamba

E. rweti

2620-3420

2720-3280

E. ruidtis

2317

—

—

—

—

—

—

—

E. rufioculis

—

—

—

1138-1750

—

2180

—

—

E. schultei

—

—

—

—

—

2400-2850

—

—

E. scivndipitiis

—

—

—

—

1700

1850

—

—

E. spiiwsiis

—

1700-2835

—

1550

—

—

—

—

E. steniotln/lnx

—

—

—

—

—

—

—

1735-1840

E. tracln/blcf'Imrif

—

950-1250

—

900-1600

—

—

—

—

E. ventriiimrmoratus''

—

1000-1740

1700

—

—

—

—

—

E. versicolor

—

2500-3100

—

665-1750

—

—

—

—

E. wieitsi

—

—

—

—

—

—

—

1600-1735

'Also at elevations of 500-1000 m in the Rio Marafion Valley
-Only at ele\'ation of 525 m in the Rio Maraiion Valley
'Also at elevation of 725 m in the Rio Maraiion Valley

on the eastern slopes of the Andes. Most species in this
group are terrestrial and capable of lengthy leaps, as evi-
denced by their long hind limbs.

The Elciitherodacti/his nigrovittatus Group consists of
five species, one of which (£. nigrovittatus) has a broad dis-
tribution in the upper Amazon Basin in Ecuador and north-
ern Peru. The other four species inhabit montane forests
and subparamo in the Andes of Colombia, Ecuador, and
northern Peru. Species in this group are small, terrestrial
frogs that hop about in leaf litter. According to Lynch et al.
(1997), the lowland E. nigrovittatus is the sister group to E.
latens + E. manipus in the Colombian Andes, and these three
species are the sister group to £. elassodiscus in the Ecua-
dorian Andes; they postulated dispersal of E. nigrovittatus
as a means of explaining the geographical and cladistic
relationships.

The seven species making up the Eleutherodactylus
orestes Group are small, terrestrial frogs that make short
hops; this mode of locomotion is correlated with the short
hind limbs. Members of this group primarily have allo-
patric distributions in humid or very humid montane for-
ests or subparamo and paramo. The distributions of £.
orestes and E. vidua are mostly allopatric, but the two spe-
cies occur sympatrically in the southern part of their ranges;
E. melanogaster and E. pataikos occur sympatrically in the
Cordillera Central in Peru.

Members of the large Eleutherodactylus unistrigatus
Group reach their greatest diversification in humid mon-
tane forests, but many species occur in humid tropical for-
ests, and a few extend upward above treeline. Although
some species are encountered on the ground by day, these
inhabitants of forests generally are arboreal, or at least
perch on leaves and limbs of bushes at night. Only a few
clades have been identified within this large group. Of the

species known from northern Ecuador, only E. galdi has
been placed in a clade (Lynch, 1996; Lynch and Rueda-
Almonacid, 1997); the four species in this clade have allo-
patric distributions in the Andes from northern Colombia
to northern Peru.

Because phylogenetic relationships have not been as-
certained among the Eleutherodactylus in the Andes of
northern Peru and their extralimital congeners, only a de-
scriptive analysis of their distributions is possible. Even
this is hampered by the absence of collections from many
areas. Nonetheless, some patterns seem to be apparent.
More species (26) inhabit the southern part of the Cordil-
lera Oriental in Ecuador than any other mountain system
under consideration here"*; this is followed by the Cordil-
lera del Condor (19) and the northern part of the Cordil-
lera Central in Peru (16). An intermediate number (11) is
known from the Cordillera de Cutucii and Cordillera de
Huancabamba, whereas many fewer species are known
from the southern part of the Cordillera Occidental in Ec-
uador (7) and the Cordillera Occidental in Peru (5). These
low numbers are indicative of the scarcity of suitable habi-
tat for most species of Eleutherodactylus. The relatively low
number of species recorded from the Cordillera de Cutucu
and Cordillera Colan probably reflect the fact that no her-
petologists have collected in those cordilleras; in fact, our
existing knowledge of the anuran faunas in those cordille-
ras comes from collections made by ornithologists.

Endemism in the various cordilleras varies from 0%
in the Cordillera del Condor to 75% in the Cordillera Cen-
tral in northern Peru (Table 5). Endemism is highest in
species that are distributed at elevations above 2000 m.

■•Seventy-two species are known from the Cordillera Occidental in Co-
lombia (J. D. Lynch, pers. comm.).

72

SciENTinc Papers, Natural History Museum, The University of Kansas

Table 5. Numerical comparisons of Elentheimlacnliis in the Andes of southern
Ecuador and northern Peru.

Region

Number
Species

Number
Endemic

Percent
Endemic

Cordillera Occidental. Ecuador

7

1

14

Cordillera Oriental. Ecuador

26

7

27

Cordillera de Cutucii

11

1

9

Cordillera del Condor

19

Cordillera Colan

5

3

60

Cordillera Central. Peru

16

12

75

Cordillera Occidental, Peru

5

2

40

Cordillera de Huancabamba

11

4

36

The apparent absence of endemism in the Cordillera del
Condor probably is related to the absence of collections
from high elevations in that cordillera. On the other hand,
the apparent high endemism in the Cordillera Central in
Peru is owing to several species that are known only from
the eastern slopes of that cordillera; once the Cordillera
Colan becomes better known, it is assumed that several
species now known in the Cordillera Central will show
up in the Cordillera Colan.

