UNi

HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

OCCASIONAL PAPERS ^

of the ^D

MUSEUM OF NATURAL HISTORY
The University of Kansas
Lawrence, Kansas

NUMBER 35, PAGES 1-51 APRIL 16, 1975

A REVIEW OF THE ANDEAN LEPTODACTYLID
FROG GENUS PHRYNOPUS

By
John D. Lynch^

In 1932 Hampton Parker described Etipsophus wettsteini from
the Peruvian Andes. In discussing the systematic position of this
species he pointed out the then ominous possibiHty that Eleuthero-
dactyltis, Etipsoplnis, and Sijnlwphus might eventually be merged
into a single genus. Parker envisioned a "Borborocoetes" group of
Eleutherodactijlus as having given rise to the annectant SyrrJwpus
simonsii, S. festae, and Eupsophiis icettsteini, and that this complex
of species had given rise to the more southern Etipsoplnis (espe-
cially E. penianus). Although he separated the species into two
genera, Parker evidently viewed them as an annectant generic
group. Subsequently, these three species and several others were
gathered into the genus Niceforonia (Lynch 1971).

Lynch ( 1971 ) recognized ten genera of leptodactylids in the
trilpe Eleutherodactylinii ( Telmatobiinae ) . Four of the genera
{Eleutherodactijlus, Sminthilhis, Syrrliophus, and Tomodoctyhis)
were characterized, in part, by the possession of digital discs and
the associated T-shaped terminal phalanges. The other six genera
{Amhlyphrynus, Euparkerella, Holoaden, Hylactophryne, Ischnoc-
nema and Niceforonia) do not have digital discs and with the ex-
ception of Euparkerella (which has complex terminal phalanges)
have knobbed terminal phalanges.

Only three eleutherodactyline genera have species represented
in the Andean frog fauna at elevations above 2000 meters. The
genus Amhlyphrynus is represented by a single species in Andean

1 Associate Professor, School of Life Sciences, The University of Nebraska,
Lincoln, Nebraska; Research Associate in Herpetology, Museum of Natural
History, The University of Kansas.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Colombia. Frogs of this genus have broad heads ( head width 50-60
% SVL) and simple digits (no pads or discs; terminal phalanges
knobbed). The genus Eleutherodactijlus is represented by many
species in the Colombian and Ecuadorian Andes and few species
elsewhere. These frogs have heads of normal width (HW 30-42%
SVL) and complex digits (discs on narrowly to broadly dilated pads;
terminal phalanges T-shaped ) .

A third supraspecific group is represented by at least two Co-
lombian, three Ecuadorian, eight Peruvian, and one Bolivian species.
These frogs have heads of normal breadth and simple digits; in-
cluded among the fourteen species are the type-species of the
nominal genera Niceforonia Coin and Cochran, 1963, Nohlella Bar-
bour, 1930, and Phrynopiis Peters, 1874. I consider the fourteen
species congeneric and a distinct generic entity; accordingly, Nice-
foronia and Nohlella are here relegated to the synonymy of Phryno-
pus.

In recent years, Niceforonia has been applied to some of the
nominate species of Phrynoptis. Since 1932, Phrynopus penianus
Peters has been considered a species of Eupsophus (Parker, 1932)
and since 1971 the type-species of Nohlella has been considered a
species of Eleutherodactylus (Lynch, 1971). Phrynoptis penianus
and Niceforonia nana so closely resemble one another that their
congeneric status cannot be seriously questioned. The third type-
species (Nohlella peruviana) is more distinctive and is treated
separately below.

Presently, ten names are applied to the species of Phrynopus
(nine are listed as Niceforonia by Lynch, 1971). One of the names
{N. festae Peracca) was incorrectly applied to the species of Phryn-
opus found in the upper Papallacta valley in Napo Province, Ecua-
dor by Lynch (1968).

Considerable material of this genus has accumulated since my
1968 paper; that material requires the recognition of four previously
unknown species and enables me to review the genus as an entity
and to discuss the variation in osteological character-states seen
among the species of the genus. Two hundred and eighty-two
specimens including 17 cleared and stained skeletons were studied.

ACKNOWLEDGMENTS

For loan of species or provision of working space at their respec-
tive institutions, I am indebted to Jose M. Cei, the late Doris M.
Cochran, William E. Duellman, Josef Eiselt, Alice C. C. Grandison,
Arnold G. Kluge, the late James A. Peters, Douglas Rossman,
Richard Thomas, Charles F. Walker, Ernest E. Williams, George
Zug, and Richard G. Zweifel.

Field work in Ecuador was supported by grants from the Com-
mittee on Systematic and Evolutionary Biology, The University of
Kansas, Sigma Xi, and the Watkins Fund of the Museum of Natural

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 3

History, The University of Kansas. Travel to museums was sup-
ported by the University of Kansas Graduate School during my
tenure as an Honors Fellow ( 1966-69 ) and The University of
Nebraska Research Council. Robert Henderson and Marsha Lynch
aided me in the field and William E. Duellman and Thomas H.
Fritts provided detailed field notes and additional information about
localities and the organisms they had collected. Todd Georgi aided
in histological study of the pectoral girdle. The aid of all these
persons has substantially contributed to my study of these frogs.
Abbreviations for collections used throughout the text are:

AMNH American Museum of Natural History
BMNH British Museum ( Natural History )
IBM/UNC Instituto de Biologia, Mendoza-Universidad

Nacional de Cuyo
KU The University of Kansas Museum of Natural

History
LSUMZ Louisiana State University Museum of Zoology
MCZ Museum of Comparative Zoology

MNW Naturhistorisches Museum zu Wien

UMMZ University of Michigan Museum of Zoology
USNM United States National Museum ( National

Museum of Natural History)
WCAB Private collection of Werner C. A. Bokermann,

Sao Paulo.

STRUCTURE OF THE DIGITS

Eleutherodactylus is a large and diverse genus; against this back-
ground of morphological diversity, Fhnjnopus can be distinguished,
at present, only on the basis of the structure of the digits. The
majority of Eleutherodactylus species have dilated digital pads
(bearing discs on their ventral surface) supported internally by
distinctly T-shaped terminal phalanges. The disc is a distinct
structure on the ventral surface of the digit and is defined by a
circumferential groove ( Fig. 1 ) . In most species having apical dila-
tion of the digit, the pad is wider than long and the dilation ratio
(pad width/digit width below pad) is 2.0-3.5. This value is lowest
when the digits bear lateral fringes (as in E. riveti). Several Eleu-
therodactylus species have narrow digits, that is, the pad is longer
than wide or the length and width are equal; the dilation ratio is
as low as 1.2-1.3. The narrow-toed Eleutherodactylus include those
with narrow finger and narrow toe pads {e.g., E. blnotatus, E. cor-
nutus, E. elassodiscus, E. octavioi, and several others) as well as
those with narrow finger pads and broad toe pads ( e.g., E. anoma-
lus). In most of the narrow-toed species, the terminal phalanges
of the fingers are more knobbed (as contrasted to T-shaped)
whereas those of the toes are more T-shaped (Lynch, 1971). The

4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

most proximal digits (I and sometimes II of the hand; I and II of
the foot) may not bear discs {i.e., no circumferential groove pres-
ent) but the more distal digits always bear discs, even in the most
narrow-toed species (e.g., E. elassodiscus, E. nigrovittatus, and
E. octavioi; Fig. 1 ) .

The simple-toed Andean frogs of the genus Fhrynopus do not
have digital discs (Fig. 1). There is apical swelling (r= bulbing)
of the digit in some of these frogs and dilation ratios of 1.0-1.2.
Earlier (1971), I characterized this genus (as Niceforonia) as having
knobbed terminal phalanges. However, some of the Peruvian taxa
have lateral processes on the terminal phalanges (Fig. 2) that are
intermediate in size between the T-shaped terminal phalanges of
some of the narrow-toed Eleutherodactylus and the knobbed char-
acter-state seen in Fhrynopus peraccai, P. favomaciihitu.s, and P.
montium.

The annectant condition seen in some Peruvian species ( no discs,

Fig. 1. — Dorsal views of distal portion of (A) fourth toe, left foot, Fhrynopus
peruvianus, (B) fourth toe, left foot, Eleutherodactyhis elassodiscus, (C)
fourth finger, left hand, E. riveti. Ventral views of same digits, (A') P. peru-
vianus, (B') E. elassodiscus, (C) E. riveti. Lateral views of same digits,
(A") P. peruvianus, (B") E. elassodiscus, (C") E. riveti. Fhrynopus peru-
vianus, KU 138959; Eleutherodactylus elassodiscus, WCAB 37915; E. riveti,
KU 120060. Lines below lateral views equal 1 mm.

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 5

rudimentary or vestigal T-shaped terminal phalanges) could be
used as evidence for combining several of the simple-toed genera
of elutherodactylines with Elentherodactyhis as once noted by
Parker (1932). To do so, howexer, does not simplify or improve
leptodactylid taxonomy. Although the phalangeal characteristic is
no longer diagnostic, the digital disc characteristic remains diag-
nostic; accordingly, I prefer to retain the simple-toed genera as
distinct from Eleutlierodactyliis.

THE STATUS OF NOBLELLA BARBOUR

Noble (1921) named SminthUhis penwianus on the basis of five
specimens from Juliaca, Puno, Peru. The species was designated the
type-species of NohJeUa by Barbour (1930), who neither diagnosed
nor defined the genus. Lynch ( 1971), following study of a partially
macerated, cleared and stained paratype of pertiviano, placed it in
the genus Eleutherodaciijhis largely because a single T-shaped
terminal phalange was found among the fragments of the skeletal
preparation.

Ill

V

Fig. 2. — Terminal phalanges of Phrtjnopus parkeri (Row A, KU 135306),
P. pemvianus (Row B, figure at far left, KU 138928; other five phalanges of
KU 138926), and P. cophites (Row C, KU 138909). Column to left (no
roman numeral ) is of tliird finger. The five columns headed by roman numerals
include terminal phalanges on toes I tlirough V.

6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Collections made in southern Peru by William E. Duellman and
Thomas H. Fritts include a large series of adults and subadults of
a simple-toed eleutherodactyline frog. Comparison of the smallest
of these with the holotype of SmintliiUus peruviana convinced me
that they are conspecific. Alizarin-Red preparations of adults of the
frog revealed that the terminal phalanges are roughly T-shaped
(Fig. 2) even though the digit tips are simple (Fig. 1). Noble
(1921) assigned the species to SminthiUiis and the Brachycephalidae
because the pectoral girdle is arcifero-firmisternal in the sense of
Boulenger and Cope. Griffiths (1959) argued that the girdle archi-
tecture is arciferal in the sense that he diagnosed the character-
state; the adult specimens are functionally firmisternal in that the
halves of the epicoracoid cartilages are fused (Fig. 3) but are
anatomically arciferal as in all other leptodactylid frogs. The clavi-
cles are oriented at approximately right angles to the sagittal plane
in contrast to the strongly curved clavicles seen in other species of
the genus Phrijnopus.

Character-states for the characteristics used in my generic re-
view of the leptodactyhds (Lynch 1971) are listed below for
Nohlella peruviana: sternum, a cartilaginous plate; omosternum
present, relatively small and elongate; eight procoelous presacral
vertebrae, non-imbricate; cervical vertebra with widely separated
cotyles, not fused to second vertebra; sacral diapophyses rounded,
deflected posteriorly; sacro-coccygeal articulation bicondylar; verte-
bral shield lacking; transverse processes of vertebrae three to eight
about as long as sacral diapophyses or slightly shorter; cranial bones
not exostosed or dermostosed; maxillary arch complete; maxilla and
premaxilla toothed, teeth blunt, pedicellate; palatal shelf of pre-
maxilla broad, weakly incised; facial lobe of maxilla relatively shal-
low; nasals small, median separation moderate; frontoparietals
complete, fontanelle not exposed, not ornamented; frontoparietal
not fused to prootic, epiotic eminences obsolete; cristae paroticae
short, broad; occipital artery passes dorsal to skull bones; zygomatic
ramus of squamosal short, widely separated from maxillary arch;
otic ramus of squamosal short, otic plate narrow; plectrum (col-
umella) present; prevomers small, edentate, dentigerous rami lost;
sphenethmoid entirely ossified, large, extending anteriorly to middle
of nasals; anterior ramus of parasphenoid short, broad, not keeled;
parasphenoid alae oriented at right angles to anterior ramus, widely
separated from short, median rami of pterygoids; pterygoids mod-
erately large, anterior rami extending nearly to palatines; occipital
condyles small, stalked, widely separated; mandibles lacking odon-
toids; phalangeal formulae 2-2-3-.3, 2-2-3-^3; terminal phalanges
bear short, irregular-shaped lateral processes; digits bulbous api-
cally, lacking discs; alary processes of hyoid plate on narrow stalks;
m. sternohyoideus and m. petrolujoideus anterior insert on lateral
edges of hyoid plate; pupil horizontal; males with median subgular

J

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS

Fig. 3. — Ventral view of pectoral girdle of Phrynopus peruvianus. Cartilage
is stippled. (A) Cross-section through epicoracoidal bridge at Point A — A.

vocal sac; toes not webbed, no lateral fringes; two metatarsal tu-
bercles present; eggs large, not pigmented, few in number; males
lacking nuptial asperities.

The type-species of Nicefownia and Phrynopus differ from one
another in skin texture (venter areolate in N. nana), condition of
the ear (plectrum, cavum tympanicum, and tympanic annulus ab-
sent in N. nana), and in coloration, proportions, and some details
of foot structure. They are similar in having widely separated
cervical cotyles ( and occipital condyles ) , in having the frontoparie-
tals complete (fontanelle not exposed), in having prevomerine
odontophores ( bearing teeth ) situated well posterior to the choanae,
in having simple digits, and in having "normal" arciferal pectoral
girdles.

The type-species of Noblella differs from the type-species of

8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Niceforonia and Phnjnopus in some tiivial features but also in lack-
ing prevomerine odontophores and teeth (as well as much of the
prevomers ) and in ha\'ing a pseudofirmisternal pectoral girdle ( epi-
coracoid cartilages partially fused; clavicles straight and at right
angles to sagittal plane ) . The absence of prevomerine odontophores
and teeth in periwiamis is not an especially significant tiait insofar
as generic distinction is concerned. Five other Bolivian and Peru-
vian species also lack prevomerine odontophores. However in these
frogs the dentigerous ramus of the prevomer has not been lost.

Thus, the only trait supporting a generic separation is the archi-
tecture of the pectoral girdle. In the absence of corroborative dis-
tinctions, I am unwilling to advocate separation of Noblella and
Plirynopus.

With two exceptions, the original descriptions of the named
species are based on adults. Werner's (1899) description of colum-
biantis is very brief and no specimens have been identified since
the original description. Noble's ( 1921 ) description of pemvianus
is incomplete in that it was based on juveniles. The acquisition of
a large sample of adults enables a redescription of the type-species
of Noblella.

