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663 SEUM OF NATURAL HISTORY NO. 69
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The Eleutherodactylus of
the Amazonian Slopes of
the Ecuadorian Andes (Anura:
Leptodactylidae)
By
John D. Lynch
and
William E. Duellman
UNIVERSITY OF KANSAS

Oynch, ID

LAWRENCE 1980

UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY

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SS

Tue UNIVERSITY OF KANSAS

MusEuM OF NATURAL HISTORY

MISCELLANEOUS PUBLICATION No. 69
August 29, 1980

The Eleutherodactylus of the Amazonian Slopes
of the Ecuadorian Andes (Anura: Leptodactylidae)

BY

Joun D. Lyncu

Professor, School of Life Sciences
The University of Nebraska, Lincoln, Nebraska, 68588 U.S.A.
Associate, Division of Herpetology,
Museum of Natural History, The University of Kansas
Lawrence, Kansas 66045 U.S.A.

AND
WILLIAM E.. DUELLMAN

Curator, Division of Herpetology
Museum of Natural History
Professor, Department of Systematics and Ecology
The University of Kansas, Lawrence, Kansas 66045 U.S.A.

THe UNIVERSITY OF KANSAS
LAWRENCE
1980

UNIVERSITY OF KANSAS PUBLICATIONS, MusEUM OF NATURAL HisToRY

Editor: E. O. Wiley
Managing Editor: Joseph T. Collins

Miscellaneous Publication No. 69
pp. 1-86; 8 figures; 10 tables
Published August 29, 1980

ae Museum oF Natura History

NEE THE UNIVERSITY oF KANSAS

LAWRENCE, Kansas 66045
U.S.A,

PRINTED BY
UNIVERSITY OF KANSAS PRINTING SERVICE
LAWRENCE, KANSAS

CONTENTS

TUSTFEETEN MID UICC TL © yet es Seo wh gS Soe ne ee eoeeene nee ieee, +)
IN ORASONIEADD CHS TAT Sy ea seis ne Sats SS SSIS eee eee psoas ete ||
CE CISUIPLEIN UN TCCAS® Soe ee ieee ozone See nia cee
Ue toy STOIC CUED AEA CSS ese ec rrereer cee 3
HRSG ee) LETC SECUESS ops es nc a 9
PAGER INS OE” ETE) SPECIES ose 11
Eleutherodactylus acerus new species —..—--——---------------------- 12
Eleutherodactylus atratus Vayneh 17
Eleutherodactylus baryecuus Lynch ---—---—-------------------------- 17
Eleutherodactylus bromeliaceus Lynch ———------------------------------- 17
Eleutherodactylus chloronotus Lynch irl
Eleutherodactylus colodactylus Lynch ~--—------------------------------------- 18
Eleutherodactylus condor new species -..-_—-----------------------—--------------- 18
Eleutherodactylus cornutus (Jiménez de la Espada) ——------------ 21
Eleutherodactylus cremnobates new species ———-------------------------------—- 21
Eleutherodactylus croceoinguinis Lynch —---------------------------—- 24
Eleutherodactylus cryptomelas Lynch ——--------------------------------~ 24
Eleutherodactylus devillei (Boulenger) ——---------------------------------- 25
Eleutherodactylus dolops new species --.--——------------------=---------- 26
Eleutherodactylus elassodiscus Lynch _—_______________-- 27
Eleutherodactylus eriphus new species 28
Eleutherodactylus galdi (Jiménez de la Espada) ———--------------------- 30
Eleutherodactylus gladiator Lynch 31
Eleutherodactylus glandulosus (Boulenger) -.-—--------------------------- 32
Eleutherodactylus ignicolor new species —---—-—------------------—---------— 32
Eleutherodactylus incanus new species — == 34
Eleutherodactylus incomptus new species — 35
Eleutherodactylus inusitatus new species — = 37
Eleutherodactylus lacrimosus (Jiménez de la Espada) —----------- 38
Eleutherodactylus lanthanites Lynch __---------—------------------------------ 39
Eleutherodactylus leoni Lynch _....----------------—---------------- 40
Eleutherodactylus leucopus Lynch _---——-—-------------------------- 4]
Eleutherodactylus lividus new species —-——————----------------- 4]
Eleutherodactylus nigrogriseus (Andersson) ——-------—----—-------------- 43

Eleutherodactylus nigrovittatus Andersson ———-------------—------------ 44

Eleutherodactylus ockendeni (Boulenger) _... 44

Eleutherodactylus pastazensis Andersson _.....--- 45
Eleutherodactylus peruvianus (Melin) 47
Eleutherodactylus petersi new species _..... 48
Eleutherodactylus prolatus new species 51
Eieuiherodactylus pirosenpens ty nc saan 52
Bleuthenodactyluspugnax Wyrchy se ee ee 52
Eleutherodacilusvpycnodermis Iaynch\ sss. 2 eee 53
Eleuthenodaciylis quaguaversus aymch, == 2s 53
Eleutherodactylus rubicundus (Jiménez de la Espada) _. 54
Eleutherodactylus spinosus \uynchy =.=) aan 56
Eleutherodactylus trachyblepharis (Boulenger) — 57
Eleutherodaciylus trepidotus, Lynch —= 2 58
Eleutherodactylus unistrigatus (Ginther) “22 ee 59
Eleutherodactylus ventrimarmoratus (Boulenger) — 60
Eleutherodactylus w-nigrum (Boettger) 61
RELATIONSHIPS OF THE WAUNA) 28 ee eee 63
DISTRIBUTION AND ECOLOGY 9.22) 5. es eee 65
IDESCRIPTION OF SINECION 225 4) ks). ee 65
PATTERNS OF [DISTRIBUTION 425.4222. 5g ee 68

EH LEUSHERODACTYEINE, (COMMUNITIES 222 a 70
RE SU MEING eee 8 Be Os ee 74
EVER RAT URE} CID ED. se 25. 6) el ee ee 75
PUBIP EE NID UGS cae oo see 77
SPECIMENS: ECXAMINED 92.260) eee ee Ue

INTRODUCTION

Field work on the Amazonian slopes
of the Andes in southern Colombia and
Ecuador has revealed, and continues to
reveal, a rich anuran fauna. Among the
124 species of frogs now known from
elevations of 1000-3000 m on the Ama-
zonian slopes in Ecuador, 12 belong to
the dendrobatid genus Colostethus, 12
to the centrolenid genus Centrolenella,
and 50 to the genus Eleutherodactylus.
Frogs of the genus Eleutherodactylus
have served as an impediment to sum-
maries or checklists of the Ecuadorian
amphibian fauna. Although some spe-
cies probably remain to be discovered,
we approach a working knowledge of
the fauna after a decade of field work.
The eleutherodactyline fauna of the
Amazonian lowlands seems to be mostly
known (Duellman, 1978; Lynch, 1980).
Lynch (1979a) summarized the eleu-
therodactyline fauna of the southern
Andes of Ecuador, including several of
the species found on the Amazonian
slopes.

In this paper we address 45 species
of Eleutherodactylus found at elevations
of 1000-3000 m on the Amazonian slopes
of the Andes in Ecuador. Records also
are given for those same species on the
adjacent slopes in Colombia; in so doing
we list more than 2800 specimens ex-
amined. The elevational limits essen-
tially correspond to the boundaries of
the lowland tropical rainforest and the
paramo. We are concerned with the
frogs inhabiting the montane rainforest
or cloud forest. These altitudinal limits
appear to have biological significance in
that only eight of the 45 species descend
below 1000 m in Ecuador; five of these
are actually lowland species that ascend
the slopes into the cloud forest. Seven
species also occur above 3000 m; one of
those is primarily a highland species that
descends into montane forest.

Twelve of the 45 species are named
as new. We also report and describe
E. pastazensis Andersson, E. rubicundus

(Jiménez de la Espada), and E. trachy-
blepharis (Boulenger), none of which
has been mentioned in the literature
(except in checklists) since their orig-
inal descriptions. Not included in this
report are E. buckleyi (Boulenger),
which descends into upper montane for-
est near El Carmelo (east of Tulcan,
Ecuador), or a species presently con-
fused with E. vertebralis (Boulenger),
also from the El Carmelo area. Eleu-
therodactylus buckleyi occurs primarily
in paramo and will be treated by Lynch
in an account of the frogs of the north-
ern Andes of Ecuador. The other spe-
cies is widespread in the Colombian
Andes, but a systematic summary is
postponed pending re-examination of the
complex of frogs now called E. verte-
bralis. Eleutherodactylus chalceus (Pe-
ters), a name long associated with the
Amazonian slopes and lowlands, will be
shown by Lynch (ms) to be identified
with a species on the Pacific slopes and
lowlands.

As part of our systematic treatment
of the frogs, we present a discussion of
taxonomic characters in the genus Eleu-
therodactylus and a key to the identifica-
tion of the species on the Amazonian
slopes of the Andes in Ecuador. In an
attempt to summarize our knowledge of
these frogs in the region, we provide dis-
cussions of the distribution of the spe-
cies and of the eleutherodactyline com-
munities. The appendices include a list
of specimens examined and a gazetteer
of localities from which specimens of
Eleutherodactylus were collected.

ACKNOWLEDGMENTS

We are especially grateful to our
field companions whose endeavors great-
ly enhanced the collections of specimens
and field data. At various times Duell-
man was accompanied by Joseph T.
Collins, Dana T. Duellman, Bruce Mac-
Bryde, Alan H. Savitzky, John E. Sim-

2 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

mons, and Linda Trueb. Lynch was
variously accompanied by David C.
Cannatella, Robert W. Henderson, Mar-
sha C. Lynch, and Gerald R. Smith.
Duellman’s field work was accomplished
as part of research supported by the
National Science Foundation (Grants
DEB 74-02998 and 76-09986), the Na-
tional Geographic Society (Grant No.
1304), and the Watkins Fund of the
Museum of Natural History, University
of Kansas. Lynch’s field work was made
possible by grants from the Society of
the Sigma Xi, Watkins Fund of the
Museum of Natural History, and the
University of Nebraska Research Coun-
cil. Personnel of Williams Brothers Con-
struction Company provided logistic
support for Duellman in 1968 and 1971-
72. Fernando Ortiz C. and Eugenia del
Pino generously provided facilities for
our use at the Universidad Catodlica,
Quito.

For the loan of specimens or the pro-
vision of working space in their respec-
tive institutions, we are grateful to
Charles W. Myers and Richard G. Zwei-
fel, American Museum of Natural His-
tory; Alice G. C. Grandison, British
Museum (Natural History); Alan E.
Leviton, California Academy of Sciences;
Gaston F. DeWitte, Institut Royal des
Sciences Naturelles de Belgique, Brus-
sels; Ernest E. Williams, Museum of
Comparative Zoology, Harvard Univer-
sity; Umberto Parenti, Museo Civico di
Storia Naturale di Torino; Javier Cas-
troviejo, Museo National de Ciencias
Naturales, Madrid; Greta Vestergren,
Naturhistoriska Riksmuseum, Stockholm;
Dorothy Smith, University of Illinois Mu-
seum of Natural History; Arnold G.
Kluge, Ronald A. Nussbaum, and
Charles F. Walker, University of Michi-
gan Museum of Zoology; the late Doris
M. Cochran, W. Ronald Heyer, the late
James A. Peters, and George R. Zug,
National Museum of Natural History;
Werner C. A. Bokermann, Sao Paulo,
Brasil; and Eugenia del Pino, Quito,
Ecuador. The American Philosophical

Society (Penrose Fund No. 5063) pro-
vided funds for Duellman to visit Euro-
pean museums; Lynch’s visit to the
British Museum was made possible by
a grant from the University of Nebraska
Research Council.

We are indebted to Debra Bennett
for executing Figs. 6-8 and to Linda
Trueb for rendering Figs. 1 and 2B.

Specimens are identified in the text
by museum number and an acronym for
the collection or institution. We have
used the acronyms suggested by Duell-
man et al. (1978).

AMNH American Museum of Natural
History, New York

ANSP Academy of Natural Sciences,
Philadelphia

BM British Museum ( Natural
History ), London

CAS-SU Stanford University collec-

tion; housed at California
Academy of Sciences,
San Francisco

EP Eugenia del Pino collection,
Universidad Catdlica,
Quito, Ecuador

GOV Gustavo Orcés V. collection,
to be catalogued at USNM

JAP James A. Peters collection,
to be catalogued at USNM
KU University of Kansas Museum

of Natural History, Lawrence

Louisiana State University

Museum of Zoology,

Baton Rouge

MCZ Museum of Comparative

Zoology, Harvard University,

Cambridge

Institut Royal des Sciences

Naturelles de Belgique,

Brussels

Museo Civico di Storia

Naturale de Torino, Torino

Naturhistoriska Museet,

Goteborg

SMF Natur-Museum Senckenberg,
Frankfurt-am-Main

LSUMZ

MRHN

MSNT

NHMG

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 3

UIMNH University of Illinois Museum USNM _ National Museum of Natural
of Natural History, Urbana History, Washington, D.C.
UMMZ University of Michigan WCAB _ Werner C. A. Bokermann
Museum of Zoology, collection, SAo Paulo, Brasil
Ann Arbor
SYSTEMATICS

Frogs of the genus Eleutherodactylus
are notorious for their taxonomic diff-
culty. This stems in large part from the
sheer size of the genus—we estimate
more than 400 species are now recog-
nized—but also because development
and discussion of taxonomically useful
characteristics has been slower than de-
scriptions of species. In addition to the
descriptions provided below with their
comparable diagnoses, we provide a dis-
cussion of the taxonomic characteristics
and a key to the species found on the
Amazonian slopes of the Andes. Lastly,
a synopsis of the taxonomic relationships
is provided even though the synopsis is
far from complete—it represents our
judgments at this stage of the assess-
ment of this large genus.

TAXONOMIC CHARACTERISTICS

Measurements and abbreviations em-
ployed are as follows: snout-vent length
(SVL)—distance from tip of snout to
vent; shank—measured on flexed hind
leg from knee to heel, including only
tibiofibula; head width (HW )—greatest
width of head, usually at level of an-
terior edge of tympanum; head length—
head length, estimation of distance from
snout tip to line between ricti (not di-
rect measure of distance between snout
tip and rictus as employed by Zweifel,
1955); upper eyelid width—distance be-
tween outer edge of upper eyelid and
inner margin of eye bulge, approximate-
ly. perpendicular to margin of jaw (not
measurable if eye retracted); interorbi-
tal distance (IOD)—shortest distance
between eyes across braincase (not a
measure of braincase width, which is
usually shorter); eye length—distance
between anterior and posterior corners

of eye (difficult to measure if eye is re-
tracted); tympanum length—distance
from anterior edge of tympanic annulus
to posterior edge of annulus; eye to nos-
tril distance (E-N)—distance between
posterior edge of nostril and anterior
edge of eye (usually reflects snout
length). Reported values for measure-
ments and proportions normally include
the range of values, mean, and number
of values. In some cases, two standard
errors of the mean are also reported
(e.9., k =. 34.8. .2:6).

The order in which characteristics
are addressed corresponds to that em-
ployed in the diagnosis. The sequence
has been employed by JDL for the past
ten years; the logic for the particular
sequence has been forgotten, but gen-
erally follows JDL’s search pattern when
examining frogs, viz., general texture of
skin, condition of tympanum, head
shape,, traits within the mouth, structure
of hands, structure of feet, color pattern,
and size.

Skin texture—The skin of the venter
is smooth or bearing wrinkles in mem-
bers of the biporcatus, discoidalis, and
fitzingeri groups but is granular (areo-
late) in members of the wnistrigatus
group. Eleutherodactylus galdi (Jimén-
ez de la Espada) has areolate skin on
the venter but also has larger more
prominent warts scattered on the belly.
Eleutherodactylus rubicundus has finely
areolate skin on the venter; the granules
are usually not visible without magnifi-
cation. The throat is smooth (or wrin-
kled) in all taxa.

Many eleutherodactylines are readily
sorted using the array of features gen-
erally described as being part of the
texture of the skin of the dorsum. Dor-

4 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

ae

solateral folds are evident in several
members of the fitzingeri group as well
as in high montane species of the uni-
strigatus group. Such folds extend from
above the tympanum to a point above
the groin. In E. conspicillatus (Gin-
ther) and E. peruvianus (Melin) the
folds are series of closely packed warts
arranged in a line.

The paravertebral folds of E. chloro-
notus Lynch are sinuous and may repre-
sent extensions of occipital folds (or
rows of tubercles) seen in many eleu-
therodactylines. The occipital folds are
frequently accented with W-shaped dark
(or pale) markings. The system of folds
and ridges of E. cornutus (Jiménez de
la Espada) includes a H-shaped occipi-
tal fold as well as many short folds
arranged in a complex but repeated pat-
tern; initial perception of the skin of
E. cornutus is that it is tuberculate.

The skin between whatever folds
and/or enlarged warts on the _ back
ranges from perfectly smooth (even un-
der magnification), to finely shagreened
(tiny, uniform-sized granules), to sha-
greened (larger granules, sometimes vis-
ible without magnification, but not
pungent), to tuberculate or spinous
(large, pungent granules). Our char-
acterizations of skin texture take the
form of one of these states plus an at-
tempt to describe whatever folds and
enlarged warts are present.

Some enlarged tubercles are more
obvious. Enlarged warts on the upper
eyelid, between the eyes, or on the fore-
arm, knee, heel, and tarsus are usually
noted in connection with traits other
than skin texture (see below).

Tympanum.—Most species of Eleu-
therodactylus have complete ears (tym-
panic annulus, cavum tympanicum, plec-
trum), but one species on the northern
Amazonian slopes lacks all of these fea-

Fic. 1—Tympanum concealment in Eleu-
therodactylus. Top to bottom: E. quaquaver-
sus, KU 123749; E. incomptus, KU 165844; E.
prolatus, KU 146165; E. eriphus, KU 155451.
Left side of head is illustrated without color
pattern or tubercles.

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 5

tures (E. pugnax Lynch). Many other
species have complete but concealed
ears, which sometimes may be detected
in desiccated individuals. When the
tympana are visible (Fig. 1), care must
be taken to recognize the state in which
the tympanum is visible through thin
skin (= partially visible) as different
from the case in most frogs having a
visible tympanum (the raised annulus
is visible along at least the anterior,
ventral, and part of the posterior mar-
gins of the tympanum; the upper border
of the annulus is normally obscured by
the supratympanic fold—extending from
the eye, above the ear, to above the
insertion of the forearm). Relatively
few taxa may be distinguished on the
basis of tympanum size; the distinguish-
ing ability is further hampered by shape
differences between sexes (usually more
round in males, higher than long in
females).

Head shape.—Rivero (1961) de-
scribed character-states for snout shape
in Venezuelan frogs. The Eleutherodac-
tylus on the Amazonian slopes exhibit
acuminate, subacuminate, ovoid, and
rounded snouts (as seen in dorsal view).
In lateral profile, fewer variations are
considered modes (rounded, acutely
rounded, truncate, and _ protruding);
Lynch (1974) provided illustrations for
some of these based on species found in
the Amazonian lowlands. Many species
have a papillum on the tip of the snout
changing the aspect of snout shape.

Rivero (1961) also diagramed dif-
ferences in the shape of the canthus
rostralis. The canthus rostralis is sharp
(approximately 100°) and_ either
straight, weakly concave, or weakly con-
vex in most species of the fitzingeri
group; this character-state is usually cor-
related with the loreal region sloping
abruptly to the lips. In those frogs in
which the loreal region slopes gradually
to the lips, the canthus rostralis is less
sharp (obtuse) but sufficiently distinct
to allow characterization as concave or
convex. Rounded canthi rostrali consti-

tute the other extreme in which the can-
thus is obsolete.

In relatively few Eleutherodactylus
(but in many members of the uwnistri-
gatus group on the northern Amazonian
slopes), the lips are flared. The flaring
is pronounced in the broad-headed frogs
of the sulcatus group and in E. cruentus
(Peters) and E. latidiscus (Boulenger)
from trans-Andean South America. Frogs
having flared lips have gently sloping
loreal regions and/or flaring evident be-
low and posterior to the eyes.

Traits of the mouth.—On the roof of
the mouth, the sizes and relative posi-
tions of the choanae and vomerine odon-
tophores are the principal characteristics.
The choanae are usually not concealed
by the palatal shelf of the maxillary arch
(condition seen in all Amazonian slope
taxa). The choanae are of various sizes,
and size is normally referenced to the
size of the frog rather than to the vom-
erine odontophores.

The vomerine odontophores (if pres-
ent) are well posteriad to the choanae
and, in all taxa except the members of
the biporcatus and discoidalis groups,
well medial to the choanae as well. In
the exceptions, the lateral edge of the
odontophore extends to the level of the
choanae. The shapes of the odonto-
phores are triangular, broad (or arched),
round, oval, or oblique. Oblique odon-
tophores are arranged directed medio-
posteriorly; the most medial (or poste-
rior) component is of the greatest ele-
vation. The teeth on the odontophores
assume transverse, slanted, or clumped
distributions (correlated with triangular
and broad, oblique, and round or oval
shapes). Odontophore size is expressed
relative to the size of the choanae.

Tongue shape has been used in frog
identification for more than a century
but is of relatively little value (as ex-
pressed by failure to include it in the
numbered traits). The degree of at-
tachment of the tongue and presence or
absence of a notch along the posterior
border appears to be of some, although
limited, value in species separation.

6 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Adult males are often recognized as
adults on the basis of the presence of
vocal slits posterolateral to the tongue
and/or the presence of an external vocal
sac. However, many species of Eleu-
therodactylus lack both vocal slits and
external vocal sacs. Few variations in
the form of the vocal sacs are expressed
in Eleutherodactylus; E. crenunguis
Lynch from the Pacific versant in Ecua-
dor has weakly bilobed vocal sacs and
several high altitude species apparently
have internal vocal sacs.

Structure of the hands.—The posterior
border of the forelimb may bear conical
(or in some cases elongate, e.g., E. in-
usitatus) tubercles. Most species have
an antebrachial tubercle but relatively
fewer have a complete series of tubercles
along the forearm (ulnar tubercles). The
ulnar tubercles may coalesce forming a
scalloped ridge (as in E. cornutus) but
are normally discrete.

The large tubercles on the palm in-
clude an inner thenar tubercle on the
thumb (normally considerably smaller
than the lateral palmar tubercle). The
palmar tubercle is bifid (or cordiform)
in all of the Andean slope Eleutherodac-
tylus but in other species of the genus
there is a round palmar tubercle (E.
zeuctotylus Lynch and Hoogmoed) or
divided palmar tubercles wherein the
bifid palmar is in two distinct elements.
The remaining palmar surface may bear
supernumerary palmar tubercles. These
are normally non-pungent and sometimes
difficult to see. They are smaller than
the subarticular tubercles and the thenar
and palmar tubercles, which are more
elevated.

The subarticular tubercles are the
most prominent tubercles on the ventral
surface of the manus. The inner two
fingers each have one tubercle whereas
each of the outer two fingers has two
tubercles. The tubercles are round or
elongate or, more rarely, broader than
long. In most eleutherodactylines the
subarticular tubercles are pungent (ele-
vated), but in some species they are
flattened and in others are conical.

In many Eleutherodactylus the lat-
eral margins of the digits bear keels or
fringes. In no case are the fringes com-
parable to those seen in elosiine lepto-
dactylids. The fingers develop false keels
or fringes if the specimen is allowed to
desiccate, so some care must be exer-
cised in the use of this character. Fringes
are easily seen but the states “keeled”
and “lacking keels or fringes” are more
difficult to distinguish. If the digit is
rotated during examination under mag-
nification, the keel becomes apparent.
However, it is desirable to inspect sev-
eral fingers along their entire margins to
gain certainty. If the pad is collapsed,
the keels are probably due to desicca-
tion.

All Eleutherodactylus bear discs on
the ventral surfaces of the digit tips of
at least the toes regardless of whether
or not the digit is expanded to form a
pad. Some species lack discs on one or
more fingers of the hand. Lynch (1975a)
reported the absence of finger discs in
E. sulcatus (Cope) and (1976a) the
loss of some discs (inner fingers) in the
pyrrhomerus assembly (unistrigatus
group). The discs are normally broader
than long but in many species having
narrow pads are as long as wide (see
Lynch 1976b, 1978).

Pad size is subject to greater varia-
tion than discs size (or presence). The
digital pads may be apically rounded
(most), emarginate, notched, or pointed.
Normally, the pad of the thumb is not
developed or only slightly broader than
the digit below the disc whereas the
pads of fingers II, III, and IV are readily
seen and those of III and IV normally
larger than that of II. However, the fin-
gers of E. dolops, E. elassodiscus, and
E. nigrovittatus (among Amazonian
slope species) lack pads on the fingers.

The relative lengths of the inner two
fingers was used by Lynch (1976b) to
separate some species groups. Frogs of
the wnistigatus group (abundant on the
Amazonian versant) have the first finger
shorter than the second, whereas frogs
of the biporcatus, discoidalis, and fitz-

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES Tf

ingeri groups have the first finger longer
than the second. To record this trait,
the inner two fingers must be adpressed
equally; if one is adpressed against the
other, finger lengths may not be correct-
ly determined.

Males of many Eleutherodactylus
have non-spinous, white (or cream)
nuptial pads on the thumbs. The pad is
distinctly glandular in some species but
not in all. Many other species lack this
secondary sex feature. Although Taylor
(1952), Rivero (1968), and Lynch
(1970a) cited the feature, it remained
for Savage (1975) to point out its taxo-
nomic utility and frequency.

Structure of the foot and tarsus.—
The heel may lack tubercles or may bear
obvious but non-conical tubercles, one
or more conical tubercles, or a single
elongate tubercle or calcar. The outer
edge of the tarsus usually bears two to
four non-conical or more rarely conical
tubercles if there is a conical tubercle
on the heel. The tubercles rarely con-
tinue along the outer edge of the foot
and toe V (but are sometimes replaced
by a ridge).

The inner edge of the tarsus normally
lacks tubercles or folds, but some species
have a single tubercle somewhere along
the distal one-half of the tarsus. Other
species have a low ridge in place of the
tubercle (usually in contact with the
inner metatarsal tubercle). The least
common variation is the presence of a
fold on the inner edge of the tarsus
(normally on the distal one-half only).

All Eleutherodactylus have two meta-
tarsal tubercles; the inner is normally
larger and longer than wide, whereas
the outer is usually round in outline.
The inner metatarsal tubercle is de-
scribed as oval (length twice width) or
elongate (length three times width).
The outer metatarsal tubercle is de-
scribed in terms of the size of the inner
and in a few Eleutherodactylus is as
large as the inner. Supernumerary plan-
tar tubercles may be entirely absent but
most species have some. These tubercles
are low (non-pungent) and smaller than

the smallest subarticular tubercle. Frogs
may have only two to four tubercles
(proximal to the basal subarticular tu-
bercles of toes II-IV) or may have the
entire plantar surface tuberculate. In a
few frogs, the plantar tubercles are more
prominent (E. sulcatus, Hylactophryne,
and Ischnocnema). The subarticular tu-
bercles of the toes are like those of the
fingers except that the inner two toes
each have one tubercle, the third and
fifth two each, and the fourth three.

The toes may or may not have lateral
fringes or lateral keels. Although most
species of the genus lack webbing,
some Eleutherodactylus (karlschmidti
and two unnamed species) have nearly
complete toe webbing and the free pha-
langes bear prominent lateral fringes to
the base of the discs. In these (and some
other species as well) prominent lateral
fringes occur on the outer edge of toe V
and the inner edge of toe I. The discs
and pads of the toes exhibit comparable
variation to that seen on the fingers ex-
cept that the variation in pad sizes
among the five toes is less pronounced.

Size and proportions.——South Ameri-
can Eleutherodactylus range in size from
the diminutive E. trachyblepharis (males
12.1-15.8, x = 13.8 mm SVL) to a series
of large, broad-headed species (adult
females of E. anomalus, E. biporcatus,
E. bufoniformis, and E. necerus exceed
95 mm SVL). On the Amazonian ver-
sant of the Andes the largest species
probably do not exceed 70 mm (E. con-
dor and E. cornutus).

Many otherwise similar species pairs
may be discriminated easily on the basis
of body size. Adult male Eleutherodac-
tylus are distinguished from juvenile
males on the basis of having vocal slits
and sacs (some species), non-spinous
nuptial pads on swollen thumbs (some
species), or enlarged testes. In some
species, the testes are black or reticu-
lated with brown in adults. Adult fe-
males are distinguished from juveniles
on the basis of enlarged convoluted ovi-
ducts and, if mature eggs are present,
large, yellow ovarian eggs. Sexual di-

8 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

morphism in body size is pronounced in
the genus. Thirty-two species found on
the Amazonian versant of the Andes in
southern Colombia and northern Ecua-
dor (south to the Rio Pastaza) exhibit
dimorphisms (female x/male x) ranging
from 1.25 (E. elassodiscus, E. trachyble-
pharis, E. trepidotus) to 1.77 (E. livid-
us). The 15 species found on the Ama-
zonian versant in southern Ecuador ex-
hibit dimorphisms ranging from 1.18
(E. proserpens) to at least 1.73 (E.
condor). Twenty-four species from the
Amazonian lowlands exhibit dimor-
phisms ranging from 1.20 (E. paululus)
to 1.97 (E. sulcatus). The degree of di-

morphism is not different among the
three areas (mean, + 2 standard errors,
and N for the three areas respectively
are 1.41 + 0.05, 32; 1.42 + 0.09, 15; and
142 + 0.07, 24). Size data for the 36
species found on the northern Amazo-
nian versant are given in Table 1, for the
16 species found on the southern Ama-
zonian versant in Table 2, and for the
24 species found in the Amazonian low-
lands in Table 3.

The only proportion suggested to be
taxonomically significant is head width
(Lynch, 1975a). With the exception of
E. cornutus (a broad-headed species),
all of the Amazonian versant frogs have

TABLE 1.—Body sizes of Eleutherodactylus found on the northern Amazonian Andean versant in Ecua-
dor. Values include range (mean + 2 standard errors) sample size.

Species Males

E. acerus 25.0-34.0( 30.2 )3

E. chloronotus 22.9-30.3( 26.4 + 0.9)25
E. cornutus 35.5-43.0( 39.2 )2

E. cremnobates 28.4-32.5(30.6 + 1.3)7
E. devillei 19.6-31.6( 26.8 + 1.2)26
E. dolops 35.8-40.3(38.0)2

E. elassodiscus 22.0-29.2(26.9 + 1.1)14
E. eriphus 18.1-25.2(21.1 + 1.6)9
E. galdi 17.1-24.8(21.0 + 1.7)8
E. gladiator 14.9-15.8(15.2)3

E. glandulosus 24.2-30.2( 26.6 + 0.6)27
E. ignicolor 18.0

E. incanus 14.9-19.4(17.5 = 1.1)9
E. incomptus 15.6-18.8(17.5 + 0.7)10
ITER 1V UES UE CLUES a

E. lacrimosus' 19.6-23.7(21.0 + 0.8) 11
E. lanthanites' 23.8-27.9(26.3 + 1.6)5
E. leoni 13.0-18.3(16.5 + 1.0)11
E. leucopus 30.0-37.8(33.3 + 1.0)18
E. lividus 18.2-20.4(19.3)2

E. nigrogriseus 19.3-26.0( 22.7 + 0.9)20
E. nigrovittatus* 17.2-24.6(21.4 + 0.6) 33
E. pastazensis 19.5

E. peruvianus* 29 .2-35.8(31.8 + 1.4)10
E. petersi* 14.5-19.9(17.2 + 0.5)35
E. prolatus 13.7-18.4(16.8 + 0.5)20
E. pugnax 22.1-28.0(25.0)2

E. quaquaversus 19.6-22.5( 20.7 + 0.4)29
E. rubicundus 31.4-35.6(32.9 + 1.5)5
E. trachyblepharis 12.1-15.8(13.8 + 0.4)20
E. trepidotus 14.4-21.3(16.9 + 2.5)5
E. unistrigatus 14.9-23.4(18.8 + 0.8)25
E. ventrimormoratus SAIS (PALS, se ILS} NS
E. w-nigrum 29.3-46.8(34.3 + 1.2)32

Females Dimorphism
45.1 1.49
31.9-38.9(35.9 + 1.1)14 1.36
S540 faa
4] .6-51.7(46.2 + 2.6)8 1.51
34.7-44.7( 39.0 + 0.9)26 1.46
56.6-57.6(57.1)3 1.50
29.7-36.9( 33.6 + 1.1)16 1.25
25.8-29.0(27.4)2 1.30
28.1-34.0(31.7 + 1.6)9 1.51
S160 (0 (0 EE
31.8-41.3(36.1 + 0.9)20 1.36
26.4 1.47
23.9-30.6(29.2 + 1.2)5 1.67
23.7-25.9(24.5 + 0.6)7 1.40
DANO =D ARA(D459)) 2
24.4-32.5(27.0 + 3.0)5 1.28
38.7-45.4( 41.2) 4 1.57
19.7-25.0(21.9 + 0.6)26 1.33
42.3-44.0(42.9)3 1.29
32.9-35.0(34.1)3 Lea
28.5-30.2(29.2)4 1.29
25.0-30.5 (27.4 + 1.7)21 1.28
30.9-31.7(31.3)2 1.60
38.6-46.4( 42.3 + 1.4)10 1.33
20.3-23.1(21.6 + 0.7)8 1.26
20.8-24.1(22.3 + 0.8)9 1.33
30.8-33.2 (32.0 )2 1.28
24.6-31.3( 27.3 + 0.8)20 1.32
46.5-51.2( 48.8) 2 1.48
15.8-19.2(17.2 + 0.4)19 1.25
NASP ( OU ae I. He 1.25
23.3-30.8( 26.3 + 0.4)46 1.40
33.3-43.8(36.9 + 1.9)9 1.69
44.4-56.6(49.9 + 1.9)16 1.45

1 Amazonian slope populations consist of larger frogs than those from the Amazon Basin (see Table 3).
2 Not including specimens from Bafios and vicinity; see Table 4

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 9

TABLE 2.—Body sizes of Eleutherodactylus found only on the southern Andean slopes of the Ama-

zonian versant. Values:

Species Males
E. atratus 17.4-24.0(21.7 + 1.7)19
E. baryecuus 27.2-30.4(29.0)5
E. bromeliaceus 16.7-22.8( 20.8 + 0.9)15
E. colodactylus 14.0-20.7(17.1 + 0.5)46
E. condor 32.1-39.5(36.5 + 1.7)9
E. cryptomelas 28.2-30.3(29.2)4
E. proserpens 15.2-21.0(18.6 + 1.0) 11
E. pycnodermis 18.0-32.3(25.1 + 1 ee
E. spinosus 16.1-25.0( 20.1 + 0.6)34
E. w-nigrum' 28.4-45.1(36.7 + 1.0)55

range (xX + 2SE) N.

Females Dimorphism
24.9-29.2(27.4 + 1.3)10 1.26
38.2-43.5(40.8 + 1.3)8 1.41
23.0-28.5( 26.6 + 1.6)6 1.28
16.5-25.8( 20.8 + 0.8)33 122,
63.2 IL)
38.6 1.32
20.2-23.5(22.0)4 1.18
32.5-44.4( 38.0 + 1.1)40 ite
28.3-34.5( 31.8 + 0.6)29 1.58
53.0-65.4(59.2 + 2.6)9 1.61

1 Specimens from the northern Amazonian slopes are significantly different in size (Table 1).

TABLE 3.—Body sizes of Eleutherodactylus' found in the Amazon Basin in Brasil, Colombia, Ecuador,
and Pert. Values: range (x + 2SE) N.

Species Males
E. acuminatus 17.1-22.6(19.4 + 0.8)14
E. altamazonicus 14.4-23.1(18.8 + 0.8)28
E. carvalhoi 13.5-14.8(14.2)2
E. conspicillatus 23.2-30.1(27.3 + 0.6)29
E. croceoinguinis 12.8-18.2(14.6 + 0.7)14
E. diadematus 21.4-27.4(23.5 + 0.8)15
E. fenestratus 23.8-34.3(28.3 + 0.9)29
E. lacrimosus* 16.1-20.0(18.5 + 0.5)18
E. lanthanites* 21.7-26.0( 23.5 + 0.6)20
E. malkini 30.4-37.2(32.9 + 1.2)11
E. martiae 11.8-16.8(15.1 + 0.4)29
E. nigrovittatus* 13.3-19.4(17.0 + 1.9)8
E. ockendeni 17.9-21.3(19.4 + 0.5)18
E. orphnolaimus 24.0
E. paululus 13.6-17.1(15.0 + 0.8)9
E. peruvianus* * 27.3-31.6(30.0 + 0.6)16
E. platydactylus 16.3-23.9(19.3 + 2.1)8
E. pseudoacuminatus 12.7-17.6(15.4 + 0.7)15
E. sulcatus 19.7-36.2 (26.0 + 4.8)8
E. variabilis 13.7-17.5(16.3 + 0.4)26
E. vilarsi 22.7-31.5(27.2 + 1.5)10
E. zeuctotylus 20.4-29.6(25.5 + 1.1)20

Females Dimorphism
25.6-31.3(28.0 + 1.0)14 1.44
23.6-33.9(27.9 + 0.8) 42 1.48
16.3-24.0(20.0)6 1.41
34.8-48.8( 40.2 + 1.7)19 1.47
18.0-21.9(19.8 + 0.6)19 1.36
35.4-44.5(39.8 + 1.2)19 1.69
34.0-50.0( 41.4 + 1.5)32 1.46
20.6-24.4(22.5 + 0.5)18 122,
QieD-AD.2 (36:2) -=alea)oll 1.54
41.5-47.9( 44.8 + 1.3)9 1.36
18.3-22.2(20.4 + 0.4)36 1.35
19.7-22.1(20.9 + 0.5)9 1.23
24.6-31.1(27.2 + 0.5)37 1.40
24.8-33.4(29.1)2 LPL
16.5-19.4(18.0)4 1.20
38.9-44.2(41.3 + 1.5)7 1.39
2816-330 (30:0 2187.) Revi
18.1-22.4(19.9 + 0.5)21 1.29
42,.4-59.7(51.2 + 2.6)15 1.97
20.0-25.3( 22.8 + 0.5)30 1.69
33.0-43.9( 39.2 + 1.0)33 1.44
30.5-43.3(37.0 + 1.2) 32 1.45

1 See Table 1 for data for E. quaquaversus, E. trachyblepharis, and E. ventrimarmoratus.
2 Amazon lowland populations consist of significantly smaller frogs than do slope populations (see Table 1).
8 Sample from Cusuime, Rio Cusuime, Provincia Morona-Santiago, Ecuador, 320m; other Amazon Basin populations

are still smaller in body size (see Lynch, 1980).

normal head widths (mean HW/SVL
0.320-0.425). Eleutherodactylus cornu-
tus has a mean HW/SVL of 0.520.

KEY TO THE SPECIES
1. First finger longer than second; skin
of venter smooth (finely tuberculate
Maiti ICUS) = aes DIES 2
First finger shorter than second; skin
of venter coarsely areolate 10

2. Pads absent on fingers (finger tip not

appreciably wider than digit below
disc) eae US, Ne BA eU ee Ihe 3

Pads present on fingers II-IV 5

. Pads of toes pointed; dorsolateral

folds present, incomplete ____-
MUL Meee ae E. nigrovittatus

Pads of toes rounded; dorsolateral
foldstabsemt 2.0 it) | eis 4

. Metatarsal tubercles approximately

equal in size; adults moderate sized
(males 22.0-30.0 mm, females 30.0-

10

10.

Ie

12.

13.

. Venter

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

37.0emmns Vib) as E. elassodiscus

Inner metatarsal tubercle 4-8 times
size of outer; adults large (males
35.0-40.0 mm, females more than
Siou(0) seman. SN VIG) ee E. dolops

. Upper eyelid bearing one or more

elongate tubercles E. cornutus

Upper eyelid lacking conical or elon-
gate tubercles — =.) 322222 6

. Heel bearing a conical or elongate

tubercle
Heel lacking enlarged tubercles __ 8

. Toes bearing narrow lateral fringes;

gular stripe absent __ E. rubicundus

Toes lacking lateral fringes; white
gular stripe present __ E. lanthanites

. Large black spots on anterior and

posterior surfaces of thighs and on
AL ANKS eee ee E. w-nigrum

Not so patterned on limbs and flanks

moderately to heavily
blotched with brown: upper eyelid
narrower than to as broad as IOD;
dorsolateral folds indistinct; adult
females more than 60 mm SVL ___..
R655 nw AVA ED tevin Dba E. condor
Venter cream with brown flecks; up-
per eyelid broader than IOD; dorso-
lateral folds present, interrupted;
adult females less than 50 mm SVL
ELAS c 2 een ae Cea a E. peruvianus
Heel bearing at least one conical or
elongate tubercle Hult
Heel lacking tubercles, or if present,
Hatteda (not conical) ====saaaaae 22
Upper eyelid bearing one or more
conical or elongate tubercle ____ 18
Upper eyelid lacking pungent tuber-
CLES ies soe to a ce 20
Forearm bearing conical ulnar tuber-
clesalongied c= =a aan 13
Forearm lacking ulnar tubercles (ex-
cept antelraclial))) esau mean 18
Skin of dorsum pustular, tuberculate,
or bearing scattered tubercles ___ 14
Skin of dorsum shagreened or finely
areolate (without enlarged tuber-

14.

15.

16.

7

18.

Ike),

20.

ai

22.

23.

24.

25.

cles )

Posterior surfaces of thighs unicolor
(cream or black)

Posterior surfaces of thighs bicolor
(black and white) ___________ 16

Posterior surfaces of thighs black _
A i doe retet op aed E. cryptomelas

Posterior surfaces of thighs cream __-

A) Ea ee nt ee E. galdi
Posterior surfaces of thighs barred
black and white ____ E. eriphus
Posterior surfaces of thighs black
with white spots E. spinosus

Tubercle on elbow and heel elongate
(caledr i eae E. inusitatus
Tubercles on elbow and heel conical
(not elongate) E. lividus
Dorsum bearing many low ridges;
groin) blacks = aes E. atratus
Dorsum shagreened or feebly warty,
lacking many low ridges; groin not

black. 19
H-shaped folds on occiput; tympan-
um not concealed E. prolatus
No folds on occiput; tympanum con-
Cealediis ste cm toa E. quaquaversus
Canthus rostralis sharp; tympanum
distinct 2.4... 21

Canthus rostralis rounded; tympan-
um partially concealed __ E. incanus
Posterior surface of thigh brown
with large white spots
Bes) her sco) Ei be 2A eee E. nigrogriseus
Posterior surface of thigh brown
with cream flecks ___ E. rubicundus
Tympanum and annulus absent or
concealed 0... ee 23
Tympanic annulus evident external-
ly and distinct (except dorsally) — 29
Tympanum and annulus absent __. 24
Tympanum and annulus concealed
beneath skin 2... | eee 26
Canthus rostralis relatively sharp _
Bia Aaah Seyi, et Met nb 2 E. baryecuus

Vomerine odontophores absent ___.
pears Sg AUR Aw E. colodactylus

26.