Each of the eight cordilleras shares species with at least
two other cordilleras (Fig. 33). Generally, the degree of
commonality of species is related to distance, as well as
elevations of intermediate regions. The two cordilleras just
east of the Cordillera Oriental in Ecuador, the Cordillera
de Cutucii and Cordillera del Condor share more species
than do any other cordilleras. However, some of these spe-
cies, as well as those shared with the Cordillera Central in
Peru (e.g., Eleiitberodactyltis actimiiintus, lanthnnitcs,
peruviamis, and ventrimarmoratus), are primarily lowland
species that ascend the slopes of the Andes. However, four
species (E. bwmeliacens , galdi , nigwgriseiis, and proserpens)
that are shared by the Cordillera Oriental in Ecuador, Cor-
dillera de Cutucu, and Cordillera del Condor are not known
to occur at elevations of less than 1000 m.

Occupation of more than one cordillera is less com-
mon in the west, where drier environments occur at the
lower elevations. However, Elentherodactylus cajamnrcensis
occurs in three cordilleras and is unknown at elevations of
less than 2200 m. Assuming that each species has changed
little, if any, physiologically and ecologically during its
existence, interpretation of present patterns of distribution

Cordillera

Occidental.

Ecuador

7

d)

Cordillera
Oriental,
Ecuador

Cordillera

de

Cutucij

11

Cordillera

Occidental,

Peru

Cordillera

del
Condor

Cordillera
de Huan-
cabamba

11

Cordillera

Central,

Peru

Fig. 33. Diagrammatic representation of eight cordilleras in southern
Ecuador and northern Peru. Numbers in boxes are the number of species
of Elentherodactylus in each cordillera; numbers in circles indicate number
of species shared by cordilleras at the ends of arrows.

must invoke past climatic change to account for present
disjunctions. If the predictions of Pleistocene climatic de-
pression in the Andes by Dollfus (1976), Hastenrath (1967),
and Sauer (1971) are anywhere near correct, regions of
humid montane forest now isolated on the various cordil-
leras would have been connected, thereby permitting dis-
persal among the cordilleras; subsequent elevation (in in-
terglacial phases) would have resulted in isolation of popu-
lations and possibly speciation. This climatic-speciation
model championed by Simpson (1979) was refined in
theory by Lynch and Duellman (1997) for Elcuthewdaciylus
in western Ecuador. Cladograms of various lineages of
Eleiithewdncti/his in the Andes of Colombia follow this pat-
tern of allopatric speciation within the same elevational
limits (Lynch et al, 1997).

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Eleutherodactylus in the Andes of Northern Peru

75

APPENDIX 1

Specimens Examined

The specimens from the Andes of northern Peru and
comparative material from elsewhere in Peru and from
Ecuador are documented below, alphabetically by species.
Localities are arranged alphabetically within
departamentos or provinces, which also are arranged al-
phabeticallv within countries. Within a locality, specimens
are listed chronologically by catalogue number following
museum abbreviations, which are arranged alphabetically.
Ek'idJiewilactyluf naiminatus (17 specimens)

Ecuador: Pwv. Monvia-Saiitingo: San Jose, 830 m, KU 146976. Proi'.
Nnpo: Limoncocha, KU 183523; Rio Yasuni, KU 175105. Pmv. Sucumbios:
Puerto Libre, 570 m, KU 123255-60; 15 km ENE Umbaqui, 720 m, KU
123254. Pmv. Pastaza: Canelos, 530 m, KU 119473.

Peru: Diyto. Lcretc: San Jacinto, 175-190 m, KU 221995. Depto. Snn
Martin: Rio Cainarachi, 33 km NE Tarapoto 330 m, KU 209466; Rio
Shilcayo, near Tarapoto, 500 m, KU 209467; 15.4 km SW Zapatero, 950 m,
KU 217308-10.

Elcnthcwdactyluf aiwnicnif (1 specimen)

Peru: Depto. Pima: El Tambo, 31 km [by road] ENE Canchaque, 2770
m, KU 219789 (holotype).

Eleutherodactylus araioiiactyhis (1 specimen)
Peru: Depto. Amazonas: 24 km [by road] SW Leimebamba, 3370 m,
UF 40764 (holotype).

Eleutherodactylus ardalom/chus (4 specimens)
Peru: Depto. San Martin: Abra Tangarana, 7 km NE San Juan de
Pacaysapa, 1080 m, KU 212299; Rio Cerranayacu, 76 km [by roadj NW
Rioja, 1200 m, KU 212301 (holotype), 212310; 8 km [by road] NE Tarapoto,
680 m, KU 212300.

Eleutherodactylus atrabracus (2 specimens)
Peru: Depto. Amazonas: Cordillera Colan, E La Peca, 2963-3330 m,
LSU 49144 (holotype), 45090.

Eleutherodactylus avicuporum (13 specimens)
Peru: Depto. Amazonas: 12 km [by trail] E La Peca, 1700 m, KU 288628,
LSUMZ 39359, 39361, 39365 (holotype), 39366-68, 39370-72, 39374-75,
45089.