SYSTEMATIC ACCOUNTS

The material available for seven of the species is too limited to
warrant a revision of the genus. The published descriptions of P.
favomaculatiis (Parker), P. laplacai (Cei), P. montium (Shreve),
and P. wettsteini ( Parker ) are considered adequate for future identi-
fication of samples when the diagnoses presented here are consid-
ered. I therefore present species descriptions for only the new
species and for P. peruanus Peters, P. pemvianus (Noble) and P.
simonsii ( Boulenger ) .

Phrynopus Peters

Phnjnopus Peters 1874:416 (Type-species by monotypy, Phrynopus peruanus
Peters, 1874).

Noblella Barbour 1930:81 (Type-species by original designation, Sminthillus
pemvianus Noble, 1921). New synonymy.

Niceforonia Coin and Cochran 1963:499. (Type-species by original designa-
tion, Niceforonia nana Coin and Cochran, 1963). New synonymy.

Diagnosis. — Eleutlierodactyline telmatobiine frogs with simple digital tips,
digits neitlier greatly shortened nor elongated, plantar surfaces lacking conical
or subconical supernumerary tubercles, having alary processes on the hyoid
plate, and having a "normal" skull architecture (i.e., not helmeted), and
normal head width (HW/SVL = 31.1-40.8%).

Content. — Fourteen species: P. hrtinneus new species, P. columhianus
(Werner), P. cophites new species, P. flavonmculatus (Parker), P. laplacai
(Cei), P. montium (Shreve), P. nanus (Coin and Cochran), P. parkeri new
species, P. peraccai new species, P. pereger new species, P. peruanus Peters,
P. pemvianus (Noble), P. simonsii (Boulenger), and P. wettsteini (Parker).

Description. — In dorsal view the snout is either rounded or

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 9

weakly acuminate. In lateral profile, the snout is sloping in favo-
maculatiis and parkeri ( Fig. 4 ) whereas it is rounded or truncate in
the other twelve species. The canthus rostralis is sharp and concave
in all species except hrunnens and peraccai which have rounded
canthi. The interorbital space is flat and cranial crests lacking in all
species except flavomoculatus. Three species (cophites, nanus, and
simonsii) are earless; the remaining eleven species have complete
ears although the tympana are concealed in loplacai, montium,
parked, peraccai, pereger, peruviamis, and luettsteini; six species
lack prevomerine teeth and odontophores; peruviamis lacks the
dentigerous processes of the prevomers; cophites, laplacai, montium,
and wettsteini have long thin dentigerous processes but lack odonto-
phores; simo7isii has large prevomers but lacks the odontophores;
the other eight species have prevomerine teeth and associated proc-
esses. The dentigerous processes lie posterior and median to the
choanae. In those species for which males are available, vocal sacs
and slits are absent in brunneus and cophites. Males are not known
for cohimbianus, nanus, peraccai, or simo7isii.

The skin of the dorsum and venter is smooth in brunneus, flavo-
macuhtus, parkeri, peraccai, pereger, peruanus, and peruviamis; the
skin of the dorsum is weakly to coarsely areolate and that of the
venter coarsely areolate in cophites, laplacai, montium, namis, simon-
sii, and loettsteini. Werner ( 1899 ) made no direct statements about
skin texture in cohimbianus, but because he compared his frog with
Ischnocnema quixensis, I conclude that cohimbianus had smooth
skin on the venter and a somewhat warty dorsum. Dorsolateral folds
are present in only simonsii.

The limbs of all species are short and robust. The limbs are
shortest in cophites (shank about 32% SVL) and longest in fJavo-
macuhtus ( shank about 50% SVL ) . No species has axillary patagia.
The fingers and toes are of moderate length (Figs. 5-7) and lack
discs although the digital tips may be weakly swollen ( perhaps due
to preservation). The subarticular tubercles are flat, round, and
simple. Most species have three palmar tubercles (innermost is the
thenar tubercle ) ; the outer palmar tubercle is absent ( probably fused
to median) in brunneus, cophites, laplacai, and peruviamis. The
thumb is longer than the second finger in brunneus, favomaculatus,
laplacai, parkeri, peraccai, pereger, and peruviamis (Figs. 5, 6a, 7b),
the thumb is equal in length to the second finger in cohimbianus
(fide Werner, 1899), and the thumb is shorter than the second finger
in cophites, montium, nanus, peruanus, simonsii, and icettsteini
(Figs. 6b, 7a).

Werner (1899) reported a single metatarsal tubercle in cohim-
bianus; all other taxa have two. Both metatarsal tubercles are large
and subequal in size in cophites, flavomaculatus, laplacai, parkeri,
peraccai, pereger, peruanus, peruviamis, and si7nonsii (Figs. 6c, 8).
The inner metatarsal tubercle is about twice the size of the outer

10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

in bmnneus, montium, nanus, and wettsteini. Most species lack
lateral fringes on the toes and basal webbing, but both features are
seen in nanus and simonsii (webbing does not enclose basal sub-
articular tubercles ) .

Males, where known, are smaller than females. The smallest
species are nanus ( a single female 20.5 mm SVL ) and peraccai ( 17
females, 18.4-20.6 mm SVL). The largest species are favomaculatus
(14 females, 34.8-42.6 mm SVL) and parkeri (10 females, 27.4-35.0
mm SVL).

Osteology. — Cleared and stained skeletal specimens (see speci-
mens examined) are available for nine of the fourteen species
(cophites, favomaculatus, laplacai, montium, porkeri, peraccai,
pereger, perumanus, and wettsteini) . Limited osteological data were
obtained through dissection of brunneus (USNM 192911, paratype),
peruanus (UMMZ 89477), and simonsii (BMNH 1947.2.15.43, lecto-
type). Aside from externally visible ti-aits, no osteological data are
available for the Colombian species {coJumhianus and nanus), and
the discussion of osteological features accordingly excludes them.

The skulls of Phrynopus species are broader than long or as
broad as long. The maxillary arch does not extend posterior to the
occipital region. The skulls are about one-half as high as long ( not
greatly flattened or conspicuously vaulted). The maxillary arch is
complete and the premaxillae and maxillae bear blunt, pedicellate
teeth. The brain case appears narrow in brwmeus, jiavomaculatus
(Fig. 9), parkeri (Fig. 10), and pereger because the edges of the
frontoparietals are approximately parallel whereas the edges diverge
posteriorly in the other species (Fig. 11, 12). The columellae are
lost in cophites and simonsii (and nanus); no other bones have been
lost in the species of the genus. Exostosis of cranial elements occurs
only in fiavomaculatus (Fig. 9) although some rugosity at the edges
of the frontoparietals is seen in montium (Lynch, 1971) and peraccai
(identified as festae in Lynch, 1968). Cranial dermostosis is not
known in the genus.

The alary processes of the premaxillae are relatively broad at
their bases and directed dorsally in species with rounded or truncate
snouts or posterodorsally in species with sloping snouts (flavomacu-
latus and parkeri ) . The premaxillary palatal shelf is broad and only
weakly dissected in cophites and peruvianus (Figs. 11-12). The
palatal process in these species is slightly longer than the maxillary
process. The premaxillary palatal shelf of laplacai is slightly more
dissected; the palatal process is prominent and longer than the
maxillary process. The premaxillary palatal shelf in brunneus, flavo-
maculatus, parkeri (Fig. 10), peraccai, pereger, peruanus, and ivett-
steini is deeply dissected and the palatal and maxillary processes
subequal in length. In montium (Lynch, 1971: Fig. 104), the palatal
and maxillary processes are greatly elongated and the dissection of
the palatal shelf greater.

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 11

The nasals are widely separated and relatively small in montium,
peruamis, and peruvianus (Fig. 11). The nasals are less widely
separated and slightly larger in cophites (Fig. 12) and peroccai.
The nasals are narrowly separated and large in lapJacai, simonsii,
and tcettsteini, and in contact and large in brunneus, fiavomacidatus
(Fig. 9), parkeri (Fig. 10), and pereger .The nasals of all species
have elongate maxillary processes that are narrowly separated from
or in tenuous contact with the pars facialis of the maxillae. The
nasals are narrowly separated from the frontoparietals in hrunneus,
■fiavomacuhtus, parkeri (Figs. 9-10), and pereger, but widely sep-
arated in the other species examined (Figs. 11-12).

The frontoparietals are in median contact in all species except
cophites which has a moderate-sized frontoparietal fontanelle (Fig.
12). The lateral edges of the frontoparietals frequently bear small
bony ridges or weak exostosis {montium, peraccai, and icettsteini)
which are not detectable externally. The frontoparietals are heavily
exostosed and bear prominent lateral crests in fiavomacnlatiis (Fig.
9). The frontoparietals are not fused to the prootics in any speci-
men examined.

The cristae paroticae are elongate and narrow in favomaculatus,
a large species (Fig. 9), short and stocky in copliites (Fig. 12),
laplacai, montium, peruvianus (Fig. 11), peraccai, and ivettsteini,
small species, and of intermediate length and robustness in parkeri
(Fig. 10) and pereger, frogs of intermediate size. Length and
breadth of the cristae paroticae appear correlated with body size,
at least in Phnjnopus. The epiotic eminesces are most prominent
in flavomaculatus and parkeri and obsolete in the other species. The
occipital condyles are widely separated in all species and weakly
stalked (not stalked in copliites or peruvianus) .

The squamosals have broad otic plates resting on the cristae
paroticae in flavomaculatus, parkeri (Figs. 9-10) and pereger, or
narrow otic plates not resting on the cristae paroticae (copliites,
laplacai, montium, peraccai, peruvianus, and icettsteini) . The zygo-
matic and otic rami of the sc^uamosals are of equal length and the
squamosomaxillary angles 50-55°. The squamosal does not contact
the maxilla.

The maxillary palatal shelf is broadest anteriorly and narrows
posteriorly. The palatal shelf tapers gradually to the end of the
dentigerous series and the point at \\'hich the pterygoid contacts
the maxilla; at this point the shelf abruptly narrows ( Figs. 10, 12 ) .
In peruvianus the shelf tapers gradually through the point of max-
illary-pterygoid contact (Fig. 11).

The sphenethmoid is very large in peraccai (calcified or ossified
to a point anterior to the nasals) and relatively small in cophites
( Fig. 12; calcified portion does not reach the nasals ) . The spheneth-
moid is of intermediate size in the other species ( Fig. 11 ) .

Prevomerine teeth are present on prominent dentigerous proc-

12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

esses in brunneus, columhianns (fide Werner), flavomaculatus,
nanus, parkeri (Fig. 10), peraccai, pereger, and peruanus. The pre-
vomer is edentate and lacks the dentigerous processes in copliites,
laplacai, monthim, penivianns, simonsii, and wettsteini (Figs. 11-
12). The prevomers are separated in all species examined. The
choanal and dentigerous rami of the prevomer are large in flavo-
maculatus, parkeri (Fig. 10), peraccai, pereger, and simonsii. The
choanal ramus is large and the dentigerous ramus thin in cophites
(Fig. 12), laplacai, montium, peniamis, and wettsteini. The den-
tigerous ramus is lost and the choanal ramus reduced to an irregular-
shaped bone anteromedial to the choana in peruvianus (Fig. 11).

Palatines are present and well developed in all species. The
anterior ramus of the parasphenoid contacts the palatines and pre-
vomers in brunneus, flavomaculatus, parkeri (Fig. 10), pereger, and
simonsii. The parasphenoid is slightly shorter in laplacai and perac-
cai and does not contact the palatines or prevomers. In cophites
(Fig. 12), montium, peruanus, peruvianus (Fig. 11), and wett-
steini, the parasphenoid is much shorter and widely separated from
the palatines. The alary processes of the parasphenoid are oriented
at right angles to the anterior ramus in all species except for a slight
posterior deflection in flavomaculatus, parkeri (Fig. 10) and pereger.
The alary processes are shortest in cophites (Fig. 12). The median
ramus of the pterygoid is in contact with the alary process (or
overlaps it) in brunneus, flavomaculatus, parkeri (Fig. 10), and
pereger. The two elements are narrowly separated in montium and
peraccai and widely separated in cophites (Fig. 12), laplacai, peru-
anus, peruvianus (Fig. 11), and wettsteini. The pterygoids are more
massive elements in flavomaculatus and parkeri than in the other
species.

Three species (cophites, nanus, and simonsii) lack the plectrum,
tympanic annulus, and cavum tympanicum; the other eleven species
have complete ears although the ears may not be visible externally.

All species have eight procoelous presacral vertebrae. The trans-
verse processes of the third and fourth vertebrae are longer than
those of the second and fifth through eighth vertebrae. The transverse
processes of vertebrae 5-8 are shorter than the sacral diapophyses
of all species, but are shortest in cophites, laplacai, montium, peru-
vianus, and wettsteini. The sacral diapophyses are weakly dilated
and deflected posteriorly. The sacrococcygeal articulation in bi-
condylar, and the coccyx lacks transverse processes. The ilia are of
the leptodactyfine type ( Lynch, 1971 ) . The limb bones lack flanges
and vary from long and thin (flavomaculatus) to short and stocky
( cophites ) .

All species have cartilaginous omosterna and sterna although
the degree of calcification varies with the size of individuals. The
omosternum is elongate and somewhat dilated anteriorly. The
sternum is broad and weakly to markedly indented posteriorly ( Cei,

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 13

1968; Peters, 1874; Fig. 13 ) . I have not found the posterior indenta-
tions of the sterna of lapJacai, montium, and peruanus as marked as
figured by Cei (1968) and Peters (1874). The sternum is much
narrower and not indented in periwianus (Fig. 3). In all species
except peril vianus, the clavicles are well-developed, thin, and mod-
erately curved (Fig. 13). The epicoracoid cartilages broadly over-
lap and are loosely attached. The attachment of the epicoracoid
cartilages is strongest anteriorly but the girdles are functionally
arciferal. The coracoids are much shorter and stockier than the
clavicles.

In periwianus, the epicoracoid cartilages overlap more narrowly
than in other Phrynopus (compare Figs. 3 and 13) and are more
firmly attached to one another. The girdle of periwianus is func-
tionally firmisternal (pulling laterally on the forearms results in no
displacement of the epicoracoid cartilages). Noble's specimens of
periwianus are all smaller than the recent material; he recorded
partial firmisterny for his small frogs but the smallest of the recently
collected specimens (which are larger than any of the types) is
equally firmisternal as the largest. The fusion of the epicoracoid
cartilages does not obscure the edges of the cartilages (Fig. 3).
The two cartilages are not fused in the posterior one-half of the
epicoracoid overlap but are tightly bound with connective tissue.
Unlike the more typically arciferal species of Fhrynopns and more
like a ranoid type of pectoral girdle, the clavicles and coracoids of
peruvir.nus are subequal in width- and the clavicles are straight and
oriented at more nearly right angles to the sagittal plane. In spite
of the firmisternal modifications of the girdle, the arciferal condition
(sensu Griffiths, 1959) of the girdle is demonstrated by the presence
of epicoracoid horns and the overlapping and only partially fused
epicoracoid cartilages (visible in serial sections of the epicoracoid
bridge).