27.

28.

29.

30.

31.

32.

33.

34.

35.

36.

37.

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES ii

i cee ed Se E. pugnax
Canthus rostralis angular (moderate-
| yegk3| NEI j05)), aoe tat we Aeon ne ee 27
@anthus rostralis round _._._.___ 28

Upper eyelid lacking tubercles ___...
a E. trachyblepharis
Upper eyelid bearing one or more
conical tubercles _____._____.- E. leoni
Venter white with black marbling
2 E. ventrimarmoratus
Venter cream... E. incomptus
Canthus rostralis round 30
Canthus rostralis angular or sharp _
consis) Ba, tS a rel 36
Venter black with white spots; pads
scarcely developed ___- E. trepidotus
Venter not spotted; pads small to
| eye a SS Sas Oe eae TNE 31
Skin of dorsum areolate
on E. proserpens
Skin of dorsum smooth to finely tu-
| SST SULA WS te lane eee 32
Upper eyelid bearing one conical
lberele: 25 E. bromeliaceus
Upper eyelid lacking tubercles, or if
present, tubercles flat 33
Posterior surfaces of thighs dark
C10} E. cremnobates
Posterior surfaces of thighs pale,
marbled or not with brown 34
Skin of dorsum very finely sha-
ereemea 2 E. lacrimosus
Skin of dorsum smooth bearing flat
Wwartsmoosteriorly 0 2 35
Tip of snout bearing papilla
vn a 8. Ft E. pastazensis
Tip of snout lacking papilla
Benen tre 8 1 bt: E. unistrigatus

Upper eyelid bearing one conical
tuberclel Stews se! E. petersi
Upper eyelid lacking pungent tuber-
GS, |) aE SA Seeds tee 37

Posterior surfaces of thighs uniform
dark brown or bearing a large dark
brownMblotehr 2.7 ee 38

Posterior surfaces of thighs brown

with large pale spots or barred with
cream) or ‘colorless’ 222 © 40

38. Large black blotch in groin ____.-
Ree 0k hes ee ee E. pycnodermis

Groin not so marked —-_._ 39
39. Skin of dorsum smooth ___ E. acerus
Skin of dorsum finely tuberculate

with scattered warts __ E. leucopus
40. Skin of dorsum bearing distinct folds
OTe ICL @ syerces t= ees ee eA 4]
Skin of dorsum bearing warts but
MORI ges = seer tienen es ee 43
41. Two folds extending from occiput to
Above eromar meets aie ieee .oe 42
Folds much shorter, more numerous
A SRR are ei ae E. gladiator

42. Paravertebral folds sinuous; posterior
surfaces of thighs barred _...--
cscs ne Mitta Cau clie a E. chloronotus
Dorsolateral folds not sinuous; pos-
terior surfaces of thighs yellow-
brown with brown reticulation

ia, Sek scenes nears Peyigee 8) lt E. devillei
43. Snout round in dorsal view; canthus
rostralis concave ___ E. glandulosus

Snout acuminate in dorsal view; can-
thus rostralis straight __ E. ignicolor

ACCOUNTS OF THE SPECIES

In the following accounts we attempt
to make comparable statements in the
diagnoses and descriptions. The diag-
noses consist of a series of 14 numbered
statements (or set of statements) fol-
lowing the format used by Lynch
(1979a). Following the numbered state-
ments is a paragraph distinguishing the
taxon from those it most nearly resem-
bles (in our opinion). Measurements
and proportions are given for each sex
when the means differ significantly (p =
0.05); otherwise these data are com-
bined.

Complete accounts are provided for
each of the twelve new species as well
as for E. pastazensis, E. peruvianus, E.
rubicundus, and E. trachyblepharis. Ab-
breviated accounts (lacking descrip-
tions) are provided for 19 species, and

12 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

very abbreviated accounts (only synony-
mies and brief remarks) are provided for
eight species (E. atratus, E. baryecuus,
E. bromeliaceus, E. colodactylus, E.
cryptomelas, E. proserpens, E. pycno-
dermis, and E. spinosus) named by
Lynch (1979a) and for E. ockendeni
(recently treated by Lynch, 1974,
1980).

Eleutherodactylus acerus new species
Fig. 2A

Holotype.—KU 165471, adult female,
taken 11 km ESE Papallacta, Provincia
Napo, Ecuador, 2660 m, 22 March 1975
by William E. Duellman and Linda
Trueb.

Paratypes—KU 165472-73 collected
at type-locality on 22 March and 27
April 1975 respectively; KU 143517, 7.8
km WNW Cuyujia, Provincia Napo, Ec-
uador, 2750 m, 27 October 1971 by
Bruce MacBryde.

Diagnosis.—1) skin of upper surfaces
smooth, that of venter coarsely areolate;
2) tympanum distinct, its length 4-%
eye length; 3) snout subacuminate in
dorsal view, pointed or protruding in
lateral profile; canthus rostralis relatively
sharp; 4) upper eyelid nearly as broad
as IOD, lacking pungent tubercles; low
cranial crests present; 5) vomerine odon-
tophores prominent, round in outline;
6) males with vocal slits and subgular
vocal sac, non-spinous nuptial pad on
thumbs; 7) first finger shorter than sec-
ond; fingers bearing large pads (largest
on II-IV); discs broader than long; 8)
fingers bearing ill-defined lateral fringes;
9) indefinite ridge of ulnar tubercles;
10) heel and tarsus lacking pungent tu-
bercles; 11) two metatarsal tubercles,
inner oval, 3-4 times size of outer; nu-
merous supernumerary plantar tubercles;
12) toes bearing indistinct lateral keels,
no webbing; toe pads smaller than those
of outer fingers; 13) nearly black above,
belly and undersides of limbs black,
throat yellow-brown; posterior surface
of thighs dark brown; 14) adults mod-

erate-sized, three males 25.0-34.0 mm,
one female 45.1 mm SVL.
Eleutherodactylus acerus is most sim-
ilar to E. glandulosus (Boulenger), E.
leucopus Lynch, and E. lividus new spe-
cies. Eleutherodactylus glandulosus has
a rounded snout in lateral profile, and
yellow spots on the flank, groin, anterior
and posterior surfaces of the thighs, and
concealed shank. Eleutherodactylus leu-
copus is probably the nearest relative of
E. acerus; it differs in having small inner
tarsal tubercles, faintly granular (not
smooth) skin on the dorsal surfaces, and
in having cream or white inner digits
of the hands and feet. Both E. acerus
and E. leucopus have protruding snouts;
both are larger than E. glandulosus.
Eleutherodactylus lividus (described be-

-low) differs from E. acerus in having a

conical tubercle on the upper eyelid and
similar tubercles along the outer edge of
the tarsus and heel.

Description—Head as wide as body,
wider than long; head width 38.8-41.9
(x = 39.8) percent SVL; snout sub-
acuminate in dorsal view, pointed or
protruding in lateral profile; snout short,
E-N 82.9-90.0 (x = 86.6) percent eye
length; nostrils not protuberant, directed
anterolaterally; canthus rostralis relative-
ly sharp, weakly concave; loreal region
weakly concave, sloping abruptly to lips;
lips not flared; upper eyelid lacking tu-
bercles (except in KU 143517, small
conical tubercle on each eyelid), its
width 81.2-100.0 (x = 91.9) percent
IOD; low cranial crests present along
lateral edges of frontoparietals; tym-
panum directed dorsolaterally, promi-
nent, round, its upper edge concealed
by heavy supratympanic fold, separated
from eye by distance 1.5 times tym-
panum length; tympanum length 36.4—
42.6 (x — 39.5) percent eye length; post-
rictal tubercles small, round, subconi-
cal; choanae small, round, not concealed
by palatal shelf of maxillary arch; vo-
merine odontophores median and _ pos-
terior to choanae, separated by distance
equal to a choanal width, round, ele-
vated, each bearing clump of 4-5 teeth;

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 13

Fic. 2——Andean Eleutherodactylus. A. E. acerus, KU 165471, adult female, 45.1 mm SVL. B.
E. chloronotus, KU 143317, juvenile, 21.8 mm SVL. C. E. condor, KU 146992, immature female,
55.5 mm SVL. D. E. cornutus, KU 168009, adult male, 55.8 mm SVL. E. E. cremnobates, KU
143459, adult female, 41.6 mm SVL. F. E. devillei, KU 143401, adult female, 36.0 mm SVL. G.
E. dolops, KU 143505, adult female, 57.6 mm SVL. H. E. elassodiscus, KU 168012, adult male,
23.1 mm SVL. C rendered by Linda Trueb; others from color transparencies by WED.

14 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Fic. 3.—Andean Eleutherodactylus. A. E. eriphus, KU 166031, adult female, 29.0 mm SVL. B. E.
gladiator, KU 143515, adult male, 14.9 mm SVL. C. E. glandulosus, KU 165435, adult male, 30.3
mm SVL. D. E. ignicolor, KU 165879, juvenile female, 21.1 mm SVL. E. E. incanus, KU 143458,
adult female, 30.6 mm SVL. F. E. incomptus, KU 143492, adult female, 25.0 mm SVL. G. E.
inusitatus, KU 166066, adult female, 24.0 mm SVL. H. E. lanthanites, KU 168834, juvenile, 23.6
mm SVL. From color transparencies by WED.

15

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES

Fic. 4——Andean Eleutherodactylus. A. E. leoni, KU 165897, adult female, 24.4 mm SVL. B. E.

lividus, KU 166005, adult female, 35.0 mm SVL. C. E. nigrogriseus, KU 146109, adult female, 29.4
mm SVL. D. E. nigrovittatus, KU 123503. E. E. peruvianus, KU 123446, adult female, 49.0 mm
SVL. F. E. petersi, KU 165993, juvenile female, 20.7 mm SVL. G. E. prolatus, KU 166008, adult
female, 22.3 mm SVL. H. E. pugnax, KU 165561, immature female, 26.5 mm SVL. From color

transparencies by WED.

16 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Fic. 5.—Andean Eleutherodactylus. A. E. rubicundus, KU 146170, adult male, 33.5 mm SVL. B.
E. trachyblepharis, KU 120180, adult female, 16.4 mm SVL. C. E. trepidotus, KU 143435, juvenile
female, 15.8 mm SVL. D. E. w-nigrum, KU 143301, adult female, 50.3 mm SVL. From color

transparencies by WED.

tongue longer than wide, posterior bor-
der feebly notched, posterior % not
adherent to floor of mouth; males with
long vocal slits (from tongue to corner
of jaw) and large subgular vocal sac.
Skin of dorsum, flanks, upper surfaces
of limbs smooth, no dorsolateral folds,
that of venter and undersides of thighs
areolate; discoidal folds prominent, end-
ing well anteriad to groin; anal opening
not extended, no para-anal warts; in-
definite ridge of low ulnar tubercles;
palmar tubercle bifid, nearly twice size
of oval thenar tubercle; numerous super-
numerary palmar tubercles, all non-
pungent; subarticular tubercles round,
non-conical, pungent; fingers bearing ill-
defined lateral fringes, all with discs
(broader than long) on dilated pads;
pad of thumb as large as tympanum,
those of fingers II-IV 1.5 times tym-
panum; thumb shorter than second fin-
ger; male with white, non-spinous nup-
tial pad on median surface of thumb.
Heel and tarsus lacking obvious tu-

bercles (inner tarsal tubercle evident in
adult female KU 165471); inner meta-
tarsal tubercle oval, length twice width;
outer metatarsal tubercle subconical,
length slightly greater than width, 4
% size of inner; numerous supernumer-
ary plantar tubercles, none prominent;
subarticular tubercles round, subconical;
lateral fringes on toes less obvious than
those on fingers; toe discs broader than
long on dilated pads, pads smaller than
those on outer fingers, rounded apically;
shank 45.0-52.7 (x = 50.4) percent SVL;
heels of flexed hind legs overlap; heel of
adpressed hind leg reaches eye.

In preservative nearly black above;
belly and undersides of limbs black or
dark brown; throat yellow-brown to
gray-brown; undersides of digits pale.
KU 143517 is brown and the pale areas
are pale golden-brown; the dorsum has
a cream dorsoconcolor pattern (Lynch,
1966) with a brown patch extending
from scapular to sacral regions.

Color in life-—Dorsum black or dark

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 17

gray; venter dark gray or dark grayish-
brown; throat of male creamy brown;
iris dark reddish-brown.

Measurements of holotype in mm.—
SVL 45.1; shank 23.4; head width 18.9;
head length 15.0; upper eyelid width
4.8; IOD 5.1; tympanum length 2.3; eye
length 5.4; E-N 4.6.

Etymology.—Greek, in reference to
the absence of tubercles on the eyelid,
heel, and tarsus.

Distribution—Known only from the
type-locality and vicinity between Pa-
pallacta and Cuyujuta in upper montane
cloud forest, 2660-2750 m.

Natural History—One _ individual
(KU 143417) was found by day under
a log in a pasture (27 October 1971).
The other three individuals were found
at night on bushes (one at base, one 2 m
above ground) on 22 March and 27
April 1975. The holotype (KU 165471)
is a gravid female.

Remarks.—Eleutherodactylus acerus,
E. glandulosus, E. leucopus, and E. li-
vidus are a small constellation of species
found in upper montane cloud forest in
the valleys of the rios Chingual and
Papallacta in northwestern Provincia
Napo, Ecuador. Lynch (1976c) sug-
gested that E. glandulosus and E. leu-
copus were geminate species but it now
appears more likely that E. acerus and
E. leucopus enjoy that relationship and
that E. glandulosus and E. lividus are
less closely related.

Eleutherodactylus atratus Lynch

Eleutherodactylus atratus Lynch, 1979, Misc.
Publ. Mus. Nat. Hist. Univ. Kansas 66:5
[Holotype—USNM 199675, Suro Rancho,
Provincia Morona-Santiago, Ecuador,
2683 m].

This species is found in subparamo
and cloud forests at elevations of 2195
to 2850 m on the Amazonian slopes of
the Andes in southern Ecuador (Cordil-
lera de Matanga to the Abra de Zamora )
(Lynch, 1979a).

Eleutherodactylus baryecuus Lynch
Eleutherodactylus baryecuus Lynch, 1979, Misc.

Publ. Mus. Nat. Hist. Univ. Kansas 66:10
[Holotype—USNM 199714, Suro Rancho,
Provincia Morona-Santiago, Ecuador,
2683 m].

This species is found in subparamo
and cloud forests at elevations of 2195
to 2988 m on the Amazonian slopes of
the Andes in southern Ecuador (Lynch,
1979a).

Eleutherodactylus bromeliaceus Lynch

Eleutherodactylus bromeliaceus Lynch, 1979,
Misc. Publ. Mus. Nat. Hist. Univ. Kansas
66:12 [Holotype —USNM 199731, from be-
tween Sapote and Suro Rancho, Provincia
Morona-Santiago, Ecuador, 2622 ml].

Lynch (1979a) reported specimens
from cloud forests at elevations of 1707
to 2622 m on the Cordillera de Matanga
in southern Ecuador. We have found
additional specimens from the Cordillera
del Condor and from the Cordillera de

Zamora.

Eleutherodactylus chloronotus Lynch
Fig. 2B
Eleutherodactylus chloronotus Lynch, 1970,

Journ. Herpetol. 3:140 [Holotype —KU

117519, 3 km E Papallacta, Provincia Napo,

Ecuador, 2900 m].

Diagnosis.—1) skin of dorsum smooth
to feebly pustulate, pair of sinuous para-
vertebral folds present, no dorsolateral
folds, that of venter coarsely areolate;
2) tympanum prominent, its length 4-
% that of eye; 3) snout subacuminate
in dorsal view, angularly rounded or
truncate in lateral profile; canthus ros-
tralis sharp, strongly concave; 4) upper
eyelid slightly narrower (to equal) than
IOD, bearing small conical warts; no
cranial crests; 5) vomerine odontophones
small, oblique to subtriangular in out-
line; 6) males with vocal slits and sub-
gular vocal sac; 7) first finger shorter
than second; fingers II-IV bearing di-
lated pads, discs broader than long; 8)
fingers lacking lateral fringes; 9) fore-
arm lacking ulnar tubercles; 10) heel
and outer edge of tarsus bearing row of
small subconical tubercles; 11) two met-
atarsal tubercles, inner oval, 5-6 times

18 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

size of round outer; 0-4 supernumerary
plantar tubercles; 12) toes lack lateral
fringes and webbing; toe pads as large
as those of outer fingers; 13) dorsum
gray to dull orange with dark brown to
black-edged hour-glass mark on back
enclosing pale gray area; flanks gray
with darker slanted bars; venter cream
spotted or reticulated with brown; limbs
yellow-brown with darker bars; poste-
rior thigh strongly barred; 14) adults
moderate sized, males 22.9-30.3 (x =
26.4, N = 25) mm, females 31.9-38.9
(x = 35.9, N = 14) mm SVL.

In life, E. chloronotus is readily rec-
ognized because the area between the
paravertebral stripes is bright (young)
to dull (old) green. Most comparable-
sized Eleutherodactylus in upper mon-
tane cloud forests have cranial crests and
lateral fringes on the digits; none has so
strongly concave canthus rostralis as E.
chloronotus.

Description—see Lynch (1970b).

Color in life —Middorsal area bright
metallic green or olive-green to orange-
brown or bronze with dark brown to
black dorsolateral ridges. Laterally
green, tan, gray, or cream usually with
brown diagonal marks. Brown or dark
green bars usually evident on hind limbs.
Venter creamy yellow to yellowish tan
with (usually) or without brown flecks
or marbling, or gray with or without
cream to bluish-white flecks. Iris bronze,
with a coppery tint in some individuals.

Distribution and ecology.—Eleu-
therodactylus chloronotus occurs in sub-
paramo and upper humid montane for-
est (2285-3350 m) along the eastern
front of the Andes from southern Colom-
bia (Narifo and Putumayo) southward
to the Pastaza depression. By day the
frogs are found beneath stones and logs,
whereas at night they are active on the
ground and perch on herbs and low
bushes. Several were observed moving
about by day after rain.

Reproduction may be seasonal. Lynch
(1970b) reported egg clutches found in
March at the type-locality. A newly
hatched frog was found at the type-

locality in mid-July 1977 but no egg
masses were found.

Remarks.—The body size of adult
females reported by Lynch (1970b) is
in error; he included juvenile and young
females in his sample. Females having
large eggs or convoluted oviducts are at
least 31 mm in SVL.

Aside from being a member of the
unistrigatus group (now containing more
than 100 species), the relationships of
E. chloronotus remain obscure. Eleu-
therodactylus chloronotus appears to be
related to E. incanus found at lower ele-
vations and, to a lesser degree, to E.
calcaratus and E. crucifer (Pacific ver-
sant). Eleutherodactylus cryptomelas
from the Amazonian slopes of southern
Ecuador likewise appears to be related
to E. chloronotus. Lynch’s (1970b) sug-
gestion of a close relationship between
E. chloronotus, E. devillei, and E. surdus
is untenable.

Eleutherodactylus colodactylus Lynch

Eleutherodactylus colodactylus Lynch, 1979,
Misc. Publ. Mus. Nat. Hist. Univ. Kansas
66:15 [Holotype—KU 142151, Abra de
Zamora, 13.5 km E Loja, Provincia Loja,
Ecuador, 2800 m].

Eleutherodactylus colodactylus oc-
curs in cloud forest and subparamo at
elevations of 2200 to 2850 m on the
Amazonian slopes of the Andes in south-
ern Ecuador and on the crest and Pacific
versant of the Cordillera between Can-
chaque and Huancabamba, Departa-
mento de Piura, Pert, at elevations of
2745 to 3100 m (Lynch, 1979a).

Eleutherodactylus condor new species
Fig. 2C

Holotype-—KU 146992, juvenile fe-
male, taken at Rio Piuntza, Cordillera
del Condor, Provincia Morona-Santiago,
Ecuador, 1830 m, on 4 January 1972 by
Bruce MacBryde and John E. Simmons.

Paratypes. — KU 146991, 146993-
147033, collected at type-locality 3-6 Jan.
O72:

Diagnosis—1) skin of dorsum sha-

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 19

greened with indistinct dorsolateral
folds, that of venter smooth; 2) tympa-
num distinct, its length % eye length;
3) snout subacuminate in dorsal view,
rounded in lateral profile, canthus ros-
tralis sharp; 4) upper eyelid narrower
than (to equal) IOD, not bearing pun-
gent warts; low cranial crests palpable
in large females; 5) vomerine odonto-
phores prominent, triangular in outline;
6) males with vocal slits, subgular vocal
sac, non-spinuous nuptial pads on
thumbs; 7) first finger longer than sec-
ond; pads of outer fingers large, those of
inner fingers narrow; discs broader than
long; 8) fingers lacking lateral fringes or
keels; 9) no ulnar tubercles except small
antebrachial tubercle; 10) no tubercles
on heel or tarsus; 11) two metatarsal
tubercles, inner elongate, 6-8 times size
of outer; 0-2 supernumerary plantar tu-
bercles; 12) toes bearing narrow lateral
fringe (keeled), not webbed; toe pads
slightly smaller than those of fingers
IIJ-IV; 13) brown with darker markings
—chevrons, interorbital bars, labial bars;
posterior thigh brown with large cream
or white spots (spots smaller than pad
of any toe); venter moderately to heavily
blotched with brown, darkest on throat
and undersides of limbs; venter of thigh
and shank cream with brown spots or
brown with cream spots; 14) adults
large, nine males 32.1-39.5 mm, five ju-
venile to young females 52.6-59.4 mm
SVL. A decapitated adult female (MCZ
- 19630) is estimated to have been 63.2
mm SVL.

Eleutherodactylus condor is one of
the largest members of the fitzingeri
group lacking toe webbing or heavy toe
fringing. Adults are probably compar-
able in size to E. lymani Barbour and
Noble and E. w-nigrum (Boettger).
Eleutherodactylus lymani has an inner
tarsal fold; E. w-nigrum has smaller tym-
pana, dark spots on the flank, and yellow
posterior thighs with brown or black
spots or reticulation. Except for some
populations of E. w-nigrum, neither spe-
cies has spotted throat and venter as
does E. condor. Eleutherodactylus con-

dor is superficially similar (in preserva-
tive) to the Amazonian slope popula-
tions of the smaller E. peruvianus (adult
females 38.6-46.4 mm SVL) but has less
intense brown markings on the throat
and chest and a narrower head.

Description—Head as wide as or
wider than body, wider than long; head
width 37.0-40.2 (x = 38.4, N = 14) per-
cent SVL; nostrils weakly protuberant,
directed laterally; snout long, E-N 85.7-
109.5 (x = 100.0, N = 9) percent eye
length in males, 102.8-127.6 (x = 114.5,
N = 5) in females; loreal region weakly
concave, sloping abruptly to lips; lips
not flared; cranial crests not present but
edges of frontoparietals slightly elevated
in large females (albeit immature); up-
per eyelid lacking tubercles, its width
64.3-105.1 (x = 90.7, N = 18) percent
IOD; tympanum directed laterally with
slight dorsal and posterior vectors, prom-
inent, higher than long, slanted anteriad,
posterodorsal edge obscured by well-
defined supratympanic fold; tympanum
length 43.2-60.0 (X = 51.2, N = 14) per-
cent eye length; postrictal tubercles
small, subconical; choanae longer than
wide, relatively small, not concealed by
palatal shelf of maxillary arch; vomerine
odontophores median and posterior to
choanae, triangular in outline, elevated,
separated by distance equal % choanal
width, each 1.5 times size of a choana,
bearing 7-9 teeth in approximately trans-
verse row.

Skin of dorsal surfaces shagreened;
ill-defined dorsolateral folds extend from
posterolateral corner of eyelid to above
groin; discoidal folds evident, areolation
below and posterodorsal to anus poorly
defined; anal opening not extended; pal-
mar tubercle bifid, larger than oval
thenar tubercle; supernumerary palmar
tubercles not pungent; subarticular tu-
bercles round, subconial; fingers lacking
lateral fringes or keels; pads of fingers
III-IV large, those of I-II weakly dilated;
discs of former broader than long, round
apically; thumb longer than second fin-
ger.

Heel and tarsus lacking tubercles or

20 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

folds; inner metatarsal tubercle weakly
compressed, its length 2.5 times width;
outer metatarsal tubercle round to elon-
gate, scarcely evident, % or less size
of inner; 0-2 supernumerary plantar tu-
bercles—if present, at base of toes III-
IV; subarticular tubercles longer than
wide, subconical to weakly elevated;
toes bearing lateral keels, webbing in-
sufficient to reach bases of basal sub-
articular tubercle (except toe V); pads
slightly smaller than those of outer fin-
gers; legs long, shank 54.9-64.2 (x =
59.5, N = 9) percent SVL in males,
63.5-66.0 (x = 64.1, N = 5) in females;
heels of flexed legs broadly overlapping;
heel of adpressed hind limb reaching
beyond tip of snout.

In preservative, brown above with
darker brown markings (chevrons, in-
terorbital bar, limb bars, oblique flank
bars, labial bars) edged with cream
halos; canthal and supratympanic stripes
prominent; posterior thigh brown with
white spots (smaller than any toe pad);
groin brown with few pale spots; throat
gray with darker blotching, especially
toward jaw and on breast; belly cream
spotted with ill-defined brown spots;
underside of thighs cream with brown
spots, of shank brown with cream spots.

Color in life—Dorsum usually tan
(reddish brown, olive-green, or grayish
green in some individuals) with darker
tan or brown markings on body. Seven
individuals (KU 146993, 147006-10,
147024) have pale cream, yellow, or
orange dorsolateral stripes. Posterior
surfaces of thighs gray or brown, with
yellow flecks in four individuals (KU
147002-4, 147023). Venter mottled
orange and yellow or orange or yellow
and gray; in juveniles (KU 146991,
147006-10, 147033) venter gray with
white flecks or mottling. Undersides of
hind limbs black in KU 147019. Canthal
and supratympanic stripes dark brown

to black.
Measurements of holotype in mm.—
SVL 55.5; shank 36.0; head width 20.7;

lead length 20.7; upper eyelid width 4.8;
IOD 6.0; tympanum length 3.3; eye
length 5.8; E-N 7.4. The oviducts of the
holotype are thin, non-convoluted tubes.

Etymology.—A noun in apposition in
allusion to the Cordillera del Condor
and the large size of species.

Distribution and ecology.—This spe-
cies is definitely known only from the
type locality in cloud forest near the
headwaters of the Rio Piuntza on the
western slopes of the Cordillera del Con-
dor, an isolated range separated from
the main Andean Cordillera by the val-
ley of the Rio Zamora. Eight adults
were perched on low vegetation at night;
juveniles and the rest of the adults were
on the forest floor by day. Three speci-
mens (MCZ 19629-30, 19632) bear im-
precise locality data: “Pastaza River,
Canelos to Marafion.” If these frogs ac-
tually were collected between Canelos
and the Rio Maranon, the species occurs
in the lowlands of eastern Ecuador.

Remarks.—Sexual dimorphism in
body size in most species of Eleuthero-
dactylus is on the order of females being
140-150 percent the size of males
(Crump, 1974; Lynch, 1976b, 1979a). If
this relationship is true for E. condor
(x size of males 36.5), females should
be 50-55 mm (mean). The largest fe-
males available are 57.3, 59.2, and 59.4
mm SVL and have slight convolutions
of the oviduct and small eggs; presum-
ably adult females are between 60 and
70 mm SVL (MCZ 19630 is estimated to
have been 63.2 mm and is an adult
female).

Eleutherodactylus condor is one of
two species of the fitzingeri group found
at moderate to intermediate elevations
on the Amazonian slopes in Ecuador.
Eleutherodactylus peruvianus has been
taken at several localities including on
the slopes of Volcan Sumaco but no
specimens of E. condor have been found
on that semi-isolated mountain; E. con-
dor may be restricted to the Cordillera
del Condor.

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 21

Eleutherodactylus cornutus
(Jiménez de la Espada)
Fig. 2D

Limnophys cornutus Jiménez de la Espada,
1870, Jorn. Sci. Math. Lisbon, 3:60 [Holo-
type.—evidently lost, taken at San Jose de
Moti, Provincia Napo, Ecuador].

Limnophys napaeus Jiménez de la Espada,
1870, Jorn. Sci. Math. Lisbon, 3:60 [Holo-
type.—evidently lost, taken at San Jose de
Moti, Provincia Napo, Ecuador].

Strabomantis cornutus: Jiménez de la Espada,
1872, Anales Soc. Espanol. Hist. Nat. 1:85.

Hylodes cornutus: Boulenger, 1882, Cat. Ba-
trachia Salientia. . . British Museum: 220.

Ctenocranius cornutus: Melin, 1941; Medd.
Goteborgs Mus. Zool., Avdel. 88:49.

Lithodytes cornutus (part): Andersson, 1945,
Arkiv for Zoologi, 37A(2):45.

Eleutherodactylus cornutus cornutus (part):
Rivero, 1961, Bull. Mus. Comp. Zoology,
126:55.

Eleutherodactylus cornutus: Gorham, 1966,
Das Tierreich (85):66; Lynch, 1975, Occas.
Pap. Mus. Nat. Hist. Univ. Kansas (38):
28-31.

Diagnosis.—1 ) skin of dorsum coarse-
ly tuberculate, bearing H-shaped folds
in center of back, that of venter smooth;
2) tympanum prominent, its length %-
% eye length; 3) snout round in dorsal
view, truncate in lateral profile; canthus
rostralis sharp; 4) upper eyelid bearing
elongate conical tubercle; heavy cranial
crests ending in bosses posteromedial to
eyes; 5) vomerine odontophores arch-
like, extending laterally to outer edge of
choanae; 6) males lacking vocal sac and
slits, lacking nuptial pads; 7) first finger
longer than second; pads small, discs as
long as wide; 8) fingers bearing thin
lateral fringes; 9) ulnar tubercles promi-
nent; 10) row of conical tubercles on
heel and outer edge of tarsus; inner
tarsal fold present; 11) two metatarsal
tubercles, inner elongate, outer promi-
nent, not elevated, % size of inner;
supernumerary plantar tubercles minute,
few in number; 12) toes bearing lateral
fringes, not webbed; toe pads slightly
larger than those of outer fingers; 13)
brown above with dark brown facial
markings; limb bars narrow; venter
cream, heavily reticulated with brown;
posterior thighs black with silvery or

golden flecks; 14) adults large, two
males 35.5-43.0 mm, one young female
54.0 mm SVL.

Eleutherodactylus cornutus is most
closely related to E. biporcatus (Peters)
and E. cerastes Lynch, neither of which
have an inner tarsal fold. Both E. ce-
rastes and E. cornutus have elongate tu-
bercles on the upper eyelid. Lynch
(1975a) included these three and E.
bufoniformis (Boulenger) and E. nece-
rus Lynch in the biporcatus group. All
are very different from most Eleuthero-
dactylus in having broad heads (head
width of cornutus 48.8-56.4 percent
SVL).

Description —See Lynch (1975a).

Color in life—Dorsum dull brown
(reddish or olive tint in some) with or
without darker brown or black markings;
posterior surfaces of thighs black with
minute cream, white, or bluish white
flecks; throat brown or reddish brown
with cream flecks; rest of venter dull
creamy gray to bluish white with brown
mottling; feet orange-tan in one indi-
vidual; iris dull bronze to brown; tongue
orange or yellow.

Distribution and ecology.—This large
terrestrial species inhabits cloud forests
at elevations of 1150-1800 m along the
Andean front from southern Colombia
(Putumayo) southward to the Cordillera
Cutucti in southern Ecuador. One indi-
vidual was on the forest floor by day;
all others were found at night on the
ground; some were on stream banks.

Eleutheradactylus cremnobates
new species
Fig. 2E

Holotype.—KU 166036, an adult fe-
male, taken 2 km SSW Rio Reventador
(Quito-Lago Agrio road), Provincia
Napo, Ecuador, 1490 m, on 6 October
1974 by William E. Duellman.

Paratypes.—KU 166037-56, topotypes
collected 19-20 March and 6 October
1974 by W. E. Duellman, A. H. Savitsky,
and L. Trueb.

Diagnosis——1) skin of dorsum finely

22 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

tuberculate, bearing short ridges and
folds, that of venter finely areolate pos-
teriorly, smooth anteriorly; 2) tympanum
distinct, its length %-% that of eye;
3) snout subacuminate in dorsal view,
rounded in lateral profile; canthus ros-
tralis rounded; 4) upper eyelid as broad
as IOD, not bearing pungent warts;
cranial crests absent, although interor-
bital space furrowed; 5) vomerine odon-
tophores prominent, triangular in out-
line; 6) males lacking vocal sac and
slits; no nuptial pad in males; 7) first
finger shorter than second; fingers I-IV
bearing large pads; discs broader than
long; 8) fingers bearing lateral fringes;
9) ulnar tubercles not distinct; 10) heel
and outer edge of tarsus bearing non-
conical tubercles, low ridge on distal
one-half of inner tarsus; 11) two meta-
tarsal tubercles, inner pungent, elongate,
4-6 times size of outer; numerous small
supernumerary plantar tubercles; 12)
toes bearing lateral fringes, not webbed;
pads of toes as large as those of fingers;
13) brown above with darker marbling;
posterior thigh dark brown, flecked with
cream; venter pale gray to cream with
brown chevron across chest separating
yellowish throat with sparse brown spot-
ting from cream belly heavily spotted
with brown; 14) adults moderately large,
seven males 28.4-32.5 mm, eight females
41.6-51.7 (x = 46.2) mm SVL.

Eleutherodactylus cremnobates _ is
similar to E. crenunguis Lynch, E. lati-
discus (Boulenger), and E. rubicundus
(Jiménez de la Espada) in that all have
large digital pads on long slender digits,
sloping loreal regions, and weakly to
strongly flared lips. Eleutherodactylus
cremnobates differs from all of these in
having apically round digital pads. Both
E. cremnobates and E. latidiscus have a
thumb that is slightly shorter than the
second finger (first slightly longer than
second in E. crenunguis and E. rubi-
cundus ).

Description.—Head as broad as or
broader than body, broader than long;
head width 36.7-41.7 (x = 39.8, N =
26) percent SVL; snout elongate; nos-

trils protuberant, directed laterally; E-N
80.5-93.3 (x = 88.2, N = 7) percent eye
length in males, 87.8-103.8 (x = 95.0,
N = 19) percent in females; loreal re-
gion weakly concave, sloping gradually
to lips; lips weakly flared; interorbital
space flat, cranial crests absent, although
edges of frontoparietals higher than cen-
ter toward anterior end of frontoparietals
and nasals dip toward midline; upper
eyelid width 91.3-124.1 (x = 105.5, N =
26) percent IOD; temporal region not
swollen; tympanic region not vertical;
tympanum distinct, much smaller than
pad of finger II, III, or IV; tympanum
length 17.1-25.0 (x = 21.7, N = 7) per-
cent eye length in males, 21.0-29.3 (x =
25.6, N = 8) percent in adult females,
and 16.7-26.5 (x = 21.4, N = 11) percent
in juvenile females (latter not different
from males, but different from adult fe-
males); supratympanic fold thickened,
concealing upper edge of tympanic an-
nulus; choanae relatively small, three-
cornered, each smaller than an odonto-
phore; vomerine odontophores median
and posterior to choanae, separated me-
dially by distance equal to choanal
breadth, bearing teeth in a transverse
row; tongue longer than wide, notched
posteriorly, the posterior 2/5 not adher-
ent to floor of mouth.

Skin of dorsum finely tuberculate and
bearing short ridges and folds; skin of
head likewise textured, that of upper
eyelid bearing slightly larger tubercles
(none elongate); postrictal tubercles
subconical; anal opening not modified;
flanks bearing flattened tubercles, most
obvious anteriorly; skin of venter finely
areolate posteriorly, that over most of
venter and on throat smooth; skin of
thighs ventral and ventrolateral to vent
coarsely areolate; discoidal folds promi-
nent; ulnar tubercles very inconspicuous,
poorly defined; palmar tubercle bifid,
slightly larger than oval thenar tubercle;
supernumerary palmar tubercles small,
numerous, subarticular tubercles round
to longer than wide, pungent, non-coni-
cal; all digits bearing discs, broader than
long and apically rounded, on pads;

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 23

pads of fingers II-IV at least twice width
of digit below pad, that on thumb only
slightly wider than digit; pads rounded
apically; fingers long and slender, thumb
shorter than second finger (when ad-
pressed, tip of I reaches proximal edge
of disc of II).

One non-conical tubercle on heel,
1-3 equal sized tubercles along outer
edge of tarsus; inner edge of tarsus
bearing a low ridge for the distal one-
half; inner metatarsal tubercle non-
compressed, pungent, its length 2.5
times its width, 4-6 times size of oval
outer; supernumerary plantar tubercles
small, non-pungent, numerous; subartic-
ular tubercles slightly longer than wide,
pungent, non-conical; toes bearing prom-
inent lateral fringes, which coalesce
basally producing very slight webs;
toes bearing broad pads and _ discs
(broader than long, apically rounded),
pads of toes III-V as large as those of
outer fingers, those of toes I-II as large
as that of finger II; hind limbs of moder-
ate length; heel of adpressed leg reaches
anterior to eye (between eye and nos-
tril); when legs are flexed and held at
right angles to sagittal plane, heels over-
lap; shank 53.1-56.8 (x = 55.2, N = 7)
percent SVL in males, 54.8-57.9 (x =
56.6, N = 11) percent in juvenile fe-
males, and 51.3-57.2 (x = 53.7, N = 8)
in adult females.

Brown on all upper surfaces with
darker brown marbling; some ridges (es-
pecially on occiput) high-lighted with
pale brown; flanks suffused with gray
and marbled with black; face pale brown
below eye, labial bars darker brown;
canthal-supratympanic stripe dark brown
or black, narrow; limb bars diffuse,
weakly oblique, as wide as interspaces;
posterior thigh dark brown, spotted
with cream (spots small); anterior sur-
face of thigh heavily reticulated with
dark brown to dark brown with cream
flecks and reticulations; underside of
shank heavily marbled with dark brown;
venter pale gray to cream, usually heav-
ily reticulated or spotted with brown;
throat dirty yellow with sparse brown

spotting, finely stippled with brown; a
diffuse to prominent brown line extend-
ing from just anterior to forearm inser-
tion posteromedially to area above
sternum where it may or may not meet
its counterpart; this “line” divides the
ventral surface into a yellowish forepart
with sparse flecking, and a cream to gray
hind part, usually heavily spotted or
marbled with brown.

Color in life-——Dorsum olive-brown
to dark brown with tips of some tuber-
cles and ridges olive-tan or orange in
some individuals. Hind limbs dark
brown to black with creamy-tan trans-
verse bars. Venter pink, rosy brown, to
dull yellow with brown or black flecks
or mottling. Iris reddish copper or cop-
pery brown. Some individuals from the
Rio Salado have a yellow spot at the
base of the thigh and in the groin. Ven-
ter in a juvenile creamy gray with brown
marbling.

Measurements of holotype in mm.—
SVL 51.7; shank 27.0; head width 20.6;
head length 18.7; upper eyelid width 4.9;
IOD 4.6; tympanum length 1.6; eye
length 6.1; E-N 6.2. The holotype is a
spent adult female having extensively
convoluted oviducts.

Etymology.— Greek (kremnos, a
cliff) and (batés, one that haunts) in
reference to microhabitat selection by
the frog.

Distribution and ecology—tThis spe-
cies is known from three localities be-
tween 1410 and 1700 m on the eastern
face of the Cordillera Oriental drained
by the Rio Coca. All individuals were
found at night at various times of the
year (March, April, July, October, De-
cember). Three of the frogs were
perched on branches or logs over small
streams in cloud forest. Seven were on
leaves of herbs along small streams, and
25 were on moss, ferns, or small herbs
on cliffs over streams. Two adult females
are spent (KU 166036, taken in October,
and KU 166044, taken in March).

Remarks.—lf E. latidiscus is the clos-
est relative of E. cremnobates, the rela-
tionship is not extremely close. Many

24 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

minor structural differences are evident
between the two but they are the only
moderate-sized species of the wnistri-
gatus group having large pads on slender
digits, flared lips, obtuse canthi rostrali,
large eyes, low cranial crests, and small
tympana. In each, males lack vocal sacs
and slits. Eleutherodactylus cruentus
(Peters) may be allied but is judged less
closely related to E. cremnobates and E.
latidiscus than either is to one another.

Eleutherodactylus croceoinguinis Lynch

Eleutherodactylus croceoinguinis Lynch, 1968,
Journ. Herpetol. 2:133 [Holotype —KU
110789, Santa Cecilia, Provincia Napo,
Ecuador, 340 m].

Diagnosis.—1) skin of dorsum tuber-
culate, that of venter coarsely areolate;
2) tympanum concealed beneath skin;
3) snout subacuminate in dorsal view,
round in lateral profile; canthus rostralis
sharp, weakly concave; 4) upper eyelid
slightly broader than IOD; no cranial
crests; 5) vomerine odontophores oval in
outline; 6) males lack vocal sac and
slits, no nuptial pads; 7) first finger
shorter than second; pads bearing discs
(broader than long), pads largest on
fingers III-IV; 8) fingers lacking lateral
fringes; 9) ulnar tubercles evident,
small; 10) tarsus bearing row of conical
tubercles along outer edge, no tubercles
or fold on inner edge; small tubercle on
heel; 11) two metatarsal tubercles, inner
elongate, 3-4 times size of rounded outer;
no supernumerary plantar tubercles; 12)
toes lacking lateral fringes and webbing;
toe pads slightly smaller than those of
fingers; 13) dorsum and venter brown,
former flecked with white and bearing
darker brown markings, latter heavily
flecked with cream; limb bars narrow;
posterior surfaces of thighs brown; a
pale spot on anterior edge of thigh (in
groin), another on adjacent flank (pale
spots yellow to orange in life); 14)
adults small, males 12.8-18.2 (x = 14.6,
N = 14) mm, females 18.0-21.9 (x =
19.8, N = 19) mm SVL.