Eleutherodactylus bearsei (11 specimens)
Peru: Depto. San Martin: Cataratas Ahuashiyacu, 14 km [by road]
NE Tarapoto, 730 m, KU 212268 (holotype), 212269-74, 217314-15; 30 km
[by road I SW Zapatero, 500 m, KU 212275-76.

Eleutherodactylus bromeliaceus (9 specimens)
Ecuador: Prov. Morona-Santiago: 8.8 km [by road] WSW Plan de
Milagro, 2370 m, KU 202316-17; Rio'piuntza, Cordillera del Condor, 1830
m, KU 146974; between Logrofio and Yaupi, 1700, ANSP 29254, 29272,
29278. Prov. Zamora-Chinchipe: 22 km [by road] W Zamora, 1730 m, KU
141772.

Peru: Depto. San Martin: East slope Abra Pardo de Miguel, 2180 m,
KU 212213-14.

Eleutherodactylus cajamarcensis (147 specimens)
Ecuador: Prov. Azuay: Luz Maria, 1880 m, KU 221706-09. Prov. Loja:
5 km NE Cariamanga, 1870 m, KU 141896-909; 12 km [by road] NE
Catacocha, 2060 m, KU 141890-95; 6.8 km [by road] E Loja, 2640 m, KU
217851-52; 9 km [by road] E Loja, 2660 m, KU 119949-50; 12 km [by road]
S Loja, KU 2250 m', KU 165190-92; 6 km [by road] N San Lucas, 2850-
2900 m, KU 165193-99, 120007-15; 7-8 km [by road] N San Lucas, 2940-
3000 m, KU 119951-120006; 13 km [by road] E Veracruz, 2250 m, KU
141860-89.

Peru: Deptto. Cajaniarca: San Andres de Cutervo, KU 221706-09. Depto.
Piura: between Canchaque and Huancabamba, 3100 m, KU 135495,
135502; 25.5 km [by road] SW Huancabamba, 3010 m, KU 181250-61; 26
km SW [by roadi Huancabamba, 3050 m, KU 196508; 29.3 km [by road]
SW Huancabamba, 3100 m, KU 181244-49.

Eleutherodactylus ceiithospitus (53 specimens)
Peru: Depto. Cajainarca: 12 km [by road] W Lamas, 1500 m, KU
212215-18. Piura: 15-16 km [by road] ENE Canchaque, 1735-1840 m, KU
181270, 181272-78, 196492-98, 219775 (holotype), 219776-85, LSUMZ
32321-31, MHNSM 15387-97; 29 mi (46 km) [by road] E Canchaque, 9200
ft (2840 m),UF 34103.

Eleutherodactylus citriogaster (22 specimens)
Peru: Depto. San Martin: Cataratas Ahuashiyacu, 14 km [by road]
NE Tarapoto, 730 m, KU 212277 (holotype), 212278-85, 217316, 217339-
42, MHNSM 6080-82; Rio Cainarachi, 33 km [by road] NE Tarapoto, 330
m, KU 209384; 8 km [by road] NE Tarapoto, 680 m, KU 212286; 18 km Iby road]
NE Tarapoto, KU 209483; 28 km Jby road] NE Tarapoto, KU 212289-90.

Eleutherodactylus colodactylus (131)

Ecuador: Prov. Loja: 14 km [by road] E Loja, 2770 m, KU 142160-61;
15 km [by road] E Loja, 2710 m, KU 142162-64. Prov. Zamora-Chichipe:
Abra de Zamora, 2800 m, KU 142151-54, 142156-59, 165219-21.

Peru: Depto. Piura: 23 mi (37 km) jby road] E Canchaque, 9300 ft
(2870 m), UF 52077-89, 52091-138, 112513-33; 31 km Jby roadJSW
Huancabamba, 3080 m, KU 181262-64; 33 km [by road] SW
Huancabamba, 3050 m, KU 196443-46; Summit of Cordillera de
Huancabamba (on road between Canchaque and Huancabamba), 3100
m, KU 135494, 135496-501.

Eleutherodactylus condor (48 specimens)
Ecuador: Prov. Moroua-Santiago: "Camp 3," Cordillera de Cutucu,

1975 m, ANSP 29235, 29255-56, 29258; Ri'o Piuntza, Cordillera del Condor,

KU 146991, 146992 (holotype), 146993-147033.

Peru: Depto. Amazonas: upper Rio Comainas, base of Cerra

Machinaza, 1750, USNM 525437.

Eleutherodactylus crytomelas (6 specimens)
Ecuador: Prov. Loja: 15 km [by road] E Loja, 2710 m, KU 141992-93;

8-9 km [by road] N San Lucas, 3000-3100 m, KU 120995-96.

Peru: Depto. Piura: El Tambo, 31 km Jby road] ENE Canchaque, 2770

m, KU 181269, MHNSM 15398.

Eleutherodactylus cuneirostris (1 specimen)

Peru: Depto. Amazonas: 12 km Iby trail] E La Peca, 1700 m, LSUMZ
39369 (holotype).

Eleutherodactylus exoristus (16 specimens)

Ecuador: Prov. Morona-Santiago: Rio Piuntza, Cordillera del Condor,
1830 m, KU 147047-50, 147051 (holotype), 147052-58.