The phalangeal formulae are 2-2-3-3, 2-2-3-4-3 in all species
and the stubbyness or slenderness of the digits is reflected in the
shapes of the phalangeal bones. Several of the species have lateral
processes on the terminal phalanges. The terminal phalanges are
knobbed in brunneus, flavomaculatus, laplacai, peraccai, and icett-
steini, whereas those of cophites, parkeri, pereger, peruvianus, and
simonsii bear short, irregularly shaped lateral processes (Fig. 2).
The lateral processes are better developed on the toes than on the
fingers. Among species of Eleutherodactylus, the most poorly de-
veloped T-shaped terminal phalanges are found on the fingers of
some species of the binotatus and fitzingeri groups (binotatus, cru-
ralis, granulosus, nigrovittatus, and octavioi) . The terminal pha-
langes of the fingers of these species match the weak lateral proc-
esses seen on the digits of Fhrynopus {cophites, parkeri, peruvianus,
QXid simonsii) .

14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

KEY TO SPECIES OF PHRYNOPUS

1. Prevomerine dentigerous processes present 2

Prevomerine teeth and dentigerous processes absent 9

2. Skin of venter smooth 3

Skin of venter areolate nanus

3. Tympanum concealed parkeri

Tympanum visible 4

4. Cranial crests present flavomaculatus

Cranial crests absent 5

5. Outer metatarsal tubercle less than one-half size of inner — . 6
Outer metatarsal tubercle more than one-half size of inner

(usually equal in size) 7

6. Toes basally webbed; venter off-white cohimhiamis

Toes not basally webbed; venter brown hmnneus

7. First finger shorter than second peruantis

First finger longer than second 8

8. Small frogs ( 9 9 19.0-21.7 mm SVL); shank less than 40%

SVL; venter uniform brown peraccai

Larger frogs ( 9 9 24.0-31.8 mm SVL); shank more than
39% SVL; venter usually spotted pereger

9. Tympanum absent 10

Tympanum present, visible or concealed beneath skin 11

10. Toes bearing lateral fringes simonsii

Toes lacking lateral fringes cophites

11. Skin of venter smooth 12

Skin of venter aerolate 13

12. Inner tarsal tubercle present peruvianus

No tubercle on tarsus periianus'*

13. First finger shorter than second 14

First finger as long as or longer than second laplacai

14. Throat black or dark brown loettsteini

Throat not distinctly darker than venter montium

ACCOUNTS OF SPECIES

In the following accounts I have presented species descriptions
only in the cases of new species or in the cases where extensive
additional material is available or where the original description
was misleading through brevity. A diagnosis, a skeletal synonymy,
and a section for miscellaneous remarks is presented for each taxon.

Phrynopus brunneus new species

Holotype.—VSNM 192909, an adult male from 10 km WNW El Camielo
( = Pun), Carchi Prov., Ecuador, 3150m, collected 30 June 1962 by James
A. Peters.

* In some peruanus, the prevomerine odontophores are concealed beneath
the tissue of the palate; such individuals would key out here.

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 15

Parattjpes.— (3) USNM 192910-12, collected syntopically with the holo-
type.

Diagnosis. — A moderate-sized Phrynopus (3 $ $ 25.8-27.6 mm, one 9
29.6 mm SVL); skin of dorsum and venter smooth; first finger longer than
second; fingers and toes not dilated at tips; toes lacking lateral fringes and basal
webbing; two metatarsal tubercles, inner much larger than outer; tarsus lack-
ing tubercles or folds; tympanum present, visible externally, one-half eye length
in males, one-third eye length in single female; snout rounded in lateral
profile; males lacking vocal sac and slits; prevomerine teeth present on promi-
nent dentigerous processes lying median and posterior to choanae; fronto-
parietals complete, no crests; nasals moderately large, narrowly separated;
anterior ramus of parasphenoid long, in contact with palatines; median ramus
of pterygoid in contact witli parasphenoid alae; body brown with indistinct
darker mottling.

Description. — Head narrower than body, head wider dian long;
head width 34.3-37.9 (x = 36.8) % SVL; snout round in dorsal
view, rounded in lateral profile (Fig. 4); snout short, eye length
1/2 times E-N (eye-nostril distance); canthus rostralis round, some-
what concave; loreal region concave, sloping gradually to lips; lips
not flared; nostrils directed dorsolaterally, not protuberant; inter-
orbital region weakly furrowed, no cranial crests; upper evelid
width 78.7-93.5 (x = 86.7) % lOD ( interorbital distance); supra-
tympanic fold thick, prominent, concealing upper edge of tym-
panum; tympanum visible, round in males, higher than long in
females, its horizontal diameter 43.1-47.8 (x = 45.8) % eye length
in males, 30.3 in single females; males lacking vocal sac and slits;
tongue large, about as long as wide, not notched posteriorly, pos-
terior one-third not adherent to Hoor of mouth; choanae small,
wider than long, not concealed by palatal shelf of maxillae; pre-
vomerine dentigerous processes present, median and posterior to
choanae, each process bearing a transverse row of 4-5 teeth; pre-
vomerine dentigerous processes three to four times as wide as a
choana.

Skin of dorsum and venter smooth except for weak areolations
along lower Hanks, posterior parts of undersides of thighs, and near
anus; no dorsolateral folds; discoidal folds obscure, ending anterior
to groin; no tubercles on eyelids; forearm lacking ulnar tubercles
or folds; palmar tubercle bifid and larger than thenar tubercle;
thenar surfaces bearing a few ill-defined supernumerary tubercles;
subarticular tubercles large, round, simple, non-conical; fingers not
fringed; first finger distinctly longer than second; digit tips round,
not bulbous.

Tarsus and heel lacking tubercles or folds; two metatarsal tu-
bercles, inner twice as long as wide, two to three times as large as
round outer tubercle; plantar surface lacking supernumerary tu-
bercles; toes lacking lateral fringes and basal webbing; subarticular
tubercles large, round, simple, non-conical; tips of toes rounded, not
dilated; hindlimbs short, shank .34.2-35.7 (x = .35.2) % SVL.

In preservative, the dorsal surfaces are brown with indistinct

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

darker brown mottling; no color pattern is evident. P. bnmneiis
lacks distinct canthal and supratympanic stripes, labial bars, and
an anal tiiangle. The ventral surfaces are paler than the dorsum
and vary from dusky gray to brown. One individual (USNM

192909) has narrow dark brown bars on the shank and tarsus.
Measurements of ihe holotijpe in mm. — SVL 25.8, shank 9.1,

head width 9.7, head length 8.8, eycHd width 1.8, lOD 2.1, tym-
panum 1.3, eye length 2.7, eye-nostril 1.8.

Etymology. — Latin, in reference to the drab coloration of browns
and grays.

Natural history. — The four specimens were collected by day
beneath logs in a wet pasture at 3150 m on the high Amazonian
slopes of the eastern Andean cordillera. The female (USNM

192910) contains large (3.5 mm in diameter), yellow, ovarian eggs
and has convoluted oviducts. The testes of the males are not
especially large or swollen and are not black (in contrast to the
black testes in male E. curtipes collected syntopically).

Distribution. — Known only from the type-locality (Fig. 14).

Phrynopus cophites new species
Fig. 15

Hohtype. — KU 138884, adult female, S slope Abra Acanacu, 14 km NNE
Paucartambo, Depto. Cuzco, Peru, 3400 m, collected 13 January 1971 by
Thomas H. Fritts.

Pflra^ypes.— (30) KU 138885-908, 138911-16, N slope Abra Acanacu, 27
km NNE Paucartambo, Depto. Cuzco, Peru, 3400 m.

Diagnosis. — A small Phrtjnopiis ( $ $ 18.0-22.7 mm, 9 9 21.9-29.3 mm
SVL ) ; skin of dorsum and venter coarsely areolate; first finger shorter than
second; tips of digits lacking apical swelling; toes lacking lateral fringes and
basal web; two metatarsal tuliercles, inner slightly larger than or equal to outer;
no tarsal tubercle or fold; tympanum, columella, and cavum tympanum absent;
snout rounded in lateral profile; males lacking vocal sac and slits; prevomerine
teeth and dentigerous processes lacking, dentigerous ramus of prevomer present;
frontoparietals not bearing crests, frontoparietal fontanelle present; nasals
separated medially, small; anterior ramus of parasphenoid not reaching pala-
tines; dorsum gray to brown with or witliout diffuse mottling; venter brown
with brovra mottling; throat usually more pale than venter.

Description. — Head narrower than body (Fig. 15), head wider
than long; head width 31.1-36.1 (x = 33.7) % SVL; snout round in
dorsal view, round to truncate in lateral profile, tip of snout level
with edge of lip; canthus rostrahs moderately sharp, concave; loreal
region flat, sloping to lip; lips not flared; nostrils weakly protuberant,
directed anterolaterally; snout very short, E-N much less than eye
length; interorbital region flat; upper eyelid width 66.6-86.7 (x =
76.2) % lOD; temporal region swollen (parotoid gland-like); no
supratympanic fold; tympanic annulus, columella, and cavum tym-
panicum absent; postrictal tubercles present, not prominent; choanae
small, round, situated well anteriad on palate, not concealed by
palatal shelf of maxillae when roof of mouth is viewed from di-

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 17

rectly above; prevomerine teeth and dentigerous processes absent;
tongue slightly longer than wide, not notched posteriorly, posterior
one-fifth to one-fourth free; males lack vocal slits and vocal sac.

Skin of dorsum finely areolate, areolation increasing in size and
prominence posteriorly and on flanks; skin of venter (including
throat) coarsely areolate; no discoidal folds; tubercles on dorsum
sometimes coalesce forming short, irregular vertebral, paravertebral,
and dorsolateral folds or ridges; dorsolateral ridges most prominent
anteriorly; no ulnar tubercles or ridge; palmar tubercle not bifid,
as large as thenar tubercle (Fig. 7); a third, outer, palmar tubercle
occasionally present, but difficult to discern from low supernumerary
tubercles on palm; subarticular tubercles low, flat, round, simple;
fingers lacking lateral fringes; tips of fingers usually lacking apical
swelling; first finger shorter than second; base of thumb swollen in
reproductively active males.

Heel bearing several small tubercles; tarsus areolate and lacking
distinct tubercles or folds on inner and outer edges; two metatarsal
tubercles, inner slightly larger than outer, both longer than wide
(Fig. 8); plantar surface areolate; subarticular tubercles low, round,
simple, more prominent than those of fingers; toes lacking lateral
fringes and basal webs; tips of toes narrow, little or no apical bulb-
ing; hindlimbs very short, heel of adpressed limb not reaching fore-
arm insertion in adults, but reaching axilla in juveniles; shank
29.2-33.8 (x = 31.7) % SVL in i 6 , 31.9-35.7 (x = 33.9) in 9 5 .

In preservative, the ground color is gray to dull brown with or
without brown to black mottling. The venter is gray to dark brown
with mottling of a darker brown. The throat is usually paler than
the venter. A few individuals have a thin sagittal and transverse
pectoral line on the venter and throat. The flanks are darker than
the dorsum; some individuals have faint, thin, cream-colored bars
on the flanks. The darker specimens have canthal stripes and labial
bars. The limbs are not barred and the posterior surfaces of the
thighs are gray to dark brown.

In life, P. coplutes is dull brown to pale tan above or has a
mottled pattern of olive-brown and black to brown and greenish-
yellow or red. The venter is dull gray. Males have an orange cast
to the throat.

Measurements of the holotype in mm. — SVL 29.3, shank 9.2, head
width 10.1, eyelid width 2.2, intcrorbital distance 2.8. The holotype
is a gravid female with convoluted o\'iducts and large yellow eggs
(2.0-2.5 mm in diameter).

EtymoJoii,ri. — From Greek, kophos, meaning deaf; in reference to
the absence of an external and middle ear.

Natural history. — Phrynopus cophites was found in paramo and
elfin forest habitats in a ravine on the Cadena de Paucartambo at
elevations of 3400 to 3450 m. The majority of the specimens were
collected syntopically with P. penwiamis. As in P. peruviantis, the

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

females of P. cophites are gravid. Males of P. cophites lack vocal
and auditory apparatus, and presumably do not call.

Distribution. — Known from both the north and south slopes of
the Abra Acanacu, a pass in the Cadena de Paucartambo, on the
northwestern end of the Cordillera Carabaya, in Departamento
Cuzco, Peru, between 3400 and 3450 m ( Fig. 14 ) .

Remarks. — Earlessness and the very short limbs readily distin-
guish P. cophites from all other known species of Phnjnopus. P
cophites has a short first finger, areolate skin, and lacks prevomerine
teeth as in P. montium, P. simonsii, and P. wettsteini. P. montium
and P. wettsteini have complete ears concealed beneath the skin
and longer legs than P. cophites. P. nanus, P. simonsii, and P.
cophites are the only earless species of the genus, but differ in
several respects. P. cophites has a frontoparietal fontanelle, small
nasals, small prevomers, and short rami of the parasphenoid. Judg-
ing from dissections, the skull of P. simonsii is similar to that of P.
parkeri; an idea of the differences between P. cophites and P.
simonsii can be obtained by comparing the skull of P. parkeri with
that of P. cophites (Figs. 10, 12). If the similarities of the skulls
are a useful index of relationship, P. cophites is allied to P. laplacai,
P. montium, P. peritanus, and P. wettsteini, and less closely to P.
peruvianus, with little relationship to the northern Phnjnopus (hrun-
neiis, flavomaculatus, parkeri, peraccai, pereger, and simonsii).

Phrynopus flavomaculatus (Parker), new combination

Eleutherodactyhis jlavoinacuhtus Parker, 1938:440 ( Ho/of [/pe.—BMNH 1935.

11.3.16/1947.2.16.11, 15 km E Loja, Prov. Loja, Ecuador, 3000 m).
Niceforonia jiavomacuhta — Lynch, 1969:273.

Diagnosis. — A large Phrynopus ( $ $ 26.0-32.5 mm, 9 9 29.2-42.6 mm
SVL); skin of dorsum and venter smooth; first finger longer than second; tips of
digits not bulbous; toes lacking basal webbing and lateral fringes; two meta-
tarsal tubercles, equal in size; tarsus lacking tubercles or folds; tympanum
visible, 37.8-51.4 % eye length; snout sloping in lateral profile; male with
vocal slits and subgular vocal sac; prevomerine teeth present on prominent
processes lying median and posterior to choanae; frontoparietals complete, .
bearing prominent lateral crests (Fig. 9); nasals large, in broad median contact;
anterior ramus of parasphenoid long, contacting palatines; median ramus of
pterygoid in contact with parasphenoid alae; dorsum brown with darker brown
blotches, venter brown with large yellow spots, throat dark bro\vn.

Description. — See Parker ( 1938).