Eleutherodactylus croceoinguinis is
mostly closely related to E. carvalhoi
Lutz and E. martiae Lynch. Eleuthero-
dactylus martiae lacks pale spots in the
groin, vomerine odontophores are usu-
ally not apparent, and exhibits pro-
nounced pattern polymorphism. FEleu-
therodactylus carvalhoi is more similar
(it has a single yellow spot in the groin)
and is allopatric.

Description—See Lynch (1968a).

Color in life——Dorsum yellow-brown
to brown, spotted with dark brown and
flecked with white or yellow; flanks
brown to black, blotched with darker
brown and flecked with white or yellow;
venter brown (gray to black) with
cream flecks; limb bars and anal tri-
angle dark brown on paler brown ground
color; spots in groin yellow to reddish-
orange; iris gray, finely reticulated with
black.

Distribution and ecology.—This spe-
cies occurs in lowland rainforests in
Ecuador and adjacent Colombia but in- -
vades low cloud forests at the base of
the Pastaza Trench. In lowlands as well
as cloud forests, E. croceoinguinis is noc-
turnal and is found on low vegetation
(0.5-1.5 m above ground) in dense for-
ests. Eleutherodactylus croceoinguinis
from low cloud forests are no larger
than specimens from the lowlands (in
contrast to E. lacrimosus, E. lanthanites,
E. nigrovittatus, and E. peruvianus) and
are as abundant in cloud forests as in
the lowlands (in contrast to E. ocken-

deni).

Eleutherodactylus cryptomelas Lynch

Eleutherodactylus cryptomelas Wynch, 1979,
Misc. Publ. Mus. Nat. Hist. Univ. Kansas
66:21 [Holotype—KU 141992, Abra de
Zamora, 15 km E Loja, Provincia Zamora-
Chinchipe, Ecuador, 2710 m].

This species occurs in subpaéramo and
cloud forests on the eastern Andean cor-
dillera in southern Ecuador at elevations
of 2470 to 3100 m (Lynch, 1979a).

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 25

Eleutherodactylus devillei (Boulenger)
Fig. 2F

Hylodes DeVillei Boulenger 1880, Bull. Soc.
Zool. France 5:47 [Holotype -—MRHN
1009, adult female taken in the “Andes of
Ecuador,” by Emile de Ville in 1874 or
1875].

Eleutherodactylus devillei Peters, 1955, Rev.
Ecuat. Ent. Par. 2:351; Lynch, 1970, Journ.
Herpetol. 3:136-38.

Diagnosis.—1 ) skin of dorsum smooth
with prominent dorsolateral folds, that
of venter areolate; 2) tympanum promi-
nent, its length %-% eye length; 3)
snout subacuminate in dorsal view,
rounded in lateral profile; canthus ros-
tralis sharp; 4) IOD broader than upper
eyelid; upper eyelid lacking tubercles;
cranial crests present, low; 5) vomerine
odontophores prominent, oblique; 6)
males lack vocal sac and slits, no nup-
tial pads; 7) first finger shorter than
second; pads bearing discs (broader than
long), pads large on fingers II-IV; 8)
fingers bearing narrow lateral fringes;
9) forearm bearing ill-defined ulnar tu-
bercles; 10) inner tarsal fold present, no
tubercles on heel or tarsus; 11) two
metatarsal tubercles, inner elongate, 3-4
times size of oval outer; no supernu-
merary plantar tubercles; 12) toes bear-
ing narrow lateral fringes, not webbed;
toe pads slightly smaller than those of
fingers; 13) dorsum brown with black or
dark brown chevrons and flecks; limb
bars narrow; posterior surfaces of thighs
yellow-brown with brown reticulations;
venter cream with brown reticulation;
14) adults moderate-sized, males 19.6-
31.6 (x = 26.8, N = 26) mm, females
34.7-44.7 (x = 39.0, N = 26) mm SVL.

Eleutherodactylus devillei is most
closely related to a series of species now
called E. vertebralis (Boulenger) and
less closely related to E. buckleyi (Bou-
lenger), E. cryophilius Lynch, and E.
curtipes (Boulenger). Eleutherodactylus
devillei differs from E. cryophilius and
E. curtipes in having visible tympana
and large digital pads as well as in col-
oration (neither has reticulation on the
venter). The constellation of frogs now

called E. vertebralis all have numerous
supernumerary plantar tubercles and less
prominent cranial crests. Eleutherodac-
tylus buckleyi lacks an inner tarsal fold
and never has a pattern consisting of
thin limb bars and dorsal chevrons but
frequently has a marbled or reticulate
venter.

Description.—See Lynch (1970b).

Color in life—Dorsum pale brown
(bronze or reddish tint common) with
dark brown to black markings. Flanks
pale gray with dark mottling, pinkish
cast in some. Throat and anterior belly
dull yellowish white, becoming pale gray
posteriorly, with dark brown to black
mottling. Iris coppery bronze.

Distribution and ecology.—This spe-
cies occurs in upper montane forests and
clearings on the eastern slopes of the
Andes in Provincia Napo, Ecuador.
With the exception of a specimen from
Santa Barbara, and one from “top of
Sumaco,” all specimens are from the
valley of the Rio Papallacta between
2350 and 3155 m. A record (USNM)
from Borja (1710 m) needs to be veri-
fied. This frog is found beneath rocks
and logs by day and on low vegetation
at night.

Remarks.—Lynch (1970b) reported
only females in a sample of 16 speci-
mens and speculated that the absence
of males might be a product of collec-
tion bias. We have seen more than 100
specimens and find no distortion from a
50:50 sex ratio.

Lynch (1970b) suggested that E.
devillei was [closely] related to E. sur-
dus (Boulenger). The failure to asso-
ciate E. devillei with E. buckleyi or E.
vertebralis was a result of not having
specimens of buckleyi or vertebralis
available (those Lynch thought to be
buckleyi are specimens of E. curtipes).
The purported relationship of E. chlo-
ronotus and E. devillei also represents a
premature association. The relationships
of E. chloronotus are obscure (see spe-
cies account above ) but are certainly not
with E. devillei. No hint of the distinc-
tive concave canthus rostralis of E. chlo-

26 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

ronotus is evident in E. buckleyi, E.
cryophilius, E. curtipes, E. devillei, or
E. vertebralis.

Eleutherodactylus dolops new species
Fig. 2G

Holotype —KU 143505, an adult fe-
male, taken at Salto de Agua, 2.5 km
NNE Rio Reventador, Provincia Napo,
Ecuador, 1660 m, on 25 October 1971
by William E. Duellman.

Paratypes—KU 143504, Rio Azuela,
Provincia Napo, Ecuador, 1740 m; KU
165866-68, 2 km SSW Rio Reventador,
Provincia Napo, Ecuador, 1490 m;
USNM GOV 8901, Loreto, Provincia
Napo, Ecuador; USNM GOV 9332-33,
Rio Villano, Provincia Pastaza, Ecuador;
KU 168811, 10.3 km W El Pepino, De-
partamento Putumayo, Colombia, 1440
m.

Diagnosis.—1) skin of dorsum pus-
tulate, no dorsolateral folds, that of ven-
ter smooth; 2) tympanum distinct, its
length 4-% eye length; 3) snout sub-
acuminate in dorsal view, round in lat-
eral profile; canthus rostralis moderately
sharp; 4) upper eyelid broader than
IOD, not bearing pungent warts; no
cranial crests; 5) vomerine odontophores
prominent, triangular in outline; 6)
males with vocal slits, subgular vocal
sac; 7) first finger longer than second;
discs as long as broad; pads narrow;
8) no lateral fringes; 9) no ulnar tuber-
cles; 10) no tubercles on heel or tarsus;
11) two metatarsal tubercles, inner elon-
gate, 4-8 times size of outer; no super-
numerary plantar tubercles; 12) toes
lacking lateral fringes and webbing; toe
pads as small as those of fingers; 13)
brown with darker blotches; posterior
thigh brown with cream flecks; venter
dirty white, reticulated with brown; 14)
adults large, two males 35.8-40.3 mm,
three females 56.6-57.6 mm SVL.

Eleutherodactylus dolops is most sim-
ilar to and apparently related to E.
cruralis (Boulenger), EE. discoidalis
(Peracca), E. elassodiscus Lynch, E.
granulosus (Boulenger), E. mantipus

(Boulenger), and E. nigrovittatus An-
dersson, but E. dolops is considerably
larger than any of these (males 21.1-
29.2 mm, females 24.0-36.9 mm SVL)
and has much smaller outer metatarsal
tubercles (at least one-half as large as
inner metatarsal tubercle in the other six
species ).

Description.—Head as broad as body,
wider than long; head width 40.2-42.0
(x = 41.3, N = 8) percent SVL; snout
subacuminate in dorsal view, rounded in
lateral profile, short, E-N 71.4-87.0 (x =
78.1, N = 8) percent eye length; loreal
region weakly concave, sloping abruptly
to lips; lips not flared; nostrils lateral,
weakly protuberant or not, much closer
to tip of snout than to eye; upper eyelid
width 100.1-152.2 (x — 122.3, N = 8)
percent IOD; no cranial crests; tympa-
num prominent, round in males, higher
than long in females, its length 32.0-
ol.8 (x = 3915, N = 8) percentieye
length; supratympanic fold heavy, con-
cealing upper edge of tympanum, not
reaching insertion of arm; postrictal tu-
bercles forming ridge anteroventral to
terminis of supratympanic fold; upper
eyelid and, to lesser extent, top of head
bearing numerous pustules, none en-
larged; tongue relatively small, round to
slightly longer than wide, its posterior
edge bearing shallow notch, posterior
one-sixth not adherent to floor of mouth;
choanae round to triangular in outline,
much smaller than a vomerine dentiger-
ous process, contained within mesial bor-
ders of jaw; vomerine odontophores
wider than long, triangular in outline,
posteromedial to choanae, separated by
% width of an odontophore width
(1.5 choanal widths), bearing a trans-
verse row of 8-13 moderate-sized teeth
along posterior border.

Skin of dorsum pustulate with short
ridges along upper flanks; flanks areo-
late; upper surface of limbs finely pustu-
lose; skin below vent areolate; discoidal
folds prominent; palmar tubercle bifid,
as large as oval thenar tubercle; super-
numerary palmar tubercles low, flat, a*
base of each digit; subarticular tubercles

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 27

elongate, subconical, simple; fingers
long, slender, thumb longer than second
finger; fingers lacking lateral fringes, but
feebly keeled laterally; fingers bearing
weakly dilated pads, all pads bearing
discs, discs as wide as long or longer
than wide; pads and discs of fingers I
and II pointed, those of III and IV
rounded apically.

Heel and tarsus lacking tubercles or
folds; two metatarsal tubercles, inner
metatarsal tubercle 2.5 times as long
as wide, not elevated or compressed,
7-8 times size of round (subconical in
KU 143504) outer metatarsal tubercle;
in KU 143505 the outer metatarsal tu-
bercle is low and flat, 4-% size of
inner; no supernumerary plantar tuber-
cles; subarticular tubercles longer than
wide, subconical, simple; no _ lateral
fringes on toes, edges weakly keeled;
toes lacking webbing; all toes with pads
and discs, tips rounded or acutely
rounded, discs longer than wide; hind
legs long, shank 59.2-69.4 (x = 63.6, N
= 8) percent SVL; heel of adpressed
hind limb extends beyond snout tip;
heels of flexed hind legs overlap.

In preservative, brown above with
darker brown markings (labial bars, can-
thal and supratympanic stripes, inter-
orbital triangle, spots on back and flanks,
limb bars); groin and posterior thigh
surfaces brown with cream flecks; venter
cream heavily reticulated with brown;
throat, chest, and underside of limbs
brown with cream flecks.

Color in life-——Dorsum dull tan to
brown with darker brown to black mark-
ings; posterior surfaces of thighs orange-
brown, reddish brown, or dark brown
with cream or pale yellow flecks. Venter
creamy yellow to pinkish tan with brown
mottling or dark brown to black with
bluish white flecks. Iris dull brown with
grayish or reddish tint.

Measurements of holotype in mm.—
SVL 57.6; shank 34.1; head width 24.2;
head length 22.0; upper eyelid width 7.0;
IOD 4.6; tympanum length 2.8; eye
length 7.7; E-N 5.5. Each vomerine
odontophore has 10 teeth. The holotype

is a gravid female with heavily convo-
luted oviducts and large (4.0-4.5 mm in
diameter), yellow ovarian eggs.

Etymology.—Greek, meaning one hid-
ing in ambush, in allusion to the frogs
being found behind waterfalls.

Distribution and ecology.—This spe-
cies is definitely known from elevations
of 1440 to 1950 m on the eastern face
of the Andes in Departamento Putu-
mayo, Colombia, and Provincia Napo,
Ecuador. Three specimens in the USNM
were received from Gustavo Orces-V.
and collected by the Olalla brothers, sup-
posedly at Loreto (550 m) and Rio
Villano (< 500 m). These localities are
questioned until additional material is
acquired from the lowlands.

Eleutherodactylus dolops inhabits
streams in cool cloud forest. Three were
sitting on boulders in streams at night,
and one was perched on a low bush
adjacent to a stream at night. By day,
two were under rocks at the edge of a
stream, one was under a rock in the
splash zone of a waterfall, and one was
in a crevice under moss behind a water-

fall.

Eleutherodactylus elassodiscus Lynch
Fig. 2H

Eleutherodactylus elassodiscus Lynch, 1973,

Copeia 1978(2):222 [Holotype-—USNM

167668, adult female from Cuyujia, Pro-
vincia Napo, Ecuador, 2380 m].

Diagnosis.—1) skin of dorsum smooth
to feebly pustulate, no dorsolateral folds,
that of venter smooth; 2) tympanum
prominent, its length %-% eye length;
3) snout subacuminate in dorsal view,
sloping in lateral profile, canthus ros-
tralis poorly to moderately well-defined;
4) upper eyelid narrower than (to
equal) IOD; no cranial crests; small
pustules on upper eyelid; 5) vomerine
odontophores prominent, broad, extend-
ing laterally to choanae; 6) males lack-
ing vocal sac and slits, no nuptial pad;
7) first finger longer than second, all
bearing narrow digital pads, rounded
apically; discs as long as wide or longer

28 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

than wide; 8) fingers lacking lateral
fringes or keels; 9) no ulnar tubercles;
10) no tubercles on heel or tarsus; 11)
two metatarsal tubercles, inner oval, ap-
proximately same size as rounded outer;
no supernumerary plantar tubercles; 12)
toes lacking fringes or webbing; toe pads
narrow, as large as those of fingers; 13)
dorsum gray to brown with or without
brown markings; limbs barred; posterior
surface of thigh gray to brown (uni-
color); venter cream with some brown
mottling on throat; 14) adults moderate-
sized, males 22.0-29.2 mm, females 29.7—
36.9 mm SVL.

Eleutherodactylus elassodiscus is a
member of the discoidalis group (Lynch,
1976b), which also includes E. cruralis
(Boulenger), E. discoidalis (Peracca),
E. dolops Lynch and Duellman, E. gran-
ulosus (Boulenger), E. mantipus (Bou-
lenger), and E. nigrovittatus Andersson.
The sloping snout of E. elassodiscus and
E. nigrovittatus distinguish that pair of
species from the other five species. Eleu-
therodactylus nigrovittatus has pointed
digital pads, the inner metatarsal tuber-
cle is twice as large as the outer, and
males have a fleshy swelling of the up-
per lip and snout tip.

Description —See Lynch (1973a).

Color in life-—Dorsum pinkish tan to
grayish brown with olive-brown to dark
brown markings; stripe below tympanum
(postrictal) creamy orange; posterior
surfaces of thighs reddish or orangish
brown; venter gray to brown, with or
without silvery flecks laterally; iris
bronze, heavily flecked with black.

Distribution and ecology.—This ter-
restrial species occurs at elevations of
2300 to 2900 m in upper humid mon-
tane forest on the eastern slopes of the
Andes in extreme southern Colombia
(Departamento Putumayo) and northern
Ecuador (Provincia Pastaza). A record
from Borja, Ecuador (1710 m) needs to
be verified. All specimens were found
under rocks and logs by day.

Eleutherodactylus eriphus new species
Fig. 3A

Holotype.-—KU 166031, an adult fe-
male, taken at the Rio Jatuntinahua, 10
km SE Cuyujua, Provincia Napo, Ecua-
dor, 2160 m, on 21 March 1976 by Alan
H. Savitsky and John E. Simmons.

Paratypes.—KU 166032-35 collected
syntopically with holotype; KU 155451-
do, from 12 km E Papallacta, Provincia
Napo, Ecuador, collected 9 January 1974
by Ronn Altig.

Diagnosis.—1) skin of dorsum bear-
ing many small spinules (small conical
tubercules ), that of venter coarsely areo-
late; 2) tympanum prominent, its length
4-% that of eye; 3) snout round in
dorsal and lateral profiles; canthus ros-
tralis sharp; 4) upper eyelid as wide as
IOD, bearing one pungent tubercle; no
cranial crests; 5) vomerine odontophores
moderately prominent, round to oblique-
ly oval in outline; 6) males with vocal
slits and subgular vocal sac; 7) first
finger shorter than second, pads largest
on fingers III-IV, discs broader than
long; 8) fingers bearing narrow lateral
fringes; 9) ulnar tubercles conical to
subconical, coalescing to form ridge; 10)
knee, heel, and outer edge of tarsus
bearing small conical tubercles; 11) two
metatarsal tubercles, inner oval, 3 times
size of subconical outer; numerous su-
pernumerary tubercles; 12) toes bearing
narrow lateral fringes, not webbed; toe
pads as large as those of fingers; 13)
brown above with indistinct darker
brown chevrons, canthal, labial, and su-
pratympanic markings; posterior surfaces
of thigh barred white and dark brown;
white spots on concealed shank; venter
brown with diffuse brown spots; 14)
adults moderate-sized, nine males 18.1-
25.2 mm, two adult females 25.8-29.0
mm SVL.

Although distinctive in having boldly
barred thighs and warty skin on the
dorsum, E. eriphus may be the Ama-
zonian slope replacement for E. calcara-
tus (Boulenger) and E. crucifer (Bou-
lenger). It differs from these species in

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 29

having a straight (or weakly concave)
canthus rostralis and in having boldly
marked thighs. The heel tubercle of E.
calcaratus is longer than that of E. eri-
phus. Eleutherodactylus crucifer has
crenulate lateral fringes on the digits.
Eleutherodactylus eriphus appears close-
ly related to E. nigrogriseus and E. spi-
nosus. All three have bold patterns on
the posterior surfaces of the thighs (bars
in E. eriphus and large spots in the other
two species). The skin texture of E.
eriphus is matched by E. spinosus Lynch
from the Amazonian slopes of southern
Andean Ecuador but E. spinosus is
slightly larger, has cranial crests (albeit
low), 2-3 enlarged tubercles on the up-
per eyelid, males lacking vocal sac and
slits, and the posterior surfaces of the
thigh and groin are black enclosing
white spots.

Description—Head as wide as or
wider than body, wider than long; head
width 34.8-40.7 (x = 37.6, N = 10) per-
cent SVL; snout shorter in small indi-
viduals, E-N 75.0-91.7 (x = 83.0, N =
5) percent eye length in males, 80.0 in
two juvenile females (18.9-19.6 mm
SVL), 100.0-106.2 (x = 103.9, N = 3)
in young and adult females; nostrils
weakly protuberant, directed dorsolat-
erally; canthus rostralis straight in males,
concave in females; loreal region con-
cave, sloping abruptly (small individ-
uals) to gently (large females) to lips;
lips not flared in small individuals, weak-
ly flared in adult females; interorbital
space flat (no cranial crests); upper eye-
lid width 84.0-122.2 (x = 96.8, N = 9)
percent IOD; one conical tubercle on
upper eyelid; supratympanic fold short,
evident above and behind tympanum,
concealing tympanic annulus; tympanum
prominent, round, separated from eye by
distance equal tympanum length; tym-
panum length 33.3-40.0 (x = 36.2, N =
10) percent eye length; choanae round,
relatively small, not concealed by palatal
shelf of maxillary arch; vomerine odon-
tophores median and posterior to cho-
anae, moderately elevated, round to
obliquely oval, separated by distance

equal their own width in small indi-
viduals, not separated in adult females;
each odontophore about size of a choana
and bearing 2-3 teeth in small individ-
uals, larger than a choana and bearing
4-6 teeth in a slanted row in adult fe-
males; tongue longer than wide, poste-
rior border shallowly notched, posterior
% not adherent to floor of mouth.

Skin of dorsal surfaces bearing many
small spinules, becoming sparse on
flanks, no dorsolateral folds; skin of ven-
ter coarsely areolate; discoidal folds
prominent, well anteriad to groin; vent
not extended by sheath; subanal tuber-
cles conical; forearm bearing conical to
subconical (adult females) ulnar tuber-
cles, partially connected in ridge; palmar
tubercle bifid, larger than oval thenar
tubercle; numerous small supernumerary
palmar tubercles; fingers bearing narrow
lateral fringes; outer edge of hand and
fourth finger bearing lateral fold; in
small individuals (all but adult female),
basal subarticular tubercles of fingers
I and IV round, non-conical, subarticular
tubercle on II, both on III, and distal
tubercle on IV, paired; in adult females
all tubercles single, distal tubercle broad-
er than long; fingers bearing discs
(broader than long) on apically rounded
pads; pads of III-IV larger than those
on inner fingers.

Knee, heel, and outer edge of tarsus
bearing small conical tubercles; inner
edge of tarsus lacking tubercles or with
two small tubercles or a thin tarsal
ridge; inner metatarsal tubercle 2-3 times
as long as wide, not compressed, outer
metatarsal tubercle round, subconical,
4-6 size of inner; numerous super-
numerary plantar tubercles; toes bearing
narrow lateral fringes, similar fringes
along inner border of toe I and outer
border of toe V and foot; all toes bear-
ing broad discs on apically round pads;
pads largest on lateral toes; heels over-
lap when hind legs flexed; heel of ad-
pressed hind leg reaches to between eye
and nostril; shank 52.0-59.2 (x = 55.3,
N = 10) percent SVL.

In preservative, pale to medium

30 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

brown with darker brown chevrons, can-
thal and supratympanic stripes, indefi-
nite labial bars; flanks slightly darker
than dorsum with white areas at base
of arm and in axilla and slightly oblique
white bars on posterior flank; venter
brown with slightly darker spotting and
marbling; inside of elbow bearing white
spot; forearms bearing diffuse brown
bars; thighs barred white and dark
brown with pale brown stripe on top of
thigh; anal triangle dark brown; con-
cealed shank dark brown with large
white spots; shank brown with indistinct
darker bars.

Color in life—Dorsum pale green
with dark olive markings; flanks and
hidden surfaces of limbs mottled cream
and black; venter greenish white heavily
suffused with black; iris reddish copper.

Measurements of holotype in mm.—
SVL 29.0; shank 15.5; head width 11.7;
head length 10.3; upper eyelid width 2.8;
IOD 2.9; tympanum length 1.2; eye
length 3.6; E-N 3.8. The holotype is a
gravid female.

Etymology.—Greek, a young goat;
name is used in allusion to the call of
the frog (goat-like).

Distribution and _ ecology. — This
small species is known from only two
localities along the valley of the Rio
Papallacta on the eastern face of the
Andes in Provincia Napo, Ecuador; the
elevational range is 2160-2630 m. At 12
km ESE of Papallacta five individuals
were found on low vegetation at night
in January 1974. At the Rio Jatuntina-
hua, a tributary of the Rio Papallacta,
five were on low herbs at night. Males
were calling: “bleep-ee-eep.” At both
localities the frogs were found in par-
tially cleared cloud forest.

Remarks.—Five specimens (WCAB
37903, 37905, 37907-09) are purportedly
from Papallacta; the specimens were
purchased from a professional collector,
and other material in the collection sug-
gests that the frogs originated from
lower in the valley of the Rio Papallacta.
The specimens are not well preserved
and differ from the type-specimens in

having a more pointed snout and flat-
tened spinules on the dorsum.

Eleutherodactylus galdi
(Jiménez de la Espada)

Pristimantis galdi Jiménez de la Espada, 1870,
Jorn. Sci. Math. Lisboa 3:62 [Types ap-
parently lost, originally in Museo Nacional,
Madrid; type-locality, San Jose de Moti,
Provincia Napo, Ecuador].

Hylodes galdi: Boulenger, 1882, Catalogue
Batrachia Salientia . . . British Mus.: 219.

Hylodes festae Peracca, 1904, Boll. Mus. Zool.
Anat. 19(465):28 [Holotype—MSNT An
413, ex. 3776, adult male collected at San
José de Cuchipamba, Provincia Morona-
Santiago, Ecuador, 1000 m, by E. Festa].

Hylodes margaritifer Boulenger, 1912, Ann.
Mag. Natur. Hist. (8)10:189 [Cotypes.—
BM 1912.11.1.54-55/RR 1947.2.16.78-79,
collected at E] Topo, Rio Pastaza, Provincia
Tungurahua, Ecuador, 4200 ft., by M. G.
Palmer].

Eleutherodactylus festae: Peters, 1955, Rev.
Ecuat. Ent. Par. 2:348.

Eleutherodactylus margaritifer: Peters, 1955,
Rev. Ecuat. Ent. Par. 2:349.

Eleutherodactylus galdi: Peters, 1955, Rev.
Ecuat. Ent. Par. 2:350; Lynch, 1969, Her-
petologica 25:270-72; Lynch, 1974, Occas.
Pap. Mus. Nat. Hist. Univ. Kansas (31):
15-16.

Diagnosis.—1) skin of dorsum smooth
or finely shagreened with scattered pus-
tules, no dorsolateral folds, that of venter
areolate and bearing much enlarged
warts scattered among smaller areola-
tions; 2) tympanum distinct, its length
4-# eye length; 3) snout acuminate in
dorsal view, truncate in lateral profile,
tip somewhat swollen; canthus rostralis
sharp, strongly concave; 4) IOD greater
than upper eyelid width; upper eyelid
bearing conical tubercle; cranial crests
present, serrate in large females; otic
ramus of squamosal also bearing a ser-
rate crest in large females; 5) vomerine
odontophores oblique; 6) males with
vocal sac and slits, no nuptial pads; 7)
first finger shorter than second; fingers
II-IV bearing broad discs on large di-
lated pads; 8) fingers bearing ill-defined
lateral fringes; 9) row of conical ulnar
tubercles present; 10) row of conical
tubercles along outer edge of tarsus,
heel and foot; tubercles along outer edge

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 31

of shank; 11) two metatarsal tubercles,
inner elongate, 5-6 times size of oval
outer; no supernumerary plantar tuber-
cles; 12) toes bearing ill-defined lateral
fringes, no webbing; toe pads slightly
smaller than those of outer fingers; 13)
body pale cream (bright green in life)
with dark brown canthal and supratym-
panic stripes, interorbital and limb bars,
sparse spotting of dorsum; venter cream,
dusted with brown; 14) adults moderate-
sized, males 17.1-24.8 (x = 21.0, N =
8) mm, females 28.1-34.0 (x = 31.7,
N = 11) mm SVL.

Eleutherodactylus inusitatus (de-
scribed below) is superficially similar to
E. galdi but lacks cranial crests and en-
larged warts on the belly, has a shorter
snout, and has elongate tubercles on the
eyelid, heel, and elbow. We are aware
of no other Eleutherodactylus having
serrate frontoparietal and squamosal
crests or having the enlarged warts on
the belly.

Descriptions.—See Jiménez de la Es-
pada (1870, 1875), Boulenger (1912),
and Lynch (1969a, 1974).

Color in life—Dorsum bright green
with or without 1) white, gold, or tan
flecks, 2) bronze-brown or black spots
and/or transverse bars on limbs; belly
white, heavily suffused with green or
not, with black flecks laterally or not;
other ventral surfaces green; yellow or
white labial stripe and bronze canthal
stripe present or not; iris green.

Distribution and ecology.—This ar-
boreal species is widely distributed along
the eastern face of the Andes in Ecua-
dor, including Volcan Sumaco and the
Cordillera del Condor. All specimens
bearing elevational data are from 1000-
1830 m. Records from Loreto (510 m)
are questioned. Our specimens were all
on vegetation (leaves of bushes 1 m
above ground to a palm frond 2 m above
ground) in cloud forest at night.

Eleutherodactylus gladiator Lynch
Fig. 3B

Eleutherodactylus gladiator Lynch, 1976, Her-

petologica 32:316 [Holotype—KU 143516,
3.3 km ESE Cuyujia, Provincia Napo,
Ecuador, 2350 m].

Diagnosis.—1) skin of dorsum smooth
with short ridges, no dorsolateral folds,
that of venter areolate; 2) tympanum
prominent, its length %~% eye length;
3) snout acuminate in dorsal view (tip
feebly pointed), round in lateral profile;
canthus rostralis moderately sharp; 4)
upper eyelid much marrower than IOD,
lacking pungent tubercles; no cranial
crests; 5) vomerine odontophores con-
cealed, oblique; 6) males with vocal
slits and subgular vocal sac; males lack-
ing nuptial pads; 7) first finger shorter
than second, discs on fingers II-IV, pads
of III-IV rounded; 8) fingers lacking
lateral fringes; 9) ulnar tubercles ob-
scure; 10) small, non-conical tubercles
on heel, outer edge of tarsus; ridge-like
inner tarsal fold; 11) two metatarsal tu-
bercles, inner elongate, three times size
of round outer; few supernumerary plan-
tar tubercles; 12) toes lack lateral
fringes, no webbing; 13) brown above
with dark brown spots; limb bars nar-
row, oblique; venter cream with brown
flecks; colorless areas in axilla, groin, an-
terior and posterior surfaces of thigh
(colorless areas orange in life); 14)
adults small, three males 14.9-15.6 mm
SVL, one immature female 15.8 mm
SVL.

Eleutherodactylus gladiator was
grouped with E. leoni Lynch and E.
pyrrhomerus Lynch but is the most dis-
tinct of the three (Lynch, 1976a), differ-
ing in having tubercles on the heel and
tarsus, less prominent tubercles on the
upper eyelid and at the corner of the
jaw, and comparatively larger digital
pads.

Description —See Lynch (1976a).

Color in life—Dorsum brown to
orange-tan with brown markings; flanks
green to orange-tan; venter gray to yel-
low-gray; throat yellow; concealed thighs
and groin orange-tan to orange; iris
bronze to gray-brown with red-brown
horizontal streak.

32 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Distribution and _ ecology. — This
small terrestrial species is known only
from the upper Rio Papallacta Valley in
Provincia Napo, Ecuador, where it has
been found by day under rocks and logs
in cloud forest and clearings at eleva-
tions of 2350-2910 m.

Eleutherodactylus glandulosus
(Boulenger)
(Fig. 3C
Hylodes glandulosus Boulenger, 1880, Bull. Soc.
Zool. France 5:47 [Holotype ——IRSNB 1010,
adult female taken in the “Andes of Ecua-
dor’ by Emile de Ville in 1874-75].
Eleutherodactylus glandulosus: Peters, 1955,
Rev. Ecuat. Ent. Par. 2:351; Lynch, 1970,
Journ. Herpetol. 3:138—39.

Diagnosis.—1 ) skin of dorsum weakly
glandular, no dorsolateral folds, that of
venter coarsely areolate; 2) tympanum
prominent, its length %~—% eye length;
3) snout round in dorsal view, truncate
in lateral profile; canthus rostralis sharp,
concave; 4) upper eyelid narrower than
IOD, lacking tubercles; low cranial
crests present; 5) vomerine odonto-
phores low, weakly slanted or subtri-
angular in outline; 6) males with vocal
slits and subgular vocal sac; no nuptial
pads; 7) first finger shorter than second;
discs broad, on broad pads, pads of II-IV
large; 8) fingers bearing ill-defined lat-
eral fringes; 9) ulnar tubercles present,
small; 10) no tubercles on heel or outer
edge of tarsus; low inner tarsal fold
present; 11) two metatarsal tubercles,
inner oval, 4 times size of round outer;
numerous small supernumerary plantar
tubercles; 12) toes bearing ill-defined
lateral fringes, no webbing; toe pads
smaller than those of fingers; 13) dorsum
yellow-tan to brown with indefinite
brown blotches; limbs not barred; venter
pale yellow with or without sparse
brown reticulation; flanks, groin, anterior
and posterior thigh surfaces, concealed
shank brown with large yellow spots;
14) adults moderate-sized, males 24.2—
30:2) (= 226'6 N27) emmy females
O41" (Xi SOaIE IN =820) samy SVE

Eleutherodactylus glandulosus is not

easily confused with any other species of
Eleutherodactylus but is most similar to
E. acerus Lynch and Duellman, E. leu-
copus Lynch, and E. lividus new species,
none of which have the distinctive yel-
low spots on the flanks and concealed
limb surfaces.

Description—See Lynch (1970b).

Color in life-—Dorsum dull tan, yel-
lowish tan, olive-tan, or brown with
yellow spots on flanks and posterior sur-
faces of thighs; venter dull cream to
gray; iris grayish bronze.

Distribution and ecology.—This spe-
cies occurs at elevations of 2105-2890 m
in the upper Rio Papallacta Valley on
the eastern slopes of the Andes in Pro-
vincia Napo, Ecuador. A record from
Borja (1710 m) is questionable. Indi-
viduals have been found under rocks and
logs in cloud forest and clearings by
day; at night they have been found
on rocks, ground and low vegetation.
Males were actively calling from low
(0.3-1.5 m) vegetation during a rain on
July 1977, 11 km ESE Papallacta; few
males were found in the woods adjacent
to the clearing.

Eleutherodactylus ignicolor new species
Fig. 3D

Holotype.—KU 165879, a juvenile fe-
male, taken at the Rio Jatuntinahua, 10
km SE Cuyujua, Provincia Napo, Ecua-
dor, 21 March 1975 by William E. Duell-
man.

Paratypes.—KU 165880, 11 km ESE
Papallacta, Provincia Napo, 2660 m; KU
177296-97, 9.2 km ESE Papallacta, Pro-
vincia Napo, Ecuador, 2750 m.

Diagnosis ——1) skin of dorsum bear-
ing low warts, especially on lower back,
that of venter smooth; 2) tympanum dis-
tinct, its length %% eye length; 3)
snout acuminate in dorsal view, rounded
in lateral profile; canthus rostralis sharp;
head longer than wide; 4) upper eyelid
narrower than IOD; not bearing pungent
warts; no cranial crests; 5) vomerine
odontophores small, oval in outline; 6)
males with vocal slits, subgular vocal

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 33

sac; 7) first finger shorter than second;
pads on IJ-IV moderate-sized; discs
broader than long; 8) fingers bearing lat-
eral keels; 9) small ulnar tubercles pres-
ent; 10) no tubercles on heel; two tu-
bercles on outer edge of tarsus; short
inner tarsal fold; 11) two metatarsal tu-
bercles, inner oval, 4-6 times size of
round outer; few (3-6) supernumerary
plantar tubercles; 12) toes not webbed,
lateral keels indistinct; toe pads smaller
than those of outer fingers; 13) brown
above with dark interorbital bar, dorsal
blotch, facial bars and stripes; posterior
thigh brown with white spots; groin red
in life; brown chevrons on throat; venter
brown with white spots; 14) adults
small, one male 18.0 mm, one adult fe-
male 26.4 mm SVL.

Eleutherodactylus ignicolor is most
closely related to E. prolatus new species
(see below). The two species are also
closely related to E. trachyblepharis
(Boulenger) and E. variabilis Lynch.
These four small frogs have elongated
heads (head length greater than head
width), acuminate snouts, and narrow
digital pads. Eleutherodactylus ignicolor
is mostly readily distinguished in having
red areas in the groin (in life) and white
spots on the posterior surfaces of the
thighs. Eleutherodactylus ignicolor and
E. prolatus have small tubercles along
the outer edge of the tarsus but E. igni-
color lacks tubercles on the heel.

Description —Head as wide as body,
longer than wide; head width 35.0-37.1
(x = 36.0, N = 3) percent SVL; snout
long; E-N 83.3-87.9 (x = 84.8, N = 3)
percent eye length; nostrils weakly pro-
tuberant, directed dorsolaterally, loreal
region weakly concave, sloping abruptly
to lips; lips not flared; upper eyelid lack-
ing prominent tubercles, its width 77.3-
100.0 (x = 89.5, N = 3) percent IOD;
interorbital space flat, no cranial crests;
supratympanic fold not prominent; tym-
panum distinct, round in males, higher
than long in females, separated from eye
by less than tympanum length; tympa-
num length 43.3-50.0 (x = 46.3, N= 3)
percent eye length; postrictal tubercles

small, prominent; choanae_ relatively
small, round, not concealed by palatal
shelf of maxillary arch; vomerine odon-
tophores median and posterior to cho-
anae, smaller than a choana, separated
medially by distance equal %—% width
of an odontophore, each odontophore
oval, bearing teeth in transverse row;
tongue broader than long, its posterior
border feebly notched, posterior one-
third not adherent to floor of mouth.

Skin of dorsum smooth with low
warts on posterior back, no dorsolateral
folds, discoidal folds well anteriad to
groin; warts on flanks larger than areo-
lations on venter; no anal sheath or tu-
bercles; skin of head smooth; palmar
tubercle bifid, much larger than oval
thenar tubercle; many supernumerary
palmar tubercles, slightly smaller than
subarticular tubercles which are round,
pungent; narrow lateral keels on fingers;
pad of I round, those of I-IV broader
than long, discs broader than long; pads
of fingers IIJ-IV narrower than tympa-
num, feebly emarginate.

No tubercles on heel; two small tu-
bercles along outer edge of tarsus; inner
edge of tarsus bearing short, low folds;
inner metatarsal tubercle oval, its length
twice width, 4-6 times size of rounded
outer metatarsal tubercle; supernumer-
ary plantar tubercles at bases of toes
II-IV, few on plantar surface proper;
subarticular tubercles round, pungent;
keels on lateral margins of toes indis-
tinct; toe pads smaller than those of
outer fingers, apically rounded; heels of
flexed hind legs overlap; shank 55.7-57.8
(x=="56'5) N= 3) percent SVL:

In preservative, tan or brown above
with darker brown markings (interorbi-
tal bar, canthal and _ supratympanic
stripes, labial bars, oblique flank bars,
blotches on back, oblique shank bars);
anal triangle brown; posterior surface of
thighs brown with white spots; throat
brown with darker brown chevrons; ven-
ter brown with white spots; groin and
posterior flank red.

Color in life—Dorsum yellowish tan
to brown with dark brown to black

34 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

markings, with or without orange mid-
dorsal stripe; groin and proximal anterior
surfaces of thighs salmon-red to dark
red, with black reticulations in former;
throat gray with black bars edged with
cream; belly pale gray to salmon with
black markings; iris dark red.

Measurements of holotype in mm.—
SVL 21.1; shank 11.8; head width 7.6;
head length 7.8; upper eyelid width 2.2;
IOD 2.4; tympanum length 1.3; eye
length 3.0; E-N 2.5. The holotype is a
juvenile female with straight, non-con-
voluted oviducts and small ovarian eggs.

Etymology.—Latin, ignis (fire) and
color (color) in reference to the flash
colors in the groin.

Distribution and ecology. — An in-
habitant of upper humid montane for-
est, this small frog is known only from
the upper Rio Papallacta Valley (2160-
2750 m) in Provincia Napo, Ecuador.
All individuals were found on bushes,
a root, and bamboo at the edges of small
cascading streams at night.

Eleutherodactylus incanus new species
Fig. 3E

Holotype—KU 143465, an adult fe-
male, from a series collected at the Rio
Azuela, Provincia Napo, Ecuador, 1740
m, 20 October 1971 by J. T. Collins and
William E. Duellman.

Paratypes.—KU 143466-83, topotypes
20-23 October 1971; KU 143457-58,
143460, 16.5 km NNE Santa Rosa, Pro-
vincia Napo, Ecuador, 1700 m.

Diagnosis.—1) Skin of dorsum bear-
ing scattered pustules, no dorsolateral
folds, that of venter coarsely areolate;
2) tympanum distinct although partially
concealed, its length 4-% eye length;
3) snout subacuminate in dorsal view,
rounded in lateral profile; canthus ros-
tralis rounded, loreal region sloping
gradually to lips; 4) upper eyelid broad-
er than IOD, especially in males, not
bearing pungent warts; no cranial crests;
5) vomerine odontophores large, oblique
(slanted); 6) males lacking vocal sac
and slits; 7) first finger shorter than sec-

ond; fingers II-IV bearing very large
pads; discs broader than long; 8) fingers
bearing narrow lateral fringes; 9) fore-
arm pustulose, ulnar tubercles present;
10) conical tubercles on knee, heel, and
outer edge of tarsus; 11) two metatarsal
tubercles, inner oval, 6-8 times size of
outer; numerous supernumerary plantar
tubercles; 12) toes bearing prominent
lateral fringes, no webbing; toe pads
smaller than those of outer fingers; 13)
dorsum brown with darker brown mark-
ings, venter cream flecked with brown;
flanks, thigh, and concealed shank spot-
ted with glossy-white; concealed flank,
thighs, and shank reddish brown with
white spots; 14) adults moderate-sized,
males 14.9-19.4 mm, females 23.9-30.6
mm SVL.

Eleutherodactylus incanus is a mem-
ber of the unistrigatus group and among
Ecuadorian species is distinctive in havy-
ing a rounded canthus. In color, habitus,
and size it resembles E. crucifer (Bou-
lenger) from comparable trans-Andean
sites but differs in lacking an enlarged
tubercle on the upper eyelid and in
lacking crenulate fringes on the digits.
The head is relatively broad for a species
of the wunistrigatus group (see Lynch,
1975a).

Description—Head_ broader than
body in males and young females, nar-
rower than body in adult females; head
broader than long; head width 39.9-
41.1 (x = 40.5, N = 4) percent SVL in
males, 41.4-45.0 (x = 42.5, N = 9) in
females; snout subacuminate in dorsal
view (less so in smaller specimens),
round in lateral profile; snout moderately
long, E-N 75.0-82.8 (x = 77.4, N = 4)
percent eye length in males, 85.3-100.0
(x = 92.1, N = 9) in females; nostrils
weakly to prominently protuberant
(more so in males), directed dorsolat-
erally; loreal region concave, sloping
gradually to lips; lips not flared; inter-
orbital space flat, slightly broader than
upper eyelid, upper eyelid broad, its
width 120.0-142.1 (x = 130.2, N = 4)
percent IOD in males, 92.6-117.7 (x =
104.1, N = 9) in females; supratympanic

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 35

fold thick, warty, concealing upper and
posterior edges of tympanic annulus;
temporal regions sloping (not vertical);
tympanum distinct, although partially
concealed, round, its diameter 25.0-31.0
(x = 27.8, N = 4) percent eye length
in males, 29.4-40.0 (x = 35.6, N = 9)
in females; tympanum directed dorso-
laterally with slight posterior vector,
separated from eye by twice tympanic
diameter; upper eyelid and skin near
tympanum bearing small conical warts;
postrictal tubercles large; choanae small,
round, not concealed by palatal shelf of
maxillary arch; vomerine odontophores
median and posterior to choanae, ele-
vated, oblique, separated by distance
equal to one-half to full width of cho-
anae, each larger than a choana, bearing
2-7 teeth in an oblique row; tongue
round, posterior edge notched, posterior
two-fifths not adherent to floor of mouth.