Peru: Depto. Amazonas: Alfonso Ugarte, upper Rio Comainas, 1138
m, USNM 525448, 525462, 525470; Puesto Vigilancia Rio Comainas, 665
m, USNM 525442.

Eleutherodactylus galdi (15 specimens)

Ecuador: Prov. Morona-Santiago: "Camp 2," 1700 m, Cordillera de
Cutucu, ANSP 29245, 29247; "Camp 3," 1975 m, Cordillera de Cutucu,
ANSP 29248; Rio Piuntza, Cordillera del Condor, KU 146977-85. Prov.
Napo: Rio Azuela, 1740 m, KU 143416, 165422.

Peru: Depto. Amazonas: 12 km [by trail] E La Peca, Cordillera Colan,
LSUMZ 39362.

Eleutherodactylus incomptus (63 specimens)

Ecuador: Prov. Napw: Rio Azuela, 1740 m, KU 143497; Rio Salado, ±
1 km upstream from Rio Coca, 1410 m, KU 146169, 165844-58, 177298-
319; 2 km [by road] SSW Rio Reventador, 1490 m, KU 165932; 16.5 km

76

Scientific Papers. Natural History Museum, The University of Kansas

[by road] NNE Santa Rosa, 1700 m, KU 143455, 1434bl-b4, 143484 (holo-
type), 143485-96. Prov. Pastnza: 9.5 km |by road] NW Mera, 1270 m, KU
179000-03.

Peru: Depto. Cajamarca: Santa Rosa de la Yunga, KU 217319-20.

Lleutherodactyhis infrnguttntus (6 specimens)
Peru: Deptto. Sni} Martin: East slope Abra Pardo de Miguel, 2180 m,
KU 212297 (holotype), 212298, 212314-16; 14 km [by road] W Venceremos,
2000 m, KU 217317.

EleuthcwdiKtylus lanthanites (50 specimens)
Ecuador: Prov. Sucumbios: Santa Cecilia, 340 m, KU 123852-77,

126215-16, 146144 (holotype), 146145-60.

Peru: Depto. Loreto: junction Rio Yanomono and Rio Amazonas, KU

220446, 220898; San Jacinto, 175-190 m, KU 222000-01. Dcpfo. Sun Martin:

Venceremos, 1630 m, KU 212225.

Eleutherodactylus lirellus (27 specimens)
Peru: Depto. San Martm: Abra Tangarana, 7 km [by road] NW San
Juan de Pacaysapa, 1080 m, KU 212240 (holotype), 212241, 212251-59;
Ponga de Shilcayo, 4 km [by road] NNW Tarapoto, 470 m, KU 212260-
63; Rio Cerranayacu, 76 km [by road] NW Rioja, 1200 m, KU 212226-32,
212234, 212236-39.

Eleutherodactylus tyinani (34 specimens)
Ecuador: Prov. Azuay: 55.9 km [by road] E Pasaje, 1000 m, KU 152009.
Prov. Lojn: Loja, 2150 m, KU 119502; 2 km [by road] E Loja, 2210 m, 3.9 km
[by road] E Loja, 2460 m, KU 202418; KU 119504-12; 7 km [by road[ E
Loja, 2500 m, KU 119503; 9 km [by road] E Loja, KU 202416; 7.6 km [by
road[ S Loja, 2210 m, KU 141962-64; 9 km [by road] S Loja, 2230 m, KU
165539^0; 12.2 km [by road] S Loja, 2275 m, KU 141292; 14.4 km [by
road] S Loja, 2260 m, KU 202417; 17 km [by road] NE Macara, 1240 m,
KU 141965.

Peru: Dtyto. Cajamarca: 28 km [by road] N Santa Cruz, 725 m, KU
196464, 196509, LSUMZ 19553. Depto. Puira: Canchaque, 1120 m, MHNSM
11172; 15 km [by road[ E Canchaque, 1850 m, KU 196465-69, 181265-67.

Eleutherodactylus niclanogaster (5 specimens)
Peru: Depto. Amazonas: north slope Abra Barro Negro, 28 km [by
road] SSW Leimebamba, 3470 m, KU 212321 (holotype), 212322-23,
218513; 25.5 km [by road] SSW Leimebamba, 3300 m, KU 181281.

Eleutherodactylus metabatcs (2 specimens)
PERu:Dt';)to. Amazonas: 20 km [by road] SW Chiriaco, 525 m, KU
196504 (holotype), LSUMZ 32460.

Eleutherodactylus muscosus (4 specimens)
Peru: Depto. San Martin: east slope Abra Pardo de Miguel, 1800 m,
KU 200479-81, 2094782 (holotype).

Eleutherodactylus nephophilus (8 specimens)
Peru: Depto. San Martin: Abra Tangarana, 7 km [by road] E San Juan
de Pacaysapa, 1080 m, KU 212311; east slope Abra Pardo de Miguel, 2180
m, KU 212305, 212306 (holotype), 212307-09, 212317; 14 km [by road] W
Venceremos, 2000 m, KU 212311.