Natural history. — P. flavomactdatus is usually found beneath
rocks or logs in paramo and subparamo habitats. At a locality north
of San Lucas (Loja Prov), I found P. flavomaculatus in the more
xeric sites within the subparamo and found Tehnatohius niger be-
neath logs in the wet cienegas of the paramo. A single egg mass
( KU 121354 ) was collected in June 1968 beneath a log with a female
P. flavomaculatus. The egg mass and adult frog were about 1 m
apart.

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 19

Distribution. — Known from between 2215 and 3100 m on the
eastern Andean Cordillera in southern Ecuador (Fig. 14).

Phrynopus laplacai (Cei), new combination

Stjrrhophus laplacai Cei, 1968:139 {Holotype.—IBN/UNC 1251-1, 3 km W

Pongo on road to Coroico from La Paz, Depto. LaPaz, Bolivia, 3400 m).
Nicefownia laplacai — Lynch, 1969:273.

Diagnosis. — A moderate sized Phrynopus ( S $ 27-31 mm, $ 9 28-33 mm
SVL); skin of dorsum and venter areolate; first finger longer than second;
tips of digits bulbous; toes lacking basal webbing and lateral fringes; two meta-
tarsal tubercles, inner slightly larger than outer; no tarsal fold or tubercle;
tympanum concealed beneatli skin; snout truncate in lateral profile; males
with vocal sac and vocal slits; prevomerine teeth and dentigerous processes
lacking; frontoparietals complete, lacking crests; nasal bones narrowly sepa-
rated; anterior ramus of parasphenoid not reaching palatines; dorsum brown,
mottled with cream; venter cream with brown flecking, throat black in males.

Description. — See Cei (1968).

Distribution. — Known only from the vicinity of the type-locahty

(Fig. 14).

Remarks. — P. laplocai, P. montium, and P. loettsteini form a
group of closely allied southern species. Males of laplacai and
loettsteini are easily recognized by the black throats; females of
tcettsteini also have black throats. Too few specimens are available
of the three taxa to permit a definitive assessment of the distinctions
presently understood. The greatest similarities are between laplacai
and wettsteini. The differences cited in the diagnoses may well
reflect the small sample sizes, age in perservative, or method of
preservation. If the type-locality for wettsteini (see species ac-
count) is actually in northern Peru, then it seems very unlikely that
the two names apply to the same population — the specimens on
which they are based were collected at localities more than 1600
kilometers (airline) apart, and at altitudes of 2000 and 3400 meters.
In view of the uncertainty concerning the type-locality of wettsteini
this argument is moot. Cei (1968) only compared laplacai to mon-
tium— these two names are certainly not synonymous.

Phrynopus montium (Shreve), new combination

Syrrhopus montium Shreve, 1938:406. {Holotype. — MCZ 22858, Cascas, near

Huashuasi, Depto. Junin, Peru).
Syrrhophus montium — Gorham, 1966:166.
Nicefownia montia — Lynch, 1968:291.

Diagnosis. — A moderate-sized Phrynopus (21-29 mm SVL); skin of dor-
sum and venter areolate; first finger shorter than second; tips of digits weakly
bulbous; toes lacking basal webbing and lateral fringes; two metatarsal tu-
bercles, inner much larger than outer; tarsus lacking tubercles or folds; tym-
panum concealed beneath skin; snout rounded in lateral profile; males with
vocal sac and slits; prevomerine teeth and dentigerous processes absent, den-
tigerous ramus of prevomer present; frontoparietals complete, lacking crests;
nasals widely separated; anterior ramus of parasphenoid short, not reaching

20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

palatines; median ramus of pterygoid narrowly separated from parasphenoid
ala; dorsum dark brown, venter cream suffused with brown; throat not con-
spicuously darker than venter.

Distribution. — Known only from the type-locality (Fig. 14).

Remarks. — The data available for P. montium are too scanty, as
for P. laplacai and P. ivettsteini, to comment beyond that given in
the remarks for those two species.

Phrynopus nanus (Goin and Cochran), new combination

Niceforonia nana Goin and Cochran, 1963:499 { Holoty pe.—USNM 150643,
Paramo de la Rusia, Depto. Santander, Colombia).

Diagnosis. — A small Phrynopus (female holotype 20.5 mm SVL); skin of
dorsum granular, that of venter coarsely areolate; first finger shorter than
second; fingers and toes not dilated at tips; toes bearing lateral fringes and
basal webbing; two metatarsal tubercles, inner slightly larger than outer; tarsus
lacking distinct tubercles or folds; tympanum absent (or greatly reduced in
size); snout rounded in lateral profile; prevomerine teeth present on small
dentigerous processes; frontoparietals not bearing crests, complete; nasals large,
in contact; body brown without distinct pattern, venter paler than dorsum.

Description. — Goin and Cochran (1963); Cochran and Goin
(1970).

Distribution. — Known only from the type-locality.

Remarks. — With only the holotype available, few additional
anatomical features can be noted for this species. Goin and Cochran
(1963) described the skin of the venter as coarsely areolate, but at
present (1973) the skin of the venter appears smooth except pos-
teriorly where very coarse granulations are evident. I suspect that
the entire venter of the specimen was once areolate and that the
now smooth appearance of most of the venter is an artifact of
preservation.

The most perplexing problem involving this species is its rela-
tionships to the other species here called Phrynopus. Comparisons
of the character mosaics of the other species of simple-toed eleu-
therodactylines with that of P. nanus prompts the conclusion that
nanus is most similar to simonsii. The similarity may be in part
spurious in that the character mosaics for each species are incom-
plete. The two species are readily distinguished in that simonsii
lacks prevomerine dentigerous processes, has glandular dorsolateral
folds, is considerably larger, and has vertical bars on the lower
flanks.

The resemblance of P. nanus to the other northern Phrynopus
is limited to the sharing of prevomerine dentigerous processes. The
Ecuadorian species of the genus uniformly have smooth skin on the
venter, the first finger longer than the second, and well-developed
ears. P. nanus may prove to be an independent offshoot of Eleu-
therodactylus, convergent with Phrynopus in digit structure. This
question cannot be answered until more material is available.

ANDEAN LEPTODACnXID FROG GENUS PHRYNOPUS 21

Phrynopus parkeri new species

Fig. 16

Holotype. — KU 135278, an adult male from tlie summit of the Cordillera
between Chanchaque and Huancabamba, Depto. Piura, Peru, 3100 m, collected
11 May 1970 by Thomas H. and Patricia R. Fritts.

Paratijpes.— (32) KU 135279-305, 135307-11, collected syntopically with
the holotype.

Diagnosis. — A moderate-sized Phrynopus { $ $ 21.4-24.2 mm, 9 $ 27.4-
35.0 mm SVL); skin of dorsum and venter smooth; first finger longer than
second; digit tips weakly bulbous; toes lacking lateral fringes and basal web-
bing; two metatarsal tubercles, inner twice size of outer to equal in size to
outer; tarsus lacking distinct tubercles or folds; tympanum present, concealed
beneath skin; snout sloping in lateral profile; prevomerine teeth present on
well-developed processes medial and posterior to choanae; frontoparietals com-
plete, lacking crests; nasals large, in median contact; anterior ramus of para-
sphenoid long, in contact with palatines; median ramus of pterygoids in
contact with parasphenoid alae; dorsum brov^Ti with darker mottling, venter
cream with brown reticulations, throat darker than venter.

Description. — (Fig. 16.) Head narrower than body, head as
wide as long or sHghtly wider than long in males, wider than long
in females, head width' 34..3-40.0 (x =: 36.6) % SVL; snout rounded
in dorsal view, sloping in lateral profile (Fig. 4); canthus rostralis
sharp, concave; loreal region concave; lips not flared; nostrils lateral,
weakly protuberant; snout short, eye length slightly greater than
E-N; interorbital region flat, no cranial crests; upper eyelid width
84.1-110.0 (x = 93.3) % lOD in males, 68.9-95.6 (x = 81.3) % in
females; tympanum concealed beneath skin; supratympanic fold
poorly-defined; tongue large, oval, not notched posteriorly, pos-
terior one-fifth free; choanae small, round, not concealed by palatal
shelf of maxillae; pre\'omerine dentigerous processes present, each
6 to 8 times size of a choana, processes slanted posteriorly, wider
than long, each bearing .3-6 teeth, processes separated by distance
equal to width of a choana; male with vocal slits and subgular
vocal sac.

Skin of body smooth except on posteroventral surfaces of thighs;
discoidal folds prominent posteriorly; no dorsolateral folds; forearm
lacking ulnar tubercles or folds; three palmar tubercles, outer minute
(Fig. 6); thenar surface bearing a few ill-defined supernumerary
tubercles, subarticular tubercles of fingers low, round, simple; fingers
not fringed, tips round and weakly bulbous; first finger longer than
second (Fig. 6).

Tarsus and heel lacking tubercles or folds (except in some
females where an idefinite inner tarsal tubercle is evident); two
metatarsal tubercles, neither enlarged nor compressed, inner oval,
relative sizes of tubercles, varies from equal in size (most individ-
uals) to inner twice size of outer; plantar surface bearing a few
indefinite supernumerary tubercles; subarticular tubercles of toes
flat, round, simple, same size as those of fingers; toes lacking lateral

22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

fringes and basal webbing; tips of toes simple and weakly bulbous;
hindlimbs short, heel of adpressed limb reaches to base of forearm
and tympanic area; shank 37.1-43.8 (x = 40.1) % SVL.

In preservative, the dorsal surfaces are gray-brown to reddish-
brown with or without a hairline cream sagittal stripe. Some indi-
viduals have a broad dorsal band of dusky cream or dull brown
and a few individuals have indefinite dorsolateral stiipes. The flanks
are darker than the dorsum and have a nebulose pattern of browns.
The lips are not marked except for an indefinite cream stripe in
some specimens. The limbs are not barred in most specimens; some
individuals do have faint barring on the shanks. The posterior sur-
faces of the thighs are unicolor brown or brown with small cream
flecks. The venter is cream or cream mottled with brown; in some
specimens the marbling is so intense that the venter and the under-
sides of the thighs are pale to dark brown. The specimens with the
cream sagittal line have another line along the posterior surface of
the thigh, underside of the shank, inside of the tarsus continuing
onto the plantar surface, and on the ulnar region continuing onto
the thenar surface.

In life, P. parkeri has a ground color of olive-brown, gray-
brown, reddish-brown, dark-brown, or metallic green-brown. The
flanks are dark-brown to gray-brown flecked with metallic green.
The posterior surfaces of the thighs are dark-brown or black with
yellow spots and blotches. The ventral surfaces are charcoal-gray
or black with large egg-yolk yellow blotches in males. The venter
of females tends to be unicolor egg-yolk yellow with indefinite
brown reticulations. Males have gray throats whereas females have
a yellow-green haze on the throat. The face is marked with an
indefinite dark-brown canthal stripe and in some individuals with
a beige-white labial stripe. The iris is dark-brown (T. H. Fritts
fieldnotes, llMay 1970).

Measurements of ihe holotijpe in mm. — SVL 22.6, shank 9.2,
head width 8.1, head length 8.1, eyefid width 2.2, lOD 2.5.

Etymology. — Named for Hampton W. Parker, the first author
to deal with the complex of frogs here called Phrytwpus.

Natural history. — All of the known specimens were collected by
day beneath rocks along the road between Chanchaque and Huanca-
bamba, Piura, Peru, at 3100 m. The west slope of the cordillera is
dry and chaparrel-like, whereas the east slope and summit of the
cordillera supports a moist, elfin forest with shrubs and shrub-like
bushes 2-3 m high. No reproductive activity was noted by the
collectors although the females in the type-series have mature
ovarian eggs.

Distribution. — Known only from the type-locality (Fig. 14).

Remarks. — P. parkeri is most closely related to P. flavomaciilatus
and P. pereger. The similarity between the three species is evident
in all characteristics cited and the three differ only in flavomaculatus

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 23

Fig. 4. — Lateral views of heads of (A) Phrijnopus brunneus, USNM 192909,
(B) P. flavomaculatus, KU 142199, (C) P. simonsii, BMNH 1947.2.15.43, and
(D) P.parkeri, KU 135298.

Fig. 5. — Palmar views of hands of (A) Phrijnopus brunneus, USNM 192911,
(B) P. peraccai, KU 117795, and (C) P. flavomaculatus, KU 119722.

24

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 6. — Palmar views of hands of ( A ) Phrtjnopus parked, KU 135282, ( B )
P. simonsii, BMNH 1947.2.15.43, and plantar view of foot of (C) P. parked,
KU 135298.

Fig. 7. — Palmar views of hands of (A) Phrtjnopus cophites, KU 138884,
and (B) P. peruvianus, KU 138138.

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS

25

Fig. 8. — Distal tarsus and proximal portion of foot (plantar views) of (A)
Phrtjnopus peruvianus, KU 138138, and (B) P. cophites, KU 138911.

•I

I

Fig. 9. — Skull of Phrtjnopus flavomaculatus in dorsal view (KU 119743).

26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 10.— Skull of Phrynopus parked (KU 135306) in dorsal (A) and
ventral (B) views.

Fig. 11. — Skull of Phrynopus peruvianus (KU 138926) in dorsal (A) and
ventral (B) views.

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 27

Fig. 12. — Skull of Phnjnopus cophites (KU 138909) in dorsal (A) and
ventral (B) views.

Fig. 13. — Ventral view of pectoral girdle of Phnjnopus peruanus (UMMZ

89477).

28

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 14. — Distribution of Phrtjnopus in Bolivia, Ecuador, and Peru. P. brun-
neus (1), P. cophites (11), P. jiavomaculatus (3-5), P. laplacai (12), P.
montium and P. peruanus (9), P. parkeri (7), P. peraccai (2), P. pereger (10),
P. peruvianus (11), P. simonsii (8), and wettsteini (6).

Fig. 15. — Dorsal and ventral views of Phrtjnopus cophites (KU 138895,
25.4mmSVL).

ANDEAN LEPTODACTiXID FROG GENUS PHRYNOPUS 29

Fig. 16. — Dorsal and ventral views of Phrijnopus parked (left, KU 135297,
31.3 mm SVL; right, KU 135298, 35.0 mm SVL).

Fig. Y1 .—Phrijnopus periivianus (left, KU 138950, 25.1 mm SVL; middle
and right, KU 138940, 20.2 mm SVL).

30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

having prominent cranial crests and reaching a larger size, and in
parked having the tympanic annulus concealed beneath the skin.

Phrynopus peraccai new species

Niceforonia festae—hynch, 1968:289-93; 1969:273; 1970:139; 1971:157.

Holotype. — USNM 160947, an adult female collected 1 km W Papallacta,
Napo Prov., Ecuador, 3155 m, 6 July 1962 by James A. Peters, P. Spoecker,
and M. Olalla.

Paratypes.— (16) ECUADOR, Napo Prov.: Papallacta, 3110 m, USNM
164397, MIZ 820; 1 km W Papallacta, 3155 m, KU 111430, USNM 160955-59,
160961, 160966-68, 164398; 4 km W Papallacta, 3300 m, KU 117795-96;
Laguna Papallacta, 3350 m, KU 109171.