Skin of dorsum and flanks pustulose,
pustules scattered; no dorsolateral folds;
discoidal folds well anteriad to groin;
anal opening not elongated by sheath;
skin of forearm pustulose, but enlarged
tubercles present along outer edge of
arm; palmar tubercle bifid, larger than
oval thenar tubercle; numerous super-
numerary palmar tubercles; subarticular
tubercles round, elevated; fingers bear-
ing narrow lateral fringes; outer edge of
hand bearing conical tubercles; all dig-
its bearing broad discs on pads; pad of
thumb scarcely enlarged, those of II-IV
very broad, round apically.

Heel and knee bearing several coni-
cal tubercles; outer edge of tarsus bear-
ing 2-3 conical tubercles, inner edge
bearing one non-conical tubercle just
proximal to inner metatarsal tubercle;
inner metatarsal tubercle oval, 1.5
times as long as wide, not compressed;
outer metatarsal tubercle subconical,
%-® size of inner; plantar surface
bearing tubercles along metatarsals; sub-
articular tubercles round, elevated,
smaller than those of fingers; toes bear-
ing prominent lateral fringes, no webs;
all toes with discs (broader than long)
on apically round pads; toe pads smaller

than those of fingers; heel of adpressed
hind leg reaches nostril; heels of flexed
hind legs overlap; shank 51.4-53.4 (x =
52.2, N = 4) percent SVL in males,
52.9-59.4 (x = 56.8, N = 9) in females.

In preservative, dorsum brown with
darker brown interorbital bar, occipital
W, chevron on sacrum (arms extended
onto flank), canthal-supratympanic
stripe, labial bars, limb bars; limb bars
oblique on shank, narrower than inter-
spaces; anal triangle brown; venter
cream flecked with dark brown; con-
cealed areas on flanks, thigh, shanks red-
dish brown (in some nearly black) with
white spots; reddish areas on upper
flanks and top of thighs; flanks, thigh,
and concealed shank appear painted
with bright enamel-white spots.

Color in life—Dorsum green with
darker green or reddish brown mottling;
flanks and posterior thighs dark red with
white spots; inner two fingers and inner
two toes yellow; venter yellow with
black spots; iris bronze with black re-
ticulations.

Measurements of holotype in mm.—
SVL 27.8; shank 15.7; head width 11.5;
head length 11.3; upper eyelid 3.2; IOD
3.2; tympanum length 1.2; eye length
3.8; E-N 3.4.

Etymology.—Latin, incanus (hoary)
in reference to white spots on the flanks
and concealed limb surfaces.

Distribution and ecology.—This dis-
tinctive frog was found at two localities
at 1700 and 1740 m on the eastern face
of the Andes in Provincia Napo, Ecua-
dor, in October 1971. All were in well-
shaded cloud forest. One was in moss
on a tree trunk by day, and 21 were on
bushes or low trees at night.

Eleutherodactylus incomptus
new species
Fig. 3F

Holotype —KU 143484, an adult
male, taken 16.5 km NNE Santa Rosa,
Provincia Napo, Ecuador, 1700 m, on 17
October 1971 by Joseph T. Collins.

Paratypes—KU 143455, 143461-64,

36 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

143485-86, collected at type-locality 17-
19 October 1971 by J. T. Collins and
W. E. Duellman.

Diagnosis——1) skin of dorsum bear-
ing low flat warts, that of venter coarsely
areolate; 2) tympanum distinct, its
length %-% that of eye; 3) snout sub-
acuminate to round in dorsal view, round
in lateral profile; canthus rostralis round-
ed; 4) upper eyelid as wide as IOD, not
bearing pungent tubercles; no cranial
crests; 5) vomerine odontophores not
visible except in large females, low,
oblique; 6) males with vocal slits and
subgular vocal sac; 7) first finger shorter
than second; pads large, that on thumb
smallest; discs broader than long; 8)
fingers bearing narrow lateral fringes; 9)
no ulnar tubercles except small ante-
brachial; 10) no tubercles on heel or
outer tarsus; 1-2 tubercles on inner edge
of tarsus; 11) two metatarsal tubercles,
inner oval, 5-6 times size of outer; nu-
merous supernumerary plantar tubercles;
12) toes bearing narrow lateral fringe,
no webbing; toe pads as large as those
of fingers; 13) above brown with darker
brown markings—interorbital bar, scap-
ular W, chevrons, facial and limb bars;
venter brown; posterior thigh rich
brown; 14) adults small, ten males 15.6—
18.8 (x = 17.5) mm, seven females 23.7-
29:9) (x = 24.5) mm SVL

Eleutherodactylus incomptus is ap-
parently related to the lowland E. car-
valhoi Lutz, E. croceoinguinis Lynch,
and E. martiae Lynch but differs from
each in be’ng larger and not having the
tympanum concealed. We are not aware
of a comparable species on the trans-
Andean slopes.

Description—Head as wide as body
in males, as wide as to narrower than in
females; head wider than long; head
width 35.2-38.4 (= 3705 Ni 10) per-
cent SVL in males, 38.0-40.1 (x = 38.7,
N = 7) in females; snout short, E-N
66.7-80.0 (x = 74.0, N = 10) percent
eye length in males, 75.0-90.0 (x = 82.8,
N = 7) in females; nostrils weakly pro-
tuberant in males, not protuberant in
females directed laterally; loreal region

weakly concave, sloping abruptly to lips;
lips not flared; interorbital space flat,
slightly broader than IOD; upper eyelid
width 831-1167 (x = 97.1.iNe— a)
percent IOD; supratympanic fold thick,
obscuring upper edge of tympanum;
tympanum distinct, round to slightly
higher than long, its length 25.0-31.8
(x = 27.9, N = 10) percent eye length
in males, 26.4-40.9 (x = 33.4, N = 7)
in females, separated from eye by dis-
tance equal tympanic diameter; temporal
region nearly vertical; no enlarged tu-
bercles on head except for some small
postrictal tubercles; choanae moderate-
sized, round, well anteriad on palate, not
concealed by palatal shelf of maxillary
arch; vomerine odontophores not visible

‘in males and small females, in adult

females posteromedial to choanae, low,
oblique, each a little smaller than a
choana, bearing 2 teeth, separated me-
dially by one odontophore width, tongue
slightly longer than wide, posterior edge
notched, posterior one-half of tongue not
adherent to floor of mouth.

Skin of dorsum bearing low, flat
warts, those toward posterior half of
body more prominent than those on an-
terior half of body; no dorsolateral folds
but indefinite, thin, fold may be present
along upper flank; discoidal folds promi-
nent; anal opening not enclosed in
sheath; forearm lacking ulnar tubercles
except for antebrachial tubercle; palmar
tubercle bifid, smaller than oval thenar
tubercle; numerous supernumerary pal-
mar tubercles, none as large as subartic-
ular tubercles; subarticular tubercles
round, elevated, non-conical; fingers
bearing narrow lateral fringes; all fingers
bearing broad discs on broadly dilated
pads, that of thumb more or less round,
others broader than long; pads rounded
apically.

No conical tubercles on heel, knee,
or outer edge of tarsus; inner edge of
tarsus bearing 1-2 elongate tubercles
just proximal to inner metatarsal tuber-
cle; inner metatarsal tubercle oval, non-
compressed, its length 1.5 times its
width, 5-6 times size of subconical outer

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 37

metatarsal tubercle; numerous plantar
supernumerary tubercles, arranged in
rows along metatarsals; subarticular tu-
bercles round, elevated, non-conical to
subconical; toes bearing narrow lateral
fringes but no basal webbing; all toes
bearing broad discs on pads, pads broad-
er than long, those on toes III-V larger
than those of I and II, pads rounded
apically; heels of flexed hind legs touch;
heel of adpressed hind leg reaches an-
terior edge of eye; shank 47.3-52.0 (x =
49.7, N = 10) percent SVL in males,
46.7-49.6 (x = 48.1, N = 7) in females.

Color in life—Dorsum tan (olive-
tan, reddish tan or brown in some)
with darker brown markings; venter gray
in most, dull brown or dirty cream with
brown flecks in some individuals; iris
dull bronze to gray with median hori-
zontal reddish brown streak. One indi-
vidual had a white snout.

Measurements of holotype in mm.—
SVL 18.8; shank 9.1; head width 7.2;
head length 7.1; upper eyelid width 2.4;
IOD 2.1; tympanum length 0.8; eye
length 3.0; E-N 2.0.

Etymology.—Latin, meaning una-
dorned; in reference to the drab pattern
and lack of distinctive features.

Distribution and _ ecology. — This
small inconspicuous frog occurs at ele-
vations of 1410-1910 m on the eastern
face of the Andes in Provincia Napo,
Ecuador; it also is known from 1270 m
in the Pastaza Valley, Provincia Pastaza,
Ecuador. All specimens have been
found in cloud forest areas, where they
are most frequently encountered at
night on leaves of herbs or low bushes
in clearings or at the forest edge. One
was on the forest floor by day.

Eleutherodactylus inusitatus new species
Fig. 3G

Holotype-—KU 166066, an adult fe-
male, taken at the Rio Jatuntinahua, 10
km SE Cuyujta, Provincia Napo, Ecua-
dor, 2160 m, on 21 March 1975 by the
incomparable John E. Simmons.

Paratypes——CAS-SU 8279, SE of Ca-

yambe, Agua Caliente, on Rio San Pe-
dro, Provincia Napo, Ecuador; KU
120099, Abitagua, ~ 8 km (airline)
NW Mera, Provincia Pastaza, Ecuador,
1300 m.

Diagnosis —1) skin of dorsum sha-
greened, that of venter coarsely areolate;
2) tympanum prominent, its length %
that of eye; 3) snout subacuminate in
dorsal view, protruding in lateral pro-
file, tip pointed; canthus rostralis sharp;
4) upper eyelid slightly narrower than
IOD, bearing a conical tubercle; no
cranial crests; 5) vomerine odontophores
oval in outline; 6) males not known; 7)
first finger shorter than second; fingers
II-IV bearing large pads; discs broader
than long; 8) fingers bearing lateral
fringes; 9) conical ulnar warts present;
10) knee, heel, and outer edge of tarsus
bearing conical tubercles; 11) two meta-
tarsal tubercles, inner oval, 6-7 times
size of conical outer; numerous super-
numerary plantar tubercles; 12) toes
bearing lateral fringes, not webbed; toe
pads as large as those of outer fingers;
13) creamy white with faint brown can-
thal mark; green above, white below in
life; 14) adults small, two adult females
24.0-24.4 mm SVL.

Superficially, E. inusitatus resembles
E. galdi (Jiménez de la Espada), but
the latter has large warts interspersed on
the areolations of the venter, cranial
crests; it is nearly twice as large as E.
inusitatus and has a truncate snout (in
lateral profile). Ignoring the prominent
conical tubercles on the eyelid, forearm,
heel, and tarsus, E. inusitatus resembles
the slightly larger E. acuminatus Shreve
of the upper Amazon Basin in Colombia,
Ecuador, and Peru.

Description. — (proportions of two
adults only): Head broader than body,
broader than long; head width 39.6-41.4
percent SVL; E-N 93.5-96.8 percent eye
length; nostrils not protuberant, directed
laterally; loreal region concave, sloping
abruptly to lips; lips not flared; upper
eyelid width 85.2-96.2 percent IOD,
bearing numerous subconical tubercles
and one large, conical tubercle; no cra-

38 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

nial crests, interorbital space flat; tem-
poral region sloping; supratympanic fold
thick, tuberculate; tympanum prominent,
higher than long, separated from eye by
1.5 times its length; tympanum length
29.0-32.3 percent eye length; skin above,
below, and behind tympanum bearing
con‘cal warts; choanae moderate-sized,
round, not concealed by palatal shelf of
maxillary arch; vomerine odontophores
prom‘nent, median and posterior to cho-
anae, oval in outline, separated by dis-
tance slightly less than an odontophore
width, each larger than a choana, bear-
ing 2-4 teeth in a clump; tongue longer
than wide, posterior edge feebly notched,
posterior one-fourth not adherent to floor
of mouth.

Skin of dorsum shagreened (smooth
in CAS-SU 8279), bearing large warts
laterally; flanks and limbs coarsely
warty; no dorsolateral folds; discoidal
folds prom/nent, well anteriad to groin;
pair of conical subanal warts ventrolat-
eral to anus; no anal sheath; three elon-
gate, conical warts proximal to subconi-
cal antebrachial tubercle, largest on
elbow; outer edge of palm bearing sub-
conical warts; thenar tubercle oval,
smaller than large round (partially di-
vided) palmar tubercle (palmar 4-5
times size of thenar); numerous small
supernumerary palmar tubercles; sub-
articular tubercles larger than palmar
supernumeraries, broader than _ long;
fingers bearing prominent lateral fringes;
all fingers bearing discs (broader than
long) on dilated pads; pads on II-IV
very large, that on thumb small; pads
apically rounded.

One or two conical tubercles on knee,
one elongate tubercle (calcar) on heel;
2-3 smaller, conical tubercles on outer
edge of tarsus; inner metatarsal tubercle
oval (length 1.5 times width), much
larger than conical outer metatarsal tu-
bercle; many supernumerary plantar tu-
bercles; subarticular tubercles round,
smaller than those of fingers; toes bear-
ing lateral fringes, broad discs on large
pads (pads of toes as large as those of
fingers); heels of flexed hind legs over-

lap; shank 50.8-53.7 percent SVL.

In preservative, cream-white with
thin brown canthal stripe, vague anal
triangle; underside of tarsus and foot,
ventral side of wrist peppered with
brown.

Color in life—Dorsum green with
faint brown bars on limbs; venter white.
One specimen (KU 120099, a juvenile
female) had a narrow yellow labial
stripe and pale lemon yellow groin, low-
er flanks, and hidden surfaces of hind
limbs. In that specimen the iris was
pale yellowish gray, whereas in the holo-
type, it was reddish brown.

Measurements of holotype in mm.—
SVL 24.0; shank 12.2; head width 9.5;
head length 9.0; upper eyelid width 2.5;
IOD 2.6; tympanum length 1.0; eye
length 3.1; E-N 3.0 The holotype is
gravid.

Etymology.—Latin, in- (negative of )
+ usitatus (usual or common), meaning
rare or uncommon.

Distribution and ecology—tThis dis-
tinctive species is known from only three
specimens from three localities: Abi-
tagua, Provincia Pastaza, Rio Jatuntina-
hua, Provincia Napo, and Agua Calienta,
on the Rio San Pedro, southeast of Cay-
ambe, Provincia Napo, Ecuador. The
first is in the Rio Pastaza valley at 1300
m at the point where the river emerges
from the Andean front. The Rio Jatun-
tinahua is a tributary at 2160 m to the
upper Rio Papallacta. The Agua Cali-
ente, on the Rio San Pedro, is presuma-
bly in the vicinity of the Rio Oyacachi.
One specimen was on a leaf of a low
herb along a small stream and another
was on the leaf of an elephant ear
(Xanthosoma); both frogs were in cloud
forest at night.

CAS-SU 8279 and KU 166066 are
gravid females. KU 120099 is a juvenile
or young female (slight convolutions of
oviduct, small eggs) 16.0 mm SVL.

Eleutherodactylus lacrimosus
(Jiménez de la Espada)

Cyclocephalus lacrimosus Jiménez de la Espada,

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 39

1875, Vert. Viaje al Pacifico . . . Batracios,
pl. 3, figs. 5, 5 a-b [Holotype apparently
lost, type-locality unknown].

Hylodes lacrimosus: Nieden, 1923, Das Tier-
reich 46:467.

Syrrhophus calceus: Lutz and Kloss, 1952,
Mem. Instit. Oswaldo Cruz, 50:647 (mis-
application ).

Eleutherodactylus lacrimosus: Gorham, 1966,
Das Tierreich, 85:80; Lynch and Schwartz,
1972, Journ. Herpetol. 5:109-12.

Eleutherodactylus chalceus: Lynch, 1968, Her-
petologica 24:292 (misapplication ).

Diagnosis.—1 ) skin of dorsal surfaces
very finely shagreened (or smooth), that
of venter areolate; no dorsolateral folds;
2) tympanum prominent, directed dorso-
laterally, round, its length 31-45 percent
eye length; 3) snout rounded in dorsal
view and lateral profile but bearing pa-
pilla at tip; canthus rostralis rounded;
4) upper eyelid narrower than IOD,
lacking tubercles; no cranial crests; 5)
vomerine odontophores prominent, oval,
obliquely oriented; 6) males with vocal
slits and subgular vocal sac; no nuptial
pads; 7) first finger shorter than second;

fingers long; pads narrow (largest on .

outer fingers), rounded apically; discs
broader than long; 8) no lateral fringe
or keel on fingers; 9) no ulnar tubercles
except for small antebrachial; 10) no
tubercles on tarsus or heel; 11) two
metatarsal tubercles, inner elongate, 5-6
times size of conical outer; few super-
numerary plantar tubercles; 12) toes
bearing lateral keels, not webbed; pads
small, outer pads as large as those on
outer fingers; 13) cream (rarely) to
golden brown (usually) above, with or
without small dark brown spots, some-
times having pale interorbital bar; ven-
ter creamy white with fine brown punc-
tations; limb bars sometimes vaguely
indicated; 14) adults small: in lowland
populations (below 800 m) males 16.1-
OOO e — Sia) ==" Ola, No = 1S) mm,
females 20.6-24.4 (x = 225 + 0.5,N =
18) mm SVL; in Amazonian slope frogs
(900-1100 m) males 19.6-23.7 (x = 21.0
+ 0.8, N = 11) mm, females 24.4-32.5
(Ce BY) Se SOS IN| =) vas VAC

Eleutherodactylus lacrimosus is most

similar to E. bromeliaceus Lynch and
E. petersi Lynch and Duellman but is
readily distinguished by lacking pungent
tubercles on the upper eyelid and heel;
the most distinctive features of E. lacri-
mosus include its round flat head and
coloration (golden-brown frog).

Description.—See Lynch and
Schwartz (1972).

Color in life—Pale golden-brown
above becoming pale yellow on flanks,
upper lip, digits, and venter; vocal sac
yellow; spotting (if present) medium
brown; iris red to bright reddish brown;
venter of juveniles and females creamy
white.

Distribution and ecology.—E. lacri-
mosus ranges from northeastern Brasil
west to Ecuador and south into northern
Peru. In east-central Ecuador it extends
to at least 1100 m (Mera, Provincia Pas-
taza); all other locality records are low-
land sites. Specimens are frequently
encountered in arboreal bromeliads by
day; at night frogs are found on vege-
tation 1-2 m above the ground.

Remarks.—Lutz and Kloss’ (1952)
record of Syrrhophus chalceus from
Iuareté, Estado Amazonas, Brasil, ap-
pears to be this frog. Specimens of E.
lacrimosus from northwestern Provincia
Pastaza, Ecuador, are significantly larger
than those found in the Amazonian low-
lands. The large examples of E. lacri-
mosus come from the only area of sym-
patry between E. lacrimosus and _ its
smaller upland relative, E. petersi.

Eleutherodactylus lanthanites Lynch
Fig. 3H
Eleutherodactylus lanthanites Lynch, 1975,
Contrib. Sci., Nat. Hist. Mus. Los Angeles
Co., (272):10 [Holotype—KU 146144,

Santa Cecilia, Provincia Napo, Ecuador,
340 m].

Diagnosis—1) skin of dorsum finely
tuberculate with scattered larger warts,
no dorsolateral folds, that of venter
smooth; 2) tympanum prominent, its
length %-% eye length; 3) snout acu-
minate in dorsal view, rounded in lateral

40 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

profile; canthus rostralis sharp; 4) upper
eyel'd narrower than IOD, lacking elon-
gate tubercles, no cranial crests; 5) vo-
merine odontophores prominent, oblique
to subtriangular in outline; 6) males with
vocal sac and slits, non-spinous nuptial
pad on thumb; 7) first finger longer than
second; digital pads large, non-emargi-
nate; 8) fingers lack lateral fringes; 9)
no ulnar tubercles; 10) conical tubercle
on heel; row of indistinct tubercles along
outer edge of tarsus; 11) two metatarsal
tubercles, inner elongate, 5-6 times size
of conical outer; few supernumerary
plantar tubercles; 12) toes lack lateral
fringes; toe pads as large as those of
outer fingers; 13) brown above with
darker brown markings; shank bars per-
pendicular to limb axis; posterior sur-
faces of thighs uniform brown; throat
heavily spotted and marbled with
brown, defining median white stripe;
14) adults moderate-sized, in lowland
populations (below 800 m) males 21.7-
26.0 mm, females 27.5-44.8 mm SVL; in
Amazonian slope frogs (900-1490 m)
males 2o:9—20.91( xe" 2610-== Ikon Ni)
mm, females 38.7-45.4 (x = 41.2, N =
4) mm SVL.

Eleutherodactylus fitzingeri (Schmidt)
and E. gutturalis Hoogmoed, Lynch and
Lescure have similar gular patterns but
lack the calcar on the heel and have
longer legs; E. fitzingeri also has pro-
nounced webbing of the toes.

Description.—See Lynch (1975b).

Color in life——(Andean slope speci-
mens): Dorsum tan (pinkish-, olive-) to
brown with or without dark brown, red-
dish brown, olive-green, or dull red
markings; in some _ individuals, lips
barred creamy tan and black; canthal
and supratympanic stripes black; pos-
terior groin and anterior and posterior
surfaces of thighs salmon to grayish
brown; venter creamy white to creamy
yellow with pinkish rose tint posteriorly
in some individuals, mottled with gray
in some individuals; bright yellow spots
on posterior flanks in some individuals;
iris bronze with median horizontal streak
to copper.

Distribution and ecology. — Eleu-
therodactylus lanthanites is an abundant
frog in the lowland rainforest in Ecua-
dor and adjacent Colombia. It ascends
the Andes in lower humid montane for-
est to 1440 m in southern Colombia and
to 1490 m in Ecuador. In the lowlands
the frogs occur on the forest floor by day
and on low vegetation at night. In the
Andes all specimens were on bushes or
low trees (to 2m) at night.

The small sample of E. lanthanites
from the cloud forest suggests that slope
frogs are significantly larger than low-
land frogs.

Eleutherodactylus leoni Lynch
Fig. 4A
Eleutherodactylus leoni Lynch, 1976, Herpeto-
logica 32:313 [Holotype—KU 130870,
taken on the N slope of the Nudo de Mo-

janda, Provincia Imbabura, Ecuador, 3400
m].

Diagnosis.—1) skin of dorsum sha-
greened, with many short ridges, scat-
tered larger tubercles, no dorsolateral
folds, that of venter areolate; 2) tympa-
num partially concealed beneath skin, its
length 4-% eye length; 3) snout sub-
acuminate in dorsal view, angularly
rounded (nearly truncate) in lateral pro-
file; canthus rostralis moderately sharp;
4) upper eyelid narrower than IOD,
bearing 2-4 conical tubercles; no cranial
crests; 5) vomerine odontophores low,
oblique; 6) males with vocal slits, sub-
gular vocal sac; no nuptial pads; 7) first
finger shorter than second; pads and
discs on fingers II-IV, pads round; 8)
fingers lack lateral fringes; 9) small ul-
nar tubercles present; 10) no enlarged
heel tubercles; outer tarsal tubercles con-
ical; ridge on inner edge of tarsus; 11)
two metatarsal tubercles, inner elongate,
twice size of round outer; supernumer-
ary plantar tubercles small; 12) toes
bearing narrow lateral keels, no web;
13) brown above with darker markings;
limb bars narrow, oblique; venter gray
to brown with cream spots; white and/or
colorless areas in axilla, groin, and con-

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 41

cealed surfaces of thigh and shank sal-
mon or bright red in life; 14) adults
small, males 13.0-18.3 mm, females 19.7—
25.0 mm SVL.

Eleutherodactylus leoni is most simi-
lar to E. pyrrhomerus Lynch but differs
in having large choanae, smaller vomer-
ine odontophores, more than one conical
tubercle on the upper eyelid, less conical
postrictal tubercles, and broader digital
pads.

Description —See Lynch (1976a).

Color in life——Dorsum brown to red-
dish brown with dark gray, black, tan,
and cream markings; lips barred cream
and brown; venter gray with white flecks
and/or small yellow spots; throat with
yellowish wash; groin and lower flanks
fleshy gray to salmon with orange yellow
spots; hidden surfaces of thighs and
groin gray to salmon in smaller individ-
uals, red in large females; iris powder
blue (with or without green tint), with
black flecks and brown horizontal streak.

Distribution and ecology.—This frog
is a small terrestrial inhabitant of upper
humid montane forest at elevations of
2710-3400 m on the Pacific slopes of the
Andes in Ecuador and 2540-2700 m on
the Amazonian slopes in Colombia and
Ecuador. On the Amazon slopes of Ecua-
dor, E. leoni is known only from the
upper Rio Chingual drainage. All speci-
mens from the Amazon slopes in Ecua-
dor were found beneath logs and rocks
in clearings in cloud forest.

Eleutherodactylus leucopus Lynch

Eleutherodactylus leucopus Lynch, 1976, Proc.
Biol. Soc. Washington 88:351 [Holotype.—
USNM_ 197927, Santa Barbara, Provincia
Napo, Ecuador, 2590 m].

Diagnosis.—1) skin of dorsum feebly
granulate, no dorsolateral folds, that of
venter coarsely areolate; 2) tympanum
prominent, its length 4—% eye length;
3) snout subacuminate in dorsal view
(tip pointed), protruding in lateral pro-
file; canthus rostralis sharp; 4) upper
eyelid slightly narrower than IOD, lack-
ing tubercles; low cranial crests present;

5) vomerine odontophores prominent,
oval to triangular in outline; 6) males
with vocal slits, subgular vocal sac; no
nuptial pads; 7) first finger shorter than
second; all fingers with broad pads; 8)
fingers lack lateral fringes; 9) small ulnar
tubercles; 10) heel and tarsus bearing
small non-conical tubercles; inner tarsal
fold ridge-like; 11) two metatarsal tu-
bercles, inner oval, outer non-conical,
4-4 size of inner; plantar surface
areolate; 12) toes bearing prominent lat-
eral fringes, no web; 13) dark gray
above, venter cream flecked with gray;
fingers and toes pale cream; posterior
thighs, groin dark gray; 14) adults mod-
erate-sized, males 30.0-37.8 mm, females
42.3-44.0 mm SVL.

Eleutherodactylus leucopus is most
similar to E. acerus Lynch and Duell-
man (described above); E. leucopus dif-
fers from E. acerus in having granular
skin (not smooth), inner tarsal tubercles
(none), and cream inner digits (brown).

DescriptionSee Lynch (1976c).

Distribution and ecology.—E. leuco-
pus is known from the valley of the Rio
Chingual, at elevations between 2440
and 2700 m in Ecuador and Colombia.
The late James A. Peters and his asso-
ciates found the frogs beneath logs in
clearings “in good forest.” At night,
specimens were found on leaves and
branches in heavy undergrowth (? sec-
ondary forest) in upper montane cloud
forest. In July 1977, Lynch collected at
El Carmelo and Santa Barbara, as well
as above El Carmelo. The forests men-
tioned in Peter’s field notes were largely
cleared; no specimens of E. leucopus
were found, although several other spe-
cies of Eleuterodactylus were encoun-
tered.

Eleutherodactylus lividus new species
Fig. 4B

Holotype.—KU 166005, an adult fe-
male, taken 11 km ESE Papallacta, Pro-
vincia Napo, Ecuador, 2660 m, on 22
March 1975 by William E. Duellman
and Linda Trueb.

42 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Paratypes—KU 155450, 166006-07,
topotypes; KU 165992, 165004, 12 km
ESE Papallacta, 2630 m.

Diagnosis.—1) skin of dorsum finely
areolate, that of venter coarsely areolate;
2) tympanum prominent, its length %-
% eye length; 3) snout rounded to sub-
acuminate in dorsal view, rounded in
lateral profile, bearing papilla at tip;
canthus rostralis sharp; 4) upper eyelid
narrower than IOD, bearing one conical
tubercle; low cranial crests present;
5) vomerine odontophores prominent,
rounded to slightly oblique; 6) males
lacking vocal sac and slits; 7) first finger
shorter than second; fingers [I-IV bear-
ing dilated pads; discs broader than
long; 8) fingers bearing lateral keels; 9)
ulnar fold bearing subconical tubercles;
10) heel and tarsus bearing pungent,
subconical tubercles; 11) two metatarsal
tubercles, inner oval, 6-8 times size of
outer; numerous supernumerary plantar
tubercles; 12) toes bearing lateral
fringes, not webbed; toe pads as large
as those of outer fingers; 13) gray above
with brown suffusion on face, sacrum
and lower limbs, dorsum flecked with
black; venter dull cream flecked with
brown; posterior thigh black with dull
yellow reticulation; 14) adults moderate-
sized, two males 18.2-20.4 mm, three
females 32.9-35.0 mm SVL.

Eleutherodactylus lividus is similar
to E. acerus Lynch and Duellman (de-
scribed above) but differs in having con-
ical tubercles on the eyelid, heel, and
tarsus, and more coarse warts on the
flanks. In addition, E. lividus is smaller
than E. acerus (three males 25.0-34.0
mm, one adult female 45.1 mm SVL)
and males lack vocal sac and slits.

DescriptionHead narrower than
body, wider than long; head width 37.2-
37.4 percent SVL in two males, 37.7—
40.3 (x = 38.6, N = 4) in females; nos-
trils not protuberant, directed dorsolat-
erally; E-N 80.8-97.4 (x = 87.1, N = 6)
percent eye length; loreal region of can-
thus rostralis concave, sloping abruptly
to lips; lips not flared; low cranial crests
(palpable); upper eyelid bearing one

conical tubercle, its width 70.8-96.8 (x
= 84.4, N = 6) percent IOD; tympa-
num directed dorsolaterally, prominent,
its upper edge concealed by thick supra-
tympanic fold, separated from eye by
tympanum length; tympanum round, its
length 29.8-41.9 (x = 34.9, N = 6) per-
cent eye length; several small, round,
subconical postrictal tubercles; choanae
small, round, not concealed by palatal
shelf of maxillary arch; vomerine odon-
tophores median and posterior to cho-
anae, each the size of a choanae, round
or somewhat oblique, elevated, bearing
a clump of 2-4 teeth, separated medially
by distance equal an odontophore width;
tongue broader than long, its posterior
edge notched, posterior two-fifths not
adherent to floor of mouth.

Skin of dorsum finely areolate (with-
out ridges) becoming more coarsely
areolate on flanks and venter; discoidal
folds not prominent; no anal sheath or
enlarged tubercles; limbs areolate except
posterior surface of thigh and underside
of shank; ulnar fold present, bearing
subconical tubercles, antebrachial larg-
est; palmar tubercle bifid, larger than
oval thenar tubercle; numerous super-
numerary palmar tubercles, all low; sub-
articular tubercles round (except for
basal on IV—longer than wide), non-
conical, elevated; fingers bearing ill-
defined lateral fringes; all fingers with
discs (broader than long) on dilated,
apically round pads; pad on thumb
scarcely wider than digit, those on I-IV
twice as wide as digit.

Heel bearing 2-3 subconical tuber-
cles, 2-3 subconical tubercles on outer
edge of tarsus; inner tarsal fold a thick-
ened ridge with a low tubercle; inner
metatarsal tubercle oval (length twice
width), not compressed, 6-8 times size
of round, subconical outer; numerous
supernumerary plantar tubercles, none
prominent except for conical tubercles
at bases of toes II-IV; subarticular tuber-
cles round, pungent, larger than any
supernumerary tubercle; toe fringes pres-
ent, more definite than those of hand,
no webbing; toes with discs (broader

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 43

than long) on dilated pads (at least
twice width of digit below pad), round-
ed apically; heels of flexed hind limbs
overlap; heel of adpressed hind leg
reaches eye; shank 47.2-52.8 (x = 49.4,
N = 6) percent SVL.

In preservative, gray above with
brown interorbital bar, some brown suf-
fusion on face, sacrum, and lower limbs;
dorsum flecked with black; venter dull
cream with brown-black flecks laterally;
underside of shank cream with black
spots; posterior thigh surfaces black
with dull yellow reticulation ventrally;
flank and anterior surfaces of thighs dark
gray with dull yellow reticulation break-
ing up into small dark spots; labial bar,
canthal-supratympanic stripe ill-defined.

Color in life——Dorsum brown (pale
green in one individual); venter cream
with brown or black flecks or pale gray;
blue or grayish blue in groin and, in
some individuals, on posterior surfaces
of thighs; iris bronze.

Measurements of holotype in mm.—
SVL 35.0; shank 16.6; head width 13.2;
head length 11.8; upper eyelid width 3.8;
IOD 4.2; tympanum length 1.8; eye
length 4.3; E-N 3.8. The holotype is
gravid.

Etymology.—Latin, meaning blue or
bluish, in reference to the concealed
colors in the groin.

Distribution and ecology.—This spe-
cies is known only from the Papallacta
Valley on the eastern face of the Andes
in Provincia Napo, Ecuador, where it
occurs at elevations of 2135-2750 m. All
individuals were found in cloud forests;
one was under a rock by day, and the
others were on the leaves of herbs and
low bushes at night.

Remarks.—Although most similar to,
and readily confused with E. acerus
(unless compared), E. lividus seems to
be.equally related to E. glandulosus and
E. leucopus. Eleutherodactylus lividus
is sympatric with E. acerus and E. glan-
dulosus; all three are allopatric to E.
leucopus. The four form a cluster of
species distributed in cloud forest on the
eastern face of the Andes in northern

Ecuador and have no other apparent
close relatives, although E. percultus
Lynch from the Abra de Zamora in
southern Ecuador may be related.

Eleutherodactylus nigrogriseus
(Andersson)

Fig. 4C

Pseudohyla nigrogrisea Andersson, 1945, Arkiv
fiir Zoologi 37A(2):87 [Syntypes—NHRM
1905 (1 adult and 5 juveniles) collected at
Banos, Provincia Tungurahua, Ecuador].

Eleutherodactylus nigrogriseus: Lynch, 1969,
Bull. So. California Acad. Sci. 68:219-24.

Diagnosis.—1) skin of dorsum sha-
greened (with scattered tubercles, no
dorsolateral folds), that of venter coarse-
ly areolate; 2) tympanum prominent, its
length #-% eye length; 3) snout sub-
acuminate in dorsal view, rounded to
truncate in lateral profile; canthus ros-
tralis sharp; 4) upper eyelid lacking
tubercles, its width about equal IOD; no
cranial crests; 5) vomerine dentigerous
processes small, triangular in outline;
6) males lack vocal sac and slits; no
nuptial pad; 7) first finger shorter than
second; fingers II-IV bear broad, api-
cally rounded pads; 8) fingers with ill-
defined lateral fringes; 9) no ulnar tu-
bercles (except antebrachial); 10) heel
with one conical tubercle, none on tar-
sus; 11) two metatarsal tubercles, inner
oval, 4 times size of round outer; many
small supernumerary plantar tubercles;
12) toes bearing lateral fringes, no web;
toe pads as large as those of fingers; 13)
dorsum brown with darker mottling;
venter tan to gray mottled with brown;
posterior flank, groin, anterior and pos-
terior surfaces of thighs, and concealed
shank dark brown to black with large
white (yellow in life) spots; iris red in
life; 14) adults moderate-sized, males
19.3-26.0 (x = 22.7, N = 20) mm, two
females 28.5-29.4 mm SVL.

In life, E. nigrogriseus cannot be con-
fused with other species but in preserva-
tive, the bold yellow spots on the flank,
groin, and thighs are white; E. nigro-
griseus might be confused with E. eri-

44 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

phus Lynch and Duellman or E. spinosus
Lynch. Both have more warty skin and
prominent tubercles along the outer
edge of the tarsus.

Description—See Lynch (1969b).

Color in life—Dorsum tan, olive-
green, or nearly black, with black mark-
ings on limbs and back; groin and an-
terior and posterior surfaces of thighs
black with bright yellow spots (discrete
in adults, diffuse in juveniles); venter
tan to gray with cream or white flecks,
brown flecks in some; iris red.

Distribution and ecology. — Most
specimens of E. nigrogriseus are from
the Rio Pastaza Valley (1180-1800 m)
and slopes of the Andes in southern
Ecuador (Provincia Morona-Santiago,
2195-2835 m), but one individual is from
the Cordillera del -Dué (1150 m) in
northern Ecuador. In these cloud for-
ests the frogs are found on low vegeta-
tion and rocks, usually along streams,
at night.

Eleutherodactylus nigrovittatus
Andersson
Fig. 4D
Eleutherodactylus nigrovittatus Andersson,
1945, Arkiv fiir Zoologi 37A(2):33 [Holo-
type—NHRM (not examined), Abitagua,
Rio Pastaza, Provincia Tungurahua, Ecua-

dor].

Diagnosis.—1) skin of upper surfaces
finely shagreened with small tubercles
and folds (including incomplete dorso-
lateral folds), that of venter smooth;
2) tympanum distinct, its length % eye
length; 3) snout acuminate in dorsal
view, pointed (males) or rounded (fe-
males) in lateral profile, fleshy swelling
of lip in adult male; canthus rostralis
obtuse, concave; 4) upper eyelid nar-
rower than IOD, lacking enlarged tu-
bercles; no cranial crests; 5) vomerine
odontophores prominent, oval, extending
laterally to level of choanae; 6) males
lacking vocal sac, slits, and nuptial pads;
7) first finger longer than second; fingers
bearing elongate discs (as long as wide),
digits not expanded into pads; 8) fingers

lack lateral fringes; 9) no ulnar tuber-
cles; 10) minute tubercles on heel and
outer edge of tarsus; 11) two metatarsal
tubercles, inner oval, twice size of round
outer; no supernumerary plantar tuber-
cles; 12) toes lack lateral fringes, no
webbing; toes bearing expanded, api-
cally pointed pads; discs as long as wide;
13) brown above with darker brown
suprainguinal spots, anal triangle, under-
side tarsi and feet; faint canthal-supra-
tympanic stripe, labial, and limb bars;
venter cream; posterior surfaces of thigh
brown with small cream spots; 14)
adults small to moderate-sized: lowland
(< 600 m) males 13.3-19.4 (x = 17.0,

= 9) mm, females 19.7-22.1 (x =
20.9, N = 9) mm SVL; slope population
(900-1935 m) males 17.2-24.6 (x = 21.4,
N = 33) mm, females 25.0-30.5 (x =
27.4, N = 21) mm SVL.

Eleutherodactylus nigrovittatus is
most likely confused with specimens of
the leptodactylines Adenomera, small
Leptodactylus, and Vanzolinius. None of
these has pointed discs and pads on the
toes as well as lacking lateral fringes on
the digits, nor does any other species of
Eleutherodactylus in northern South
America.

Description See Andersson (1945).

Color in life —Dorsum gray, tan, red-
dish brown, or orange; inguinal spots
and anal triangle black; labial and limb
bars dark brown; venter gray or lemon
yellow with black and white mottling on
throat; orange suffusion in groin, iris dull
bronze heavily suffused with black.

Distribution and_ ecology. — Eleu-
therodactylus nigrovittatus occurs in the
upper Amazon Basin in Ecuador and
Pert; in Ecuador it ascends the Andes
in cloud forest to 1935 m: The frogs are
active on the forest floor by day; a few
have been found on low herbs at night.

Eleutherodactylus ockendeni
(Boulenger)
Hylodes ockendeni Boulenger, 1912, Ann. Mag.

Nat. Hist. (8)10:187 [Syntypes—BM
1907.5.7.19-21/RR 1947.2.16.88-90, La

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 45

Union, Rio Huacamayo, Carabaya, Depar-
tamento Puno, Perti, 600 m].

Hylodes hylaeformis Melin, 1941, Medd. Géote-
borgs Mus. Zool. Avdel. 88:48 [Holotype
(not examined), Roque, Departamento San
Martin, Peri]. Synonymy fide Lynch
(1980).

Syrrhophus calcaratus Andersson, 1945, Ark.
Zool. 37A:27 [Holotype—NHRM 1941, Rio
Cosanga near Archidona, Provincia Napo,
Ecuador, 800 m]. Synonymy fide Lynch
(1974).

Eleutherodactylus melini Bokermann, 1958,
Herpetologica 14:95 [replacement name for
Hylodes hylaeformis Melin, 1941, non Phyl-
lobates hylaeformis Cope, 1975].

Eleutherodactylus anderssoni Lynch, 1968,
Herpetologica 24:292 [replacement name
for Syrrhophus calcaratus Andersson, 1945,
non Hylodes calcaratus Boulenger, 1908].

Eleutherodactylus ockendeni is a
frog of lowland rainforests in the upper
Amazon Basin from southern Colombia
to southern Peri (Lynch, 1974, 1980).
Like E. croceoinguinis, E. ockendeni in-
vades very low cloud forests in the lower
Pastaza Trench. Unlike E. croceoinguin-
is, E. ockendeni is uncommon in low
cloud forests.

Eleutherodactylus pastazensis
Andersson

Eleutherodactylus pastazensis Andersson, 1945,
Arkiv for Zoologi 37A(2):37 [Syntypes.—
NHRM 1918, 2 individuals, the larger from
Yauguilla, the smaller from Mount Tungura
(= ? Tungurahua), 1840 m, Rio Pastaza,
Provincia Tungurahua, Ecuador, collected
in December 1937 by Wm. Clarke-Mac-
Intyre].