Eleutherodactylus ockendeni (71 specimens)
Peru: Depto. Amazonas: 4 km [by road[ SW Chiriaco, 725 m, KU
196470. Depto. Cuzco: Atalaya, 650 m, KU 154797-99, 154805. Depto.
Hudnuco: Finca Panguana, Rio LluIIapichis, KU 154761-78, 154804,
171831^8; south slope Serrania deSira, 690-1280 m, KU 154779-85. Depto.
Loreto: junction Rio Sucusari and Rio Napo, KU 220356-57, 220449; San
Jacinto, 175-190 m, KU 222021; Teniente Lopez, 200 m, KU 222022; 1.5
km N Teniente Lopez, 31 0-340 m, KU 222023. 0177(0. Madre dc Dios: Cocha
Cashu, 400 m, KU 154788-96; Manu, 365 m, KU 154786, 154866. Depto.
San Martin: Rio Cainarachi, 33 km [by road] NE Tarapoto, 330 m, KU
209468-71.

Eleutherodactylus patalkos (1 specimen)
Peru: Depto. Amazonas: north slope Abra Barro Negro, 28 km Jby
road] SSW Leimebamba, 3470 m, KU 212320 (holotype).

Eleutherodactylus pecki (9 specimens)

Ecuador: Prov. Morona-Sautiago: Rio Piuntza, 1550 m, Cordillera del
Condor, KU 147040 (holotype), 147041-42; trail between Logroiio and
Yaupi, 1700 m, ANSP 29271, 29273-77.

Peru: Depto. Amazonas: Alfonso Ugarte, Rio Comainas, 1138 m,
USNM 525476.

Eleutherodactylus pycrcuopterus (24 specimens)
Peru: Depto. Amazonas: Alfonso Ugarte, Rio Comainas, 1138 m,
USNM 525445-46, 525449-63; 20 km [by road] SW Chiriaco, 525 m, KU
196506; 33 km [by road] SE Ingenio, 1830 m, KU 196505, LSUMZ 32462-
63; 5 km [by road] NW Mendoza, 2400 m, KU 209472; upper Rio
Comainas, base of Cerro Machinaza, 1750 m, USNM 525443; Santa Rosa
de la Yunga, 1300 m, KU 217318 (holotype).

Eleutherodactylus peruvianus (79 specimens)
Ecuador: Prov. Morona-Santiago: "Camp 2," Cordillera de Cutucii,
ANSP 29234, 29237-44, 29249-50, 29259, 29264; "Camp 3," Cordillera de
Cutucii, ANSP 29286; Rio Pintza, Cordillera del Condor, 1830 m, KU
147034-38; trail between Logroho and Yaupi, ANSP 29236.

Peru: Depto. Amazonas: Puesto Vigilancia, Rio Comainas, 665 m,
USNM 525440; upper Rio Comainas, base of Cerro Machinaza, 1750 m,
USNM 525438-39. Depto. Ayacucho: Estero Ruana in Tambo-Valle de
Apurimac trail, 1800 m, KU 196473; Huanuaehayocc on Tambo-Valle de
Apurimac trail, 1650 m, KU 196472, 196474. Depto. Huanuco: Finca
Panguana, Rio LluIIapichis, KU 154835-47, 154858-62, 1718667-91; South
slope Serrania sw Sira, 690 m, KU 154848-52, 154863-68. Depto. Loreto:
Quebrada Oran, 5 km N Rio Amazonas, 85 km NE Iquitos, KU 206094;
Quebrada Vainilla, 10 km SSW mouth of Rio Napo, KU 206093; Teniente
Lopez, 200 m, KU 222030; 1.5 km N Teniente Lopez, 310-340 m, KU
222024-29. Depto. Pasco: Santa Cruz, 9 km [by road] SSE Oxapampa, 2050
m, KU 206095-97. Depto. San Martin: Abra Tangarana, 7 km [by road] NE
San Juan de Pacaysapa, 1080 m, KU 212291; Rio Cainarachi, 33 km [by
road] NE Tarapoto, 330 m, KU 208475-77; 20 km (by road] NE Tarapoto,
KU 209473-74; 40 km [by road] NE Tarapoto, KU 217312; 15.4 km [by
road] SW Zapatero, 950 m, KU 217313.

Eleutherodactylus pctrohardus (7 specimens)
Peru: Depto. Cajamarca: 2 km [by road[ W Huambos, 2500 m, KU
212292 (holotype), 212293-96, MHNSM 6196-97.

Eleutherodactylus phoxocephalus (36 specimens)
Ecuador: Prov. Azuay: 10 km [by road[ SW Victoria del Portete, 2700
m, KU 131281-82. Prov. Canar: 18.4 km [by road] NW EI Tambo, 2860 m,
KU 142118-31. Prov. Loja: Rio Zamora, 6.5 km [by road] N Loja, 2060 m,
KU 142113; Saraguro, 2500 m, KU 135460-62; 3.3 km [by road] NNE
Saraguro, 2400 m, KU 142117; 2 km [by road] S Saraguro, 2680 m, KU
142114-16. Prov. Zamora-Chinchipc: 15 km [by road] E Loja, 2710 m, KU
142104-12.

Peru: Depto. Cajamarca: San Andres de Cutervo, KU 221710-13. Depto.
Piura: 15 km [by road] ENE Canchaque, 1850 m, KU 181271; El Tambo,
31 km [by road] ENE Canchaque, 2770 m, MHNSM 15399.