Diagnosis. — A small Phrynopus (adult 9 9 19.0-21.6 mm SVL); skin of
dorsum smooth with flat warts on eyelids and flanks, that of venter smooth;
first finger longer than second; tips of digits not bulbous; toes lacking lateral
fringes and basal webbing; two metatarsal tubercles, equal in size or outer
larger; tarsus lacking tubercles or folds; tympanum visible externally, not
prominent, its length 1/3-2/5 tliat of eye; snout rounded to truncate in lateral
profile; males not known; prevomerine dentigerous processes prominent, bearing
teedi; frontoparietals complete, lacking crests; nasals separated, moderately
large; anterior ramus of parasphenoid long but not reaching palatines; median
ramus of pterygoid in contact wdth parasphenoid alae; coloration brown, dorsum
darker than venter and bearing indefinite blotches.

Description. — Head narrower than body, head wider than long;
head width 35.0-40.7 (x = 37.5) % SVL; snout round in dorsal view,
rounded to truncate in lateral profile; snout short, E-N 60.0-100.0
(x = 77.4) % eye length; canthus rostralis moderately sharp, con-
cave; loreal region weakly concave, sloping gradually to non-flared
lips; nostrils directed dorsolaterally, weakly protuberant; interorbital
space flat, no cranial crests; upper eyelid width 63.4-90.9 (x = 75.1)
% lOD; supratympanic fold indistinct, obscuring upper edge of
tympanum; tympanum visible externally, not prominent, higher than
long, separated from eye by 1/2 times t\nnpanum width; tympanum
length 35.1-51.3 (x = 41.8) % eye length; males not known; tongue
about as long as wide, not notched posteriorly, posterior one-fourth
to one-third not adherent to floor of mouth; choanae small, round,
not concealed by palatal shelf of maxillae; prevomerine dentigerous
processes present, bearing 0-7 (x = 4.4) teeth /process, wider than
long, median and posterior to choanae, separated medially by dis-
tance equal to width of a choana, teeth in a transverse row; pre-
vomerine dentigerous process 3-4 times width of a choana.

Skin of dorsum smooth except for low, obscure warts on eyelids
and sometimes flanks, skin of venter smooth; no dorsolateral folds;
discoidal folds obscure; forearm lacking ulnar tubercles or folds;
palmar tubercle bifid, as large as thenar tubercle (Fig. 5); thenar
surface lacking supernumerary tubercles; subarticular tubercles
round, non-conical, simple; fingers not fringed; first finger longer
than second; tips of digits pointed, not bulbous.

Heel and tarsus lacking tubercles and folds; two metatarsal

ANDEAN LEPTODACTi'LID FROG GENUS PHRYNOPUS 31

tubercles, inner longer than wide, outer about as long as wide; meta-
tarsal tubercles equal in size or outer larger than inner; no super-
numerary plantar tubercles; subarticular tubercles not distinct,
round, simple, smaller than those of fingers; toes lacking lateral
fringes and basal webbing; tips of toes rounded, not bulbous; hind-
limbs short, shank 33.7-40.4 (x = 37.0) % SVL.

In preservative, P. peraccai is brown. The dorsum is darker than
the venter. The limbs are faintly barred and the dorsum bears in-
distinct darker brown markings ( interorbital bar, scapular and sacral
chevron, anal triangle ) . Labial bars are weakly evident and a dark
supratympanic stripe is present bordered below by a cream postric-
tal stripe. The venter is finely punctated with cream.

In life, P. peraccai is gray-brown with darker brown markings
on the dorsum and cream spots on the venter.

Measurements of the Iwlofype in mm. — SVL 2L7, shank 7.6,
head width 8.0, head length 6^9, eyelid width L7, lOD 2.1, tym-
panum length 0.9, eye length 2.4, eye-nostril 1.8. The holotype has
4 and 5 prevomerine teeth on the right and left processes respec-
tively, and is a gravid female with heavily convoluted oviducts and
large yellow eggs (2.5-3.0 mm in diameter).

Etymology. — Named for Mario G. Peracca.

Natural history. — I have examined 21 specimens of this species,
all of which were collected within 5 km of Papallacta, Ecuador. All
of the specimens are females. The four smallest individuals (USNM
160958, 160966-68) are 16.1-18.3 mm SVL and have weakly convo-
luted oviducts and small ovarian eggs (less than 1.0 mm in diam-
eter). The next smallest individual (KU 117796, 19.0 mm SVL)
contained large eggs (2.0-3.0 mm in diameter) and has weakly
convoluted oviducts. Indi\'iduals more than 20.0 mm SVL have large
eggs and heavily convoluted oviducts. Gravid females have been
collected in March, July, and November but no egg clutches have
been found. All specimens were collected beneath rocks in the
grassy paramo or subparamo habitats that surround Papallacta.

Distribution. — Known only from the vicinity of Papallacta, Napo
Prov., Ecuador, between 3000 and 3350 m (Fig. 14).

Remarks. — Since 1968 this species has been called Niceforonia
festae (Peracca). I have now examined 12 syntypes of festae
BMNH 1926.12.4.1/1947.2.15.47, MIZ 428(9), 819 ( lectoholotype
of Paluclicola festae, here designated), 820. Eleven of the speci-
mens are the frog I ( 1968 ) named EleutJierodactyhis trepidotus.
The other specimen is a Phrynopus peraccai. The description of
Paludicola festae clearly applies to the species with narrow, but
complex, digits (E. trepidotus) and cannot be appHed to the simple-
toed species (P. peraccai).

Pahidicola festae Peracca is the oldest name for Eleutherodac-
tyhis trepidotus Lynch, but is not available because in the same
paper naming Pahidicola festae, Peracca (1904) named Hylodes

32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

festae, identical with Eleutherodactylus galdi (Lynch, 1974). Pa-
ludicola festae Peracca is thus a secondary homonym, and the correct
name for the narrow-toed Eleutherodactylus at Papallacta is E.
trepidottis Lynch.

I have previously (1970) mentioned that three species of elu-
therodactyline frogs found in the vicinity of Papallacta were known
only from females. In spite of large samples, several different col-
lectors with presumably different techniques, and collections at dif-
ferent times of the year, I had not found male E. trepidotus and
had begun to consider it all female. However, two of the specimens
in the syntypic series of Pahidicola festae are males (vocal sac and
slits, testes). The sex ratio remains impressively biased (2 males:
81 females) for this species. In light of this sex ratio, the absence
of males in a sample of only 21 Phrynopiis peraccai takes on less
significance and may simply reflect secretiveness of the males.

A close relationship between P. hrunneus, P. favo7nacuJatiis, P.
parkeri, P. peraccai and P. pereger is suggested by the commonality
of smooth skin of the venter, first finger longer than the second, and
prominent prevomerine dentigerous processes. The sloping snouts
of P. flavomactdatits and P. parkeri are distinctively different from
the truncate or rounded snouts seen in P. hrunneus and P. peraccai.
Little osteological data are available for P. hrunneus, but what is
available suggests that P. hrunneus is intermediate between P.
peraccai and the sloping-snout species. P. hrunneus is probably the
closest ally of P. peraccai. The two species differ in size ( peraccai is
smaller), relative sizes of the metatarsal tubercles (inner much
larger than the outer in hrunneus) , and head shape (subjective) in
addition to the meager osteological differences gleaned by dissection
of one specimen of hrunneus and three cleared and stained skele-
tons of peraccai.

Phrynopus pereger new species

Holotijpe. — LSUMZ 26101, adult female, part of a series taken between
Mitupucuru and Yuraccyacu, Depto. Ayacucho, Peni, by Richard Thomas,
11-13 May 1971.

Paratypes.— LSUMZ 26102-10, KU 151908-09, topotypes; LSUMZ 26117-
23, 26255, Yuraccyacu. on Tambo-Val'e del Apurimac t'ai!, 2650 m; LSUMZ
26124-31, Ccarapa, below Tambo on Ayacucho-Valle del Apurimac trail, 2460
m, Depto. Ayacucho, Peni.

Diagnosis. — A moderate-sized Phrynopus ( S S 20.5-25.4, 2 5 24.2-31.8
mm SVL); skin of dorsum and venter smooth; first finger longer than second;
digit tips bulbous; toes lacking lateral fringes and basal webbing; two meta-
tarsal tubercles, inner slightly larger than outer; tarsus lacking tubercles or folds;
tympanum visible, not prominent, its length 31.2-50.0 % eye length; snout
rounded in lateral profile; prevomerine teeth present on well-developed proc-
esses posteromedial to choanae; frontoparietals complete, lacking prominent
crests; nasals large, in median contact; anterior ramus of parasphenoid in
tenuous contact with palatines; median pterygoid ramus in contact with
parasphenoid alae; brown above with diffuse dark blotches, usually with a

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 33

pale interorbital bar, limbs not barred, venter brown with indefinite pale brown
spots to cream with brown suffusion or indefinite brown spots.

Description. — Head narrower than body, broader than long ex-
cept in juveniles; head width 35.8-40.8 (x = 37.4) % SVL; snout
subacuminate in dorsal view, rounded in lateral profile; snout short,
E-N slightly less than eye length; canthus rostralis sharp, concave;
nostrils weakly protuberant, directed dorsolaterally; loreal region
concave, sloping abruptly to non-flared lips; interorbital space flat
(no cranial crests), broad; upper eyelid width 74.1-95.8 (.x = 87.5)
% lOD; upper eyelid bearing one or two tubercles and /or ridges
running posteromedially onto occiput; supratympanic fold obsolete,
covering upper edge of ear; ear visible, higher than long, its length
38.6-50.0 (x =r 43.4) % eye length in males, 31.2-45.4 (x = 39.4) %
in females; choanae moderate-sized, wider than long, about size of
a prevomerine dentigerous process, well within maxillary arch; pre-
vomerine dentigerous processes posteromedial to choanae, round,
each bearing 3-6 teeth; tongue longer than wide, not notched pos-
teriorly, posterior one-third not adherent to floor of mouth; males
with vocal slits, subgular vocal sac.

Skin of dorsum finely shagreened, some specimens with thin
sagittal and dorsolateral ridges; skin of venter smooth, that below
and lateral to vent coarsely areolate; no discoidal folds; no ulnar
tubercles or folds; palmar tubercle bifid and larger than thenar
tubercle; subarticular tubercles round, flat; fingers lacking lateral
fringes; digits with moderate apical swelling, no discs; first finger
longer than second.

Shank 39.0-48.6 (x = 43.6) % SVL; no tarsal or heel tubercles;
two prominent metatarsal tubercles, inner longer than wide, 1/2
times size of outer (also longer than wdde); no supernumerary
plantar tubercles; subarticular tubercles round or slightly longer
than wide, less flattened than those of fingers; toes lacking lateral
fringes and basal webbing; digit tips swollen; lacking discs.

In preservative, P. pereger is medium to dark brown above with
diffuse darker blotches; the limbs are not or only weakly barred;
canthal and supratympanic stripes are dark brown, the latter edged
below by a cream postrictal stripe; lips not or only weakly barred;
flanks paler than dorsum and occasionally appearing reticulated,
especially in groin; a large dark brown spot or bar is present on the
proximal surface of the anterior face of the upper arm; anterior and
posterior surface of thighs dark brown without spots but some indi-
viduals have cream spots on the posterior surface of the thighs;
black or dark brown anal patch present, marked in some specimens
by cream lines extending along the posterior surface of the thighs;
in most specimens the venter is uniform brown with indefinite paler
brown areas; in some specimens the venter is largely cream with a
suffusion of brown or indefinite brown spots; the extreme ventral
coloration is cream densely marbled with brown. The throat tends

34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

to be darker than the rest of the venter. Most specimens have a
pale interorbital bar or spot.

In hfe, P. pereger has a dark gray, reddish-brown, or dark gray-
brown ground color with little indication of pattern. The flanks are
yellowish brown to dull yellow with an orange suffusion ventrally.
The venter varied from brown with orange spots to off-white with
black to gray spotting and/or suffusion. If pale canthal lines were
present, they were pale yellow. (R. Thomas field notes.)

Measurements of holotype in mm. — SVL 28.8, shank 12.9, head
width 10.7, head length 10.7, eyelid width 2.4, lOD 2.6, tympanum
length 1.2, eye length 3.0, eye-nostril 3.0.

Etymology. — Latin, pereger, on a journey, in reference to the
wide geographic hiatus between P. pereger and the other members
of the predominantly northern flavomacuJatiis group.

Natural history. — All of the specimens of P. pereger were col-
lected by day in terrestrial situations (beneath rocks, in leaf litter,
and in moss along a road cut). All of the localities at which the
frogs were found are described by the collector as dwarf cloud
forest {monte chico of Terborgh, 1971), characterized by the small,
microphyllous trees.

A total of six males ( 2 immature, no vocal slits ) and 23 females
(4 immature, straight oviducts) comprise the total sample of this
species. The 19 adult females contain large, yellow eggs. In two
females, the largest eggs are only 1.5-1.7 mm in diameter contrast-
ing strongly with the much larger eggs (2.6-4.0 mm) in the other
females.

The cleared and stained females (KU 26112, 26111) had egg
complements of 18 and 20 eggs respectively.

Distrilmtion. — Known only from the Andean Cordillera Oriental
west of the Rio Apurimac and Cordillera Vilcabamba in Depto.
Ayacucho, Peru, between 2460 and 2650 m (Fig. 4). A single
specimen was evidently collected at 1650 m.

Remarks. — P. pereger is probably most closely allied to P. park-
eri. Both have the most T-shaped terminal phalanges of the species
of the flavoinaculatus group. The two species are readily separable
in that the tympanum is visible in pereger and concealed in parkeri,
the snout is rounded in pereger whereas it is sloping in lateral pro-
file in parkeri, and the parasphenoid is shorter in pereger (not
reaching the palatines) than in parkeri (Fig. 10). The two frogs
are about the same size and have similar proportions.

The discovery of a fiavomaculatiis group species so far south in
Peru suggests that there are probably several other populations
distributed along the higher elevations of the Amazonian versant of
the Cordilleras Central, Oriental, and possibly Vilcabamba. Once
these populations become known, it may prove to be the case that
fiavomaculatiis, parkeri, and pereger, are merely vicariant popula-
tions now separable in our ignorance.

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 35

Phrynopus peruanus Peters

Phnjnopus peruanus Peters, 1874:416 (Type-locality. — Maraynioc, Depto.

Junin, Peru; holotijpe not seen by author).
Borborocoetes peruanus — Boulenger, 1882:255.
E[upsophus] peruanus — Parker, 1932:344.

Diagnosis. — A moderate-sized Phrynopus ( 9 ? 23.3-30 mm SVL); skin of
dorsum and venter smooth with scattered low pustules; first finger shorter than
second; digit tips not bulbous; toes lacking lateral fringes and webbing; two
metatarsal tubercles, equal in size; tarsus lacking tubercles or folds; tympanum
visible, prominent, its length 2/5-1/2 that of eye; snout rounded in lateral
profile; prevomerine teeth present on small odontophores posteromedial to
choanae; frontoparietals complete, lacking prominent crests; nasals large, sepa-
rated medially; anterior ramus of parasphenoid long, not reaching palatines;
medial pterygoid ramus widely separated from parasphenoid ala; brown above
and below; venter may be reticulated with dark bro'WTi.