Diagnosis.—1) skin of upper surfaces
smooth with non-pungent warts on pos-
terior back, that of venter coarsely areo-
late; 2) tympanum distinct, its length
¥-% eye length; 3) snout subacumi-
nate in dorsal view, protruding or angu-
larly rounded in lateral profile, papilla
at tip; canthus rostralis obtuse; 4) upper
eyelid narrower than IOD, lacking pun-
gent tubercles; no cranial crests; 5) vo-
merine odontophores low, not prominent,
oblique; 6) males with vocal slits and
subgular vocal sac, non-spinous nuptial
pads on thumb; 7) first finger shorter
than second; fingers bearing large pads

(largest on III-IV); discs broader than
long; 8) fingers bearing indistinct lateral
fringes; 9) ulnar tubercles absent; 10)
heel and tarsus lacking pungent tuber-
cles; 11) two metatarsal tubercles, inner
oval, 4-5 times size of round outer; super-
numerary plantar tubercles numerous,
ill-defined; 12) toes bearing lateral
fringes, no webbing; toe pads smaller
than those of outer fingers; 13) brown
above with indefinite darker brown fleck-
ing, sometimes forming pattern of chev-
rons, interorbital bar, and limb bars;
venter cream or brown; concealed thigh
uniform pale brown or bearing indefinite
cream and brown marbling; 14) adults
small to moderate in size, one male 19.5
mm, two females 30.9-31.7 mm SVL.

Eleutherodactylus pastazensis is a
remarkably nondescript frog. We are
aware of only four examples, all collect-
ed 40 years ago, lacking color descrip-
tions in life or while freshly preserved.
In many respects, E. pastazensis resem-
bles E. unistrigatus but is readily dis-
tinguished in having a papilla at the tip
of the snout, smaller tympana, non-
spinous nuptial pad in males, and more
distinct fringes on the digits. Although
the color pattern of E. unistrigatus is
variable, the lack of pattern (or reduced
pattern) of E. pastazensis is beyond the
range of variation in E. wnistrigatus; the
nearly uniform brown posterior thigh
of E. pastazensis (except in CAS-SU
5084) is unlike the mottled concealed
thigh of E. unistrigatus (occasionally
unpigmented ).

Description —Head slightly narrow-
er than body; head broader than long;
head width 38.0-39.4 (x = 38.7) percent
SVL; canthus rostralis obtuse, relatively
straight; nostrils weakly protuberant, di-
rected dorsolaterally; loreal region weak-
ly concave, sloping abruptly to lips; lips
not flared; upper eyelid 73.0-87.5 (x =
81.4) percent IOD, bearing small, non-
pungent warts; interorbital space flat,
no cranial crests; supratympanic fold
present, not obscuring tympanic annulus;
tympanum distinct, round in outline, its
length 25.9 percent eye length in male,

46 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

31.6-46.2 in two females; tympana of
male (CAS-SU 5065) less distinct than
in three females; tympanum smaller than
pad of third finger, separated from eye
by tympanum diameter; choanae rela-
tively small, round, not concealed by
palatal shelf of maxillary arch when
viewed from directly above; vomerine
odontophores median and posterior to
choanae, oblique, low, bearing clump
or slanted row of 1-4 teeth; odontophores
w_dely separated medially; tongue long-
er than wide, posterior 2/5 not adherent
to floor of mouth.

Skin of dorsum and head smooth,
that on posterior back bearing scattered
or dense low, non-pungent warts; no
dorsolateral folds; discoidal folds promi-
nent posteriorly, well anteriad of groin;
no ulnar tubercles (even antebrachial
not apparent); palmar tubercle bifid,
slightly larger than oval thenar tubercle;
supernumerary palmar tubercles numer-
ous, small, non-pungent; subarticular
tubercles round, pungent, distal tuber-
cles on fingers III-IV least pungent;
fingers bearing lateral fringes (most dis-
tinct in CAS-SU 5084, somewhat desic-
cated); fingers bearing discs (broader
than long) on dilated pads, pads of III
and IV largest (at least twice digit
width), that on thumb smallest (II in-
termediate in size); pads apically
rounded.

Heel and tarsus lacking tubercles;
inner metatarsal tubercle not com-
pressed, length twice width; outer meta-
tarsal tubercle round i-% size of
inner; plantar surface bearing indistinct
supernumerary tubercles arranged in
rows; subarticular tubercles round, pun-
gent (distal tubercles much less so),
smaller than subarticular tubercles of
fingers; toes bearing lateral fringes, not
webbed; toe pads like those of fingers
but smaller than those of fingers III-IV;
hind limbs short, shank 46.9-50.8 (x =
48.3) percent SVL.

In preservative, E. pastazensis is pale
brown to reddish brown above with
dark-brown or black flecking forming ill-

defined interorbital bar, chevrons, limb
bars, and diffuse marbling on upper
flanks. Canthal and supratympanic
stripes are present but not distinct. The
venter is cream, stippled (punctations )
with brown. The posterior flanks and
concealed surfaces of thighs and shanks
are heavily peppered with brown. CAS-
SU 5084 has some pale cream on the
upper edge of the posterior thigh; the
cream area is reticulated with reddish
brown markings extending from the top
of the thigh.

Measurements of larger syntype.—
SVL 30.9; shank 14.5; head width 12.0;
head length 9.8; upper eyelid 3.1; IOD
3.7; tympanum 1.2; eye length 3.8; E-N
3.3. The specimen is a gravid female.

Natural history——Both adult females
are gravid. CAS-SU 5084 was collected
in September 1939; NHRM 1918 (larger)
was collected in December 1937. The
smaller syntype is a juvenile female 23.3
mm SVL. The male (CAS-SU 5065)
has a distended vocal sac and nuptial
pads; the testes are relatively small and
cream colored.

Remarks.—The larger individual in-
cluded under NHRM 1918 was described
and illustrated by Andersson (1945).
This individual is here designated as the
lectoholotype.

The rarity of ‘E. pastazensis is inex-
plicable. All four known individuals
were collected by Wm. Clark-MacIntyre.
The University of Michigan Museum of
Zoology also purchased specimens from
him, but no specimens were found in a
survey of that collection. Most collectors
operating in Ecuador in the past 20 years
have neglected the Bafios area in the
Pastaza valley while opting for the richer
habitats at lower elevations or along
other routes into the Amazon Basin. Col-
lecting by individuals searching for frogs
may well reveal E. pastazensis to be
restricted to the upper Pastaza valley.

Distribution —Known only from the
upper Rio Pastaza valley in the vicinity
of Banos at elevations of 1800-1840 m.

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES AT

Eleutherodactylus peruvianus (Melin)
Fig. 4E

Hylodes peruvianus Melin, 1941, Medd. Gote-
borgs Mus. Zool. Avd. 88:43 [Holotype.—
NHMG 490, adult female, taken at Roque,
Departamento San Martin, Peri].

Eleutherodactylus peruvianus: Gorham, 1966,
Das Tierreich 85:91.

Eleutherodactylus conspicillatus (part): Lynch,
1975, Contrib. Sci., Nat. Hist. Mus. Los
Angeles Co. (272):3.

Lynch (1980) and Duellman and
Toft (1979) advocated recognition of a
lowland frog found in western Brasil,
southern Ecuador, and Pert, that is
similar to E. conspicillatus; they applied
the name E. peruvianus (Melin) which
Lynch (1975b) had placed in the sy-
nonymy of E. conspicillatus. Lynch
(1980) noted that the two putative
species are very similar but presumably
sympatric over central Peri. We have
assigned specimens found on the lower
Andean slopes in southern Colombia and
Ecuador, as well as from several ques-
tionable localities in lowland eastern
(Napo and Pastaza provinces) Ecuador
to E. peruvianus rather than describe
the frogs as a new species. Resolving
the specific status of these frogs will
require considerable field work in areas
of southern Ecuador and northern Peru;
assigning them to E. peruvianus is not
entirely satisfactory but seems prefer-
able to proposing another name at pres-
ent. The following account applies only
to what we refer to as “slope popula-
tions.”

Diagnosis.—( based on Andean slope
specimens): 1) skin of dorsum sha-
greened with scattered warts and inter-
rupted (albeit complete) dorsolateral
folds, that of venter smooth; 2) tympa-
num prominent, its length %-% eye
length; 3) snout rounded to subacumi-
nate in dorsal view, rounded in lateral
profile; canthus rostralis sharp; 4) upper
eyelid broader than IOD, not bearing
pungent warts; no cranial crests; 5) vo-
merine odontophores prominent, trian-
gular in outline; 6) males with vocal
slits and subgular vocal sac, white, non-

spinous nuptial pads on thumbs; 7) first
finger much longer than second; fingers
III-IV bearing small pads; discs broader
than long; 8) fingers bearing lateral
keels; 9) no ulnar tubercles; 10) no
tubercles on knee, heel, or tarsus aside
from slight tubercle on inner edge of
tarsus; 11) two metatarsal tubercles, in-
ner oval, 6 times size of outer; 2-3 super-
numerary plantar tubercles; 12) toes
bearing lateral keels, not webbed; toe
pads slightly larger than those of fingers
IlJ-IV; 13) dorsum brown with darker
chevrons, facial and limb bars; venter
cream with brown flecks, throat heavily
peppered with brown, bearing thin gular
stripe; posterior thigh brown with small
cream flecks to brown with large cream
spots; undersides of limbs cream dusted
with brown; 14) adults moderate-sized,
males 29.2-35.8 (Gs ——w Sl. Opa leae ee
10) mm, females 38.6-46.4 (x = 42.38 +
AS N= 10) mom SVE:

These frogs differ from E. conspicil-
latus (found at lower elevations) in
having moderate to heavy brown mark-
ings on the throat, the labial bars and
canthal-supratympanic stripes are evi-
dent through the masking on the face,
and in being slightly larger. They also
differ in having scattered enlarged tu-
bercles on the shagreened skin of the
dorsum.

Description. — (slope populations):
Head wider than body, wider than long;
head width 37.0-44.0 (x = 40.1, N =
20) percent SVL; E-N 86.1-108.3 (x =
96.2, N = 19) percent eye length; nos-
trils weakly protuberant, directed lat-
erally; loreal region weakly concave,
sloping abruptly to lips; lips very weakly
flared; upper eyelid bearing small tuber-
cles, none pungent; upper eyelid width
(QA 122:01(x— SONG 19) epercent
IOD; cranial crests not present but edges
of frontoparietals slightly elevated; tym-
panum round to slightly higher than
long, directed laterally with slight dorsal
vector, upper edge concealed by supra-
tympanic fold; tympanum length 38.8-
a9) (a ==. 47-2) Ni = 19)) percentweye
length, separated from eye by its length;

48 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

postrictal tubercles small, conical; cho-
anae moderate-sized, not concealed by
palatal shelf of maxillary arch; vomerine
odontophores medial and posterior to
choanae, elevated, triangular in outline,
separated medially by distance equal
one-half width of an odontophore, each
15 times size of a choana, bearing
slanted row of 8-9 teeth; tongue longer
than wide, posterior border notched, pos-
terior 45-4 not adherent to floor of mouth.

Skin of dorsum shagreened and bear-
ing slightly larger warts and interrupted
dorsolateral fold (series of warts from
eye to above groin); several small warts
behind eye (above supratympanic fold )
and on flanks; skin of limbs shagreened;
no supra-anal warts or anal sheath; dis-
coidal folds prominent; skin below and
lateral to vent bearing flat warts; no
ulnar tubercles; palmar tubercle bifid,
larger than oval thenar tubercle, super-
numerary palmar tubercles not pungent,
scarcely evident; subarticular tubercles
longer than wide, conical; fingers bear-
ing lateral keel; small pads, largest on
fingers III-IV, those of I-II scarcely ex-
panded; discs broader than long; pads
rounded apically.

No tubercles on heel or outer edge
of tarsus; inner edge of tarsus bearing
indistinct tubercle; inner metatarsal tu-
bercle oval, length twice width, not com-
pressed, 6 or more times size of subconi-
cal outer; 2-3 supernumerary plantar
tubercles (base of toes II, III, sometimes
IV), indistinct; subarticular tubercles
longer than wide, non-conical; toes bear-
ing lateral keels, no web; pads of toes
slightly larger than those of outer fingers
(pads of toes I-II about same size as
pads of fingers III-IV); hind legs long,
heels of flexed hind legs broadly over-
lap; heel of adpressed hind leg reaches
beyond snout tip; shank 50.9-69.8 (x =
63.1, N = 23) percent SVL.

In preservative, brown (pale to dark)
above with darker brown chevrons and
slanted flank bars, interorbital bar, can-
thal-supratympanic stripe, and_ labial
bars; lower flanks pale brown; limb bars
narrow (less than half width of inter-

spaces), oblique; anal triangle promi-
nent; posterior surfaces of thighs brown
with small cream flecks to relatively
large cream spots; throat heavily dusted
with brown and bearing narrow pale
gular stripe; breast and belly more
sparsely peppered with brown (appearing
cream in most individuals; throat paler
in many individuals as well); undersides
of limbs dusted with brown, underside
of shank bearing colorless spots.

Color in life-—(specimens from Rio
Azuela and Rio Salado, Provincia Napo,
Ecuador): Dorsum brown to reddish
brown with dark brown chevrons on
body and diagonal bars on limbs; an-
terior and posterior surfaces of thighs
dark brown with small orange-red spots;
belly white or creamy yellow with black
or brown flecks; throat, groin, and ven-
tral surfaces of thighs with reddish buff
tint; iris reddish-copper.

John E. Simmons’ specimens from
the Cordillera del Condor were as fol-
lows: dorsum tan or brown with dark
brown to black markings; post-tympanic
spot black; limbs barred with white,
venter mottled gray and tan or white.
Simmons’ brief and seemingly contrasting
notes probably stem from confusing E.
condor and E. peruvianus in the field
with the attendant assumption that the
smaller E. peruvianus were juveniles of
the larger frog that he found (E. con-
dor).

Distribution and ecology. — Eleu-
therodactylus peruvianus occurs in the
upper Amazon Basin in western Brasil,
southern Ecuador, and eastern Pert and
in cloud forests on the eastern slopes of
the Andes to 1910 m in Ecuador and
Peru. It is primarily a terrestrial species,
individuals being found in the forest
floor by day and night; a few have been
found on low vegetation at night.

Eleutherodactylus petersi new species
Fig. 4F
Holotype-—KU 143508, adult male,

taken 16.5 km NNE Santa Rosa, Provin-
cia Napo, Ecuador, 1700 m, 17 October

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 49

1971 by William E. Duellman.

Paratypes.—KU 143509-12, topotypes
collected 18-19 October 1971 by W. E.
Duellman and J. T. Collins.

Diagnosis.—1) skin of dorsum
smooth, no dorsolateral folds, that of
venter coarsely areolate, discoidal folds
present; conical tubercle on eyelid; 2)
tympanum prominent, round, its length
4 that of eye; 3) snout round in dorsal
view, truncate in lateral profile, bearing
papilla at tip; snout short; 4) interor-
bital space flat, broader than upper eye-
lid; 5) vomerine odontophores low,
oblique; 6) males with large, subgular
vocal sac and vocal slits; 7) first finger
shorter than second; all fingers long,
bearing discs on broadly dilated, round
apically, pads; 8) fingers bearing nar-
row lateral fringes; 9) no ulnar tuber-
cles; 10) no tubercles on knee, heel, or
outer edge of tarsus; inner tarsal fold
bearing 2-3 tubercles; 11) two metatarsal
tubercles, inner oval, 4 times size coni-
cal outer; plantar surface areolate; 12)
toes bearing narrow lateral fringes, discs
on broad pads; 13) pale cream with
brown interorbital bar, canthal and su-
pratympanic stripes; venter white; pos-
terior thigh pale brown; pale green in
life; 14) adults small, 35 males 14.5-
19.9 mm, eight females 20.3-23.1 mm
SVL.

Eleutherodactylus petersi is readily
distinguished from all other species in
the unistrigatus group on the basis of
its very short snout, conical tubercles
on the snout and upper eyelid (but none
on the heel), and its long, slender digits
bearing large pads. The frog is most
similar to, and related to, E. bromelia-
ceus Lynch and E. lacrimosus (Jiménez
de la Espada). Eleutherodactylus bro-
meliaceus has conical tubercles on the
heel and outer edge of the tarsus; E.
lacrimosus lacks tubercles on the upper
eyelid.

Description.—Head as broad as body,
wider than long; head width 35.6-42.5
(x = 37.6, N = 25) percent SVL; snout
very short, E-N 66.1-100.0 (x = 80.2, N
= 25) percent eye length; nostrils weak-

ly protuberant, directed dorsolaterally;
loreal region concave, sloping abruptly
to lips; lips not flared; interorbital space
flat, no cranial crests; upper eyelid width
67.7-100.0 (x = 89.9, N = 24) percent
IOD; upper eyelid bearing one conical
tubercle; supratympanic fold narrow,
concealing uppermost edge of tympa-
num; tympanum prominent, round, its
length’ 28:8-43.5) (x = 26,0), Ni= 7)
percent eye length in males, 30.4-46.2
(x = 40.2, N = 8) in females; tympa-
num separated from eye by 1.5 tym-
panic diameters; tongue longer than
wide, posterior border not notched, pos-
terior one-half not adherent to floor of
mouth; choanae relatively large, round
not concealed by palatal shelf of maxil-
lary arch when roof of mouth is viewed
from directly above; vomerine odonto-
phores posteromedial to choanae, low,
oblique, widely separated (by distance
equalling 1.5 choanal widths), bearing
2-3 teeth. Skin of dorsum smooth to
finely areolate; no dorsolateral folds; dis-
coidal folds obscure; anal opening not
extended by skin flap; no ulnar tubercles
or folds except for small antebrachial
tubercle; palmar tubercle bifid, much
larger than oval thenar tubercle; subar-
ticular tubercles small, round, non-coni-
cal, simple; fingers bearing narrow lat-
eral fringes, even along outer edge of
palm and finger IV; all digits bearing
discs on broadly dilated pads, pads
round at tip; outer pads larger than
inner pads; fingers long and slender.

No tubercles on knee or heel; outer
edge of tarsus lacking tubercles; inner
edge of tarsus bearing fold running
length of tarsus and bearing 2-3 small
tubercles; inner metatarsal tubercle 2-3
times as long as wide, non-compressed,
4 times size of subconical outer meta-
tarsal tubercle; plantar surface areolate;
subarticular tubercles round, elevated,
non-conical, simple; toes bearing narrow
lateral fringes; all toes bearing discs on
dilated pads; pads of toes slightly larger
than those of fingers; hind legs moder-
ately long, shank 48.4-57.6 (x = 52.7,
N = 25) percent SVL.

50 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Pale cream above with brown inter-
orbital bar, canthal and supratympanic
stripes; no facial bars; limbs not barred
or there are indefinite brown bands at
wrist and near knee; posterior surface of
thigh pale brown; belly creamy white.
KU 143510 has the face washed with
brown. KU 143512 has a pale spot an-
terior to eye.

Color in life—Dorsum usually pale
green to dull green with no distinct
markings; in some individuals dorsum
tan or brown with or without brown
markings; venter grayish or creamy yel-
low to bright yellow; vocal sac bright
yellow; iris reddish bronze to reddish
copper.

Measurements of holotype in mm.—
SVL 19.9; shank 10.2; head width 7.4;
head length 7.0; upper eyelid width 2.2;
IOD 2.4; tympanum length 1.0; eye
length 2.6; E-N 2.0.

Etymology—Named_ for the late
James A. Peters who pioneered the her-
petolog'cal exploration of the Amazonian
slopes and along whose trails we now
search.

Distribution and ecology.—This small
green Eleutherodactylus occurs in cloud
forests at elevations of 1410-1950 m on
the eastern face of the Andes in De-
partamento Putumayo, Colombia, and
Provincia Napo, Ecuador. Five individ-
uals referred to this species from San
Agustin, Departamento Huila, Colom-
bia, are from an elevation of 1750 m in
cloud forest on the eastern slope of the
Cordillera Central in the upper drainage
of the Rio Magdalena.

All individuals were found at night.
Most were on the leaves of herbs or
bushes within 1 m of the ground, but
two were on leaves and one on a twig
more than 1.5 m above the ground. Call-
ing males were found in March, May,
July, September, and October. The call
consists of a single “peep.”

The late James A. Peters recorded
the following notes on these frogs (notes
27 June 1962, specimens from 2 km SE
La Bonita, Provincia Napo): “4862-80
were taken in the base of elephant ear
leaves, where water collects. The lighter
areas on body and legs were a golden
or yellow in life; the toe and finger disks
were coppery. The dorsal dark areas
ranged from a chocolate brown to a light
brown. They react in one of two ways to
the stem being pulled down. Some hop
up the stem and dive into the underbrush,
while others scoot backwards down
deeply into the overlapping stems, into
the water, where their big eyes are visi-
ble. In life there is a distinct tho small
horn over each eye and one on the
snout.”

Remarks.—E. petersi is considered to
be a northern replacement for E. bro-
meliaceus and both are considered up-
land replacements for the lowland E.
lacrimosus. Both Amazonian slope spe-
cies have tubercles on the upper eyelid.
Based on limited data, E. petersi ex-
hibits pronounced geographic variation
in size (Table 4). The sample of males
from the upper Pastaza trench (Banos,
Rio Blanco, and Yungilla) at 1600-2000
m consists of frogs as large as E. brome-

TABLE 4.—Variation in body size in Eleutherodactylus petersi.

Locality (collection )

Calling Males Gravid Females*

La Bonita (JAP)
Rio Quijos (KU) and Loreto (JAP)
Mera (KU)

Bafios (JAP, UMMZ)

14.5-17.2( 16) 19.3-22.8(2)
16.1+0.4 21.0
16.7-19.9( 14) PA le PS ILS)
18.4 + 0.6 22.0
16.9-18.0(5) 20.3-21.9( 4)
17.3 + 0.4 21.0
O'0=23:5(610) eee
2lAs 09. 4) = See

1 Each sample of females includes young (nearly mature) individuals; this fact plus small sample sizes mean that

mean size estimates are probably low estimates.

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 51

liaceus (Table 2) or slope populations
of E. lacrimosus (Table 1) but consid-
erably larger than males of E. petersi
from the lower reaches of the Pastaza
trench (Mera, 1100 m), the vicinity of
the type-locality (1410-1700 m), or the
Rio Chingual at La Bonita (1920 m).
Although populations of E. lacrimosus,
E. lanthanites, E. nigrovittatus, and E.
peruvianus consist of larger frogs at
higher elevations (see Tables 1 and 3)
this explanation is unsuitable for E.
petersi. The frogs from the upper Pas-
taza trench may represent yet another
species of this complex; these frogs have
larger digital pads than are seen in other
specimens here assigned to E. petersi.

Eleutherodactylus prolatus new species
Fig. 4G

Holotype—KU 166008, an adult fe-
male, taken 2 km SSW Rio Reventador,
Provincia Napo, Ecuador, 1490 m, on
6 October 1974 by William E. Duellman
and John E. Simmons.

Paratypes——KU 166009-15, collected
at the type-locality.

Diagnosis.—1) skin of dorsum feebly
warty, bearing H-shaped ridges on occi-
put, that of venter areolate; 2) tympa-
num prominent, its length 4% eye
length; 3) snout acuminate in dorsal
view, nearly truncate in lateral profile;
canthus rostralis moderately sharp; head
longer than wide; 4) upper eyelid nar-
rower than IOD, bearing one pungent
tubercle; no cranial crests; 5) vomerine
odontophores prominent, oblique and
oval; 6) males with vocal slits and sub-
gular vocal sac; 7) first finger shorter
than second; fingers bearing moderately
large pads (II-IV); discs broader than
long; 8) fingers lacking lateral fringes;
9) no ulnar tubercles; 10) heel and tar-
sus bearing row of conical tubercles; 11)
two metatarsal tubercles, inner elongate,
8-10 times size of faint outer; no super-
numerary plantar tubercles; 12) toes
lacking lateral fringes and webbing; toe
pads as large as those of fingers; 13)
gray to brown above with cream H-

shaped occipital mark; venter cream
spotted with brown; throat streaked with
brown and creamy white; posterior
thigh stippled with brown; 14) adults
small, males 13.7-18.4 mm, females 20.8—
24.1 mm SVL.

Eleutherodactylus prolatus is super-
ficially similar to juvenile E. rubicundus,
especially in throat pattern, but is read-
ily distinguished in having the H-shaped
occipital mark, long head, and being
smaller. Eleutherodactylus prolatus
shares the long head and several other
minor features with E. ignicolor Lynch
and Duellman, E. trachyblepharis (Bou-
lenger), and E. variabilis Lynch. Eleu-
therodactylus prolatus has a more tuber-
culate skin than E. trachyblepharis or E.
variabilis. Eleutherodactylus ignicolor
has red flash colors in the groin (color-
less in preservative) and lacks the tu-
bercles on the heel.

Description—Head narrower than
body, longer than wide; head length
110.0-115.0 (x = 111.6, N = 7) percent
head width in males, 103.6-115.6 (x =
109.0, N = 12) percent in females; head
width) 33:6=39:6 (x =436.5, N) = 31)
percent SVL; E-N 70.4-91.3 (x = 80.8,
N = 18) percent eye length in males,
80.0-103.3 (x = 93.7, N = 10) percent
in females; tip of snout swollen and
elongated; nostrils protuberant, directed
laterally; loreal region concave, sloping
to lips; lips not flared; interorbital space
flat, no cranial crests; upper eyelid width
82.6-126.7 (x = 99.2, N = 26) percent
IOD; upper eyelid bearing one or two
conical tubercles; temporal region near-
ly vertical; supratympanic fold obscure
or absent; tympanum distinct, higher
than long, separated from eye by dis-
tance equal tympanum length, its length
24ST -0ii(xe=_ SIRSIING =903)5 percent
eye length in males, 28.6-44.4 (x = 35.2,
N = 10) percent in females; postrictal
tubercles conical as are those along mar-
gin of lower jaw; choanae small, round,
not concealed by palatal shelf of maxil-
lary arch; vomerine odontophores ob-
lique and oval, well elevated, median
and posterior to choanae, separated me-

52 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

dially by distance equal to choanal
width, each odontophore one-half size
of a choana, bearing a slanted row of
3-4 teeth; tongue slightly longer than
wide, posterior edge feebly notched, pos-
terior 4-% not adherent to floor of
mouth.

Skin of dorsum feebly warty, that of
venter coarsely areolate; no dorsolateral
folds; discoidal fold prominent, well an-
teriad to groin; H-shaped ridge on occi-
put; anal opening not extended; no
obvious ulnar tubercles; small antebra-
chial tubercle present; palmar tubercle
bifid, larger than oval thenar tubercle;
if present, supernumerary palmar tuber-
cles low, few in number; subarticular
tubercles round, low, simple; fingers
lacking lateral fringes or keels; all fin-
gers bearing broad discs; thumb lacking
digital pad, those of II-IV dilated; api-
cally round.

Tarsus bearing row of conical tuber-
cles along outer edge; inner edge of
tarsus lacking tubercles or bearing one
indistinct tubercle; one conical tubercle
on heel; inner metatarsal tubercle elon-
gate (length three times width), not
compressed, 8-10 times size of small,
round, low outer metatarsal tubercle;
plantar surface lacking supernumerary
tubercles; subarticular tubercles low, flat,
round; toes lacking lateral fringes of
keels and webbing; all toes bearing pads
(that of inner toe narrowest), with
broad (broader than long) discs, round-
ed apically; heels of flexed hind legs
overlap; heel of adpressed hind leg
reaches nostril; shank 48.1-60.1 (x =
50.4, N = 30) percent SVL.

In preservative, gray above with
brown blotches, sacral chevron, 2—3 bars
on flank, and cream H-shaped occipital
mark; brown canthal, supratympanic,
and labial markings; limb bars narrow,
oblique on shank; anal triangle brown,
diffuse; anterior and posterior surfaces
of thighs and concealed shank stippled
with brown; venter cream, spotted with
brown; anterior chin white bordered
posteriorly by brown bar and median
gular brown streak; lateral to gular stripe

are two slanted brown bars extending
onto lip.

Color in life——Dorsum in most indi-
viduals is tan to brown (reddish brown
or olive-tan in some) with dark brown
to black markings and orange occipital
ridges. Two individuals were green with
brown marks; another was gray anterior-
ly and posteriorly and pale green medi-
ally. Posterior thighs brown. A few indi-
viduals have yellow flecks dorsally and
a golden tint to upper lips. The venter
varies from dull cream with dense brown
flecking to gray, flecked or not with
black. Throat dull green to brown or
gray with darker streaks (cream streaks
in two). Iris gray-brown.

Measurements of holotype in mm.—
SVL 22.3; shank 12.6; head width 8.4;
head length 8.7; upper eyelid width 2.1;
IOD 2.3, tympanum length 1.0; eye
length 3.0; E-N 2.8. The holotype is a
gravid female.

Etymology.—Latin, meaning drawn
out, in reference to the apparent elonga-
tion of the head.

Distribution and ecology.—This spe-
cies is known only from lower cloud for-
ests (1140-1490 m) on the eastern face
of the Andes in Ecuador. Except for
one individual found on the forest floor
by day, all were on low vegetation at
night; most were in ravines near streams.

Eleutherodactylus proserpens Lynch

Eleutherodactylus proserpens Lynch, 1979,
Misc. Publ. Mus. Nat. Hist. Univ. Kansas
66:32 [Holotype—USNM 198484, from be-
tween Sapote and Suro Rancho, Provincia
Morona-Santiago, Ecuador, 2622 m].

Eleutherodactylus proserpens occurs
in cloud forests at elevations of 1707 to
2622 m on the Cordillera del Condor and
the Cordillera de Matanga in the Andes
of southern Ecuador (Lynch, 1979a).

Eleutherodactylus pugnax Lynch
Fig. 4H
Eleutherodactylus pugnax Lynch, 1973, Bull.

So. California Acad. Sci. 72:107 [Holotype.
—KU 146466, an adult female, taken at

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 53

Salto de Agua, 2.5 km NNE Rio Reventa-
dor, Provincia Napo, Ecuador, 1660 m].

Diagnosis.—1) skin of dorsum bear-
ing flattened, ill-defined warts, flanks
and venter areolate; no dorsolateral
folds; 2) tympanum absent; 3) snout
semicircular in dorsal view, truncate in
lateral profile; canthus rostralis round;
4) upper eyelid as wide as IOD, lacking
pungent tubercles; no cranial crests; 5)
vomerine odontophores round, promi-
nent; 6) males with vocal sac and slits,
thumb bearing non-spinous nuptial pad;
7) first finger shorter than second; fin-
gers bearing large pads (largest on H-
IV); discs broader than long; 8) fingers
bearing lateral fringes; 9) low ulnar
warts present; 10) no tubercles on outer
edge of tarsus or heel; inner tarsal fold
along distal one-third of tarsus; 11) two
metatarsal tubercles, inner oval, 5-6
times size of indistinct outer; supernu-
merary plantar tubercles indistinct, few
in number; 12) toes bearing -lateral
fringes and basal webbing; webbing
formula I 2—2 II 2—3+ TIT 2 1/2—4
IV 3 3/42 1/3 V; toe pads enlarged,
slightly smaller than pads of outer fin-
gers; 13) brown with darker markings
(interorbital bar, blotches on back, fa-
cial stripes and bars, transverse limb
bars); posterior surfaces of thighs brown
with cream flecks; venter cream with
brown flecks or spots; 14) adults mod-
erate-sized, two males 22.1—28.0 mm, two
females 30.8-33.2 mm SVL.

Eleutherodactylus pugnax is readily
distinguished from all other members of
the unistrigatus group in having toe web-
bing and lacking the ear. It is most
similar to an undescribed species found
at elevations between 1900 and 2700 m
on the Pacific Andean slopes of northern
Ecuador; that species has less prominent
toe webbing.

Description—See Lynch (1973b).

Color in life —Gray-brown, dull olive-
green, or yellow-tan above with brown
markings; throat cream with tan flecks;
venter gray with brown flecks or bluish-
white flecks; iris reddish brown, olive-

brown, or pale gold with diffuse red
streak.

Distribution and ecology.—The few
specimens of E. pugnax have been at
the edges of streams in cloud forest at
1660-2540 m on the eastern face of the
Andes in Provincia Napo, Ecuador. Two
were under rocks at the edge of a cas-
cading stream by day. One was sitting
on a rock in a stream at night and two
others were on leaves of herbs at night.

Eleutherodactylus pycnodermis Lynch

Eleutherodactylus pycnodermis Lynch, 1979,
Misc. Publ. Mus. Nat. Hist. Univ. Kansas
66:35 [Holotype—USNM 199754, San Vi-
cente, Provincia Morona-Santiago, Ecuador,
2805-2835 m].

Eleutherodactylus pycnodermis seems
to be primarily an inhabitant of paramo
and ranges down into meadows and
pastures in upper cloud forests. It is
known at elevations of 2652-3384 m on
the Cordillera de Matanga in southern
Ecuador (Lynch, 1979a).

Eleutherodactylus quaquaversus Lynch

Eleutherodactylus quaquaversus Lynch, 1974,
Occas. Pap. Mus. Nat. Hist. Univ. Kansas
(31):9 [Holotype—KU 123745, an adult
female from a series collected on the south
slope Cordillera del Dué above Rio Coca,
Provincia Napo, Ecuador, 1150 m].

Diagnosis—1) skin of dorsum sha-
greened, that of venter coarsely areolate;
no dorsolateral folds; 2) tympanum con-
cealed (or partially exposed); 3) snout
subacuminate in dorsal view, rounded or
weakly pointed in lateral profile; 4) up-
per eyelid bearing conical tubercle; up-
per eyelid width slightly narrower than
IOD; no cranial crests; 5) vomerine
odontophores triangular in outline; 6)
males with subgular vocal sac and vocal
slits; no nuptial pad; 7) first finger
shorter than second; all fingers bearing
pads and discs, largest on III-IV; 8)
fingers bearing narrow lateral keels; 9)
no ulnar tubercles except antebrachial;
10) tarsus bearing small tubercles on
inner and outer edge; one large, conical

54 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

tubercle on heel; 11) two metatarsal tu-
bercles, inner elongate, 4-5 times size of
round outer; few supernumerary plantar
tubercles; 12) toes bear narrow lateral
fringes, not webbed; toe pads as large
as those of outer fingers; 13) dorsum
pale brown to reddish brown marked
with brown chevrons, interorbital bar,
and spots; never with occipital W-
shaped mark; no canthal or supratym-
panic stripes; posterior surfaces of thighs
reddish, reticulated with brown (or pig-
mentless ); venter white, frequently spot-
ted with brown; 14) adults moderate-
sized, males 19.6-22.5 (x = 20.7, N =
29) mm, females 24.6-31.3 (x = 27.3, N
= "20)) mS V Le

Eleutherodactylus quaquaversus is
most similar to E. frater (Werner) and
E. ockendeni (Boulenger) but differs
from both in having the conical tuber-
cles on the eyelid and heel and in hav-
ing reticulations on the posterior surfaces
of the thighs (uniformly brown in E.
frater and E. ockendeni). The three spe-
cies are easily recognized in that each
lacks a canthal stripe.

Description—See Lynch (1974).

Color in life——Pale tan at night, be-
coming pale brown, olive-tan, or dark
brown by day; markings brown; in ju-
veniles, venter black anteriorly and yel-
low posteriorly (continuing onto under-
sides of legs); in adults, venter white to
cream-bronze with gray to brown spots;
lower flanks and groin white to pale pink
with black flecks or pink to purplish
red with white to yellow flecks; posterior
surfaces of thighs are pale rose to pur-
plish red and may bear white spots; iris
silvery gray to cream without a_hori-
zontal streak.

Distribution and ecology.—Although
E. quaquaversus is known from eleva-
tions as low as 320 m in the upper Ama-
zon Basin in Ecuador, it reaches its
greatest abundance in cloud forests be-
tween 1150 and 1830 m on the eastern
face of the Ecuadorian Andes. Nearly
all specimens were perched on low
bushes and herbs at night.

Eleutherodactylus rubicundus
(Jiménez de la Espada)
Fig. 5A

Hylodes rubicundus Jiménez de la Espada,
1875, Vert. Viaje al Pacifico . . . . Batracios,
plate 3, figs. 4, 4a—b [Holotype lost, type-
locality unknown].

Eleutherodactylus rubicundus: Gorham, 1966,
Das Tierreich, 85:99.

Diagnosis.—1) skin of dorsum coarse-
ly tuberculate bearing vague outline of
W-shaped occipital ridges, that of venter
finely areolate; 2) tympanum prominent,
its length % eye length; 3) snout sub-
acuminate to rounded in dorsal view,
rounded in lateral profile; canthus ros-
tralis moderately sharp, loreal region
sloping gradually to lips; 4) upper eye-
lid broader than IOD, not bearing pun-
gent tubercles; furrowed _interorbital
region in adult females; 5) vomerine
odontophores prominent, triangular in
outline; 6) males lacking vocal slits and
sac; 7) first finger slightly longer than
second; fingers long and slender, bearing
large, weakly emarginate pads (III-IV);
discs broader than long; 8) fingers lack-
ing lateral keels; 9) ulnar tubercles
small, subconical; 10) heel bearing elon-
gate, conical tubercle; conical tubercles
on outer edge of tarsus; 11) two meta-
tarsal tubercles, inner elongate, 4 times
size of subconical outer; several indis-
tinct supernumerary plantar tubercles;
12) toes bearing narrow lateral fringes,
no webbing; toe pads slightly smaller
than those of outer fingers; 13) brown
with darker brown markings above; ven-
ter cream blotched and marbled with
brown; posterior thighs rich brown with
cream flecks; 14) adults moderate-sized,
four adult males 32.0-35.6 mm, two adult
females 46.5-51.2 mm SVL.

Eleutherodactylus rubicundus is most
similar to and most closely related to E.
crenunguis Lynch of the Pacific Andean
versant of central Ecuador. The two are
readily distinguished in that E. crenun-
guis has a shagreened dorsum, less pun-
gent ulnar and tarsal tubercles, flared
lips, and bilobed subgular vocal sac.

Description.—Head as wide as body,

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 55

as long as wide; head width 33.9-39.7
(— 37.5, Ni =" 7) "percent SVL; snout
subacuminate in dorsal view in males
(rounded in females ), rounded in lateral
profile; snout long, E-N 94.5-105.9 (x =
100.8, N = 6) percent eye length; nos-
trils weakly protuberant, directed later-
ally; loreal region weakly concave, slop-
ing gradually to lips (slope steepest in
small individuals; lips not flared; eyes
large, upper eyelid width 110.0-140.0 (x
= 125.5, N = 5) percent IOD; inter-
orbital space narrow, flat in males, weak-
ly furrowed in adult females; temporal
region nearly vertical; supratympanic
fold tuberculate, not concealing tym-
panic annulus; tympanum prominent,
higher than long, its length 32.4—40.2
(x = 36.0, N = 6) percent eye length,
separated from eye by distance equal to
1.5 tympanum lengths; postrictal -tu-
bercles prominent; choanae moderate-
sized, round or slightly longer than wide,
not concealed by palatal shelf of maxil-
lary arch; vomerine odontophores me-
about size of a choana, separated medial-
dian and posterior to choanae, each
ly by distance equal to %-% odon-
tophore width, triangular in outline,
elevated, each bearing a weakly sloping
transverse row of 4-9 blunt teeth; tongue
large, a little longer than wide, posterior
border feebly notched, posterior two-
fifths adherent to floor of mouth.

Skin of dorsal surfaces of head and
body coarsely tuberculate; one large tu-
bercle on snout just anterior to eyes,
another between eyes; upper eyelid bear-
ing 1-2 large, non-conical tubercles;
vague outline of W-shaped ridges on
occiput; no other folds on dorsum; upper
surfaces of limbs less tuberculate than
dorsum; discoidal folds prominent, well
anteriad to groin; skin posterior and
posterolateral to vent coarsely areolate;
vent not extended in sheath; two promi-
nent subanal tubercles; antebrachial tu-
bercle largest in series of subconical ulnar
tubercles; palmar tubercle bifid, outer
lobe nearly isolated, smaller than oval
thenar tubercle; supernumerary palmar
tubercles prominent, those above outer

metacarpal forming an elongate tuber-
cle, all smaller than subarticular tuber-
cles which are round, simple, non-coni-
cal; no lateral fringes on fingers; all
digits bearing discs (wider than long)
on pads; dilation ratios of pads (pad
width/digit width below pad) [range
follows digit number, mean in paren-
theses] I 1.5-1.7 (1.6), II 2.1-2.3 (2.2),
III 3.2-3.6 (3.5), IV 3.4-3.5 (3.5); ungual
flaps indented, pads on fingers III-IV ap-
pear emarginate, slightly on J-II.

Heel tuberculate, all tubercles round,
some long, one large conical tubercle at
upper edge of heel; series of 2-4 conical
outer tarsal tubercles; 1-2 small inner
tarsal tubercles on _ ill-defined tarsal
ridge; outer metatarsal tubercle subconi-
cal, round to slightly longer than wide,
4% size of elongate (length three times
width), non-compressed inner metatarsal
tubercle; series of conical tubercles along
outer edge of foot; 1-2 round supernu-
merary plantar tubercles at bases of toes
II-V; several low, flat, additional super-
numerary plantar tubercles; subarticular
tubercles longer than wide, subconical
to conical, simple; toes bearing narrow
lateral fringes coalescing at bases of toes
into nearly indistinguishable basal web;
toes bearing discs (broader than long)
on dilated pads (smaller than those of
fingers III-IV ); ungual flaps not notched;
heels of flexed hind legs overlap broadly;
heel of adpressed hind leg reaches be-
yond snout tip; shank 54.8-61.9 (x =
57.8, N = 7) percent SVL.

In preservative, brown above with
darker brown markings—occipital W,
sacral chevron, postsacral bar, interor-
bital bar, canthal-supratympanic stripe,
labial bars, limb bars; flanks less dark
than dorsum; venter cream, blotched and
marbled with brown (least distinct in
groin and on lower venter); throat
barred with brown; anal triangle brown
posterior surfaces of thighs rich brown
with cream flecks; shank bars oblique,
narrower than interspaces; W-shaped
occipital mark, scapular spots, and some

limb bars edged with black.

Color in life—Dorsum green or

56 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

brown variously marked with reddish
brown, dark brown, olive-green, or
orange; venter dull greenish yellow to
grayish white or orange-tan, mottled
with brown or gray; hidden surfaces of
thighs dark brown to reddish salmon;
iris bronze with brown or black reticu-
lations.

Distribution and ecology.—This spe-
cies has a restricted distribution in the
lower part of the Pastaza trench (1080-
1300 m) and to the north in the Cordil-
lera del Dué (1150 m). All individuals
have been found on herbs, bushes, or
low (up to 2 m) limbs of trees at night
in cloud forest.