Eleutherodactylus pinguis (4 specimens)
Peru: Depto. Cajamarca: 57 km [by road] N Cajamarca, 3760 m, UF
40766; 23 km [by road] SW Celendin, 3050 m, KU 181282, 181283 (holo-
type); 33 km [by road] SW Celendin, 3200 m, KU 181284.

Eleutherodactylus proserpcns (10 specimens)
Ecuador: Prov. Loja: Abra de Zamora, 15 km [by road) E Loja, 2800

m, KU 202130-31. Prov. Morona-Santiago: "Camp 2," 1700 m, Cordillera

de Cutucu, ANSP 29230-33; Rio Piuntza, 1830 m, Cordillera del Condor,

KU 147044-46.

Peru: Depto. Amazonas: Upper Rio Comainas, base of Cerro

Machinaza, 1750 mm, USNM 525447.

Eleutherodactylus quaquaversus (52 specimens)
Ecuador; Prov. Morona-Sautiago, "Camp 2," 1700 m, Cordillera de
Cutucu, ANSP 29257, 29263, 29265-66, 29270, 29281, 29283-85; Rio

Eleutherodactylus in the Andes of Northern Peru

77

Piuntza, 1830 m, Cordillerd del Condor, KU 146'387-')0. Pwv. Napo: Rio
Azuela, 1740 m, KU 143448-54; 2 km |by road] SSW Rio Reventador, 1490
m, KU lb5563-65; Rio Salado, ± 1 km upstream from Rio Coca, 1410 m, KU
165566-85; 16.5 km [by road] NNE Santa Rosa, 1700 m, KU 143441^7.

PtKL': Dc/'hi. Aiiin20)inf: Upper Rio Comainas, base of Cerro
Machinaza, 1750 m, USNM 525444. Depto. Loreto: 1.5km N Teniente Lopez,
3U)-340m,KU 222031.

Eleuthcwdach/lus rhodoplichuf (24 specimens)

Perl': Dcf'tc. Pima: El Tambo, 31 km [by road ) ENE Canchaque, 2770
m, KU 219786 (holotype), 219787-91, MHNSM 15400-04; 12.7 km [by
road] NE El Tambo, 2820 m, KU 219792; crest of Cordillera de
Huancabamha, 33 km [by roadi SW Huancabamba, 3050 m, KU 196499-
503, LSUMZ 32417, 32428-33.

Elcuthcwilactyhif iltodosticlnif (4 specimens)
Peru: Depto. San Martin: West slope Abra Tangarana, 7 km [by road[
NE San Juan de Pacaysapa, 1080 m, KU 212264 (holotype), 212265-67.

Eleutherodactylus rufioculis (10 specimens)
Peru: Depto. Amazcnas: Alfonso Ugarte, Rio Comainas, 1138 m,
USNM 525474-75; Upper Rio Comainas, base of Cerro Machinaza, 1750
m, USNM 525464-65, 525467-68, 525473-74. Depto. San Martin: East slope
Abra Pardo de Miguel, 2180 m, KU 212312, 212313 (holotype).

Eleutherodactylus schultei (16 specimens)
Peru: Depto. Amazonas: 5 km [by road]N Levanto, 2850 m, KU

212220-21, 212222 (holotype), 212223-24, 209496-97, MHNSM 6172-73;
5 km [by road] NW Mendoza, 2400 m, KU 209498-504.

Eleutherodactylus serendipitus (5 specimens)
Peru: Depto. Ainazonas: 8 km [by road] NNE Balzapata, 1850 m, KU
181279 (holotype), 181280; 12 km (bv trail] E La Peca, Cordillera Colan,
LSUMZ 39360, 39363, 39377.

Eleutherodactylus sternothylax (49 specimens)
Peru: Depto. Piura: 15 km [by road I ENE Canchaque, 1735 m, 196479-
91, LSUMZ 32320, 32332-50, 32352, 32418-27, 32458; 16 km [bv road|
ENE Canchaque, 1840 m, KU 219793 (holotype), 212794, MHNSM 15405-06.

Eleutherodactylus versicolor (69 specimens)
Ecuador: Prov. Loja: 13.5 km [by road] E Loja, 2800 m, KU 1198858

(holotvpe), 119859-71, 119911-44, 141449-60, 141465-66; 8-9 km [by road|

N San Lucas, 3000-3100 m, KU 119945.

Peru: Depto. Ainazonas: Alfonso Ugarte, Rio Comainas, 1138 m,

USNM 525469, 525471-72; Puesto Vigilancia Rio Comainas, 665 ni, USNM

525441; Upper Rio Comainas, base of Cerro Machinaza, 1750 m, USNM

525466, 525477.

Eleutherodactylus wiensi (7 specimens)
Peru: Depto. Piura: 12.7 km [by road] ENE Canchaque, 1600 m, KU
219795 (holotype), 219706-97, MHNSM 15407-08; 15 km [by road[ ENE
Canchaque, 1735 m, KU 196510-11.