Description (based on 2 females). — Head narrower than body,
longer than wide; head width 32.2-38.2 % SVL; snout subacuminate
to rounded in dorsal view, rounded to weakly sloping in lateral
profile; snout short, E-N 59.1-63.6 % eye length; canthus rostralis
sharp, concave; nostrils not protuberant, directed laterally; loreal
region conca\'e, sloping gradually to lips; lips not flared; interorbital
space flat ( no frontoparietal crests ) , broad, upper eyelid width 74.4
% lOD; upper eyelid not tuberculate; supratympanic fold present,
glandular, not obscm-ing tympanum; tympanum visible, higher than
long, its length 41.8-52.3 % eye length; choanae small, wider than
long, slightly larger than a prevomerine odontophore, at edge of
palate but not concealed by palatal shelf of maxillary arch; pre-
vomerine odontophores well posterior and slightly medial to cho-
anae, bearing a clump of 2-4 teeth; tongue longer than wide, not
notched posteriorly, posterior one-half not adherent to floor of
mouth; males not seen.

Skin of dorsum smooth with low scattered warts ( not areolate ) ;
dorsum not bearing ridges or folds; skin of venter smooth; discoidal
folds obscure, well posterior on venter; no ulnar tubercles or folds;
one or two palmar tubercles (when two are present, outer is no
larger than a supernumerary tubercle); larger palmar tubercle
round, twice size of elongate thenar tubercle; a few supernumerary
tubercles distal to palmar tubercle(s); subarticular tubercles broader
than long, low, flat, non-conical; fingers lacking fringes or webbing;
fingers lacking pads or discs, not dilated apically; first finger slightly
shorter than second.

Shank 31.2-33.0 % SVL; no tarsal or heel tubercles; two promi-
nent metatarsal tubercles, both longer than wide, equal in size; no
supernumerary plantar tubercles; subarticular tubercles round, low,
flat, non-conical, smaller than those of fingers; toes not fringed or
webbed, tips not bulbous, lacking pads or discs.

In preservative, P. peruanus is brown above and below; the
venter is paler than the dorsum. The limbs are not or only feebly

36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

barred and the dorsal pattern, if present, consists of indefinite fleck-
ing of darker brown. A canthal stripe may be present or not. The
lips are not barred. The venter is uniform brown with occasional
cream flecks to creamy-brown with brown reticulation.

Distribution. — Known only from the type-locality, Maraynioc,
Depto. Junin, Peru (Fig. 14).

Remarks. — I have seen only two specimens of this species. The
smaller individual was partially dissected in order to gain some
knowledge of the skeletal features of this species. The frontoparietal
bones are complete and no fontanelle is exposed. The frontoparietal
bones are not in contact with the nasal bones. The latter are rela-
tively large and distinctly separated. The sphencthmoid is large
and extensive, paralleling the situation illustrated for P. peraccai
(Lynch, 1968, as Niceforonia festae) or P. montium (Lynch, 1971,
as Niceforonia montium). The palatal process of the premaxilla is
very long. The prevomers resemble those of P. montium except that
odontophores are present and bear teeth. The parasphenoid of P.
peruanus is like that of P. montium. The medial ramus of the ptery-
goid is shorter in P. peruanus than in P. montiu7n. The occipital
condyles are not stalked and are widely separated in P. peruanus.

Osteologically, P. peruanus is very similar to P. laplacai, P. mon-
tium, and P. wettsteini, but is easily differentiated in retaining pre-
vomerine teeth. The external features of P. peruanus do not readily
support this association. The nearly smooth skin of P. peruanus
suggests association with the frogs of the favomaculatus group or
with P. peruvianus. However, all of these species have a thumb
that is longer than the second finger. The apparently smooth skin
of P. peruanus may be an artifact of preservation. The areolations
lateral and ventral to the anus are nearly obliterated in the speci-
mens available to me. This suggests to me that the skin texture on
the venter and dorsum of the specimens I have seen may not be
reflective of the condition in living or carefully preserved frogs.
Peters' ( 1874) characterization of the skin texture as "etwas uneben"
suggests that the skin texture of his specimens was not clearly
smooth or areolate. At present, I consider P. peruanus closely related
to P. laplacai, P. montium, and P. loettsteini.

Phrynopus peruvianus (Noble), new combination

Sminthillus peruvianus Noble, 1921:1 (Holotype. — AMNH 14526, near Juliaca,

Depto. Puno, Pern ) .
Nohlella peruviana — Barbour, 1930:8L
(Eleutherodactylus) peruvianus — Lynch, 1971:148.

Diagnosis. — A small Phrynopus ( $ $ 16.4-21.7 mm, 9 9 20.6-25.1 mm
SVL); skin of dorsum and venter smootli; first finger slightly longer than or
equal to second; fingers and toes with bulbous tips; toes without lateral fringes
and basal webbing; two metatarsal tubercles, inner slightly larger than or equal
to outer; inner tarsal tubercle present; tympanum present, concealed (partially)
beneath skin; snout truncate in lateral profile; males with vocal sac and slits;

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 37

prevomerine teeth, dentigerous processes, and dentigerous ramus of prevomer
lacking; frontoparietals complete, not bearing crests; nasals widely separated;
anterior ramus of parasphenoid not reaching palatines; dorsum gray to brown,
blotched with dark brown; venter dark brown flecked with cream; throat of
males dull gray-brown.

Description. — Head narrower than body (Fig. 17), slightly wider
than long; head width 34.4-40.4 (x = 36.6) % SVL in c? c5 , 32.9-36.5
(x = 34.7) in 9 9 ; snout subovoid in dorsal view, truncate in
lateral profile; tip of snout extends further anteriad than anterior
edge of upper jaw; canthus rosti'alis sharp, concave; loreal region
concave, sloping abruptly to lip; lips not flared; nostrils not pro-
tuberant, directed laterally; snout short, E-N slightly less than to
equal to eye length; interorbital region flat, upper eyelid width
66.4-86.3 (x = 76.6) % lOD in ^ ^ , 71.5-93.8 (x = 81.2) in 9 9 ;
supratympanic fold glandular, not prominent; tympanum present,
concealed beneath skin but e\idenced externally by a depression in
the temporal region; postrictal tubercles prominent; choanae small,
round, situated well anterolaterally on palate, concealed by palatal
shelf of maxillae when roof of mouth is \iewed from directly above;
prevomerine teeth and dentigerous processes lacking; tongue nar-
row, much longer than wide, not notched posteriorly, posterior one-
fourth free; males with short, posterolateral vocal slits and a median,
external, subgular vocal sac ( Fig. 17 ) .

Skin smooth except for postrictal tubercles and scattered warts
on flanks, posteroventral surfaces of thighs, and tarsus; dorsolateral
folds are weakly evident in scapular region; no discoidal folds;
forearm lacking ulnar tubercles; palmar tubercle not bifid, larger
than thenar tubercle; palmar surface with a few prominent super-
numerary tubercles (Fig. 7); subarticular tubercles of fingers round,
flat, simple; fingers lacking lateral fringes; tips of fingers bulbous,
lacking pads; first finger slightly longer than second, or equal in
length to second.

Heel lacking tubercles; outer edge of tarsus bearing a row of
weakly developed tubercles; inner edge of tarsus bearing a promi-
nent, elongate, sigmoid-shaped tubercle (Fig. 8) not contiguous
with inner metatarsal tubercle; two metatarsal tubercles, both prom-
inent and subconical, inner slightly larger and longer than wide,
outer round; subarticular tubercles of toes conical, round, simple;
toes without lateral fringes and basal webbing; tips of toes bulbous,
lacking pads; hind limb short, heel of adpressed limb reaches to
between tvmpanum and posterior edge of eye; shank 38.2-45.2
(x = 42.9) % SVL in £ i, 38.0-46.0 (.x = 41.7) in 9 9.

In preservative the ground color is usually gray to brown with
dark brown markings; one indixidual is dull yellow above with
dark brown markings. The dorsal pattern is irregular and variable.
Eleven indixiduals have a mottled dorsum with no distinct pattern.
Ten others have a broad middorsal yellow stripe and a disruption

38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

of the brown markings which form a dark brown border with
irregular edges (see Noble's, 1921, Fig. 2); the dorsal stripe bifur-
cates at the vent and continues along the posterior surface of the
thigh. Thirty-seven others have a dorsal pattern of an interorbital
triangle, a large scapular X, and a sacral chevron.

The flanks and sides of the head are darker than the dorsum and
the facial markings consist of a canthal stripe, supratympanic stripe,
and labial bars. The limbs are barred and the bars are as wide as
the paler interspaces. All markings are edged with cream. Seven-
teen specimens have a hairline middorsal stripe from the sacrum
to the vent. The posterior surface of the thighs is black. The venter
is dull brown to black and weakly to heavily flecked with pale
cream. The throat is darker than the venter and rarely flecked with
cream. In the specimens with a broad middorsal stripe, the venter
has a sagittal stripe with a crossbar in the pectoral region; the
ventral lines are cream.

In life, the ground color varies from dull brown, tan, tan-orange,
to reddish and if bearing dorsal markings, they are dull red, tan,
or gray. The venter is bluish-gray with brown reticulations. The
throats of the males are dull grayish-brown. The middorsal stripe,
when present, is orange-yellow. The iris is dark brown with gold
flecks.

Natural history. — The KU specimens were collected beneath
rocks in ravines within the puna. In the ravines, shrubby vegetation
(2 m in height) was found. Most specimens of peruvianus were
found beneath moss-covered rocks in seepage areas within the
ravines. The adult females are gravid and calling males were en-
countered on the afternoon of 16 January 1971. The call was de-
scribed as "a series of short, quickly repeated, moderately high
notes, resembling a slow-calling Pseudacris triseriata" (William E.
Duellman field notes 16 January 1971 ) . The altitudinal range of
the KU sDecimens is from 3270 m to 3450 m.

Distrilnition. — Thnjnopus peruvianus is known only from the
localities cited in the appendix. The Abra Acanacu is a pass in
the Cadena del Paucartambo on the northwestern end of the Cordil-
lera Carabaya. The localities lie in moist paramo in conti-ast to the
stark habitat seen around Juliaca (Puno). Dr. Thomas Fritts is of
the opinion that Keays' collection was made in the more mesic
habitats to the northeast of Juliaca and may have come from a broad
assortment of habitats, elevations and localities in southern Peni
(Fig. 14).

Remarks. — Fhnjnopus peruvianus is osteologically isolated from
the other species of the genus. The extensive reduction of the pre-
vomers and fused pectoral girdle readily distinguish it from all
other species. No other species has a tarsal tubercle but the other
external features of P. preuvianus are not especially distinctive. In
view of the variation prevomers (especially reduction) seen else-

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 39

where among frogs, I am not willing to use that trait in conjunction
with the pseudofirmisterny to generically distinguish penwianus;
in other respects the species is too similar to other southern Andean
simple-toed eleutherodactylines.

Phrynopus simonsii (Boulenger), new combination

Paludicola simonsii Boulenger 1900:182 ( Cotypes.— BMNH RR 1947.2.15.43-
44, paramo; here restricted to 8 km E Cajamarca, Depto Cajamarca, Peni,
3000 m; BMNH 1947.2.15.43 is here designated the Lectoholotype of
Paulicola simonsii Boulenger).

Syrrhophus simonsii — Parker, 1927:451.

Niceforonia simonsii — Lynch, 1968:291.

Diagnosis. — A moderate-sized Phrynopus (single adult female 30 mm SVL);
skin of dorsum and venter areolate; dorsolateral folds present; first finger
shorter than second; digital tips bulbous; toes bearing lateral fringes and basal
webbing; two metatarsal tubercles, inner twice as large as outer; tarsus lacking
tubercles, indefinite inner tarsal fold present; tympanic annulus, columella,
and cavum tympanum absent; snout rounded in lateral profile; prevomerine
teetli and dentigerous processes absent, dentigerous ramus present, large;
frontoparietals complete, lacking crests; nasals large, narrowly separated; an-
terior ramus of parasphenoid in contact with palatines; dorsum tan, flanks
spotted with brown, groin with vertical bro\\Ti bars, venter cream.

Description of the lectoholotype (BiMNH 1947.2.15.43).— Head
as wide as body, as wide as long, head width 35.4 % SVL; snout sub-
acuminate in dorsal view, rounded in lateral profile ( Fig. 4 ) ; canthus
rostralis sharp, slightly concave; loreal region weakly concave, slop-
ing abruptly to non-flared lips; nostrils not protuberant, directed
dorsolaterally, much closer to tip of snout than to eye; snout short,
eye length greater than E-N; interorbital region flat, no cranial
crests; width of upper eyelid 93.3 % lOD; supratympanic fold promi-
nent; tympanum absent; tongue large, thick, fleshy, not notched
posteriorly, posterior one-half free; prevomerine teeth and dentig-
erous processes absent; choanae small, completely visible when
roof of mouth is viewed from directly above, just inside palatal
shelf of maxillae.

Skin of head and dorsum warty, at present warts are loosely
defined; glandular dorsolateral folds present from scapular region
to above groin; skin of limbs warty, venter areolate, throat less
areolate; skin on anterior surface of thighs smooth; shank 36.7 % SVL;
heel of adpressed limb reaches to posterior edge of head; forearm
bearing indefinite ulnar ridge; three palmar tubercles, outer poorly
defined (Fig. 6); no supernumerary thenar tubercles; subarticular
tubercles low, flat, round, simple; fingers lacking lateral fringes;
first finger shorter than second; tips of fingers swollen into pad-like
structures.

Inner edge of tarsus bearing indefinite fold; outer edge of tarsus
and heel lacking tubercles or folds; two metatarsal tubercles, inner
twice as large as outer, both ovoid, not compressed, non-conical;

40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

plantar surface bearing indefinite supernumerary tubercles; sub-
articular tubercles low, round, flat, simple; toes bearing lateral
fringes and basal webbing; tips of toes bulbous.

The ground color is tan. The flanks are spotted with brown and
on the upper flank is an elongated dark brown stripe-like spot. The
supratympanic fold is edged with brown. The venter is dirty cream
without spotting or reticulations. The groin is marked with a dark
brown vertical bar against a cream background. Dark brown
patches are present behind the knee and inside the heel. The limbs
are not barred.

Distribution. — Known only from the type-locality (Fig. 14).
During my stay at the British Museum I examined correspondence
between P. O. Simons, the collector, and Dr. Oldfield Thomas, then
curator of mammals and director of the British Museum. In a letter
from Simons to Thomas dated 20 November 1899, Simons refers to
a paramo locality some 5 miles east of Cajamarca. The type-locality
is here revised from "Paramo, Cajamarca " to 8 km E Cajamarca.