Remarks.—Lynch and Schwartz
(1972) summarized the characteristics
evident in Jiménez de la Espada’s (1875)
illustration but deferred to associate the
name with any known population of
Eleutherodactylus. The arrangement of
tubercles on the body and limbs of E.
rubicundus is reasonably congruent with
that seen in E. appendiculatus (Wer-
ner), E. calcaratus (Boulenger), E. cru-
cifer (Boulenger), E. galdi (Jiménez de
la Espada), and E. orocostalis Rivero,
but on the basis of color patterns and
proportions, none of these can be con-
sidered as identical with E. rubicundus
(and all are associated with distinctive
frog populations). The frogs here asso-
ciated with E. rubicundus from the lower
Andean slopes in eastern Ecuador are
not separable from the illustrations of
the frog Jiménez de la Espada (1875)
called Hylodes rubicundus. This tenuous
argument is joined by the equally tenu-
ous one that several of the species re-
ported by Jiménez de la Espada (1875)
are found together in the upper Amazon
Basin in central Ecuador.

Lynch (1976b, d) placed E. crenun-
guis in the fitzingeri group of Eleuthero-
dactylus. Eleutherodactylus rubicundus
is without doubt closely allied to E.
crenunguis but its association with the
fitzingeri group would strain definition
of the group. The indented ungual flap

of the digits of the two species are dis-
similar to the apically rounded flaps
seen in members of the fitzingeri group.
In order to reflect the similarities of E.
crenunguis and E. rubicundus and to
preserve the integrity of the fitzingeri
group, E. crenunguis and E. rubicundus
are here assigned to an assembly within
the wnistrigatus group. This arrangement
is not entirely satisfactory, but until
other allies are identified, it seems to be
the best course. This assembly is de-
fined as follows: skin of venter smooth
or feebly areolate; first finger longer than
second; tympanum prominent, annulus
not concealed; vomerine odontophores
prominent, triangular in outline; head
narrow (HW/SVL .339-.418); low cra-
nial crests in large females; interorbital
space narrow; ungual flap indented (at
least on fingers); digits bearing pads;
toes not webbed.

Lynch (1976d) noted that E. cruen-
tus (Peters) and especially E. latidiscus
(Boulenger), both members of the uni-
strigatus group, resembled E. crenunguis
in having flared lips and indented ungual
flaps (and digital pads) but that the
unistrigatus group species have areolate
skin on the venter and the thumb is
shorter than the second finger. The
Amazonian slope species, E. rubicundus,
bridges much of the apparent gap. With
E. cremnobates, the species form a series
as follows: crenunguis — rubicundus —
cremnobates — latidiscus — cruentus.
The series will be expanded when the
fauna of the Pacific versant of Ecuador
is more completely described.

Eleutherodactylus spinosus Lynch

Eleutherodactylus spinosus Lynch, 1979, Misc.
Publ. Mus. Nat. Hist. Univ. Kansas 66:43
[Holotype—USNM 199891, Sapote, Pro-
vincia Morona-Santiago, Ecuador, 2470 m].

Eleutherodactylus spinosus occurs in
cloud forests at elevations of 1707-2835
m on the Cordilleras del Condor and
Matanga in southern Ecuador (Lynch,
1979a).

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 57

Eleutherodactylus trachyblepharis
(Boulenger)
Fig. 5B

Hylodes trachyblepharis Boulenger, 1918, Ann.
Mag. Nat. Hist. (9) 2:429 [Syntypes—BM
1912.11.1.58-60/RR 1947.2.17.2-4, collected
at El Topo, Provincia Tungurahua, Ecuador,
4200 feet].

Eleutherodactylus trachyblepharis: Peters, 1955,
Rev. Ecuat. Ent. Par. 2:349.

Diagnosis.—1) skin of dorsum smooth
or with low warts on posterior back, no
dorsolateral folds, that of venter coarsely
areolate; 2) tympanum partially con-
cealed beneath skin, its length %-%
eye length; 3) snout acuminate in dorsal
view, rounded in lateral profile, swollen;
canthus rostralis distinct; 4) upper eye-
lid narrower than IOD, lacking tuber-
cles; no cranial crests; 5) vomerine odon-
tophores very low, oblique; 6) males
lack vocal sac and slits; no nuptial pad;
7) first finger shorter than second; pads
enlarged, smaller than tympanum; discs
broader than long; 8) fingers lack lateral
fringes; 9) no ulnar tubercles; 10) no
tubercles on heel or tarsus; 11) two
metatarsal tubercles, inner oval, 3-4
times size of round outer; 2—4 indistinct
supernumerary plantar tubercles; 12)
toes lack lateral fringes, no webbing; toe
pads equal to or slightly larger than
those of outer fingers; 13) tan above
with sparse brown markings; venter
cream, stippled with brown, concealed
surfaces of thighs brown; 14) adults mi-
meemales 121-156 (x = 13.8, N =
20) mm, females 15.8-19.2 (x = 17.2, N
a9) sam SV 1h,

Eleutherodactylus trachyblepharis is
the smallest species of the genus found
in South America. It is probably most
closely related to the slightly larger E.
ignicolor Lynch and Duellman, E. pro-
latus Lynch and Duellman, and E. vari-
abilis Lynch; all four have heads that
are longer than wide. Eleutherodactylus
trachyblepharis lacks a pattern on the
throat (unlike E. ignicolor and E. pro-
latus) and venter and groin (unlike E.
variabilis). Eleutherodactylus ignicolor
and E. prolatus have tubercles on the

outer edge of the tarsus and have larger
digital pads. The pads of E. variabilis
are truncate unlike the round pads of
E. trachyblepharis.

Description—Head narrower than
body; head longer than wide [head
length 111.4-117.8 (x = 114.9, N = 9)
percent head width in males, 101.7-113.6
(x = 106.8, N = 7) in females]; head
width 32.9-37.0 (x = 34.3, N = 16) per-
cent SVL; tip of snout swollen, accentu-
ating head length; E-N 72.7-88.2 (x =
77.7, N = 9) percent eye length in
males, 78.3-90.5 (x = 86.1, N = 7), in
females; nostrils weakly protuberant, di-
rected laterally; loreal region nearly flat,
sloping abruptly to lips; lips not flared;
upper eyelid 73.5-93.8 (x = 81.6, N =
9) percent IOD in males, 77.8-94.4 (x =
89.0, N = 7) in females, lacking tuber-
cles; no cranial crests; supratympanic
fold obsolete; tympanum partially con-
cealed beneath skin, anterior and ventral
edges of tympanic annulus distinct; tym-
panum round in males, higher than long
in females, separated from eye by half
its length in males, its length in females;
tympanum length 33.3-42.9 (x = 37.8,
N = Q) percent eye length in males,
26.1-38.1 (x = 33.4, N = 7) in females;
postrictal tubercles not evident; choanae
relatively small, longer than wide, near
edge of palate; vomerine odontophores
absent in some adults, when present,
median and posterior to choanae, sepa-
rated medially by distance equal 1.5-2
times odontophore width, oblique, about
size of a choana, bearing 0-4 teeth
in slanted row; tongue longer than wide,
posterior border feebly notched, poste-
rior half not adherent to floor of mouth.

Skin of dorsum smooth or with vague
flat warts on lower back, flanks smooth,
no dorsolateral folds; discoidal folds well
anteriad to groin; no anal sheath or anal
warts; no ulnar tubercles; palmar tuber-
cle bifid, 1.5 times size of oval thenar
tubercle; supernumerary palmar tuber-
cles low, numerous; subarticular tubercle
round, non-conical; fingers lack lateral
fringe or keel; discs nearly round but
wider than long; pads apically rounded,

58 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

largest on III-IV (1.5-2 times width of
digit), all expanded, all smaller than
tympanum.

No tubercles on heel or tarsus; inner
metatarsal tubercle low, oval (length
2.5 times width); outer metatarsal
tubercle round, pungent, #- size of
inner; supernumerary tubercles flat, at
bases of toes II-IV (also few on plantar
surface proper ), large (nearly as large as
subarticular tubercles which are round,
pungent); no lateral fringe or keel on
toes; no webbing; pads on toes about
same size as those of outer fingers; heels
broadly overlap when hind legs are
flexed; heel of adpressed hind leg
reaches to between eye and _ nostril;
shank 49.0-55.8 (x = 52.9, N = 16) per-
cent SVL.

In preservative, tan or pale brown
above with brown interorbital bar, dark
occipital W, scapular spots, diverging
bars from sacrum to flank, canthal-supra-
tympanic stripe, labial bars, limb bars;
limb bars about as wide as interspaces,
oblique on shank; anal triangle dark
brown; cream or tan line above vent;
posterior surfaces of thighs uniform
brown; throat and venter cream pep-
pered with brown.

Color in life-—Dorsal surfaces pale
yellow to light brown or reddish brown
(sometimes with pale green wash) with
brown and/or black markings; flanks yel-
low with brown area behind insertion of
arm; occasional examples have orange
wash in groin; anal triangle light to dark
brown, cream or white bar above vent;
posterior surfaces of thigh gray to
brown; venter off-white to gray, throat
yellow; iris pale gray with reddish
brown; horizontal streak (iris in some
individuals has red wash).

Measurements of adult male and fe-
male in mm.—(KU 120200 and 120203,
respectively): SVL 14.9, 17.5; shank
7.3, 9.0; head width 4.9, 5.9; head length
5.7, 6.7; upper eyelid width 1.4, 1.4; IOD
1.7, 1.6; tympanum length 0.8, 0.8; eye
lene thy2-20 273-2 = Ne leaeles:

Distribution and ecology.—Most spec-
imens of E. trachyblepharis are from the

valley of the Rio Pastaza (950-1250 m)
where special efforts were made to se-
cure specimens. The other records are
from low elevations (320-530 m) in Pro-
vincia Pastaza. The presently known dis-
tribution (low to moderate elevations
along the Rio Pastaza) may be real in
view of the occurrence of a similar spe-
cies (E. prolatus) at slightly higher ele-
vations to the north or may be an artifact
of most collectors failing to scrutinize
the low herbaceous layer of the forest
floor at night and the reluctance of many
collectors to pick up what appear to be
juvenile Eleutherodactylus.

Most specimens have been found on
small herbs 10-30 cm above the ground
in primary and secondary forest. KU
120254 was found in a recently drained
swamp in a cleared area at Sarayacu.
Although E. trachyblepharis lacks vocal
slits and vocal sac, males call. KU
120199-200 were collected as they were
calling on vegetation along a small for-
ested stream at Abitagua. The call is a
“sharp chirp-like click, one every 3-5
seconds.”

The ovarian complement is small in
so small a frog. KU 120203 has three
large ovarian eggs as well as many small-
er eggs. The left ovary contains two
large eggs, 1.8 & 1.8 and 1.8 « 2.3 mm;
the right ovary contains one large egg,
Sc 2:5 mame

Eleutherodactylus trepidotus Lynch
Fig. 5C

Paludicola festae Peracca, 1904, Boll. Mus.
Zool. Anat. Comp. 19:32 [Lectoholotype.—
MSNT 820, an adult female from Papa-
llacta, Provincia Napo, Ecuador; lectoholo-
type designated by Lynch, 1975c:31; pre-
occupied in Eleutherodactylus by Hylodes
festae Peracca (= E. galdi, fide Lynch,
1974) ].

Eleutherodactylus trepidotus Lynch, 1968, Her-
petologica 24:295 [Holotype.—USNM
164399, an adult female from 1 km W
Papallacta, Provincia Napo, Ecuador, 3155
m].

Diagnosis.—1) skin of dorsal surfaces
smooth with small pointed warts and
short ridges and folds, that of venter

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 59

coarsely areolate; no dorsolateral folds;
2) tympanum prominent, round, its
length %-% eye length; 3) _ snout
rounded in dorsal and lateral profiles;
canthus rostralis rounded; 4) upper eye-
lid much narrower than IOD; no pun-
gent tubercles on eyelid; no cranial
crests; 5) vomerine odontophores low,
oblique; 6) males with vocal slits and
subgular vocal sac, no nuptial pads; 7)
first finger shorter than second; digits
bearing long discs (length equal width),
lacking pads; 8) fingers lack lateral
fringes; 9) no ulnar tubercles; 10) no
tubercles on heel or outer edge of tarsus;
no supernumerary plantar tubercles; 11)
two metatasal tubercles, inner oval, twice
as large as round outer; 12) toes bearing
lateral keels, no web; toes bearing nar-
row pads; 13) dorsum dark brown to
black; facial markings distinct; flanks
and venter black with pale cream or
white spots (spots red in life); 14)
adults small, males 14.4-21.3 (x = 16.9,
N = 5) mn, females 17.3-25.5 (x =
21.2, N = 21) mm SVL.
Eleutherodactylus trepidotus is dis-
tinguished from most species of the uni-
strigatus group by virtue of its lack of
digital pads. None of the other species
lacking pads have dark venters with pale
spots. Eleutherodactylus trepidotus is
most frequently confused with speci-
mens of the disc-less genus Phrynopus,
but all species of Phrynopus in Ecuador
have digits that appear pointed without
magnification whereas those of E. trepi-
dotus appear to be round. Eleutherodac-
tylus elassodiscus also has narrow digits,
but the skin of the venter is smooth and
the first finger longer than the second.
Description.—See Lynch (1968b).
Color in life —Dorsum brown washed
with green; dorsal markings dark green,
dark brown, or black; venter brown to
black; spots on throat white to yellow;
those on venter and in groin white with
pale red centers to red; iris gray-brown.
Distribution and ecology.—Eleuthe-
rodactylus trepidotus is known in para-
mo and subparamo habitats from the
region of Nevado Cayambe south to the

Llanganti mountains (N side of Pastaza
valley) at elevations between 2360 and
3650 m. All individuals have been taken
by day beneath rocks or logs in pastures
or paramos. Males are taken _infre-
quently.

Eleutherodactylus unistrigatus (Giinther)

Hylodes unistrigatus Giinther 1859, Proc. Zool.
Soc. London, 1859:416 [Syntypes—BM
60.6.16.97-98, 60.6.16.105/RR 1947.2.17.
7-9, collected in western Ecuador by Mr.
Fraser].

Phyllobates equatorialis Barbour, 1908, Bull.
Mus. Comp. Zool. 51:320 [Cotypes—MCZ
2261, collected at “Equator’].

Eleutherodactylus wunistrigatus: Barbour and
Noble, 1920, Bull. Mus. Comp. Zool. 63:
401.

Eleutherodactylus unistrigatus [unistrigatus]:
Cochran, 1948, Amer. Mus. Novitat.
(1375):1.

Diagnosis.—1 ) skin of dorsum smooth
to shagreened with small scattered warts,
flanks bearing flattened warts, that of
venter coarsely areolate; no dorsolateral
folds; 2) tympanum distinct, its length
about half eye length; 3) snout round or
subovoid in dorsal view, round in lateral
profile; canthus rostralis rounded; 4) up-
per eyelid much narrower than JOD;
upper eyelid not bearing pungent tuber-
cles; no cranial crests; 5) vomerine odon-
tophores oblique, indistinct in small indi-
viduals; 6) males with vocal slits and
subgular vocal sac; no nuptial pads; 7)
first finger shorter than second; enlarged
pads on fingers II-IV, discs broad; 8)
fingers lack lateral fringes; 9) ulnar tu-
bercles poorly defined; 10) heel and
tarsus lacking tubercles except for poor-
ly defined inner tarsal tubercle; 11) two
metatarsal tubercles, inner oval, 6-7 times
size elongate outer metatarsal tubercle;
supernumerary plantar tubercles few in
number; 12) toes bearing narrow lateral
fringes, not webbed; toe pads as large
as those of outer fingers; 13) pattern
polymorphic; most individuals cream,
tan, or brown with brown or black in-
terorbital bar, dorsal chevrons, suprain-
guinal bar, slanted bars on _ flanks,
canthal-supratympanic stripe, labial bars;

60 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

limbs marbled with brown (bars indis-
tinct); posterior surfaces of thighs and
groin cream with brown marbling or not;
venter cream; throat of male yellow,
that of female cream peppered with
brown; 14) adults small, males 15.5-23.4
(x = 8.8, N = 26) mm, females 23.3-
30.8 (x = 25.4, N = 45) mm SVL (Ama-
zonian slopes only).

Eleutherodactylus unistrigatus is
closely related to several species found
at moderately high elevations in Colom-
bia and Ecuador [E. bogotensis (Peters),
E. cajamarcensis Barbour and Noble, E.
coeruleus (Andersson), E. erythropleu-
rus (Boulenger), E. lehmanni (Boett-
ger), E. orcesi Lynch, E. pastazensis
Andersson, E. riveti (Despax), and E.
thymelensis Lynch]. Eleuthterodactylus
unistrigatus is most easily recognized be-
cause it is a small tan, squat-bodied frog
having brown or black chevrons and
bars on the dorsal surfaces. It is not
separable from E. coeruleus ( Andersson )
or E. lehmanni (Boettger), but placing
those taxa in synonymy is deferred to
Lynch’s forthcoming paper on the frogs
of the genus found in the northern
Andes.

Description—The only usable de-
scriptions are those by Andersson (1945)
for Syrrhophus coeruleus and Cochran
and Goin (1970) for Eleutherodactylus
lehmanni. Considerable variation is evi-
dent among highland populations, and
that variation is evident when comparing
the two descriptions.

Color in life—Dorsum creamy tan,
yellow, orange brown, or brown with
brown to black markings; middorsal
stripe (if present) yellow; venter white
to cream; throat yellow in males, dirty
cream in females; posterior surfaces of
thighs pale fuscus brown; iris bright
copper brown above, same or reddish
brown below, usually with reddish
brown horizontal streak.

Distribution and ecology—Through
most of its distributional area, E. unistri-
gatus is a frog of high altitude grasslands
and Eucalyptus groves (2300-3200 m),
occurring primarily in the inter-Andean

hoyas from Pasto in southern Colombia
south to Riobamba in central Ecuador.
Eleutherodactylus unistrigatus also oc-
curs on the Andean slopes. Presumably
isolated populations are known from the
vicinity of Mindo and Nono (Provincia
Pichincha, Ecuador, 1500-2500 m) and
Pilal6 (Provincia Cotopaxi, Ecuador,
2580 m) on the Pacific slopes. Similar
populations are known from the Ama-
zonian slopes (Colén, Departamento Pu-
tumayo, Colombia, 2200 m; E] Carmelo
to Santa Barbara, Provincia Napo, Ecua-
dor, 2600-2800 m; and Banos, Provincia
Tungurahua, Ecuador, 1800 m). The
Banos population is contiguous with
thcse populations found along the Rios
Chambo and Patate (and their tribu-
taries) between Latacunga and _ Rio-
bamba.

By day individuals are found be-
neath rocks, dirt clods, wood, bricks,
paper, or under the cover of grass. At
night, during December through March,
males are actively calling on low vege-
tation (herbs within one meter of the
ground). The call is a “quonk,” some-
times given in a series of 5-6 notes. Egg
masses are deposited beneath rocks or
dirt clods and require 39 days to hatch
(Nina and del Pino, 1977). Breeding is
seasonal (December through March).
During the non-breeding season, males
do not call, but frogs do venture from
beneath cover at night to forage for
insects. Circumstantial evidence suggests
that both sexes mature in a single year.

Eleutherodactylus ventrimarmoratus
(Boulenger)

Hylodes ventrimarmoratus Boulenger, 1912,
Ann. Mag. Nat. Hist. (8)10:187 [Syntypes.
—BM 1911.1.1.51-53/RR 1947.2.15.74-76,
from El] Topo, Provincia Tungurahua, Ecua-
dor, 1280 m, collected by M. G. Palmer,
and BM 1901 12:12)77/7RiR” O44 e2elsaias
from Chanchamayo, Departamento Junin,
Pert, collected by G. Shunke].

Eleutherodactylus _ ventrivittatus | Andersson,
1945, Ark. Zool. 37A:33 [Holotype.—
NHRM (not examined), from Ambitagua
(= Abituagua), Rio Pastaza, Provincia
Tungurahua or Pastaza, Ecuador, collected

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 61

by W. Clarke-Macintyre in September
1937]. New synonymy.

Eleutherodactylus ventrimarmoratus: —Anders-
son, 1945, Ark. Zool. 37A:30.

Diagnosis—1) skin of dorsum, flanks,
and limbs tuberculate, that of venter
aerolate; no dorsolateral folds; 2) tym-
panum concealed; 3) snout rounded in
dorsal and lateral profiles; canthus ros-
tralis rounded; 4) interorbital space
broader than upper eyelid; upper eyelid
lacking enlarged tubercles; no cranial
crests; 5) vomerine odontophores trian-
gular in outline; 6) males lacking vocal
sac, vocal slits, and nuptial pads; 7) first
finger shorter than second; pads large,
largest on IIJ-IV, not emarginate; 8)
fingers bearing narrow lateral fringes; 9)
ulnar tubercles present; 10) no tubercles
on heel or outer edge of tarsus; fold on
inner edge of distal one-half of tarsus;
11) two metatarsal tubercles, inner oval,
six times size of round outer; supernu-
merary plantar tubercles not pungent;
12) toes bearing lateral fringes, no web;
toe pads large but smaller than those of
outer fingers; 13) brown above with dif-
fuse gray flecks or vermiculation; flanks,
anterior and posterior surfaces of thighs,
and concealed shank bearing black spots
or bars on colorless background; belly
white, spotted or marbled with black;
throat cream with pale brown marbling;
14) adults moderate-sized, males 17.8-
25.5 (x = 21.8, N = 8), females 33.3—
43.8 (x = 36.9, N = 9) mm SVL.

The pattern on the venter and con-
cealed surfaces of the limbs precludes
confusion of E. ventrimarmoratus with
any other species. Eleutherodactylus alta-
mazonicus Barbour and Dunn has simi-
lar markings in the groin and on the
concealed surfaces of the limbs but does
not have bold marbling on the venter
and is smaller (males 14.4-23.1 mm,
females 23.6-33.9 mm SVL). Eleuthero-
dactylus diadematus (Jiménez de la Es-
pada) is slightly larger (males 21.4-27.4
mm, females 35.4-44.5 mm SVL), has
diffuse marbling on the venter and con-
cealed surfaces of the limbs, and tuber-

culate skin, but has a prominent tym-
panum.

Description.—See Andersson’s (1945)
description of E. ventrivittatus.

Color in life—Rich brown above
with faint black markings edged with
white flecks; flanks, anterior and poste-
rior surfaces of thighs, concealed shank,
and venter black and white; throat and
undersurfaces of limbs red; iris pale
gold.

Distribution and ecology.—E. ventri-
marmoratus ranges from below 100 m
(western Brasil) to 1740 m (Rio Azuela,
Provincia Napo, Ecuador). Its geo-
graphic distribution is from the cloud
forests in northern Ecuador and the rain-
forests in western Brasil south to south-
ern Peru. Specimens have been found
on low vegetation at night and in a rot-
ten log in swamp forest by day.

Remarks.—Lutz and Kloss (1952)
suggested that E. ventrimarmoratus and
E. ventrivittatus are synonymous. An-
dersson’s (1945) description and figures
of E. ventrivittatus are in agreement with
the cotypes of E. ventrimarmoratus. An-
dersson separated his specimens from E.
ventrimarmoratus on the basis of slight
proportional differences, tongue shape,
and skin texture. The difference in skin
texture may well be due to preservation.
The coloration, body size, concealed tym-
panum, digit pad size, and proportions
of E. ventrivittatus are in agreement
with E. ventrimarmoratus and the two
are here proposed as synonyms.

Eleutherodactylus w-nigrum (Boettger)
Fig. 5D

Hylodes w-nigrum Boettger, 1892, Ber. Senck.
Naturf. Ges., 1892:28 [Holotype —SMF
3804, taken at Zurucuchu, approximately 15
km W Cuenca, Provincia Azuay, Ecuador,
3500 m, in 1890 by C. F. Lehmann].

Hylodes buergeri Werner, 1899, Verh. Zool.-
Bot. Ges. Wien 49:476 [Syntype—BM
1900.2.7.4/1947.2.15.71, Alto de Sibaté
near Bogota and Fusagasuga, Departamento
Cundinamarca, Colombia. (This may be
the only surviving syntype) J.

Eleutherodactylus buergeri: Dunn, 1944, Rev.
Acad. Colombiana Cienc. Exacta Fis. Nat.
6:73.

62 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Eleutherodactylus w-nigrum: Peters, 1955, Rev.
Ecuat. Ent. Par. 2:337; Cochran and Goin,
1970, Bull. U.S. Natl. Mus. (288) :395.

Diagnosis——1) skin of dorsum sha-
greened with granules and warts on up-
per flanks, that of venter smooth; no
dorsolateral folds; 2) tympanum promi-
nent, its length 35-52 percent eye length;
3) snout rounded or ovoid in dorsal
view, round in lateral profile; canthus
rostralis sharp; 4) upper eyelid width
usually slight less than IOD; no pun-
gent tubercles on eyelid; no cranial
crests; 5) vomerine odontophores promi-
nent, triangular in outline; 6) males with
vocal slits and subgular vocal sac, white
non-spinous nuptial pads; 7) first finger
longer than second; all bearing broad
discs on pads, pads largest on fingers
IIJ-IV; 8) fingers bearing lateral fringes;
9) ulnar tubercles indistinct or absent;
10) heel and tarsus lacking pungent
tubercles; 11) two metatarsal tubercles,
inner elongate, 6-8 times size of round
outer; supernumerary plantar tubercles
few or absent; 12) toes bearing lateral
fringes, no webbing; pads of toes as
large as those of outer fingers; 13) dor-
sum tan to brown with brown interorbi-
tal bar, occipital W, sacral chevron;
flanks bearing dark brown to black spots;
canthal-supratympanic stripe and labial
bars prominent; posterior surfaces of
thighs tan with brown to black spots or
reticulations; groin spotted or marbled;
venter cream to yellow with gray to
brown spots; 14) adults large, males
29.3-46.8 (x = 34.2, N = 32) mm, fe-
males 44.4-56.6 (x = 49.9, N = 15) mm
SVL (Amazonian slopes of southern Co-
lombia and northern Ecuador only).

Eleutherodactylus w-nigrum is most
easily recognized by virtue of having
round dark spots on the flank. Round
spots occur on the posterior surfaces of
the thighs in the populations on the
Amazonian Andean slopes. Casual in-
spection of the frog allows it to be con-
fused with most members of the fitz-
ingeri group. For example, Cochran and
Goin (1970) reported examples of E.
w-mgrum as E. brederi, E. conspicillatus,

and E. longirostris, as well as E. w-
nigrum. Some of the examples of Hy-
lodes conspicillatus reported by Bou-
lenger (1882) are E. w-nigrum, and
those misidentifications probably con-
tributed to much of the confusion of the
identity of E. conspicillatus. Eleuthero-
dactylus w-nigrum is probably most
closely related to E. lymani Barbour and
Noble, from which it differs in lacking
an inner tarsal fold.

Description.—See Cochran and Goin
(1970).

Color in life——Dorsum light brown
to brown (sometimes becoming rusty
brown above) with brown markings
edged with cream; venter white, cream,
or yellow (yellow in juveniles and most
males); groin, anterior and posterior sur-
faces of thighs, and concealed shank yel-
low with black markings; iris bright yel-
low-bronze above, reddish bronze be-
low, usually with reddish horizontal
streak, reticulated with black. The pale
areas on the concealed limbs and flanks
are white in adults in populations from
the Amazonian slopes.

Distribution and ecology.—Eleuthero-
dactylus w-nigrum occurs in lower mon-
tane and cloud forests between 800 and
3300 m on both the Pacific and Ama-
zonian slopes in Colombia and Ecuador.
In Colombia, E. w-nigrum ranges as far
north as the Medellin Valley; in Ecuador
it ranges south to provincias E] Oro and
Zamora-Chinchipe. By day specimens
are found beneath logs, rocks, or in dead
vegetation. At night frogs are especially
active along streams, but may be found
in forests adjacent to streams. Adult
females are seldom encountered very far
from streams, but young individuals and
males may be extremely abundant along
the forest edge (such as a road cut).
Observing 40 to 50 individuals in a night
is not uncommon; up to 250 individuals
have been observed in a four-hour period
in cloud forest.

Breeding occurs in January to April;
young frogs are abundant in May to
July but have disappeared by December.
The call of E. w-nigrum is very much

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 63

like that of many Gastrotheca—‘braaack,
bonk, bonk, bonk, bonk.”

Frogs from the Amazonian versant of
southern Ecuador are significantly larger
than those from northern Ecuador (see
Tables 1 and 2). Those from northern

Ecuador are approximately the same size
as frogs from northern, central, and
southern Colombia whereas those from
southern Ecuador (Amazonian versant)
are as large as E. w-nigrum from the
Pacific versant of northern Ecuador.

RELATIONSHIPS OF THE FAUNA

The 47 species (including two species
not treated above, E. buckleyi and an
undescribed species confused with E.
vertebralis, belong to four species groups
as recognized by Lynch (1976b) and
listed below.

I. biporcatus group: E. cornutus is
closely related to E. cerastes found at
comparable altitudes on the Pacific ver-
sant of Colombia and Ecuador; three
other species occur in trans-Andean
South America and Central America
(biporcatus, bufoniformis, and necerus).

II. discoidalis group: E. dolops and
E. elassodiscus occur on the Amazonian
versant as does E. nigrovittatus; the lat-
ter also is distributed into the Amazon
Basin. One species of the group (E.
mantipus) occurs on the Pacific versant
of the Cordillera Occidental in Colom-
bia; two occur on the Amazonian versant
in southern Pert and Bolivia (E. cru-
ralis and E. grandulosus), and one in the
mountains of northern Argentina (E. dis-
coidalis ).

Ill. fitzingeri group: E. condor, E.
lanthanites, E. peruvianus, and E. w-
nigrum occur on the Amazonian versant
but only E. condor is endemic. Eleu-
therodactylus w-nigrum is widely dis-
tributed in cloud forests in Colombia
and Ecuador and not closely related to
the other three (its closest allies include
E. actites and E. lymani; E. lanthanites
and E. peruvianus are broadly distrib-
uted over the Amazonian lowlands
where they are rarely sympatric; on the
Amazonian slopes they do not co-occur.

IV. unistrigatus group: The remain-
ing 39 species belong to the largest spe-
cies group of the genus (Lynch, 1976b),
including well over 100 species. Lynch
(1979a) suggested that the 18 species

of the wnistrigatus group in southern An-
dean Ecuador were members of eight
or nine intragroup assemblies. Some of
those 18 species are members of the
Amazonian slope fauna. Expanding that
earlier summary of apparent relation-
ships to the Amazonian slope fauna re-
quires 16 intragroup units:

1. acuminatus assembly: atratus and
inusitatus on the Amazonian slopes seem
most closely related to acuminatus of
the Amazon Basin.

2. celator assembly: colodactylus
and proserpens on the Amazonian slopes
seem most closely related to celator of
the upper cloud forests of the Pacific
versant of Ecuador.

3. crucifer assembly: chloronotus
and incanus on the Amazonian slopes ap-
pear to be closely related, and crypto-
melas may be allied to these species; the
assembly is represented on the Pacific
lowlands and slopes of Colombia and
Ecuador by E. calcaratus and E. cruci-
fer, respectively.

4, curtipes assembly: buckleyi in-
vades the upper edge of cloud forest on
the northern Amazonian slopes but is
primarily a frog of the paramo, as are
the closely related E. cryophilius and E.
curtipes. A second series of species in-
cludes trepidotus (invading upper cloud
forests on the Amazonian slopes), a
paramo frog, as are the related ginesi,
myersi, and nicefori.

5. devillei assembly: devillei occurs
in upper cloud forests in the Papallacta
Valley, an undescribed species (current-
ly confused with vertebralis) occurs in
similar habitats as well as paramo in the
headwaters of the Rio Chingual; the
closest allies of these two species are
vertebralis (cloud forests on the Pacific

64 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

versant of northern Ecuador) and an-
other undescribed species from cloud
forests on the Pacific versant of central
Ecuador.

6. diadematus assembly: incomptus
and ventrimarmoratus of the Amazonian
slopes are allied to lowland species alta-
mazonicus, carvalhoi, croceoinguinis, dia-
dematus, martiae, and platydactylus (the
last occurs on the Amazonian slope in
southern Pert).

7. frater assembly: quaquaversus
occurs in low cloud forests on the Ama-
zonian slopes; its allies are lowland for-
est frogs—frater in the Chocé and ocken-
deni in the upper Amazon Basin.

8. glandulosus assembly: acerus,
glandulosus, leucopus, and lividus from
upper cloud forests on the Amazonian
slopes of northern Ecuador and adjacent
Colombia; percultus from subpdramo of
southern Ecuador may be closely related.

9. lacrimosus assembly: bromelia-
ceus and petersi on the Amazonian slopes
of Ecuador are allied to lacrimosus
(Amazonian Basin) and mendax (slopes
and lowlands in southern Pert).

10. nigrogriseus assembly: eriphus,
nigrogriseus, and spinosus on the Ama-
zonian slopes in Ecuador.

ll. pugnax assembly: pugnax on the
Amazonian slopes of northern Ecuador
is closely related to an undescribed spe-
cies from upper cloud forests on the
Pacific versant of northern Ecuador.

12. pyrrhomerus assembly: gladiator
(Papallacta Valley), leoni (Rio Chingual
and Pacific cloud forests of northern
Ecuador ), and pyrrhomerus (high cloud
forests on Pacific versant in central
Ecuador; the distribution of this assem-
bly parallels that of the devillei assem-
bly, except that one species (leoni)
occurs on both Amazonian and Pacific
slopes of the Andes.

13. rubicundus assembly: rubicun-
dus (low cloud forests on the Amazonian
slopes) is closely related to crenunguis
and an undescribed species (low cloud
forests on the Pacific versant of northern
Ecuador); cremnobates (low cloud for-
ests on the Amazonian slopes) is related

to latidiscus and more remotely to cru-
entus (both from lowlands and lower
cloud forests ) of the Chocé.

14. surdus assembly: baryecuus
(high cloud forests of the Amazonian
slopes in southern Ecuador) is related to
surdus (high cloud forests on the Pacific
versant of the north and central Ecua-
dor.

15. trachyblepharis assembly: igni-
color, prolatus, and trachyblepharis from
the cloud forests on the Amazonian
slopes of Ecuador are related to varia-
bilis (Amazonian lowlands); galdi (Am-
azonian slopes in Ecuador ) is apparently
allied to this group.

16. unistrigatus assembly: this as-
sembly includes two series; the first in-
cludes pastazensis and unistrigatus found
in upper cloud forests on the Amazonian
slopes as well as bogotensis, erythropleu-
rus, and lynchi from Colombia and caja-
marcensis and versicolor from southern
Ecuador; the second series includes pri-
marily paramo frogs (balionotus, orcesi,
pycnodermis, riveti, ruidus, and thymel-
ensis), but pycnodermis invades upper
cloud forests on the Amazonian slope in
southern Ecuador, and ruidus is a cloud
forest frog on the Pacific versant in
southern Ecuador.

The members of the curtipes and
unistrigatus assemblies are primarily
frogs distributed above tree-line (alti-
tudinal range 1800 to at least 4100 m).
Only E. erythropheurus, E. pastazensis,
and E. ruidus are restricted to cloud for-
ests, but the limits of the unistrigatus
assembly remain to be established on
the Pacific versant and in Colombia.

The members of the celator, crucifer,
devillei, glandulosus, nigrogriseus, pug-
nax, pyrrhomerus, and surdus assemblies
are frogs of cloud forests. Only E. cal-
caratus (crucifer assembly ) invades low-
land forest. At present, only the glandu-
losus and nigrogriseus assemblies are en-
demic to the Amazonian slope cloud
forests; representatives of the other six
assemblies occur on both Amazonian and
Pacific slopes, but the absence of the
glandulosus and nigrogriseus assemblies

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 65

from the Pacific slopes is probably due
to incomplete study of the Pacific slope
fauna.

Frogs of the rubicundus assembly in-
clude one series of species found in very
low cloud forests (below 1500 m) on
both Amazonian and Pacific slopes, as
well as a second series having one spe-
cies in low cloud forests on the Ama-
zonian versant and a pair of frogs in low
cloud forests and lowland rainforests on
the Pacific versant.

Five assemblies (in addition to the
rubicundus assembly) occur in cloud
forests, as well as lowland forests (acu-
minatus, diadematus, frater, lacrimosus,
and trachyblepharis assemblies). The
frogs of the acuminatus, diadematus,
lacrimosus, and trachyblepharis assem-
blies occur in the Amazon lowlands, as

well as in the cloud forest of the Ama-
zonian slopes, whereas frogs of the frater
assembly include one in the trans-
Andean lowlands (frater), one in the
lowlands of the upper Amazon Basin
(ockendeni), and one in lower cloud for-
ests on the Amazonian slopes in Ecuador
( quaquaversus ).

Of the 71 species included in these
16 units, only 12 occur in paramo and
subparamo (all species in the curtipes
and unistrigatus assemblies) and only 15
occur in lowland forests (representatives
of 6 assemblies ). The majority (44) are
cloud forest animals (1000-3000 m) con-
firming Lynch’s (1979a) observation that
most species in southern Ecuador (and
now the Amazonian slopes as well) have
their closest allies in adjacent (or the
same altitudinal) strata.

DISTRIBUTION AND ECOLOGY

The delimitation of distributions of
species and recognition of communities
frequently is a difficult task in regions of
high relief and complex topography. The
Amazonian slope of the Andes in Ecua-
dor is no exception. Prior to the analysis
of the distribution and the discussion of
the communities we provide the reader
with a description of the region and a
physiographic map showing localities
where specimens were collected (Fig. 6).

DESCRIPTION OF REGION

Viewed from the lowlands of the
upper Amazon Basin, the Cordillera Ori-
ental of the Ecuadorian Andes rises
much like an irregular wall to the clouds
and beyond to a series of volcanic peaks,
some active and many perpetually cov-
ered with snow. West of this range is
a complex system of intermontane val-
leys separating, and ridges connecting,
the Cordillera Oriental and the Cordil-
lera Occidental. In Ecuador the Cordil-
lera Oriental is contiguous with the
Colombian cordillera and continues
southward with elevations above 3000
m from the Colombian border for about

275 km to the Pastaza trench. South of
the Pastaza trench, elevations of more
than 3000 m are continuous for another
150 km to the Rio Paute Valley. From
the Colombian border to the valley of
the Rio Paute are extensive areas of
more than 4000 m and many snow-cov-
ered volcanoes exceeding 5000 m; these
are (north to south) Volcan Cayambe
(5789 m), Volcan Antisana (5705 m),
Volean Cotopaxi (5697 m), Cerro Altar
(5319 m), and Volcan Sangay (5323 m).
The latter, Volcan Reventador (3483 m),
and Volcan Tunguragua (5005 m) have
been active in the past decade. South of
the Rio Paute elevations above 3000 m
are less extensive than to the north, and
the highest peak is Cerro El Picacho
(3850 m). One continuous range with
elevations of more than 3000 m extends
130 km south-southwest from the Rio
Paute to the Rio Zamora Valley, and
another extends from that valley south-
ward for about 125 km to the Huanca-
bamba Depression in northern Pert.
Three highland areas lie east of the main
cordillera. Volc4n Sumaco (3900 m) is
situated on an eastern extension of the
foothills and connected to the Cordillera

66 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Oriental by elevations of more than 2000
m. The Cordillera Cutucu reaching ele-
vations in excess of 1500 m is separated
from the Cordillera Oriental by the val-
leys of the Rio Upano and Rio Naman-
gosa at elevations of less than 1000 m.
The Cordillera del Condor is separated
from the Cordillera Oriental by the val-
leys of the Rio Nangariza and Rio Za-
mora at elevations of less than 1500 m.

Countless small streams cascade
down the eastern slopes of the Cordil-
lera Oriental and coalesce to form small
rivers, which in turn join to form larger
rivers, all of which are tributaries of
major rivers flowing into the Amazon—
from north to south—Rio Putumayo, Rio
Coca, Rio Napo, Rio Pastaza, Rio San-
tiago, and Rio Marafién. Three rivers
have their head-waters in intermontane
basins; the valleys of these rivers (Rio
Pastaza, Rio Paute, Rio Zamora) cut
through the Cordillera Oriental thereby
creating discontinuities in the high
ranges. Of these, the Rio Pastaza is the
largest and most impressive, lying in a
deep gorge between the Cerro Los Llan-
ganati (4639 m) to the north and Vol-
can Tungurahua (5005 m) to the south.
This valley, frequently referred to as the
Pastaza trench, forms a lowland incision
far into the Cordillera Oriental.

Only three climatological stations
exist on the eastern slopes of the Andes
in Ecuador; two of these (Bafios and
Pastaza) are in the Pastaza trench, and
one (Papallacta) lies above 3000 m
(Table 5). These data demonstrate the
decline of temperatures (about 1°C for
each 200 m) with increasing elevation

and show reduced rainfall but essen-
tially equal relative humidity from lower
to higher elevations. Bafios, situated in
the upper part of the deep Pastaza
trench, certainly receives less rainfall
than sites at the same elevation but sit-
uated on the montane slopes. In general
the eastern slopes of the Andes from
1500 to 3000 m are bathed in clouds
daily; usually mornings are relatively
clear, but light rain may fall at any time
of the day or night.

Between 1000 and 3000 m on the
eastern slopes of the Andes the nature
of the vegetation changes from tropical
to subtropical. According to Holdridge’s
(1947, 1964) system of classification, the
lower regions are humid subtropical for-
est and the upper reaches are very hu-
mid subtropical forest, as shown on an
ecological map of Ecuador (Vivanco de
la Torre et al., 1963). This is the altitudi-
nal belt frequently referred to as cloud
forest or montane rainforest. Below 1000
m the cloud forest gives way to tropical
rainforest (humid tropical forest or very
humid tropical forest in the classification
of Holdridge). Above 3000 m the mon-
tane rainforest becomes dwarfed and at
elevations of 3500-4000 m gives way to
alpine formations, principally paramo.
On the basis of the works of Acosta
Solis (1968), Espinal and Montenegro
(1963), Grubb and Whitmore (1966),
and Grubb et al. (1963), the common
genera of trees and shrubs at elevations
below 2000 m include Adenaria, Byrso-
nima, Calliandra, Cedrela, Guarea, Gy-
nerium, Rapanea, and Salix. Between
2000 and 3000 m the common genera

Fic. 6.—Map of the Amazonian slopes of the Andes in Ecuador. The area on the Amazonian
slopes between 1,000 and 3,000 m is shaded. Numbers correspond to the following localities, ar-
ranged from northwest to southeast (see Gazetteer for details): 1. El Carmelo; 2. Santa Barbara;
3. La Alegria; 4. La Bonita; 5. Agua Caliente; 6. Cordillera del Dué; 7. Rio Reventador; 8. Rio
Azuela; 9. Salto de Agua; 10. Rio Salado; 11. Santa Rosa; 12. Papallacta; 13. Chalpi; 14. Cuyujia;
15. Borja; 16. Jatintinahua; 17. Baeza; 18. Oritoyacu; 19. Rio Bermejo; 20. Cerro Sumaco; 21. San
José; 22. Rio San José; 23. Paracayacu; 24. Rio Jorge; 25. El Golpe; 26. Los Llanganatis; 27. Bafios;
28. Runtun; 29. Rio Negro; 30. El Topo; 31. Mirador; 32. Abitagua; 33. Mera; 34. Rio Alpayacu;
35. Puyo; 36. Macas; 37. Méndez; 38. Mirador; 39. Cerro Negro; 40. Pailas; 41. Cordillera de Cu-
tucu; 42. Sevilla de Oro; 43. General Plaza; 44. San Vicente; 45. Loma de Puerco; 46. Suro Rancho;
47. Sapote; 48. San Juan Bosco; 49. El. Cruzado; 50. Plan de Milagro; 51. Rio Pitintza; 52. Saba-

nilla; 53. Abra de Zamora; 54. Zamora.