APPENDIX 2

Gazetteer

Following is a list of localities in the Andes of northern
Peru where specimens of Elcuthewdactifliis were collected.
After each place name the departamento is given in pa-
rentheses, followed by geographic coordinates, elevation,
and vegetation type. Except for localities on the eastern
slope of the Cordillera del Condor documented by
Reynolds and Icochea (1997), coordinates were determined
from maps (Mapa Ffsico PoUtico del Peru, 1 : 1,000,000, 1973,
and Carta Nacional del Peru, 1:100,000, 1986). Elevations
were obtained from altimeter readings or from maps. When
known, specific sites and habitats are given followed by
names of collectors (only those responsible for field notes)
and the month and years that they collected at the sites.
Names of collectors are abbreviated: AM - Alfonso
Miranda, ERW = Erik R. Wild, ERG = Fred G. Thompson,
GKN = G. K. Noble, KLC = Kenneth L. Campbell, LSU =
Louisiana State University ornithologists; RPR = Robert P
Reynolds, RAM = Russell A Mittermeier, RS = Rainer
Schulte, RT = Richard Thomas, WED = William E.
Duellman. All localities are shown on the map (Fig. 1).

Abra Barro Negro (Amazonas) — 06 '41' S, 77 51 W, 3500 m; very hu-
mid montane forest. A pass in the Cordillera Central on the road be-
tween Balsas and Leimebamba. Collections made on the north slope along
the road at 25,5 km SSW of Leimebamba, 3300 m, and 28 km SSW of
Leimebamba 3470 m in January 1989 by WED. At both sites there were
few trees; dominant vegetation consisted of small shrubs (Baccharis),
bunch grass, ferns, scattered flower herbs, and mosses 10-20 cm deep.

Abra Pardo de Miguel ( Amazonas-San Martin)— 05 ■46' S, 77 '47 W, 2210
m; very humid montane forest. A pass in the eastern range of the Cordil-
lera Central. Collections made on the east slope at about 1800 m in Janu-
ary 1981 by RS and just below the crest on the road to La Rioja, at an
elevation of 2180 m in January 1989 by WED. Many trees laden with

bromeliads, tree ferns, spiny bamboo (Chiisquea spicata) and broad-leafed
Cunnera; rocky cliffs covered with mosses and lichens.

Abra Tangarana (San Martin)— 06 °1 2' S, 76 "44' W, 1120 m; humid sub-
tropical forest. A pass in a low ridge of the Cordillera Central, 7 km NE of
the small village San Juan de Pacaysapa on the road between Moyohamha
and Tarapoto. Collections were made in a rocky ravine on the western
slope by WED in February 1989. Trees in the forest held many bromeli-
ads; elephant-ear plants (Xanthosoma) were abundant along the stream.

Alfonso Ugarte (= Puesto Vigilancia 3) (Amazonas)— 03 °54' S, 78°25' W,
1138 m; humid montane forest. Military post on the upper Rio Comainas,
eastern slope of Cordillera del Condor. Collections made in July-August
1994 by RPR.

Alva (Amazonas) — Approximately 05°53' S, 78 "56' W, 1000 m; pre-
sumably thorn forest. Site on slopes of Cordillera Central above Pedro
Ruiz Gallo in Rio Utcubamba Valley. Collection in May 1974 by RAM.

Balzapata (Amazonas) — 05°46'S, 77*51' W, 1640 m; humid montane
forest. A village just west of the crest of the Cordillera Central. Collec-
tions made in disturbed and partly cultivated forest 8 km NNE (1850 m)
on the road to La Rioja in March 1979 by WED.

Cajamarca (Cajamarca) — 07°12' S, 78°30' W, 2750 m; montane dry for-
est. City in Cordillera Occidental. Collection from grassland in humid
montane forest at 3750 m, 57 km N of Cajamarca in April 1972 by FGT.

Canchaque (Piura)— 0572' S, 7936' W, 1120 m; tropical dry forest. A
town near the western base of the Cordillera de Huancabamba. Collec-
tions were made in dry forest near the town and humid montane forest
at varying distances from the town on the road across the Cordillera de
Huancabamba to the town of Huancabamba in April 1970 by KLC, De-
cember 1974 by RT, March 1979 by WED, and January 1991 by ERW.

Cataratas Ahuashiyacu (San Martin)— 0630' S, 76 "20' W, 730 m; humid
subtropical forest. Waterfall and rocky stream in deep ravine 14 km NNE
of Tarapoto on the road to Yurimaguas. Collections in February 1989 by
WED.

Celeiidin (Cajamarca)— 06 °52' S, 78"08 W, 2625 m; humid montane
forest. City in the Cordillera Occidental. Collections from the east slope
of Abra Comulica, 23 km SW (3050 m) and 33 km SW (3200 m) of Celendin

78

Scientific Papers, Natural History Museum, The University of Kansas

in March 1979 by WED. Both sites heavily disturbed, mostly grassy marsh
(3050 m) and bunch-grass-BiJcdinns association.

Chirmco (Amazonas)— 05 °09' S, 78 71 ' W, ± 450 m; thorn forest. Town
on the Rio Maranon a the confluence of the Rio Chiriaco. Collections
made in dry forests 4 km SW and 20 km SW (525 m) on road to Bagua in
December 1974 by RT

El Tambo (Plura)— 05°21'S, 7933' W, 2770 m; humid montane forest.
Settlement on the western slope of the Cordillera de Huancabamba, 31
km NNE of Canchaque on the road to Huancabamba. Collections in rela-
tively undisturbed forest in March 1979 by WED and January 1991 by
ERW.