Remarks. — Three species of Plirynopus lack ears (cophites,
ruimis, and simonsii). The three also agree in having the first finger
shorter than the second, and in having coarsely areolate skin on the
venter. Although nanus differs from cophites and simonsii in having
visible dentigerous processes on the prevomer, I consider nanus and
simonsii closely related. The limited knowledge of the osteology of
nanus and simonsii reveals considerable similarity in contrast to the
marked dissimilarity of copliites. The osteological data suggests
that simonsii is more closely allied to the frogs of the fiavomaculatus
group than to the more southern peruanus group (which it resem-
bles in external characteristics ) .

Phrynopus wettsteini (Parker), new combination

Eupsophiis wettsteini Parker 1932:43. {Holotijpe. — NMW 15845, Untuaro or
Pongo, possibly Depto. Amazonas, Peru, 2000 m. Not examined hij author).
Niceforonia wettsteini — Lynch, 1969:273.

Diagnosis. — A small to moderate-sized Phnjnopus (known specimens 21-28
mm SVL); skin of dorsum and venter areolate; first finger shorter than second;
tips of fingers not bulbous; toes lacking lateral fringes and basal web; two
metatarsal tubercles, inner much larger than outer; tarsus lacking tubercles or
folds; tympanum partially concealed beneath skin; snout rounded in lateral
profile; males with vocal sac and slits; prevomerine teeth and dentigerous proc-
esses lacking, dentigerous ramus of prevomer present; frontoparietals complete,
lacking crests; nasals narrowly separated; anterior ramus of parasphenoid short,
not reaching palatines; dorsum dark brown, venter brown with darker reticu-
lations; throat black.

Descriptions. — See Parker ( 1932) .

Distribution. — Known only from the type-locality, presumably
on the Amazonian slopes of one of the Andean Cordilleras ( Fig. 14 ) .

Remarks. — The only specimen of this species which I have
examined in detail relative to its relationships to other Fhnjnopus

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 41

species is the cleared and stained example. Although the other four
paratypes were studied, my notes on them deal only with the generic
affinities of this small frog and my data are thus inadequate for
species-level comparisons.

Osteologically, P. ivettsteini agrees well with P. laphcai and P.
montium, both of which occur at much higher elevations than that
recorded for P. wettsteini. There is considerable external similarity
between these three frogs as well. The principal differences, judg-
ing from the unsatisfactory position of small samples, involve rela-
tive sizes of the metatarsal tubercles, relative lengths of the inner
two fingers, and coloration. Aside from the striking altitudinal and
geographic differences in type-localities, and the very small samples
available, I might argue that all three are conspecific.

The type-locality of P. wettsteini is not known with certainty.
Dr. Josef Eiselt informed me that in the records at Vienna, the word
"Pongo" was associated with or situated near Borja, a small town on
the Rio Maraiion near the frontier between Depto. Amazonas and
Loreto. The mountains in that \icinity reach 2000 m and the type-
locality of wettsteini is presumed to be in the vicinity of Borja.

One other nominate species has been referred to this generic
grouping — Borhorocoetes columhiana Werner. The holotype of the
species is now lost and no other specimens have been assigned to
the species. If Werner's (1899) description and my inferences from
the description concerning character states not specifically men-
tioned by Werner are accurate, the frog may be a Phnjnopus allied
to P. brunneiis and P. peraccai. A partial diagnosis of the frog is
presented below; the character states in italics are suppositions on
my part, not data given by Werner.

Phrynopus columbianus (Werner), new combination

Borhorocoetes columhiana Werner 1899:480 (Holotype now lost; Monte
Redondo, Buenavista, Depto. Cundinamarca or Meta, Colombia, 1000-
1300 m).

Niceforonia columhiana — Lynch, 1968:292.

Diagnosis. — A small Phnjnopus (an individual of unknown sex and age,
18 mm SVL); skin of dorsum warty, that of venter smooth; first finger equal
in length to second; fingers and toes not dilated at tips; toes basally webbed;
inner metatarsal tubercle small, outer absent (unlikely; outer prohahly ohscure
hut much smaller than inner); inner edge of tarsus lacking tuhercle; tympanum
prominent, M length of eye; snout rounded or truncate in lateral profile; pre-
vomerine dentigerous processes present, posteromedial to choanae; dorsum dark
brown with little or no pattern other than pale interorbital bar; venter white
with brown reticulation.

Remarks. — Cochran and Coin (1970) and Werner (1899) sug-
gested that the frog was immature; the suggestion is based on the
small size of the type. However, it may be an adult {nanus and
peraccai are only slightly larger frogs). If the frog is a Plirynopiis,
it is most similar to the two Ecuadorian species found at higher

42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

elevations on the Amazonian versant of the northern Andes (P.
brunneiis and P. peraccai). I doubt that it is identical to either of
the Ecuadorian species; this conclusion is based on Werner's ( 1899)
description of the color pattern and the purported presence of basal
webbing of the toes of the holotype.

INTRAGENERIC RELATIONSHIPS

If P. columhianus is included, the genus Fhnjnopus contains
fourteen nominate species distributed in the Andes from northern
Colombia to western Bolivia at elevations of 1000-4000 m (Fig. 18).
A single case of sympatry is known (P. copluies and P. penwianns)
for the otherwise dispersely distributed genus. The relationships of
the frogs of the genus Fhnjnopus to other eleutherodactylines are
obscure, but the genus is probably most closely related to Eleuthero-
dactylus (Lynch, 1971). The simple digits characteristic of
Fhrijnopiis could represent the primitive character state for eleu-
therodactyline frogs or may represent a retrograde evolutionary
step. The digital character state seen in several Fhnjnopus species
(no discs, terminal phalanges intermediate between knobbed and
T-shaped ) could support either argument.

The fourteen species are divided into four species groups; these
are defined below:

flavomacidatiis group: first finger longer than second; skin smooth;
toes lacking lateral fringes and basal webbing; no tarsal tubercle;
prevomerine teeth present on processes lying posteromedial to
choanae; median pterygoid ramus in contact with parasphenoid
ala; anterior parasphenoid ramus in contact (or nearly so) with
palatines; ear fully developed; pectoral girdle functionally arcif-
eral. Frogs of this group also have comparatively large nasal
bones.

Content: P. hrunneus, P. cohimbianus, P. fiavomaculatus, P.
parkeri, P. peraccai, and P. pereger.

penwianns group: first finger as long as second (or slightly longer);
skin smooth; toes lacking lateral fringes and basal webbing;
inner tarsal tubercle present; prevomerine teeth, dentigerous
processes, and dentigerous ramus absent; pterygoid not in con-
tact with parasphenoid; anterior parasphenoid ramus short, not
reaching palatines; ear fully developed; pectoral girdle func-
tionally firmisternal. The nasal bones are relatively small and
widely separated medially.
Content: P. peruviamis.

simonsii group: first finger shorter than second; skin areolate; toes
bearing lateral fringes and basal webbing; no tarsal tubercle;
prevomerine teeth and processes present {nanus) or absent
(simonsii), when absent, dentigerous ramus not reduced in size;

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 43

Fig. 18. — Clinal variation in prevomerine bones in Phrijnopus. (A,B)
flavomaculatus and simonsii groups, (C) peruanus group, and (D) peruvianus
group.

44 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

median pterygoid ramus in contact with parasphenoid ala
(simonsii); anterior parasphenoid ramus long, in tenuous con-
tact \^dth palatines ( simonsii ) ; ear absent ( possibly only greatly
reduced in nanus); pectoral girdle functionally arciferal. The
nasals are large and in contact or narrowly separated.
Content: P. nanus and P. simonsii.

peritanus group: first finger not longer than second (usually
shorter); skin areolate (condition uncertain in peruanus, see
description); toes lacking lateral fringes and basal webbing; no
tarsal tubercle; prevomerine teeth and dentigerous processes
lacking (except in peruanus) , dentigerous ramus thin and sliver-
like; median pterygoid ramus narrowly (laplacai, nwntium,
peruanus, wettsteini) or broadly (copliites) separated from
parasphenoid ala; anterior parasphenoid ramus short, not reach-
in palatines; ear fully developed (except cophites, ear absent);
pectoral girdle functionally arciferal. The nasal bones are
moderate-sized and narrowly separated.

Content: P. cophites, P. laplacai, P. montium, P. peruanus, and
P. wettsteini.

On the basis of external characteristics (including those of the
mouth), I consider the jlavomaculatus and peruvianus groups more
similar to one another than to the other two species groups, which
form a compact unit. The osteological data suggest that the jlavo-
maculatus and simonsii groups are closely related and that veru-
vianus and peruanus groups are closely related. This lack of con-
cordance is suggestive that the genus Phrynopus may not be
diphyletic but that early in its evolution gave rise to two lineages,
one leading to the jlavomaculatus group and the second giving rise
to the predominantly southern component.

In an attempt to test my subjectively determined grouping of
Phrynopus, I analyzed the intrageneric relationships with the
method described by Camin and Sokal ( 1965 ) . Eleven character-
istics (number 1-11), each with two to four character-states, were,
used. Data are incomplete for three species (P. columhianus, P.
nanus, and P. peruanus) and these taxa have accordingly been ex-
cluded from the analysis. The characteristics, character-states, and
coding (0 = primitive, 1 or 1' derived, 2 derived from 1, and 3
derived from 2 ) are listed below :

1. — Prevomers. Four character-states. 0: prevomer large, toothed;
1: prevomer large, toothless; 2: prevomer moderate-sized,
edentate or not, dentigerous ramus thin (sliver-like); 3: pre-
vomer minute, dentigerous ramus lost.

2. — Ears. Two character-states. 0: ear present, visible or con-
cealed; 1: ear absent.

3. — Frontoparietals. Three character-states. 0: no cranial crests.

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 45

no fontanelle (complete frontoparietals); 1: no cranial crests,

fontanelle present; 1': cranial crests present, no fontanelle.
4. — Parasphenoid ala. Three character-states. O: ala in contact

with median pterygoid ramus : 1 : ala narrowly separated from

median pterygoid ramus; 2: ala widely separated from median

pteiygoid ramus.
5. — Anterior parasphenoid ramus. Two character-states. 0: not

reaching palatines; 1 : in contact with palatines.
6. — Nasal bones. Two character-states. 0: large nasals, in broad

median contact; 1 : nasals smaller, narrowly to relatively widely

separated medially.
7. — Skin of venter. Two character-states. 0: texture smooth; 1:

texture coarsely areolate.

8. — Finger lengths (I and II). Two character-states. 0: thumb
longer than second finger; 1: thumb not longer than second
finger, usually shorter.

9. — Toe fringes. Two character-states. 0: present; 1: absent.

10. — Snout shape. Two character-states. 0: rounded or truncate in
lateral profile; 1 : sloping in lateral profile.

11. — Metatarsal tubercles. Two character-states. 0: outer meta-
tarsal tubercle much smaller than inner; 1: outer metatarsal
tubercle more than one-half size of inner ( equal in most cases ) .

Because both derived character-states of character 3 are unique
{i.e., a derived character-state is evident in less than two taxa),
characteristic 3 was not used in the analysis. The derived states are
indicated on the cladogram (Fig. 19) by a dash accompanied by
the number of the characteristic. In determining evolutionary di-
rection within characteristics, I considered widespread character
states ("normal" among frogs as a whole, but especially among the
archaic groups) as well as character states occurring in putative
primitive eleutherodactyline genera (viz., Hylactophryne and Isch-
nocnema) to be primitive character-states. Those character states
found among Phrijnopus, but not seen in the species of more primi-
tive telmatobiines (Grypiscini, Telmatobiini, Batrachylini, Odonto-
phrynini) or among most eleutherodactylines, are considered to be
derived.

Inspection of the cladogram (Fig. 19) reveals a concordance of
groupings to those proposed above (flavomaculatus, peruanus, and
simonsii species groups) in the cladogram using subcladograms A
or A', and B. Using the equally parsimonious subcladogram B'
requires considering the penivianiis and peruanus groups not de-
fensibly separable on the basis of the characteristics and character-
states employed here. Subcladogram B emphasizes characteristic 11
(metatarsal tubercle size) at the expense of an extra evolutionary
step for characteristics 4 and 8; subcladogram B' emphasizes 8
(finger lengths) at the expense of extra steps for 4 and 11; emphasis

46

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

b a c d e

Fig. 19. — Most parsimonious cladogram (27 evolutionary steps) for eleven
species of Phrynopus. Equally parsimonious subcladograms for the jiavomacu-
latus group ( A', upper left ) and the peranus-peruvianus groups ( B', top right )
are also illustrated. The dashes across lines (and the attending number)
represent evolutionary shifts from a primitive to a derived character state.

of characteristic 4 or 7 results in less parsimonious cladograms ( the
number of steps is increased from 27 to 28).

My preference among the equally parsimonious cladograms is
subcladogram B, as indicated in the recognition of a peruvianus
group. In support of this, I cite the functionally firmisternal pec-
toral girdle of peruvianus, a feature not duplicated elsewhere in the
genus, and the likewise extensive reduction of the prevomers.

INTERGENERIC RELATIONSHIPS

In the course of my studies of South American Eleutherodac-
tylus, I have concluded that there are two major groupings of South
American species — (1) the binotatus and fitzingeri groups, con-
taining those frogs having the first finger longer than the second
and having smooth or feebly granulate skin of the venter, and (2)
the unistrigatus group, containing species having the first finger
much to slightly shorter than the second and having coarsely areo-
late skin on the venter. In excess of 95% of the nominate species
from South America fall into the two contingencies. The species
of the Phrynopus flavomaculatus group bear considerable external
as well as osteological similarity to the frogs of Eleutherodactylus
binotatus group. The recently described E. elassodiscus (Lynch,
1973) is almost perfectly annectant between other narrow-toed

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 47

species of the E. hinotatus group and the larger species of the P.
flavomaculafus group, except that it has digital discs as do other
Eleiitherodacfyhts. The similarity between these frogs is so exten-
sive that I am convinced that the P. favomactilatus group is an
upland derivati\'e of the E. fitzingeri group (a predominantly low-
land group — below 1000 m ) .

Parker's ( 1932) suggestion that some of these frogs (the simonsii
and peruanus groups) were derived from a "Borhowcoetes" group
of Eleittherodacfylus is plausible if the two groups of EUutherodac-
tylus mentioned above are natural groups. The E. tinistrigatus
group and the P. simonsii and P. peruanus groups agree in relative
finger lengths and the texture of the skin of the venter. In addition,
many of the Andean species of the E. tinistrigatus group in southern
Ecuador have narrowly separated nasal bones, as do the frogs of
the P. peruanus group. Several high-altitude species of the E. uni-
strigatus group have reduced digital pads and small prevomerine
dentigerous processes (thus approaching the P. simonsii and P.
peruanus groups), but differ in having discs on the digits.