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES

"Nie
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RIO.

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SAAB RO

V. Cayambe @_*

IG
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4 ay
a

V. Antisana®

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SY ‘ sy 7
V. Cotopaxi @—

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67

68 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

are Albizzia, Alchornia, Cupania, Inga,
Miconia, Ochroma, Persea, Trema, and
Trichanthera. Tree ferns are abundant
throughout the elevational range, where-
as bromeliads are most abundant below
2000 m. Above 1800 m the viney bam-
boo, Chusquea, becomes more abundant,
especially along streams, at high eleva-
tions. Physiognomically, the forest
changes from lower to higher elevations
(Grubb et al., 1963). In the lower mon-
tane rainforest (1000-+2000 m), the
trees reach heights of 15-33 m and have
small, if any, buttresses; leaves are noto-
phyllic or mesophyllic, frequently have
drip tips, and occasionally are com-
pound. In the upper montane rainforest
(2000-3000 m), the trees reach heights
of 15-18 m and usually have no but-
tresses; leaves are microphyllic, few or
none have drip tips, and few are com-
pound. In the upper reaches of the for-
est (subalpine rainforest) the trees reach
heights of 1.5-19 m and have no but-
tresses; leaves are nanophyllic, and none
is compound or has a drip tip. Locally,
due to winds and precipitation, a low
(<1.5 m) subalpine type of elfin forest
occurs on ridges above 2700 m; the low,
gnarled bushes support heavy growths of
mosses and large bromeliads.
Generally in the Neotropics many of
the humid montane slopes are heavily
cultivated in coffee, but on the Ama-
zonian slopes of the Ecuadorian Andes,
relatively little land is in cafetales; most
of these are in the Pastaza trench, where
cultivation is more extensive than else-
where on the slopes. Other agricultural
use of the land includes citrus planta-

tions, naranjilla (Solanaceae), dairy cat-
tle, and subsistence farming.

PATTERNS OF DISTRIBUTION

To one unfamiliar with the diversity
of Eleutherodactylus on the Andean
slopes and with the topographic com-
plexity of the region, the recognition of
45 species in a region about 500 km in
length and 2000 m in elevation may be
incomprehensible. Although Eleuthero-
dactylus is the most speciose anuran
genus in the region, Colostethus (Ed-
wards, 1974) and Centrolenella (Lynch
and Duellman, 1973) also are repre-
sented by many species there. Analysis
of distributional data reveals that many
species have rather restricted ranges lati-
tudinally and/or altitudinally (Table 6).
The major topographic barrier on the
eastern face of the Andes in Ecuador is
the Pastaza trench; thus, distributions
were analyzed as: 1) north of the trench,
2) in the trench, and 3) south of the
trench. Of the 31 species from north
of the trench, 10 are endemic to the
region and 10 are also known from Co-
lombia; six of the latter are among 14 of
the northern species also found in the
Pastaza trench. Five of the 20 species
in the trench are not found elsewhere
on the Amazonian slopes of Ecuador,
but three of these (E.. croceoinguinis,
lacrimosus, and ockendeni) are widely
distributed at lower elevations; of the
other two E. trachyblepharis occurs at
elevations of 320-1300 m in the Pastaza
Valley, whereas E. pastazensis is restrict-
ed to the upper part of the valley. Five

TABLE 5.—Summary of Climatic Date for Three Stations on Amazonian Slopes of Andes in Ecuador
for 1971-1972.
(Data from INAMHI, 1973, 1975)

Rainfall (mm) Atmospheric
Station Mean An. Driest Wettest Temperature (°C) Humidity
Papallacta 1310 Dec. 64 182 = 7.3-18.0 (x = 9.1) 72-93 (x = 88)
3160 m
Bajfios 1350 Nov. 46 287 13.2-218(x=16.8) 64-97 (x= 86)
1843 m
Pastaza 4980 Sep. 140 Jun. 768 16.4-24.5 (x= 20.0) 68-96 (x = 86)

1043 m

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 69

species in the trench are among the 15
species known from south of the trench,
where eight species are endemic and
three are shared with Pert. Only four
species are found in and north and south
of the trench (E. galdi, nigrogriseus,
quaquaversus, and w-nigrum); the last
species also occurs in Colombia.

Species that occur at lower elevations
(<2000 m) tend to be more widespread
than do those at higher elevations.
Among the 13 species not ranging below
2000 m to the north of the trench, only
two (E. trepidotus and E. unistrigatus)
occur in the trench; those species are
chiefly at higher elevations (>3000 m)

TABLE 6.—Distributional summary of 45 species of Eleutherodactylus on the easten slopes of the
Andes in Ecuador.
(* = also on Andean slopes in Colombia and ** = also in Pert)

North of

Species Pastaza Trench
E. acerus 2660
FAMQENQUUSS es
(, DEG EHTS. i. ih a i aan eee ee
leepromenacus, jf j uns
E. chloronotus* 2285-3000-+
(ecolodactyluss = 9 = = ~~
condone) § | © F © FF se
E. cornutus* 1150-1740
E. cremnobates 1410-1700
E. croceoinguinis
HERCTUDLOMCLOS 2 tcc
E. devillei 2195-1980
E. dolops* 1100-1950
E. elassodiscus* 2300-2610
E. eriphus 2160-2600
E. galdi 1700-1740
E. gladiator 2360-2910
E. glandulosus 2105-2890
E. ignicolor 2160-2750
E. incanus 1700-1740
E. incomptus* 1410-1910
E. inusitatus 2160
RSIGGTUMOSUS,
E. lanthanites* —1000-1490
E. leoni 2130-3000-+
E. leucopus* 2400-2590
E. lividus 2135-2750
E. nigrogriseus 1150
E. nigrovittatus —1000-1935
pemockendenie eee
iEenastazensisn PM VM Oy ee ee
E. peruvianus** 1410-1910
E. petersi* 1100-1935
E. prolatus 1150-1490
E. proserpens ge
E. pugnax 1660-2540
1D. FOG ROGIS, os ee Oe ee eee
E. quaquaversus —1000-1740
E. rubicundus 1150
ERESINOSUSEE ee et
Betrachyblepharis- =
E. trepidotus 2360-3000-++
E. unistrigatus* 2710-3000-+
E. ventrimarmoratus* * 1740
E 1410-2540

In South of
Pastaza Trench Pastaza Trench
seas 2195-2850
Seanad 2195-2990
eae 1710-2620
I a 2195-3000+
— 1830
EO O=31!50 0s
—1000-1100 sen
ee 2470-3000-+-
1000-1280 1000-1830
DOOM Wy) eee
130 Oke eer ead ee ee
—1000-1100 uu
—1000-1100 — _uue
1180-1800 2195-2835
—1000-1100
—1000-1100 Wwe
1800-1840 8
+ 1830
1000-1800 ones
1140 ie ae
1700-2620
eit 9s 2650-3000+
—1000-1140 —1000-1830
1000-1300 anees
ae 1700-2835
—1000-1300
2925-3000-+ ——_
1800-3000-+- kat ald
—1000-1300 —1000-1100
1100-1600 2220

. w-nigrum*

70 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

3000

2500

2000

1500

1000

Fic. 7.—Altitudinal distribution of 26 species of Eleutherodactylus along a transect from Papallacta
to the Cordillera del Dué. Each line represents one species. Arrows indicates that the range extends

above 3000 m or below 1000 m.

and entered the trench from above. None
of the six southern species occurring only
above 2000 m is found in the trench.

Five species are primarily inhabitants
of the upper Amazon Basin; E. croceo-
inguinis, lacrimosus, and ockendeni reach
the slopes only in the Pastaza trench
(elevations to 1100 m), whereas E. lan-
thanites occurs to elevations of 1490 m
in Colombia and northern Ecuador, and
to 1100 m in the Pastaza trench, and E.
ventrimarmoratus ranges to 1300 m in
the trench and to elevations of more
than 1500 m in northern and southern
Ecuador and in Peru.

It is evident that there are notable
elevational changes in the Eleutherodac-
tylus fauna on the eastern slopes of the
Andes. In a transect in northern Ecua-
dor, 26 species occur from 1000 to 3000
m (Fig. 7); a transect in southern Ecua-

dor contains 11 species; in the former
transect no more than 14 species have
been found at any given elevation (1700-
1800 m), whereas in the latter transect
six species occur at 2200-2300 m. A more
realistic analysis compares the fauna at
specific sites along the northern transect
(Table 7; Fig. 8). The maximum num-
ber of species at any given site is 11 at
the Rio Azuela, 1740 m; four other sites
have seven species, and one has six
species.

ELEUTHERODACTYLINE
COMMUNITIES

Several eleutherodactyline communi-
ties are apparent on the Amazonian
slopes to the Andes. There are listed
below.

1. Upper cloud forest, Provincia Mo-

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES (ut

oO
~
O
ise)
i)
Q
{a0}
oO

|
Santa Rosa

ills
Lago i

?.

Lago Agrio

Fic. 8.—Map and profile of transect from Quito to Lago Agrio showing sites compared in Table 5.
1. 11-12 km ESE Papallacta, 2660 m. 2. 16.5 km NE Santa Rosa, 1700 m. 3. Rio Salado, 1410
m. 4. 2 km WSW Rio Reventador, 1490 m. 5. Rio Azuela, 1740 m. 6. Cordillera del Dué, 1150 m.

rona-Santiago (2000-3000 m): E. atra-
tus, E. baryecuus, E. bromeliaceus, E.
colodactylus, E. cryptomelas, E. nigro-
griseus, E. proserpens, E. pycnodermis,
E. spinosus, and E. w-nigrum.

2. Upper cloud forest, Provincia
Napo north to Nevado Cayambe (2000-
3000 m): E. acerus, E. chloronotus, E.
devillei, E. elassodiscus, E. eriphus, E.
gladiator, E. glandulosus, E. ignicolor,
E. lividus, E. pugnax, E. trepidotus, and
E..w-nigrum.

3. Upper cloud forest, northern Pro-
vincia Napo and southern Colombia
(2000-3000 m): E. buckleyi, E. chloro-
notus, E. elassodiscus, E. leoni, E. leuco-
pus, E. pugnax, E. unistrigatus, E. sp.
(vertebralis-like), and E. w-nigrum.

4. Lower cloud forest, Pastaza valley
north into Colombia (1300-2000 m): E.
cornutus, E. cremnobates, E. dolops, E.
galdi, E. incanus, E. incomptus, E. inu-
sitatus, E. nigrogriseus, E. pastazensis,
E. peruvianus, E. petersi, E. prolatus, E.
quaquaversus, and E. w-nigrum.

5. Lowest cloud forest (1000-1300
m): E. rubicundus, E. trachyblepharis,
and E. ventrimarmoratus. These are
joined by some predominately lowland
rainforest species at some localities—E.
croceoinguinis, E. lacrimosus, E. lantha-
nites, E. nigrovittatus, and E. ockendeni.

Eleutherodactylus w-nigrum is com-
mon to four (1-4) of the communities.
Communities 2 and 3 share E. chlorono-
tus, E. elassodiscus, and E. pugnax in ad-

72 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

dition to E. w-nigrum; they remain amply
distinct—only part of the distinction is ex-
plicable in terms of B-diversity (species
turnover between localities). It seems
most unlikely that several of the species
have been missed by collectors, although
that may well be the case for such spe-
cies as E. eriphus and E. ignicolor.
Except for E. pycnodermis, all spe-
cies of community 1 were collected at
Sapote (locality 47). All but E. crypto-
melas and E. pycnodermis were found
at El Cruzado (locality 49). Eleuthero-
dactylus pycnodermis was found at near-
by Suro Ranch (locality 46) with four
other members of the community. All
but E. pugnax of community 2 have
been collected in the immediate vicinity
of Cuyujua, Prov. Napo (locality 14).
The species of community 3 have been
collected from several localities between
FE] Carmelo and Santa Barbara, but no
site has yielded all. No locality has pro-
duced all the species for community 4;
nine of the species were found at the Rio
Azuela. Eleutherodactylus inusitatus
and E. pastazensis are both rare (three
and four specimens respectively); E. ni-
grogriseus is common to the south but
quite rare north of the Pastaza trench.
Community 5 may not be distinct from
community 4 but does include five low-
land species that extend up into the
cloud forest (six if E. ventrimarmoratus
is considered a lowland species). The
overlap of communities 4 and 5 is espe-
cially evident in the eleutherodactyline
fauna found at Mera, Provincia Pastaza
(locality 33): E. croceoinguinis, E. lacri-
mosus, E. lanthanites, E. nigrovittatus,

E. ockendeni, E. peruvianus (old speci-
mens, UMMZ), E. petersi, E. prolatus,
E. quaquaversus, E. rubicundus, E.
trachyblepharis, and E. ventrimarmora-
tus (four species of community 4 and
eight of community 5).

Habitat partitioning by these species
essentially occurs along spatial and
trophic dimensions, because with the
exception of E. nigrovittatus all appear
to be nocturnal. During the daylight
hours all others are found in sites of
concealment (beneath rocks and logs or
in axillae of plants).

Spatial (microhabitat) separation is
not pronounced. In community 2, E.
elassodiscus and E. trepidotus are terres-
trial, E. pugnax and w-nigrum are very
closely associated with streams, and the
remaining eight species are found on low
vegetation (not necessarily close to
streams). In community 3, E. elassodis-
cus, E. pugnax, and E. w-nigrum behave
as in community 2, and the remaining six
species are found on low vegetation.

In community 4, E. cornutus and E.
dolops are terrestrial, E. cremnobates, E.
nigrogriseus, and E. w-nigrum are closely
associated with streams (on vegetation),
and the other nine species are active on
vegetation (presumably E. pastazensis
is active on vegetation at night—no data
are available).

In community 5, E. nigrovittatus is
terrestrial and diurnal, E. rubicundus
usually is found on vegetation above or
beside streams and seeps, and the re-
maining six species are found on low
vegetation.

TaBLeE 7.—Comparison of the numbers of species of Eleutherodactylus at six sites along a transect
from Papallacta to the Cordillera del Dué, Ecuador

(11-12 km ESE Papallacta, 2660 m)
(Rio Azuela, 1740 m)

(16.5 km NE Santa Rosa, 1700 m)

(2 km WSW Rio Reventador, 1490 m)
(Rio Salado, 1410 m)

(Cordillera del Dué, 1150 m)

C2 UIC oN iS

Sree ered |

2. 3. 4. 5. 6.
00 .00 .0O .00 00
11 67 44 00 23
6 7 43 Rots) 15
4 3 U O7 3]
5 6 4 7 5
2 iL 2 1 6

1 Boldface numbers are actual numbers of species at a site;

Roman numbers are the number of species in common

between sites, and italicized numbers are the coefficients of community (Whittaker, 1970).

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 73

No direct data on trophic separation
are available. Duellman (1978) found
little trophic separation at Santa Cecilia
but such a finding might have been pre-
dicted because his food classes were not
precise. Indirect data are available in
the form of size differences between spe-
cies. Lynch (1976d, 1979a) compared
communities and ecological contingen-
cies on the basis of ¢ values (¢ = mean
size of larger/mean size of smaller; see
Hutchinson and MacArthur, 1959, for
theory). The frogs in these five commu-
nities exhibit comparable size differences
in each community (Table 8). Mean ¢
values range from 1.08 to 1.15. Separat-
ing members of the communities by both
size and microhabitat elevates mean ¢
values principally because the species
associated with streams as well as the
terrestrial taxa exhibit more pronounced
size differences among themselves than
do the species found on vegetation (con-
tingency 3, Table 8).

Frogs of the wnistrigatus group are
predominant in each of the five commu-
nities (Table 9) but few close relatives
are sympatric. Only four assemblies in
the five communities have more than one
species per community (diadematus,
glandulosus, nigrogriseus, and trachyble-
pharis assemblies ). The sympatric mem-
bers of the diadematus assembly (com-
munity 5) are E. croceoinguinis and E.
ventrimarmoratus; they differ markedly

in size (Table 3). As many as five spe-
cies of this assembly (EF. altamazonicus,
E. croceoinguinis, E. diadematus, E. mar-
tiae, and E. ventrimarmoratus ) may co-
occur in the vicinity of Puyo, Provincia
Pastaza (900 m); the size differences be-
tween adjacent-sized species is greater
(Table 10) than those for entire commu-
nity assemblages on the Amazon slopes
(Table 8) or in the lowlands (Lynch,
1979a ).

Three species of the glandulosus as-
sembly are sympatric in community 3
(E. acerus, E. glandulosus, and E. li-
vidus). These frogs are moderately well-
separated by size (acerus-glandulosus,
@ 1.14 male, 1.25 female; glandulosus-
lividus, g 1.38 male, 1.06 female), but
only E. glandulosus is sufficiently abun-
dant for us to have confidence in its
body size. Two species of the nigro-
griseus assembly (E. nigrogriseus and E.
spinosus ) are sympatric in community 1;
they are nearly equal in size (x 6 =
1.11). Two species of the celator assem-
bly are also sympatric in community 1;
their size difference is likewise slight
(x g = 1.08). The two species of the
trachyblepharis assembly sympatric in
community 4, E. galdi and E. prolatus,
differ markedly in size (x ¢ = 1.34).

Our current knowledge of these frogs
leaves the impression of only slight habi-
tat partitioning in communities in the
cloud forests and lowland forests. Parti-

TABLE 8.—¢ values for Amazonian Slope Eleutherodactyline Communities. Values include range (N)
mean.

Contingency
3. Size and micro-
habitat; orboreal
species only

2. Size and micro-
habitat; all species

1. Size alone;
all species

Community 1 4 4 MOMMIGGIKOS = , -idbides Tg © use

Community 2 ¢ 4 1.00-1.18(10)1.08 1.00-1.59(9)1.19 1.00-1.25(7)1.10
Community 3 é 1.00-1.33(8)1.10 1.06-1.37(6)1.19 1.06-1.33(5)1.15
Community 4 ¢@ 1.00-1.34(11)1.08 1.00-1.51(10)1.12 1.00-1.51(7)1.10
Community 5 ¢é 1.01-1.33(8)1.12 1.01-1.33(7)1.14 1.01-1.33(7)1.14
Community 1 ? 9 OSS OMSK e ee) ee Re oY epee ee

Community 2 2 2 1.00-1.24(10)1.09 1.00-1.58(8)1.21 1.00-1.24(6)1.10
Community 3 ¢ 9 1.00-1.22(8)1.11 1.00-1.56(6)1.23 1.00-1.36(5)1.16
Community 4 9 2 1.00-1.33(12)1.08 1.00-1.58(10)1.14 1.00-1.33(8)1.09
Community 5 9 9 1.00-1.36(8)1.15 1.00-1.36(7)1.17 1.00-1.36(7)1.17

74

tioning seems to be more complete in
paramo communities comprised of few
species (Duellman, 1979). The large
number of species of Eleutherodactylus,
as well as many other kinds of frogs, in
cloud forests (and in lowland rainforest
in the upper Amazon Basin) may have
both historical and ecological explana-
tions. According to the refugia theory
(Haffer, 1969, 1974), the lower Ama-
zonian slopes and adjacent Amazon Ba-
sin in Ecuador supported tropical rain-
forest throughout the Pleistocene, where-
as forests in much of the Amazonian
region diminished during arid phases of
the Pleistocene. Lynch (1979b) showed
a high fidelity of Eleutherodactylus to
humid forests and high degrees of en-
demism to area of proposed refugia. Our

TABLE 9.—Distribution of species groups and
assemblies in five Amazonian slope eleuthero-
dactyline communities.

Community

Pe eee Ae
biporcatus group 0 0 0 1 0
discoidalis group OFet a ie= xis
fitzingeri group eet Soe mel
unistrigatus group 9 10 7 10 6
acuminatus assembly 1 1
celator assembly 2 ar
crucifer assembly 1 cr lige. ght |
curtipes assembly ce ape reel
devillei assembly ee Cr ae ee
diadematus assembly eg es eres bE)
frater assembly Pa ples ene BW BE is |
glandulosus assembly I, See aN) chee eet
lacrimosus assembly ieee fe wee ll
nigrogriseus assembly DF weal see eee aes
pugnax assembly Oe aly ee ae
pyrrhomerus assembly Prey ae Cees
rubicundus assembly reat Ties errr a Oe
surdus assembly igcaoymulen Araactor aphie) © jatar
trachyblepharis assembly _. 1 —. 2 1
unistrigatus assembly Pr ae

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

limited data on habitat partitioning
among Eleutherodactylus parallels that
presented for a lowland Amazon site by
Duellman (1978), who interpreted the
low g values and broad niche overlap
values in a large anuran community as
being indicative of bountiful resources
and little interspecific competition. Al-
though we have no precise measure-
ments of the structural heterogeneity of
the forests on the Amazonian slopes of
the Andes, we are impressed with the
general trend from greater heterogene-
ity at lower elevations (<1800 m) to
less heterogeneity at higher elevations
(> 1800 m) and diminution of the for-
est above 3000 m. In general there is a
decline in the numbers of species of
Eleutherodactylus from low to high ele-
vations. It has yet to be demonstrated
whether frogs are responding directly to
heterogeneity of the habitat or both frogs
and vegetation are responding to physi-
cal environmental factors.

RESUMEN

E] género Eleutherodactylus esta rep-
resentado en las laderas amazdénicas de
los Andes Ecuatorianos por no menos de
47 especies. En este informe damos reg-
istros de 45 de las 47 especies, incluyen-
do las descripciones de 12 especies
nuevas (E. acerus, E. condor, E. cremno-
bates, E. dolops, E. eriphus, E. ignicolor,
E. incanus, E. incomptus, E. inusitatus,
E. lividus, E. petersi, E. prolatus), y
descripciones igualmente detalladas de
cuatro especies poco conocidas (E. pas-
tazensis, E. peruvianus, E. rubicundus,
E. trachyblepharis). Las especies nuevas
incluyen un miembro del grupo discoi-
dalis (E. dolops), un miembro del grupo

Taste 10.—Size differences among sympatric members of the diadematus assembly. Overall x ¢
(8 comparisons) = 1.16.

x 9 SVL Eo) x ¢ SVL a6 Xp
E. diadematus 39.8 1.08 23.5 1.08 1.08
E. ventrimarmoratus 36.9 1.32 21.8 1.16 1.24
E. altamazonicus 27.9 1.37 18.8 1.24 1.30
E. martiae 20.4 1.03 15.1 1.03 1.03
E. croceoinguinis 1S )} seca i= a Seer AIG) SS eee =

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 75

fitzingeri (E. condor), y 10 miembros
del grupo unistrigatus. Sinonimias, diag-
nosticos, coloracién en vida, distribu-
ciones, y notas ecoldgicas, asi como ref-
erencias a descripciones previamente
publicadas son dadas para E. chlorono-
tus, E. cornutus, E. croceoinguinis, E.
devillei, E. elassodiscus, E. galdi, E.
gladiator, E. glandulosus, E. lacrimosus,
E. lanthanites, E. leoni, E. leucopus, E.
nigrogriseus, E. nigrovittatus, E. pugnax,
E. quaquaversus, E. trepidotus, E. uni-
strigatus, E. ventrimarmoratus, and E.
w-nigrum. Las poblaciones de E. lacri-
mosus, E. lanthanites, E. nigrovittatus,
and E. peruvianus de las laderas ama-
zonicas son de mayor tamafnio a aquellas
poblaciones de las tierras bajas.

Las 47 especies incluyen represen-
tantes de cuatro grupos de especies (los
grupos biporcatus, discoidalis, fitzingeri,
y unistrigatus). El grupo unistrigatus
esta representado por 16 subgrupos; 71
especies del grupo unistrigatus pertene-
cen a estos 16 subgrupos—44 son ani-
males de bosque nublado, 12 son espe-
cies de paramo y subparamo, y 15
especies son de la selva de tierras bajas.

E] Valley del Rio Pastaza es la bar-
rera mas importante de las laderas ama-
zonicas, y su effecto es mas notorio en
las especies que se encuentran en alturas
medias (2000-3000 m). Cinco unidades
ecogeograficas son aparentes y estan par-
cialmente correlacionadas con tres es-
tratas altitudinales y de vegetacién. Las
comunidades de ranas eleutherodacty-
linas en cada una de las unidades eco-
geograficas son distintas entre si. Las

cinco comunidades que reconocemos
son: 1) bosque nublado superior, Pro-
vincia Morono-Santiago, 2000-3000 m, 10
especies; 2) bosque nublado superior,
Provincia Napo, hacia el norte hasta el
Nevado Cayambe, 2000-3000 m, 12 es-
pecies; 3) bosque nublado superior al
norte de la Provincia Napo y de Colom-
bia adyacente, 2000-3000 m, 8 especies;
4) bosque nublado inferior al norte del
Rio Pastaza, 1300-2000 m, 14 especies;
y 5) bosque nublado bajo, 1000-1300 m,
8 especies.

Entre estas comunidades hay una
ligera separacién ecoldgica, excepto una,
son nocturnas. Solamente una o dos es-
pecies terrestres (o una o dos especies
estrechamente relacionadas con arroyos)
por comunidad son separadas por micro-
habitat. Gran parte de los miembros de
cada comunidad son ranas arboricolas
las cuales se encuentran en vegetaciones
bajas. Diferencias en tamafio (refleja-
das por diferencias del tamafio de la
preza) son muy sutiles (especies de
tamanios similares en las cuales la mayor
es aproximadamente 130 porciento del
tamano de la menor, promedio = 110%).
Si consideramos simultaneamente el ta-
mano y el microhabitat hay una ligera
separaciOn la cual es aparente (110-
120%).

E] numero de especies generalmente
disminuye a medida que aumenta la
elevacion. Hay disminucidn en la hetero-
genidad estructural del bosque. Las
ranas responden a esa heterogenidad o
ambos pueden estan respondiendo a gra-
dientes en los factores fisicas del medio.

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APPENDIX A: SPECIMENS EXAMINED

Eleutherodactylus acerus (4 spec.)

ECUADOR, Prov. Napo: 7.8 km WNW
Cuyujia, 2750 m, KU 143517; 11 km ESE
Papallacta, 2660 m, KU 165471-73.

Eleutherodactylus atratus (44 spec.)

ECUADOR, Prov. Morona-Santiago: El]
Cruzado, 2195 m, USNM 199690-99; San Vi-
cente, 2835 m, USNM 199712; Sapote, 2470
m, USNM 199701-10, 199713 (cleared and
stained skeleton); 2 km W Sapote, 2561 m,
USNM_ 199711; between Sapote and Suro
Rancho, 2604-2622 m, USNM 199683-89; Suro
Rancho, 2683 m, USNM 199675-82; 0.5 km
W Suro Rancho, 2744 m, USNM 199700; Prov.
Zamora-Chinchipe: Abra de Zamora, 2850 m,
KU 165236-38, 166265 (cleared and stained
skeleton).

Eleutherodactylus baryecuus (17 spec.)
ECUADOR, Prov. Morona-Santiago: Cerro

Negro, 2927 m, USNM 199715; El Cruzado,
2195 m, USNM 199723-24; San Juan Bosco,
2195 m, USNM 199725; between San Juan
Bosco and E] Cruzado, 2226 m, USNM 199717-
22; San Vicente, 2805-2835 m, USNM 199729;
3 km W San Vicente, 2988 m, USNM 199730;
Sapote, 2470 m, USNM 199726-28; between
Sevilla de Oro and Mendez (probably between
the crest and Pailas), USNM 199716; Suro
Rancho, 2683 m, USNM 199714.

Eleutherodactylus bromeliaceus
(23 spec. )

ECUADOR, Prov. Morona-Santiago: El
Cruzado, 2195 m, USNM 199738-39; Mirador,
1982 m, USNM 199735; Pailas, 2195 m, USNM
199732-34, 199743-45; Plan de Milagro, 1707
m, USNM 199736-37, 199746-47; Rio Piuntza,
Cordillera de] Condor, 1830 m, KU 146974-75;
0.5 km E Sapote, 2393 m, USNM 199740,
199748-49: 1 km E Sapote, 2332 m, USNM
199741; mountain above Sapote (to the south),

78 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

2500 m, USNM 199742, 199750; between Sa-
pote and Suro Rancho, 2622 m, USNM 199731;
Prov. Zamora-Chinchipe: 22 km W Zamora,
1730 m, KU 141772.

Eleutherodactylus chloronotus
(231 spec. )

COLOMBIA, Depto. Narifio: 12 km E
Pasto, 3050 m, KU 167972-8002, 170127-29
(cleared and stained skeletons); Depto. Putu-
mayo: 13 km W Santiago, 2900 m, KU
168003-08.

ECUADOR, Prov. Napo: 2 km E Chalpi,
2700 m, USNM JAP 5547-48, 5550: 4 km E
Chalpi, 2590 m, USNM JAP 5553, 5558; Chaspi
Chico bridge, 4.6 km E Papallacta, 2870 m,
MCZ 92117; Cuyujia, UIMNH 55823-24,
55846; 1 km E Cuyujua, 2285 m, USNM JAP
5531; 7.8 km WNW Cuyujia, 2750 m, KU
143317-25; Laguna del San Marcos, NE of
Cayambe, CAS-SU 8287; Llanganati moun-
tains, El Golpe, 3000 m, CAS-SU 17433:
Llanganati mountains, Rio Jorge, 3000 m, CAS-
SU 17444; Llanganati mountains, at Rio San
Jose, 3049 m, CAS-SU 10385-89, 10391, 10401-
08, 17395-425, 17444; Papallacta, 3110-3130
m, KU 143331-49, USNM JAP 5108, 5110,
5113-14; 0.5 km E Papallacta, 3095 m, USNM
JAP 5128; 1.6 km E Papallacta, 2980 m, KU
177242-44; 3 km E Papallacta, 2900 m, KU
117516-49, 118103(5), 118104-06 (lots of
eggs), 118130-33 (cleared and stained skele-
tons); 5.7 km E Papallacta, 2910 m, KU
143326-30; 9.2 km ESE Papallacta, 2750 m,
KU 177246; 9.6 km E[SE] Papallacta, 2700 m,
MCZ 92116; 11 km ESE Papallacta, 2660 m,
KU 155436-40, 165208-18, 166267-68 (cleared
and stained skeletons), 177245; 12 km ESE
Papallacta, 2630 m, KU 165202-07; 15 km
E[SE] Papallacta, 2600 m, KU 127007-11; 1
km W Papallacta, 3200 m, KU 111385-86;
2 km W Papallacta, 3270 m, KU 177247: 4
km W Papallacta, 3300 m, KU 117550-56;
north side Laguna Papallacta, 3350 m, KU
109071-72; Rio Bermejo, Cordillera Guacamayo,
WCAB 35651; 0.8 km NW Santa Barbara,
Quebrada El] Corazon, 2540 m, KU 177248.

Eleutherodactylus colodactylus
(113 spec.)

ECUADOR, Azuay Prov.: 8 km ESE Sevilla
de Oro (on Azuay, Morona-Santiago frontier),
3140 m, USNM_ 198458. Morona-Santiago
Prov.: El Cruzado, 2195 m, USNM 198436:
Pailas, 2195 m, USNM 198431-33, 198459: San
Juan Bosco, 2195 m, USNM_ 198434-35,
198460-61; immediate environments of San
Vicente, 2790-2820 m, USNM 198465-79; 1
km W San Vicente, 2851 m, USNM 198462-64:
1 km E Sapote, 2332 m, USNM 198457; moun-
tain above Sapote to the south, 2500 m, USNM
198456; 1/2 km W Sapote, 2546 m, USNM

198437-55. Zamora-Chinchipe Prov.: Abra de
Zamora, 2800 m, KU 142151-59 (142155 is
cleared and stained skeleton), 165219-21; 14
km E Loja, 2770 m, KU 142160-61; 15 km E
Loja, 2710 m, KU 142162-64.

PERU, Depto. Piurd: 33 kn SW Huanca-
bamba, 2745-3050 m, LSUMZ 32368-414.

Eleutherodactylus condor (46 spec. )

ECUADOR, Prov. Morona-Santiago: Pas-
taza River, Canelos to Maranon, MCZ 19629-
30, 19632; Rio Piintza, Cordillera del Condor,
1830 m, KU 146991-7033.

Eleutherodactylus cornutus (46 spec.)

COLOMBIA, Depto. Putumayo: 10.3 km
W El Pepino, 1440 m, KU 168009.

ECUADOR, no specific locality: USNM
167431; Prov. Morona-Santiago: Cutucti moun-
tains, E of Macas, 1800-2000 m, AMNH 39499:
Macuma, USNM_ 167401-08; Miazal, USNM
167409-27; Prov. Napo: Cerro Sumaco, AMNH
22265-70; S slope Cerro Sumaco, USNM
167429; S slope Cordillera del Dué above Rio
Coca, 1150 m, KU 123447-48: Loreto, USNM
167428; Rio Azuela, 1740 m, KU 143350; 2 km
SSW Rio Reventador, 1490 m, KU 165222:
Prov. Pastaza: Canelos, USNM 167430; 10 km
S Veracruz, MCZ 89972; Prov. Tungurahua:
El Topo, 1270 m, BM 1912.11.1.57; Mirador,
Banos, 1500 m, BM 94.10.30.44.

Eleutherodactylus cremnobates
(40 spec. )

ECUADOR, Prov. Napo: 7 km SW Rio
Azuela, KU 155456-60; 2 km SSW Rio Reven-
tador, 1490 m, KU 166036-56: Rio Salado +
1 km upstream from Rio Coca, 1410 m, KU
146164, 177249-60: 16.5 km NNE Santa Rosa,
1700 m, KU 143459.

Eleutherodactylus croceoinguinis
(66 spec. )

Only Amazonian slope specimens are listed
here; see Lynch (1980) for lowland localities.
200 lowland specimens also examined.

ECUADOR, Prov. Pastaza: Mera, 1140 m,
KU 119527-30, 119534-43, 119546-72, 121352
(eggs), 121353 (juveniles), 177261-74; 1 km
W Puyo, 1000 m, MCZ 90008-11; 25 kn NNE
Puyo, 1040 m, KU 146093; Rio Alpayacu, 1 km
E Mera, 1100 m, KU 119531-33.

Eleutherodactylus cryptomelas (9 spec.)

ECUADOR, Prov. Morona-Santiago: Sapote,
2470 m, USNM 198480-82; 2 km W Sapote,
2560 m, USNM 198483; Prov. Loja: 7 km N
San Lucas, 2840 m, KU 177735; 8-9 km M San
Lucas, 3000-3100 m, KU 120095-96; Prov.

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 79

Zamora-Chinchipe: Abra de Zamora, 2710 m,
KU 141992-93.

Eleutherodactylus devillei (146 spec. )

ECUADOR, “Andes of Ecuador’: MRHN
1009; Prov. Napo: Baeza, WCAB 35531; Borja,
1710 m, USNM JAP 5670; 2 km E Chalpi,
2700 m, USNM JAP 5535, 5546, 5549; 4 km
E Chalpi, 2620 m, USNM JAP 5156-59, 5552,
5556-57, 5559, 5565-74, 5576-83; Chaspi Chico
bridge, 4.6 km E Papallacta, 2870 m, KU
143408-14, MCZ 92118-23; Cuyujua, 2360 m,
UIMNH 55825-35 (55833, cleared and stained
skeleton), USNM JAP 5209, 5295, 5298, 5314,
5316, 5607, 5615, 5699-5700, 5762-64; 0.4 km
ESE Cuyujia, 2460 m, KU 178999; 1 km E
Cuyujiia, Rio Victoria, 2420 m, KU 146095;
1.5 km E[SE] Cuyujia, 2440 m, USNM JAP
5262-73; 3 km E[SE] Cuyujia, 2285 m, USNM
JAP 5257; 3.3 km ESE Cuyujuia, 2350 m, KU
143393; 7 km E[SE] Cuyujuia, 2195 m, USNM
JAP 5358; 1 km W Cuyujta, 2500 m, USNM
JAP 5303-05; 2.5 km WNW Cuyujua, 2550 m,
KU 143394-99; 5 km W[NW] Cuyujia, 2745
m, USNM JAP 5746-52; 7.8 kn WNW Cuyu-
jua, 2750 m, KU 143400; “top” of Mt. Sumaco,
AMNH 22192; 1.6 km E Papallacta, 2980 m,
KU 177275; 3 km E[SE] Papallacta, 2900 m,
KU 117570-71; 5.7 km ESE Papallacta, 2910
m, KU 143401-07; 8.3 km ESE Papallacta, Rio
Chalpi, 2750 m, KU 177276-77; 9.2 km ESE
Papallacta, 2750 m, KU 177278; 9.6 km ESE
Papallacta, 2700 m, MCZ 92124; 11 km ESE
Papallacta, 2660 m, KU 165408-14, 165933-34,
166271-72 (cleared and stained skeletons),
JDL 9422; 12 km ESE Papallacta, 2630 m, KU
155441, 166269-70 (cleared and stained skele-
tons); 13.1 km ESE Papallacta, 2610 m, KU
177279-80; 15 km E[SE] Papallacta, 2600 m,
KU 127012-13; 1 km W Papallacta, 3155 m,
USNM JAP 5150-51; Rio Bermejo, Cordillera
Guacamayo, WCAB 35653; Santa Barbara,
2590 m, USNM JAP 4478.

Eleutherodactylus dolops (11 spec.)

COLOMBIA, Depto. Putumayo: 10.3 km
W EI Pepino, 1440 m, KU 168811; 35 km SE
San Francisco, 1950 m, KU 168809-10.

ECUADOR, Prov. Napo: Loreto, USNM
GOV 8901; Rio Azuela, 1740 m, KU 143504; 2
km SSW Rio Reventador, 1490 m, KU 165866-
68; Salto de Agua, 2.5 km NNE Rio Reventa-
dor, 1660 m, KU 143505; Prov. Pastaza: Rio
Villano, USNM GOV 9332-33.

Eleutherodactylus elassodiscus
(44 spec., only 17 extant)

COLOMBIA, Depto. Putumayo: 13 km W
Santiago, 2900 m, KU 168012-13.

ECUADOR, Prov. Napo: Borja, 1720 m,
JAP 5671-72; Cuyujia, 2380 m, JAP 5206,

5210-13, 5228, 5294, 5299-300, 5315, 5472-74,
5701-02, KU 130271-72, USNM_ 167668-69;
0.4 km ESE Cuyujia, 2460 m, KU 177282-85;
1 km E[SE] Cuyujia, 2300 m, JAP 5236,
5238-42, USNM 167670 (cleared and stained
skeleton); 3 km E[SE] Cuyujia, 2300 m, JAP
5252-55; 3.3 km ESE Cuyujia, 2350 m, KU
143415, 146097; Papallacta, WCAB 37915;
Santa Barbara, 2610 m, USNM 192907-08; 1
km NW Santa Barbara, 2610 m, USNM
192906. [All specimens identified as JAP were
lost in the mails in 1970 (Lynch, 1973a)].

Eleutherodactylus eriphus (15 spec.)

ECUADOR, Prov. Napo: Papallacta, WCAB
37903, 37905, 37907-09; 12 km E[SE] Pa-
pallacta, KU 155451-55; Rio Jatuntinahua, 10
km ESE Cuyujia, 2160 m, KU 166031-35.

Eleutherodactylus galdi (39 spec.)

ECUADOR, no other locality: AMNH
17444; Prov. Morona-Santiago: El Cruzado,
2195 m, USNM JAP 7276; Miazal, USNM GOV
8947; Pailas, 2195 m, USNM JAP 6567-69,
6621-22; Rio Pitintza, Cordillera del Condor,
1830 m, KU 146977-85; San José de Cuchi-
pamba, MSNT 513(3776); Prov. Napo: Loreto,
CAS-SU 13155, USNM GOV 8945-46, 8944
(cleared and stained skeleton), JAP 3893, 8828,
WCAB 39910; Mt. Sumaco, side of, AMNH
22334; Mt. Sumaco, S slope, 800-1000 m, CAS-
SU 13154; “top” of Mt. Sumaco, AMNH 22201,
22205-08; Rio Azuela, 1740 m, AMNH 89726,
KU 143416, 165422; Prov. Pastaza: Rio Upano,
Macas, 1000 m, AMNH 21496; Prov. Tungura-
hua: El Topo, 1280 m, BM 1912.11.1.54-55
(reregistered as 1947.2.16.78-79); Prov. Za-
mora-Chinchipe: Sabanilla, AMNH 13531.

Eleutherodactylus gladiator (6 spec. )

ECUADOR, Prov. Napo: 5 km E[SE]
Chalpi, 2560 m, USNM JAP 5256; Cuyujua,
2360 m, USNM JAP 5608; 3.3 km ESE Cuyu-
jua, 2350 m, KU 143516; 5.7 km ESE Papal-
lacta, 2910 m, KU 143513-15.

Eleutherodactylus glandulosus
(187 spec. )

ECUADOR, “Andes of Ecuador’: MRHN
1010; Prov. Napo: Borja, 1710 m, USNM JAP
5673-83; 2 km E Chalpi, 2700 m, USNM JAP
5289-90; 4 km E Chapli, 2620 m, USNM JAP
5560; Cuyujia, 2360 m, UIMNH 55836-45,
55847-49, USNM JAP 5301-02, 5589-93, 5595-
603, 5616-20, 5684-97, 5766-69; 0.4 km ESE
Cuyujia, 2460 m, KU 177291-95; 1 km E[SE]
Cuyujia, 2285 m, USNM JAP 5233-35, 5321-
30; 3 km E[SE] Cuyujiia, 2285 m, USNM JAP
5249: 3.3 km ESE Cuyujua, 2350 m, KU
143417-18, 146099; 6.6 km ESE Cuyujia, Rio
Molando, 2360 m, KU 146098; 7 km E[SE]

80 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Cuyujua, 2195 m, USNM JAP 5351-57; 9 km
E[SE] Cuyujia, 2105 m, USNM JAP 5317-18;
1 km W Cuyujuia, 2500 m, USNM JAP 5308;
1.5 km W Cuyujia, 2440 m, USNM JAP
5260-61; 2.5 km WNW Cuyujta, 2550 m, KU
143419-30; 7.8 km WNW Cuyujia, 2750 m,
KU 143431; 4.6 km E[SE] Papallacta Rio Pa-
pallacta (probably equals Chaspi Chico bridge),
2890 m, KU 143432-33; 11 km ESE Papallacta,
2660 m, KU 155442-47, 165435-67, 166273-74
(cleared and stained skeletons), 177286; 12
km ESE Papallacta, 2630 m, KU 165423-34,
166275-76 (cleared and stained skeletons);
13.1 km ESE Papallacta, 2610 m, KU 177287-
90

Eleutherodactylus ignicolor (4 spec.)