Huambos (Cajamarca)— 06°27' S, 78 "57' W, ± 2200 m; montane dry for-
est. Town on the western slopes of the Cordillera Occidental. Collections
from "pre-Incan ruins" in 1916 by GKN and in scrubby forest with ter-
restrial bromeliads 2 km W (2500 m) in February 1989 by WED.

Huancahambn (Piura)— 05"14' S, 79°28' W, 1957 m; dry forest. City on
the Rio Huancabamba at eastern base of the Cordillera de Huancabamba;
area in immediate vicinity mostly cultivated. Collections from very hu-
mid montane forest just east of and at the crest of the Cordillera de
Huancabamba, 25-26 km SW (3010-3050 m) in March 1979 by WED and
31-33 km SW (5050-3100 m) in December 1974 by RT and in March 1979
by WED.

Ingenio (Amazonas)— 05 °55' S, 78°55' W, 1280 m; thorn forest. Settle-
ment in the Ri'o Uti.icubamba Valley. Collection from dry forest on slopes
of Cordillera Central 33 km SE (1830 m) on road to Chachapoyas in De-
cember 1974 by RT

Lamns {Llama) (Cajamarca)— 0631' S, 79 "07' W, ±2200 m; thorn forest.
Village in valley on west slope of Cordillera Occidental. Collection from
12 km W (1500 m) in December 1974 by RT

La Peca (Amazonas)— 0536' S, 78"22' W, ± 1400 m; thorn forest. Vil-
lage near western base of Cordillera Colan. Collections from the Cordil-
lera Colan — humid montane forest 12 km (by trail) E (1700 m) and very
humid montane forest E (2963-3330 m) in August-September 1978 by
LSU.

Lcimebaniba (Amazonas) — 0633'S, 77°49' W, 2200 m; humid montane
forest. Town in the upper Rio Utcubamba drainage in the Cordillera Cen-
tral. Collection from 24 km (by road) SW, 3370 m in very humid montane
forest in April 1 972 by FGT

Levanto (Amazonas) — 06''17'S, 77 "21' W, 2650 m; humid montane for-
est. Village south of Chachapoyas in the Cordillera Central. Collection
from bromeliad-laden trees in highly disturbed forest 5 km N (2850 m) in
January 1989 by WED.

Mendoza (Amazonas) — 06°I8'W, 77°25' W, 1650 m; humid montane
forest. Town in the Cordillera Central. Collection from pass (headwaters
of Rio Chiriaco) 5 km NW (2400 m) in July 1981 by RS.

Palambla (Piura)— 05 °23'S, 7936' W, 1120 m; tropical dry forest. Vil-
lage near the western base of the Cordillera de Huancabamba. Collec-
tion in 1916 by GKN.

Perico (Cajamarca)— 0572' S, 78"47' W, ±1000 m; thorn forest. Village
in Rio Marafion Valley; collection in 1916 by GKN.

Pucsto Vigilancia Coinainas, Rio Coiuamaf (Amazonas) — 04 '06' S, 7873'
W, 665 m; humid subtropical forest. Military camp on the west bank of
the Rio Comainas, eastern slope of Cordillera del Condor. Collections
made in July-August 1994 by RPR.

Rib Cenanayacu (San Martin)— 05 •46' S, 7777' W, 1200 m; humid
montane forest. Tributary of the Rio Mayo on the eastern flank of the
Cordillera Central. Collection from narrow valley near crossing of the
Balzapata-La Rioja road in February 1989 by WED.

Rio Comainas, base of Cerro Machinaza (Amazonas) — 03°53' S, 7875'
W, 1750 m; humid montane forest. Site on the eastern slopes of the Cor-
dillera del Condor. Collection in July-August 1994 by RPR.

San Andres de Ciitervo (Cajamarca)— 05 T4' S, 78°42' W, ± 2500 m; hu-
mid montane forest. Village in the Cordillera Occidental. Collection in
June and December 1992 by AM.

Santa Cruz (Cajamarca)— 06 °05' S, 78 "51' W, ± 1300 m; thorn forest.
Village in the Rio Chamaya Basin. Collection from 28 km N (725 m) in
December 1974 by RT.

Santa Rosa de la Yunga (Cajamarca)— 06 ''05' S, 78'43' W, 1000 ms; dry
forest. Village on the southern slope of Cordillera del Condor. Collection
from disturbed dry forest immediately above village in July 1989 by RS.

Tarapoto (San Martin)— 0631' S, 7673'W, 370 m; humid tropical for-
est. Town near confluence of Rio Mayo and Ri'o Huallaga just east of
Cordillera Central. Collections from various distances and elevations in
February 1989 by RS and WED.

Venceremos (San Martin) — 05"44' S, 7731' W, 1630 m; humid montane
forest. Former road camp and now small settlement on the road between
Balzapata and La Rioja. Collections from immediate vicinity in February
1989 by WED and from 14 km W (2000 m on road to Balzapata) in July
1981 by RS.

Zapatero (San Martin)— 0634' S, 7630' W, 320 m; humid tropical for-
est. Village at base of hills north of the Rio Huallaga. Collections from
disturbed humid montane forest 15.4 km SW (950 m) in June 1989 by RS
and from ravine in cutover humid tropical forest 30 km SW in February
1989 by WED.

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