One difficulty with deriving the flavomacuhtus group from the
E. fitzingeri group and the simoivsii and peruanus groups from the
E. tinistrigatus group is the disposition of P. peruviamis. Osteologi-
cally, P. penwianus is much more similar to the frogs of the P.
peruanus group than to the P. flavomaculatus group or the P. simon-
sii group, whereas, on the basis of external characteristics, P. peru-
viamis more closely resembles the frogs of the P. favomacuhfus
group. The lack of concordance between species groups of Phryno-
piis and the major groups of South American Eleutlierodactyhis does
not support the idea that Phrynopus is diphyletic. Although I have
not yet examined the assumption cladistically, I assume Phrynopus
to be derived from an EJeutlierodactylus stock not unlike the group
of species now represented by E. crtiralis, E. elassodiscus, E. granu-
losus, and E. nigrovittatus. This complex of species occurs along the
eastern base of the Andes in Ecuador and in southern Peru and
adjacent Bolivia.

SUMMARY

The Andean eleutherodactyline frogs ha^'ing narrow heads and
lacking digital discs are placed in the genus Phrynopus. The genera
Niceforonia and Nohlella are synonyms. Fourteen species are recog-
nized; five of them are named as new species. Phrynopus hrunneus
is named from the high Amazonian Andean slopes (valley of the
Rio Chingual) in Ecuador. Phrynopus peraccai is named from the
high Amazonian Andean slopes (\alley of the Rio Fapallacta) in
Ecuador; this species was previously known as Niceforonia festae.
Phrynopus parheri is named from a mountain crest in the Huanca-
bamba pass in northern Peru. Phrynopus pereger is named from

48 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

intermediate elevations on the eastern slopes of the Cordillera
Oriental in south-central Peru. Fhnjnopus copliites is named from
high elevations in the Cordillera Carabaya in southern Peru.

Four species groups are recognized. The flavomacidatus group
includes six species (brunneus, cohimhianus, fiaxjomacidatus, park-
eri, peraccai, and pereger), five occurring in central Colombia, Ecua-
dor, and extreme northern Peru, and one (pereger) found in south-
ern Peru. The generotype of NobJella (penwiamis) is the only
species included in the penwiamis group (southern Peru). The
simonsii group included nanus (generotype of Niceforonia) from
north-central Colombia and simonsii from northern Peru. The peru-
anus group includes five species, four from southern Peru and
Bolivia {cophites, Japlacai, montium and peruanus) and one ap-
parently found on the Amazonian slopes of northern Peru (tvett-
steini).

Some characteristics used to distinguish genera exhibit variation
among the species of Fhnjnopus. Three species {cophites, nanus,
and simonsii) lack ears. Prevomerine teeth and odontophores are
absent in the species of the peruviamis and peruanus groups as well
as in simonsii, but are uniformly present in species of the fiavomacu-
latus group. One species {cophites) has a large frontoparietal fonta-
nelle; one other species (flavomaculatus) has prominent cranial
crests. Phnjnopus peruvianus is unique in having a functionally
firmisternal pectoral girdle.

The distribution of Fhrynopus is diperse. A single case of sym-
patry is known {cophites and peruvianus) . Only flavomaculatus is
known to occur over any modest area; the other thirteen species
occur at single localities or at a few clustered localities. A major
distributional gap occurs from northern Peru to central Peru and
many additional populations are expected in the 600 kilometer dis-
tributional hiatus.

RESUMEN

Las ranitas andinas eleutherodactilinas que tienen la cabeza
angosta y carecen de discos digitales se colocan en el genero
Fhrynopus. Los generos Niceforonia y Nohlella son sinonimos. Se
reconocen catorce especies denti-o de este grupo de ranitas, cinco
de las cuales son nuevas para la ciencia. Se nombra a Fhrynopus
brunneus de las faldas andinas altoamazonicas del Ecuador (Valle
del Rio Chingual). Se nombra a F. peraccai de las faldas andinas
altoamazonicas del Ecuador ( Valle del Rio Papallacta ) ; esta especie
se le conocio previamente como Niceforonia festae. Se nombra a
Fhrynopus parkeri de la cresta montanosa del paso Huancabamba
en el norte de Peru. Se nombra a Fhrynopus pereger de elevaciones
intermedias en las faldas del este de la Cordillera Oriental del sur-
centro de Peru. Se nombra a Fhrynopus cophites de las altas eleva-
ciones de la Cordillera Carabaya en el sur de Peru.

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 49

Se reconocen cuatro grupos de especies. El giupo jiavomacu-
latus incluye seis especies {hrunneus, columhianus, favomaculatus,
parkei'i, peraccai, y pereger), cinco se distiibuyen en el centre de
Colombia, Ecuador, y en el extremo norte de Peru, y una (pereger)
se encuentra en el sur de Peru. El generotipo de NohleUa {peru-
vianus) es la unica especie que se incluye en el grupo penwianus
(sur de Peru). El grupo simonsii incluye a nanus (generotipo de
Niceforonki) de la parte nor-central de Colombia y a simonsii del
norte de Peru. El grupo peruanus incluye cinco especies, cuatro
del sur de Peru y Boli\'ia {cophites, laplacai, montium, y peruanus)
y una que aparentemente se encuentra en las faldas amazonicas del
norte de Peru {wettsteini) .

Algunas de las caracteristicas usadas para separar a los generos
muestran variacion entre las especies de Fhnjnopus. Tres especies
carecen de oido {cophites, nanus, y simonsii). Las especies de los
grupos peruanus y peruvianas, asi como del grupo simonsii, carecen
de dientes prevomerianos y odontoforos, pero estan presentes en las
especies del grupo flavomaculatus. Una especies {cophites) tiene
una fontanela frontoparietal grande; otra especie {flavomaculatus)
tiene crestas craneales prominentes. Phnjnopus peruvianas sobre-
sale del resto de las especies por tener una cintura pectoral firmi-
sterna funcional.

La distribucion geografica de Phnjnopus es dispersa. Solo se
conoce un caso de simpatridas {cophites y peruvianus) . Solamente
flavomaculatus presenta una distribucion modesta; las otras trece
especies se conocen de una sola localidad o de pocas pero muy
juntas Unas de las otras. Hay gran discontinuidad distributiva desde
el norte hasta el centio de Peru, y es de esperarse muchas pobla-
ciones adicionales en este hiato de 600 kilometros.

APPENDLX: SPECLMENS EXAMINED

Phnjnopus hrunneus. — ECUADOR: Carchi: 10 km WNW El Carmelo,
3150 m, USNM 192909-12.

Phnjnopus cophites. — PERTJ: Cuzco: S slope Abra Acanacu, 14 km NNE
Paucartambo, 3400 m, KU 138884; N slope Abra Acanacu, 27 bn NNE Paucar-
tambo, 3450 m, KU 138885-908, 138909-10 (cleared and stained skeletons),
138911-16; N slope Abra Acanacu, 29 km NNE Paucartambo, 3400 m, KU
138918.

Phnjnopus jiavomaculatus . — ECUADOR: Loja: 13.2 km E Loja, 2770 m,
KU 141474; 13 5 km E Loia. 2800 m, KU 141475; 15 km E Loja, 3000 m.
BMNH 1947.2.16.11-15; 8-9 km N San Lucas, 3000-3100 m, KU 119737-42,
121354 (eggs); 10 km S Saraguro, 3100 m, KU 142201-02. Morona-Santiago:
San Juan Bosco, 2215 m, USNM 195393-994; between Sevilla de Oro and
Mendez, 2460-2580 m, USNM 195395, 19S425-27; between Suro Rancho and
Sapote, 2650 m, USNM 195396-97; Suro Rancho, 2710 m, USNM 195398-411;
Yi km W Suro Rancho, 2770 m, USNM 195412-17; San Vicente, 2860 m,
USNM 195418, 195419 (cleared and stained skeleton), 195420-23; Cerro
Negro, 2950 m, USNM 195424. Zamora-Chinchipe: 13-14 km E Loja, 2800
m, KU 119721-24, 119743 (cleared and stained skeleton); 14 km E Loja,

50 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

2780 m, KU 141476-77; 15 km E Loja, 2710 m, KU 142198-200; 15 km E
Loja, 2800 m, KU 119716-20, 119725-36.

Phrynopus laplacai.— BOLIVIA: La Paz: 50 km NE La Paz, KU 154555-
59; 3 km W Pongo, 3400 m, IBN/UNC 1251.6 (cleared and stained skeleton).

Phrynopus montium. — PERU: Jiinin: Cascas, MCZ 24352 (2 specimens,
one a cleared and stained skeleton ) .

Phrynopus nanus. — COLOMBIA: Santander: Paramo de la Rusia, USNM
150643.

Phrynopus parkeri. — PERU: Piura: summit of cordillera between Chancha-
que and Huancabamba, 3100 m, KU 135278-305, 135306 (cleared and stained
skeleton), 135307-11.

Phrynopus peraccai. — ECUADOR: Napo: Papallacta, 3110 m, MIZ 820,
USNM 164397; 1 km W Papallacta, 3155 m, KU 111430, USNM 160944 and
160950 (cleared and stained skeletons), 160954-59, 160961, 160966-68,
164398; 4 km W Papallacta, 3300 m, KU 117795-96, 118137 (cleared and
stained skeleton); Laguna Papallacta, 3350 m, KU 109171.

Phrynopus pereger. — PERU: Ayacudw: Ccarapa, below Tambo on Aya-
cucho-Valle del Apurimac road, 2460 m, LSUMZ 26124-31; Huanhuachayocc,
1650 m, LSUMZ 26100; Mitupucuru, 2425 m, LSUMZ 26115-16; between
Mitupucuru and Yuaccyacu, LSUMZ 26101-10, KU 151907 (cleared and
stained skeleton), 151908-09; Yuraccyacu on Tambo- Valle del Apurimac
trail, 2650 m, KU 151906 (cleared and stained skeleton), LSUMZ 26117-23,
26255.

Phnjnopus peruanus. — PERU: Junin: MarajTiioc, UMMZ 89477(2).

Phryrwpus peruvianus. — PERU: Cuzco: N slope Abra Acanacu, 27 km
NNE Paucartambo, 3450 m, KU 138919-25, 138926 (cleared and stained
skeleton) 138927-35; 29 km NNE Paucartambo, 3400 m, KU 138917, 138936-
64; 31 km NNE Paucartambo, 3270 m, KU 138968-69 (cleared and stained
skeletons), 138970-78. Puno: near Juliaca, AMNH 14526, and one uncata-
loged cleared and stained skeleton.

Phrynopus simonsii. — PERU: Cajamarca: paramo, ca 8 km E Cajamarca,
3000 m, BMNH 1947.2.15.43-44.

Phrynopus tvettsteini. — PERU: ? Amazonas: Untuaro or Pongo, 2000 m,
BMNH 1932.4.30.1-2, 1932.4.30.3 (cleared and stained skeleton), NMW
15846.1-2.

LITERATURE CITED

Barbour, T. 1930. A list of Antillean reptiles and amphibians. Zoologica,
11:61-116.

BouLENGER, G. A. 1882. Catalogue of tlie Batrachia Salientia s. Ecaudata in
the collections of the British Museum. 2nd ed. British Museum (Nat,
Hist.), 503 p.

BouLENGER, G. A. 1900. Descriptions of new batrachians and reptiles collected
by Mr. P. O. Simons in Peru. Ann. Mag. Nat. Hist., (7)6:181-86.

Camin, J. H., SoKAL, R. R. 1965. A method for deducing branching sequences
in phylogeny. Evolution, 19:311-26.

Cei, J. M. 1968. A new frog of the genus Syrrhophus from the Bolivian plateau.

J. Herpetol., 2:137^1.
Cochran, D. M., Coin, C. J. 1970. Frogs of Colombia. Bull. U.S. Natl. Mus.,

(288): 1-655.

Coin, C. J., Cochran, D. M. 1963. Two new genera of leptodactylid frogs
from Colombia. Proc. California Acad. Sci., 31:499-505.

Gorham, S. W. 1966. Liste der rezenten Amphibien und Reptilien. Ascaphi-
dae, Leiopelmatidea, Pipidae, Discoglossidae, Pelobatidae, Leptodactyli-
dae, Rhinophrynidae. Das Tierreich, 85:1-222.

ANDEAN LEPTODACTYLID FROG GENUS PHRYNOPUS 51

Griffiths, I. 1959. The phylogeny of Sminthillus limbatus and the status of
the Brachycephahdae (Amphibia, Sahentia). Proc. Zool. Soc. London,
132:457-87.

Lynch, J. D. 1968. Systematic status of some Andean leptodactyhd frogs with
a description of a new species of Eleutherodactylus. Herpetologica, 24:
289-300.

Lynch, J. D. 1969. Taxonomic notes on Ecuadorian frogs (Leptodactyhdae:
Eleutherodactylus) . Ibid., 25:262-74.

Lynch, J. D. 1970. Identity of two Andean Eleutherodactylus with the de-
scription of a new species (Amphibia: Leptodactyhdae). J. Herpetoh,
3:135-43.

Lynch, J. D. 1971. Evolutionary relationships, osteology, and zoogeography of
leptodactyloid frogs. Univ. Kansas Mus. Nat. Hist., Misc. Pubis., (53):
1-238.

Lynch, J. D. 1973. A new narrow-toed frog from Andean Ecuador (Lepto-
dactyhdae: Eleutherodactylus). Copeia, 1973(2) :222-25.

Lynch, J. D. 1974. New species of frogs (Leptodactyhdae; Eleutherodactylus)
from the Amazonian lowlands of Ecuador. Occ. Pap. Mus. Nat. Hist.
Univ. Kansas, (31): 1-22.

Noble, G. K. 1921. Five new species of Salientia from South America. Amer.
Mus. Novitates, ( 29 ) : 1-7.

Parker, H. W. 1927. A revision of the frogs of the genera Pseudopaludicola,
Physalaemus, and Pleurodema. Ann. Mag. Nat. Hist., (9)20:450-78.

Parker, H. W. 1932. The systematic status of some frogs in the Vienna
Museum. Ibid., (10)10:341-44.

Parker, H. W. 1938. The vertical distribution of some reptiles and amphibians
in southern Ecuador. Ibid., (11)2:438-50.

Peracca, M. G. 1904. Rettili et amfibii. Viaggio del Dr. Enrico Festa nell'
Ecuador e regioni vicine. Boll. Mus. Zool. Anat. Comp., 19:1-41.

Peters, W. 1874. Zwei Giftschlangen aus Afrika und iiber neue oder weniger
bekannte Gattungen und Arten von Batrachiem. Monatsb. der konig-
lich preussichen Akademie der Wissenschaften zu Berlin, 1873:411-18.

Shreve, B. 1938. A new Liolaemus and two Syrrhopus from Peru. J. Wash-
ington Acad. Sci., 28:404-407.

Terborgh, J. 1971. Distribution on environmental gradients: theory and a
preliminary interpretation of distributional patterns in the avifauna of
Cordillera Vilcabamba, Peru. Ecology, 52:23—40.

Werner, F. 1899. Ueber Reptilien und Batrachier aus Colombien und Trini-
dad. Verb. Zool.-Bot. Ges. Wien, 49:471-84.

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