ECUADOR, Prov. Napo: 9.2 km ESE Pa-
pallacta, 2750 m, KU 177296-97; 11 km ESE
Papallacta, 2660 m, KU 165880; Rio Jatuntina-
hua, 10 km SE Cuyujia, 2160 m, KU 165879.

Eleutherodactylus incanus (22 spec.)

ECUADOR, Prov. Napo: Rio Azuela, 1740
m, KU 143465-83; 16.5 km NNE Santa Rosa,
1700 m, KU 143457-58, 143460.

Eleutherodactylus incomptus (68 spec. )

ECUADOR, Prov. Napo: Rio Azuela, 1740
m, KU 143497; Rio Salado + 1 km upstream
from Rio Coca, 1410 m, KU 146169, 165844-
58, 177298-319; 2 km SSW Rio Reventador,
1490 m, KU 165932; 16.5 km NNE Santa Rosa,
1700 m, KU 143455, 143461-64, 143484-96;
Prov. Pastaza: 9.5 km NW Mera, 1270 m, KU
179000-03.

Eleutherodactylus inusitatus (3 spec. )

ECUADOR, Prov. Napo: Agua Caliente on
Rio Pedro, SE of Cayambe, CAS-SU 8279; Rio
Jatuntinahua, 10 km SE Cuyujia, 2160 m, KU
166066; Prov. Pastaza: Abitagua, 8 km NW
Mera, 1300 m, KU 120099.

Eleutherodactylus lacrimosus (43 spec. )

Only Amazonian slope records are given be-
low; see Lynch (1980) and Lynch and
Schwartz (1972) for lowland records. 99 low-
land specimens were also examined.

ECUADOR, Prov. Pastaza: Mera, 1140 m,
KU 119513-17, 119522-23, 132631-32, 177361-
68, 177376-82; Puyo, 960 m, KU 146100; 1
km N Puyo, 945 m, USNM 167372-78; 1 km
W Puyo, 1000 m, MCZ 92030, 92037; Rio
Alpayacu, 1 km E Mera, 1100 m, KU 119518-
21; 2 km S Shell Mera, 1006 m, USNM
167383-84; 3 km S Shell Mera, 960 m, USNM
167385-87.

Eleutherodactylus lanthanites (30 spec.)

Only specimens from the Amazonian slopes are

listed here; see Lynch (1975b, 1980) for low-
land records. 431 lowland specimens also ex-
amined.

COLOMBIA, Depto. Putumayo: 10.3 km
W El Pepino, 1440 m, KU 168834-45, 170172.

ECUADOR, Prov. Napo: Cerro Caleras (=
Galeras ), CAS-SU 10362; 7 km SW Rio Azuela,
KU 155448; 2 km SSW Rio Reventador, 1490
m, KU 165885-93, 165931; Prov. Pastaza: 1 km
W Puyo, 1000 m, MCZ 90037-48, 92034-35,
92128-30, 92132.

Eleutherodactylus leoni (65 spec.)

COLOMBIA, Depto. Narino: La Victoria,
2700 m, KU 140311.

ECUADOR, Prov. Carchi: El] Carmelo,
2715 m, USNM JAP 4951; 5.7 km NW El
Carmelo, 2910 m, KU 177341-42; 14 km (air-
line) SE Maldonado, 2500 m, KU 177320-40;
Prov. Imbabura: N slope Nudo de Mojanda,
3400 m, KU 130870-72; La Delicia, 2710 m,
KU 132799; Prov. Pichincha: Palma Real,
WCAB 35662; Prov. Napo: Santa Barbara,
2590 m, USNM 197941(6), 197944(11), GOV
7172-73, 7175, 7178, 7188, 7205, JAP 4482,
4484, 4486, 4541; 0.8 km NW Santa Barbara,
Quebrada El Corazon, 2540 m, KU 177348;
1 km SW Santa Barbara, 2590 m, USNM
197942(4), JAP 4496, 4589; between Santa
Barbara and Alegria, 2130-2440 m, USNM
197943.

Eleutherodactylus leucopus (32 spec.)

COLOMBIA, Depto. Narino: La Victoria,
2700 m, KU 140303-07, 140312.

ECUADOR, Prov. Napo: Santa Barbara,
2590 m, USNM 197927-36, 197940(7), JAP
7192, 7194; 1 km NW Santa Barbara, 2590 m,
USNM 197938(2); 1 km SW Santa Barbara,
2590 m, USNM 197939(2); 3 km SW Santa
Barbara, 2440 m, USNM 197937.

Eleutherodactylus lividus (17 spec.)

ECUADOR, Prov. Napo: 4 km E Chalpi,
2590 m, USNM JAP 5554-55; 3 km E[SE]
Cuyujia, 2135 m, USNM JAP 5251; 9.2 km
ESE Papallacta, 2750 m, KU 177344; 9.6 km
E[SE] Papallacta, 2700 m, MCZ 92115; 11
km ESE Papallacta, 2660 m, KU 155450,
165935-36, 165937-40, 166005-07; 12 km ESE
Papallacta, 2630 m, KU 165992, 166004.

Eleutherodactylus nigrogriseus
(46 spec. )

ECUADOR, Prov. Morona-Santiago: El
Cruzado, 2195 m, USNM JAP 7283-86; Loma
de Puerco, 2226 m, USNM JAP 7359-62;
Pailas, 2195 m, USNM JAP 6663; San Vicente,
2835 m, USNM JAP 7653; Sapote, 2470 m,
USNM JAP 7616-20; 2 km W Sapote, 2561 m,
USNM JAP 7656; Prov. Napo: S slope Cor-

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 81

dillera del Dué above Rio Coca, 1150 m, KU
123501; Prov. Pastaza: Abitagua, 1330 m, KU
120157-63, 120164(2); 3.5 km W Mera, 1180
m, KU 146109-11; 8.1 km NW Mera, 1270 m,
KU 177345-48; 9.5 km NW Mera, 1270 m,
KU 177349-55; Prov. Tungurahua: Banos,
NHRM 1905(6).

Eleutherodactylus nigrovittatus
(82 spec. )

Only specimens from the Amazonian slopes are
listed here; see Lynch (1980) for lowland
records. 49 lowland specimens also examined.
ECUADOR, Prov. Morona-Santiago: Rio
Yuquipa, Macas, USNM GOV 7210; Prov.
Napo: S slope Cordillera del Dué above Rio
Coca, 1150 m, KU 123503-42, 123544-65; La
Bonita, 1935 m, USNM JAP 4831; slopes of
Mt. Sumaco, AMNH 22284, 22287-89, 22293-
98, 22327, 22335-37, USNM GOV 8899-900;
Prov. Pastaza: Mera, 1100 m, KU 177356-57.

Eleutherodactylus ockendeni (11 spec.)

Only Amazonian slope specimens are listed be-
low; see Lynch (1974, 1980) for lowland
records. 314 lowland specimens were also
examined.

ECUADOR, Prov. Napo: S slope Cordillera
del Dué above Rio Coca, 1150 m, KU 123809-
11; Prov. Pastaza: Mera, 1140 m, KU 120261,
120269, 177358-59.

PERU, Depto. Hudnuco: S slope Serrania
Sira, Laguna, 1280 m, KU 154783-85; S slope
Serrania Sira, Pata Rojo, 1100 m, KU 154782.

Eleutherodactylus pastazensis (4 spec. )

ECUADOR, Prov. Tungurahua: Runttin
(Banos), CAS-SU 5084; Volcan Tungurahua,
1840 m, NHRM 1918 (small specimen); Yau-
guilla, CAS-SU 5065, NHRM 1918 (larger
individual).

Eleutherodactylus peruvianus (52 spec.)

Only specimens from the Amazonian slopes are
listed below; see Lynch (1980) for lowland
records. 346 lowland specimens were also ex-
amined.

ECUADOR, Prov. Morona-Santiago: Rio
Pitntza, Cordillera del Condor, 1830 m, KU
147034-38; Pastaza River, Canelos to Maranon,
MCZ 19635-36, 19639, 89314; Prov. Napo:
“top” of Mt. Sumaco, AMNH 22202-04; 3.2
km NNE Oritoyacu, 1910 m, KU 177360; Rio
Azuela, 1740 m, AMNH 89743-48, KU 143502-
03; Rio Salado + 1 km upstream of Rio Coca,
1410 m, KU 165859-65; 16.5 km NNE Santa
Rosa, 1700 m, KU 143498-501; Prov. Pastaza:
Mera, 1000 m, UMMZ 92124; near Mera, 1000
m, UMMZ 92126(5); Prov. Tungurahua: Abi-
tagua, 1100-1200 m, CAS-SU 5070, 5079,
UMMZ 92119(2), 92120, 92121(3); El Topo,

UMMZ 51264. Province unknown: Mapoto,
1300 m, UMMZ 89069.

PERU, Depto. Junin: Valle de Perene, 1500
m, AMNH 38598-601.

Eleutherodactylus petersi (71 spec.)

COLOMBIA, Depto. Huila: Parque Arche-
ologico San Agustin, 1750 m, KU 168818-22;
Depto. Putumayo: 10.3 km W El Pepino, 1440
m, KU 168812; 35 km SE San Francisco, 1950
m, KU 168813-17.

ECUADOR, Prov. Napo: La Bonita, 1935
m, USNM JAP 4637; 2 km SE La Bonita, 1920
m, USNM 204704, JAP 4862-68, 4870-80; Lo-
reto, USNM JAP 9367-68; Rio Azuela, 1740
m, KU 143506-07, 165993; Rio Salado, + 1
km upstream from Rio Coca, 1410 m, KU
165994-6003, 177383; 16.5 km NNE Santa
Rosa, 1700 m, KU 143508-12; Prov. Pastaza:
Mera, 1140 m, KU 120247-49, 177369-75; Prov.
Tungurahua: Banos, 1829 m, USNM JAP 5840-
47; Rio Blanco, UMMZ 92136; Yanguilla,
UMMZ 92138.

Eleutherodactylus prolatus (91 spec. )

ECUADOR, Prov. Napo: S slope Cordillera
del Dué above Rio Coca, 1150 m, KU 123755-
61; 2 km SSW Rio Reventador, 1490 m, KU
166008-15; Rio Salado, 1 km upstream from
Rio Coca, 1410 m, KU 146163, 146165-68,
166016-30, 177384-438; Prov. Pastaza: Mera,
1140 m, KU 120176.

Eleutherodactylus proserpens (27 spec.)

ECUADOR, Morona-Santiago Prov.: El]
Cruzado, 2195 m, USNM 199751; Plan de
Milagro, 1707 m, USNM 198485, 198504; 1 km
S Plan de Milagro, 1707 m, USNM 198503;
Rio Pitntza, Cordillera del Condor, 1830 m,
KU 147044-46; 1/2 km E Sapote, 2393 m,
USNM 199752; 1 km E Sapote, 2332 m, USNM
198499-501, 199753; mountain above Sapote
(to the south), 2500 m, USNM 198496-98;
1/2 km W Sapote, 2546 m, USNM 198486-91:
between Sapote and Suro Rancho, 2622 m,
USNM 198484, 198492-95, 198502.

Eleutherodactylus pugnax (9 spec.)

ECUADOR, Prov. Napo: Rio Azuela, 1740
m, EP 70/978, KU 155449; Rio Jatuntinahua,
10 km SE Cuyujia, 2160 m, KU 165561-62;
Salto de Agua, 2.5 km NNE Rio Reventador,
1660 m, KU 146466-67; 0.5 km NW Santa
Barbara, 2540 m, KU 177440-41; 3.7 km NW
Santa Barbara, 2410 m, KU 177439.

Eleutherodactylus pycnodermis
(137 spec. )

ECUADOR, Azuay Prov.: 3 km E Sevilla
de Oro, 2713 m, USNM 199851-53; 5 km ESE

82 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Sevilla de Oro, 2957 m, USNM_ 199854,
199867; between Sevilla de Oro and Cerro
Negro, 3110 m, USNM 199868; crest at Azuay-
Morona Santiago frontier, ca. 8 km SE Sevilla
de Oro, 3354 m, USNM 199855-59, 199866;
crest at Azuay-Morona Santiago frontier, ca.
8 km SE Sevilla de Oro, 3384 m, USNM
199758-59, 199860-65. Morona-Santiago Prov.:
between Cerro Negro and Pailas, 2652 m,
USNM_ 199755-57, 199869-71; San Vicente,
2805-35 m, USNM 199754, 199815-50, 199884,
199888-90; San Vicente, 2851 m, USNM
199813-14, 199885-87; 3 km W San Vicente,
2988 m, USNM 199794-812, 199882-83; 6 km
W San Vicente, 3100 m, USNM 199787-93,
199880-81; Suro Rancho, 2683 m, USNM
199760-66; 1/2 km W Suro Rancho, 2744 m,
USNM 199767-86, 199872-79.

Eleutherodactylus quaquaversus
(176 spec. )

ECUADOR, Prov. Morona-Santiago: Cu-
suime, Rio Cusuime, 320 m, AMNH 93646;
Rio Piuntza, Cordillera del Condor, 1830
m, KU _ 146987-90; Prov. Napo: Bermejo
No. 4 (well site), 15 km ENE Umbaqui,
740 m, KU 123690-701; S slope Cordillera
del Dué above Rio Coca, 1150 m, KU
123731-53; Loreto, USNM GOV _ 9371-72,
9557, JAP 9199, 9204; side of Mt. Sumaco,
AMNH 22338; Puerto Libre, Rio Aguarico, 570
m, KU 123706-30, 123778-80; Quebrada Paca-
yacu, tributary of Rio Cotapino, 600-650 m,
CAS-SU 13153; Rio Azuela, 1740 m, KU
143448-54; 2 km SSW Rio Reventador, 1490
m, KU 165563-65; Rio Salado, + 1 km up-
stream from Rio Coca, 1410 m, KU 165566-85,
166281-82 (cleared and _ stained skeletons);
Santa Cecilia, 340 m, KU 104593-96, 111181,
123702-05, 146161, 149197-99, 149232-37; 16.5
km NNE Santa Rosa, 1700 m, KU 143441-47;
Prov. Pastaza: Mera, 1140 m, KU 120165,
120169, 177442-50; Puyo, 960 m, UMMZ
164323; 3 km S Puyo, 920 m, KU 127034, 1
km W Puyo, 1000 m, MCZ 90116-38; Rio
Alpayacu, 1 km E Mera, 1100 m, KU 120166-
68, 120170-72; headwaters, Rio Bobonaza, 685
m, USNM JAP 2131; Shell Mera, UIMNH
93562; 10 km ESE Veracruz, MCZ 90139.

Eleutherodactylus rubicundus (19 spec. )

ECUADOR, Prov. Napo: S slope Cordillera
del Dué above Rio Coca, 1150 m, KU 123754,
123762-64; Prov. Pastaza: Abitagua, 1100 m,
CAS-SU 5077; Abitagua, 1200 m, UMMZ
92115; Abitagua, 8 km NW Mera, 1300 m,
KU 120173; Mera, 1140 m, KU 177452-56,
UMMZ 92116; 9.5 km NW Mera, 1270 m,
KU 177457-58; 5 km E, 6 km S Puyo, 925 m,
MCZ 92027; Rio Alpayacu, 1 km E Mera, 1100
m, KU 120174-75; Prov. Tungurahua: 11 km
E Rio Negro, 1170 m, KU 146170.

Eleutherodactylus spinosus (100 spec.)

ECUADOR, Morona-Santiago Prov.: be-
tween Cerro Negro and Pailas, 2652 m, USNM
199916; between Cerro Negro and Pailas, 2439-
2561 m, USNM 199917; El Cruzado, 2195
m, USNM 199945-66; Loma de Puerco, 2226
m, USNM 199967-71; Pailas, 2195 m, USNM
199918-35, 199973-77; 1 km S Plan de Milagro,
1707 m, USNM 199978; Rio Piuntza, CordiJ-
lera del Condor, 1830 m, KU 147039; San Juan
Bosco, 2195 m, USNM 199936-44, 199979-82:
San Vicente, 2835 m, USNM 199972, 199988-
89; Sapote, 2470 m, USNM_ 199891-915,
199986-87; between Sapote and Suro Rancho,
2604-2622 m, USNM 199983-85.

Eleutherodactylus trachyblepharis
(79 spec. )

ECUADOR, Prov. Morona-Santiago: Cu-
suime, Rio Cusuime, 320 m, AMNH 93668-
69; Prov. Pastaza: Abitagua, 1200 m, UMMZ
92117(2), 92118; Abitagua, 8 km NW Mera,
1300 m, KU _ 120199-200; Canelos, 530
m, KU 120253; Mera, 1140 m, KU 120179-
80, 120182-98, 120205-09, 120210(2), 120442,
177458-68; Puyo, 1000 m, USNM JAP
6184; 5 km SSE Puyo, 975 m, USNM
JAP 2022, 2024; 5 km E, 6 km S Puyo,
925 m, MCZ 92022-26; 1 km W Puyo, 1000
m, MCZ 92029; Rio Alpayacu, 1 km E Mera,
1080-1100 m, KU 120181, 120204; Sarayacu,
400 m, KU 120254; Veracruz, 950 m, KU
120203, USNM JAP 6145, 6147; 10 km ESE
Veracruz, MCZ 90143-44; Prov. Tungurahua:
El Topo, 1280 m, BM 1912.11.1.58-60 (re-
registered as 1947.2.17.2-4); Rio Negro, 1260
m, KU 120211-14, USNM JAP 6075, 6077,
6079-81, 6083-87.

Eleutherodactylus trepidotus (112 spec.)

ECUADOR, Prov. Napo: Cuyujia, 2360
m, USNM JAP 5214-15, 5609-14; 2.5 km
WNW Cuyujta, 2550 m, KU 143434; Laguna
de Papallacta, 3330 m, KU 143435-40; Laguna
del San Marcos, NE of Cayambe, CAS-SU
8300; Llanganati mountains, El] Golpe, 3000 m,
CAS-SU 13191-92, 17428; Papallacta, BM 1926.-
12.4.1 (reregistered as 1947.2.15.47), MSNT
428(9), 819, UIMNH 55869-91 (55874 cleared
and stained skeleton), 55916-17; 3.4 km ESE
Papallacta, Quebrada San Pedro, 2960 m, KU
177882; 1 km W Papallacta, 3155 m, KU
106938-42, USNM_ 160940-43, 160945-46,
106948-49, 160951-53, 106960, 106962-65,
106969, 164399-406, JAP 5153-54; 2 km W
Papallacta, 3270 m, KU 177883-87; 4 km W
Papallacta, 3300 m, KU 117618-22, 118134-35
(cleared and stained skeletons); 4.7 km W
Papallacta, 3360 m, KU 177881; 5 km W
Papallacta, 3250 m, USNM 164407-11; east
slope, Paso de Guamani, 3650 m, KU 109061-
63; Rio Bermejo, Cordillera de Guacamayo, KU

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 83

106937, WCAB 35641; Prov. Tungurahua: 10
km W Banos, 2925-3050 m, USNM JAP 5860.

Eleutherodactylus unistrigatus
(29 spec. )

Only specimens from the Amazonian slopes are
listed below; Lynch (ms) will report highland
material later. More than 1300 highland speci-
mens were also examined.

COLOMBIA, Depto. Putumayo: Coldén,
2220 m, KU 168618-24; 4 km SE San Fran-
cisco, 2320 m, AMNH 84837-38.

ECUADOR, Prov. Carchi: El Carmelo,
2710 m, KU 177521-28; 5.7 km NW EI Car-
melo, 2910 m, KU 177529-32; Prov. Napo:
ca. 25-30 km SE Cayambe, 3200 m, CAS-SU
8280; Santa Barbara, USNM GOV 7183, 7200,
7207; Prov. Tungurahua: Banos, 1900 m, CAS-
SU 5083, KU 141379-80, UIMNH 55722.

Eleutherodactylus ventrimarmoratus
(31 spec. )

ECUADOR, Prov. Morona-Santiago: Ashura
village on Rio Macuma, 300 m, AMNH 94685-
86; General Plaza, 1100 m, USNM JAP 6954,
6956; Miazal, USNM GOV 9657; Prov. Napo:
Loreto, USNM GOV 9364-65, JAP 3854; Rio
Azuela, 1740 m, EP 68/976; Prov. Pastaza:
Abitagua, 1100 m, CAS-SU 5078, UMMZ
92131-33; Abitagua, 8 kn NW Mera, 1300 m,
KU 119810-11; Mera, 1140 m, KU 119805-06,
119809; near Mera, 1000 m, UMMZ 92127;
Rio Alpayacu, 1 km E Mera, 1100 m, KU
119807-08; Prov. Tungurahua: El] Topo, 1280
m, BM 1911.1.1.51-53 (reregistered as 1947.-
254-16).

PERU, Depto. Amazonas: headwaters of
Rio Caterpisa, Manseriche range, 457 m,
AMNH 42435; Depto. Hudnuco: Finca Pan-
guana, Rio Llullapichis, 4-5 km upstream from
Rio Pachitea, 200 m, KU 154801; Depto. Junin:
Chanchamayo, BM 1911.12.12.77 (reregistered
as 1947.2.15.73); Depto. Loreto: Rio Uto-
quinia, AMNH 43376; Tipishca, opposite Con-
tamana, AMNH 42938; Depto. Madre de Dios:
Cocha Cachu, Rio Mant between Rio Cachiri
and Rio Panagua, 400 m, KU 154803; Manu,
365 m, KU 154802.

Eleutherodactylus w-nigrum
(377 spec. )

Only specimens from the Amazonian slopes are
listed below. 1444 other examples have been
examined from Colombia and Ecuador.
COLOMBIA, Depto. Cundinamarca: east
base of Cordillera de Bogota, ANSP 24394;
Depto. Putumayo: 10.3 km W E] Pepino, 1440

m, KU 168735-41; Mocoa, CAS 85176; ca. 15
km (airline) SW Mocoa, 1180-1480 m, AMNH
84851-52; 4 km (airline) SE San Francisco,
2320 m, AMNH 84839-46; 35 km SE San
Francisco, 1950 m, KU 168721-34.
ECUADOR, Prov. Morona-Santiago: Agua
Rica, 1890 m, USNM JAP 7072; El Cruzado,
2195 m, USNM JAP 7209-17; Gallineros, 2195
m, USNM JAP 7226-27; Loma de Puerco, 2226
m, USNM JAP 7334-47, 7358; between Loma
de Puerco and Sapote, 2317-2408 m, USNM
JAP 7401-05, 7322-33; Pailas, 2195 m, USNM
JAP 6546-53, 6662, 6675-81, 6725, 6777-93;
between Pailas and Mirador, 1982-2195 m,
USNM JAP 6705, 6707-10; San Juan Bosco,
2195 m, USNM JAP 7066-69, 7083-86, 7088,
7223-25; slightly downtrail from San Juan
Bosco, 2113 m, USNM JAP 7256-59; slightly
downtrail from San Juan Bosco, 2192 m, USNM
JAP 7229-33; between San Juan Bosco and
El Cruzado, 2226 m, USNM JAP 7177-91,
7193-96; Sapote, 2470 m, USNM JAP 7507-11,
7542-60; 1 km W Sapote, 2564-2591 m, USNM
JAP 7411-16; between Sapote and Suro Rancho,
2604 m, USNM JAP 7762; Prov. Napo: Alto
Rio Quijos, E] Chaco, 1600 m, USNM GOV
8007; Baeza, 1870 m, KU 146063-64, USNM
JAP 5728-29; Borja, 1707 m, USNM JAP 5386,
5437, 5442; 6.6 km ESE Cuyujia, Rio Mo-
lando, 2360 m, KU 146066-70; 7 km E[SE]
Cuyujia, 2195 m, USNM JAP 5207, 5359;
9 km E[SE] Cuyujua, 2104 m, USNM JAP
5219, 5319; La Alegria, 1900 m, USNM JAP
4608-09, 4621-22, 4626-28, 4662-63, 4665-80,
4718-19, 4721-30, 4732-48, 4848-57, 4897-
99; La Bonita, 1650 m, USNM JAP 4715-
17, 4858-60, 4915; Loreto [probably in error],
USNM GOV 7090; 3.2 km NNE Oritoyacu,
1910 m, KU 177568-81; Papallacta, WCAB
37923, 37930; Rio Azuela, 1740 m, KU 143303-
16, 165659-62; Rio Jatuntinahua, 2160 m, KU
165683-88; 2 km SSW Rio Reventador, 1490
m, KU 165663-80, 166286 (cleared and stained
skeleton); Rio Salado, 1 km upstream from
Rio Coca, 1410 m, KU 146065, 165681-82,
177582-91; Santa Barbara, 2591 m, USNM
GOV 8542-43, 8545, JAP 4542-54, 4556; 0.5
km NW Santa Barbara, 2540 m, KU 177592;
1 km SW Santa Barbara, 2591 m, USNM JAP
4586-87; between Santa Barbara and L’Ale-
gria, 2134-2439 m, USNM JAP 4592-96; 16.5
km NNE Santa Rosa, 1700 m, KU 143301-02;
between Sebundoy and La Bonita, 1524-1829
m, USNM JAP 4630-31; Prov. Pastaza [prob-
ably in error]: Rio Villano, USNM JAP 6257-59,
6261-62; Prov. Tungurahua: Abitagua, 1100 m,
UMMZ 92119 (one individual); 18.5 km E
Banos, 1600 m, KU 141291; Prov. Zamora-Chin-
chipe: 23 km E Loja, 2220 m, KU 141966.

84 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

APPENDIX B: GAZETTEER

Following is a list of localities on the east-
ern slopes of the Andes where specimens of
Eleutherodactylus were collected. After each
place name the province is given in paren-
theses, followed by geographic coordinates and
elevation. Coordinates were determined from
the 1974 Edition of Mapa de Ecuador, Instituto
Geografico Militar, Quito (1:1,000,000). Most
of the elevations were determined from altim-
eter readings at the sites. When known, spe-
cific sites and habitats are given, followed by
names of collectors and months and years that
they collected at the sites. Names of some
collectors are abbreviated: JAP = James A.
Peters, JDL = John D. Lynch, WED = Wil-
liam E. Duellman.

Abitagua (Pastaza)—01°26’ S, 78°07’ W,
1300 m. Region on Banos-Puyo road in Pastaza
Trench, 8 km NW of Mera. Small streams in
lower montane rainforest. Wm. Clarke-Mac-
Intyre, Sep. 1939; JDL, Jul. 1968, Jul. 1977.
32

Agua Caliente (Napo)—00°22’ S, 77°52’
W, =~ 3000 m. Place on the Rio San Pedro,
a tributary of the upper Rio Azuela on the
eastern slope of Volcan Cayambe. I. L. Wig-
gins, Jul. 1944. 5

Alpayacu, Rio (Pastaza)—01°28’ S, 78°05’
W, 1100 m. Small tributary of Rio Pastaza,
1 km E Mera; remnants of lower montane rain-
forest. Wm. Clarke-MacIntyre, Sep. 1939; JDL,
Jul. 1968, Jul. 1977. 34

Azuela, Rio (Napo)—00°07’ S, 77°37’ W,
1740 m. Tributary of Rio Coca east slope of
Volcan Cayambe. Collections from slopes above
(NW) of point where Papallacta-Lago Agrio
road crosses river; lower montane rainforest
(1971), partly cultivated in naranjilla (1974).
WED, Oct. 1971, Oct. 1974, Mar. 1975; C. W.
Myers, Feb. 1974; F. Ortiz C, Jan. 1976. 8

Baeza (Napo)—00°28’ S, 77°53’ W, 1990
m. Town on bluff just southwest of confluence
of Rio Papallacta and Rio Cosanga to form Rio
Quijos. Collections from vicinity of town in
partly cleared lower montane rainforest; many
large trees with abundant bromeliads. JAP,
Jul. 1962; WED, Oct. 1971, Apr. 1972. 17

Banos (Tungurahua )—01°23’ S, 78°24’ W,
1800 m. Town in upper part of Pastaza Trench;
dry subtropical forest. Collections from vicinity
of town and valley (many labelled only as
“Banos”) by Wm. Clarke-MacIntyre Jul. 1937,
and Olallas, 1932-43; WED, Jul. 1971, Apr.
1972. Collections in partly cleared lower mon-
tane rainforest, 8.4 km E of Banos, 1590 M,
WED, Apr. 1972. Collections from grazed
subalpine woodland, 10 km W of Banos, 3000
m, WED, Jul. 1972. 27

Bermejo, Rito (Napo)—00°31’ S, 77°53’
W, 1910 m. River draining north slope of
Cordillera de Guacamayo; Baeza-Tena road
crosses river 12.5 km S of Baeza, at settlement

of Bermejo at junction of Rio Bermejo and
Rio Cosanga. 19

Borja (Napo)—00°25’ S, 77°51’ W, 1710
m. Village just west of Rio Quijos, just off of
Papallacta-Lago Agrio road; mostly cleared
lower montane rainforest with scattered large
trees in pastures. JAP, Jul. 1962. 15

Cerro Negro (Azuay)—02°45’ S, 78°32’
W, 2925 m. Place just east of Sevilla de Oro
on trail to Méndez; subalpine rainforest. JAP,
Aug. 1962. 39

Chalpi (Napo)—00°23’ S, 78°04’ W, 2590
m. Finca 8.3 km ESE of Papallacta on road
to Lago Agrio; pastures amidst remnants of
upper montane rainforest, JDL, Jul. 1977. Col-
lections from short distances east and west,
JAP, Jul. 1962. 13

Cutucti, Cordillera (Morona-Santiago )——~
02°45’ S, 78°05’ W, 2000+ m. Isolated moun-
tain range east of Méndez. E. Feyer, 1921. 41

Cuyujtia (Cuyuja) (Napo)—00°24’ S, 78°
02’ W, 2460 m. Small village 17 km ESE of
Papallacta on road to Lago Agrio. Pastures
with remnants of upper montane rainforest in
ravines. Collections from vicinity of village,
JAP, Jul. 1962; from 7.8 km WNW (2750 m),
3 km E (2135 m), Rio Victoria, 1 km E (2420
m), and Rio Molando, 6.6 km ESE (2360 m),
WED, Oct. 1971, Apr. 1972. 14

Dué, Cordillera del (Napo)—00°03’ S, 77°
35’ W, 1150 m. Ridge between Rio Coca (S)
and Rio Dué (N); lower montane rainforest.
WED, Aug. 1968. 6

El Carmelo (El Pun) (Napo)—00°40’ N,
77°37’ W, 2750 m. Small town on upper Rio
Chingual; patches of upper montane rainforest
in otherwise cultivated areas; JAP, Jun. 1962;
JDL, Jul. 1977. Same habitat at 5.7 kn NW
of El Carmelo on road to Tulcan; JDL, Jul.
1ST tf

El Cruzado (Morona-Santiago )—03°03’ S,
78°31’ W, 2195 m. Ranch on trail from Guala-
ceo to General Plaza, 0.5 hr up trail from San
Juan Bosco; “montane forest”; JAP, Aug. 1962.
49

El Golpe (Napo)—01°10’ S, 78°20’ W,
3000 m. Place name (along Rio El Golpe ?)
on higher eastern slope of Los Llanganatis. R.
Blomberg, Dec. 1951. 25

El Topo (Tungurahua)—01°25’ S, 78°10’
W, 1220 m. Settlement 32 km E of Banos on
road to Puyo; lower montane rainforest, citrus
groves, and pasture. M. G. Palmer, 1912. 30

General Plaza (Morona-Santiago )—02°58’
S, 78°26’ W, 1100 m. Town on Rio Yunganza;
lower montane rainforest. JAP, Aug. 1962. 43

Jattintinahua (Jattin Tinajua), Rio (Napo)—
00°26’ S, 77°57’ W, 2160 m. Tributary of Rio
Papallacta, 27.2 km ESE of Papallacta on road
to Lago Agrio; upper montane rainforest and
pasture. WED, Apr. 1975. 16

Jorge, Rio (Napo)——~01°08’ S, 78°17’ W,

ELEUTHERODACTYLUS OF THE ECUADORIAN ANDES 85

3000 m. Stream on high eastern slopes of Los
Llanganatis. R. Blomberg, Dec. 1951. 24

La Alegria (Napo)—00°33’ N, 77°31’ W,
1930 m. Settlement on Rio Chingual, 20 km N
of La Bonita. Presumably lower montane rain-
forest. JAP, Jun. 1962. 3

La Bonita (Napo)—00°29’ N, 77° 32’ W,
1920 m. Small village on Rio Chingual; pre-
sumably lower montane rainforest. JAP, Jun.
1962. 4

Loma de Puerco (Morona-Santiago )—03°
02’ S, 78°32’ W, 2225 m. Rancho between
Sapote and El] Cruzado on Gualaceo-General
Plaza trail. JAP, Aug. 1962. 45

Los Llanganatis (Tungurahua, Napo)—~
01°10’ S, 78°12’ W, 4000+ m. A highland
region immediately to the north of the Pastaza
Trench. R. Blomberg, Dec. 1950, Dec. 1951.
26

Macas (Morona-Santiago )—02°19’ S, 78°
07’ W, 1015 m. Town on Rio Upano in foot-
hills of Andes. Cultivated with few remnants
of rainforest. Olallas. 36

Méndez (Morona-Santiago )—02°43’ S, 78°
19’ W, 550 m. Town on the lower Rio Paute.
Collections from various elevations between
Méndez and Sevilla de Oro (2550 m). JAP,
Aug. 1962. 37

Mera (Pastaza)—01°28’ S, 78°06’ W, 1000
m. Town in lower Pastaza Trench, 18 km
WNW of Puyo on road to Bafios. Cultivated
with remnants of rainforest in ravines and on
steep slopes N of town; JDL, Jul. 1968, Jul.
1977. Collections from forested ravines 6.6 km
NW (1200 m), 8.1 km NW (1270 m), and
9.5 km NW (1270 m) of Mera on road to
Banos; WED, Apr. 1972. 33

Mirador (Morona-Santiago )—02°44’ S, 78°
22’ W, 1982 m. Rancho on Sevilla de Oro-
Méndez trail. JAP, Aug. 1962. 38

Mirador (Tungurahua)—01°26’ S, 78°15!
W, 1530 m. Finca on N slope of Rio Pastaza,
8.4 km E of Banos on road to Puyo. Second
growth lower montane rainforest and cultivated
fields. Olallas, P. O. Simons, 1899. 31

Oritoyacu (Napo)—00°29’ S, 77°52’ W,
1910 m. Settlement at crossing of Baeza-Tena
road of Rio Oritoyacu, 10.1 km S of Baeza.
Collections from Quebrada Amarilla, 3.2 km
NNE Oritoyacu; lower montane rainforest.
JDL, Jul. 1977. 18

Pailas (Azuay )—02°45’ S, 78°31’ W, 2195
m. Rancho on Sevilla de Oro-Méndez trail.
JAP, Aug. 1962. 40

Papallacta (Napo)—00°20’ S, 78°05’ W,
3150 m. Village near headwaters of Rio Pa-
pallacta; subalpine rainforest. Collections from
immediate vicinity of village; JAP, Jul. 1962,
JDL, Jul. 1967, Feb. 1968, Jul. 1977; WED,
Mar. 1967, Oct. 1971. Collections from Que-
brada San Pedro, 3.4 km ESE (pastures), JDL,
Feb. 1968, Jul. 1977; upper montane rainforest
and pastures, 9.2 km ESE, JDL, Jul. 1977;

11-12 km ESE, R. Altig, Dec. 1973, Jan. 1974;
WED, Oct. 1974, Mar.-Apr. 1975; Finca Santa
Fe, 13.1 km ESE, JDL, Jul. 1977. 12

Paracayacu (Rio Parcayacu ?) (Napo)—01°
07’ S, 78°15’ W, 2700 m. Stream on high
eastern slope of Los Llanganatis. R. Blomberg,
Dec. 1951. 23

Piuntza, Rio (Morona-Santiago )—03°52’ S,
78°27' W, 1830 m. Stream on western slope
of northern part of Cordillera del Condor, flow-
ing into Rio Zamora about 30 km SSW of
Yuquianza. Collections from upper montane
rainforest near crest of cordillera. J. E. Sim-
mons, Jan. 1972. 51

Plan de Milagro (Morona-Santiago )—03°
03’ S, 78°29’ W, 1710 m. Rancho about 10
km WSW of General Plaza on trail to Guala-
ceo. JAP, Aug. 1962. 50

Puyo (Pastaza)—01°29’ S, 77°59’ W, 1000
m. Town in Andean foothills; pastures, citrus
groves, and tea plantations. Olallas; JAP; WED,
Apr. 1972. 35

Reventador, Rio (Napo)—00°07’ S, 77°38
W, 1490 m. Tributary of Rio Coca draining
south slope of Volcdn Reventador. Collections
from vicinity of two small streams, 2 km WSW
of Puente Reventador on Papallacta-Lago Agrio
road; lower montane rainforest on steep slopes.
R. Altig, Dec. 1973; WED, Oct. 1974, Mar.
1975.7

Rio Negro (Tungurahua)—01°25/ S, 78°13’
W, 1240 m. Village in Pastaza Trench, 29.7
km E of Banos on road to Puyo. Collections in
secondary forest and abandoned naranjilla plan-
tations in immediate vicinity of village; JDL,
Jul. 1968. Collections from citrus groves and
extensively disturbed lower montane rainforest
11 km E of village (1170 m); WED, Apr.
1972. 29

Runttin (Tungurahua)—~01°24’ S, 78°35/
W, 1840 m. Northeast shoulder of Volcan Tun-
gurahua, south of Banos. Wm. Clarke-MacIn-
tyre, Jun. 1939. 28

Sabanilla (Zamora-Chinchipe )—03°57’ S,
79°03’ W, 1750 m. Village on Loja-Zamora
road. Badly cutover lower montane rainforest.
H. E. Anthony, Nov. 1920. 52

Salado, Rio (Napo)—00°13’ S, 77°44’ W,
1410 m. River draining east slope of Volcan
Cayambe. Collections made in vicinity of small
settlement of Rio Salado at point where Papal-
lacta-Lago Agrio road crosses Rio Salado, about
1 km upstream from its confluence with the
Rio Coca; lower montane rainforest with
swamp and many small streams. WED, Apr.
1972, Oct. 1974, Mar. 1975; JDL, Jul. 1977.
10

Salto de Agua (Napo)—00°08’ S, 77°38’
W, 1660 m. Waterfall on Papallacta-Lago
Agrio road, 2.5 km NNE of Puente Reventador;
upper montane rainforest. WED, Apr. 1972. 9

San José (Napo)—00°36’ S, 77°35’ W,
1000 m. Finca owned by Olalla family on

86 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

lower eastern slopes of Volcan Sumaco. Olallas,
1920’s-1950’s. 21

San José, Rio (Napo)—01°07’ S, 78°15’ W,
3000 m. Stream on high eastern slopes of Los
Llanganatis. R. Blomberg, Nov. 1950. 22

San Juan Bosco (Morona-Santiago )—03°03’
S, 78°31’ W, 2195 m. Rancho on Gualaceo-
General Plaza trail. JAP, Aug. 1962. 48

Santa Barbara (Napo)—00°35’ N, 77°33!
W, 2625 m. Town on upper Rio Chingual.
Upper montane rainforest and pastures (1962),
but essentially cleared (1977). Collections from
vicinity of town; JAP, Jun. 1962; JDL, Jul.
1977. Collections from 3.7 km NW of Santa
Barbara on road to Tulcan (2410 m); JDL,
Jul. 1977. 2

Santa Rosa (Napo)—00°18’ S, 77°43’ W,
1600 m. Village on Rio Santa Rosa just up-
stream from Rio Coca, 61 km from Papallacta
on road to Lago Agrio. Collections from 16.5
km by road NNE of Santa Rosa, 1700 m; lower
montane rainforest on upper slopes of Rio
Salado valley; WED, Oct. 1971. 11

San Vicente (Morona-Santiago )—03°02’ S,
78°33’ W, 2835 m. Rancho on Gualaceo-
General Plaza trail; subalpine rainforest. JAP,
Aug. 1962. 44

Sapote (Morona-Santiago )—03°02’ S, 78°
32’ W, 2470 m. Rancho on Gualaceo-General

Plaza trail; patches of upper montane rainforest
amidst pastures. JAP, Aug. 1962. 47

Sevilla de Oro (Azuay)—02°47’ S, 78°34!
W, 2650 m. Village on upper Rio Negro on
trail from Paute to Méndez; collections from
various elevations between Sevilla de Oro and
Méndez. JAP, Aug. 1962. 42

Sumaco, Cerro (Napo)—00°35’ S, 77°37’
W, 3900 m (crest). Isolated volcano rising
from Andean foothills. Olallas, 1920’s-1950’s.
20

Suro Rancho (Morona-Santiago )—03°02’
S, 78°32’ W, 2745 m. Rancho on Gaulaceo-
General Plaza trail. JAP, Aug. 1962. 46

Zamora (Zamora-Chinchipe )—04°05’ S, 78°
57’ W, 1000 m. Town in Rio Zamora valley.
Collections from extensively cleared lower mon-
tane rainforest on eastern slope of Andes, 22
km NW of Zamora on road to Loja (1730 m).
WED, Jul. 1971. 54

Zamora, Abra de (Zamora-Chinchipe )—04°
00’ S, 79°07’ W, 2800 m. Crest of Cordillera
Oriental, 13.5 km E of Loja on road to Zamora;
also known as Nudo de Sabanilla. Subalpine
forest with many large bromeliads on low
bushes; JDL, Jun. 1968, Jan. 1978; WED, Jul.
1971, Mar. 1975. Collections from eastern
slope, 15 km E of Loja (2710 m); WED, Jul.
1971; 23 km E Loja (2220 m); WED, Jul.
1971. 53

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RECENT MISCELLANEOUS PUBLICATIONS
UNIVERSITY OF KANSAS MUSEUM OF NATURAL HISTORY

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