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w

OCCASIONAL PAPERS

of the

MUSEUM OF NATURAL HISTORY

The University of Kansas

Lawrence, Kansas

MU9. t

NUMBER 16, PAGES 1-66 SEPTEMBER 7, 1973

A REVIEW OF THE CENTROLENID FROGS

OF ECUADOR^^^
WITH DESCRIPTIONS OFiWgjKf^pf lES

K) By

o John D. Lynch^ and William E. Duellman"

Since the description of Centrolene geckoidcum by Jimenez de
la Espada (1872), there have been few contributions to our knowl-
edge of Ecuadorian centrolenid frogs. Boulenger ( 1882, 1898, 1899)
named three Ecuadorian species as Hyla or Hijlella. Noble (1924)
recorded another species. Goin ( 1961 ) named three new species,
and Goodman and Goin (1970) commented on additional Ecua-
dorian specimens of Centrolene geckoicleum.

The generic status of these frogs has been reviewed by Noble
(1920), Taylor (1951), Goin (1964) and Savage (1967).' Taylor
(1951) perceived the distinctive features of the assemblage and pro-
posed the recognition of the Centrolcnidae, a family occurring from
southern Mexico to Peru and Surinam and with an additional pro-
liferation of species in southeastern Brasil.

The Middle American species have been reviewed by Taylor
(1949, 1952, 1958), Duellman and Tulecke (1960), Savage (1967),
and Savage and Starrett (1967). Ta>'lor and Cochran (1953) sum-
marized the Brasilian species; Cochran and Goin (1970) accounted
for the known Colombian species, and Rivero (1968) published on
the Venezuelan centrolenids. Additional species have been named
from Peril (Boulenger, 1918), Surinam (Goin, 1966), Brasil (Lutz
and Kloss, 1952), and Guyana (Goin, 1968).

^ Associate in Herpetology, Museum of Natural History, University of Kan-
sas, and Associate Professor, Department of Zoology, University of Nebraska,
Lincoln, Nebraska 68508.

" Curator, Division of Herpetology, Museum of Natural History, University
of Kansas.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Currently the Centrolenidae is composed of two genera — Cen-
trolene Jimenez de la Espada, 1872, and Centrolenella Noble, 1920;
approximately 50 species are recognized in the family. Eight species
have been recorded from Ecuador: Centiolene geckoideum and
Centrolenella buckleyi, cochranae, fleischmanni, griffithsi, ocellifera,
parabambae, and petersi.

Field work in Ecuador from 1967 through 1972 has resulted in
the accumulation of nearly 200 specimens, representing all of the
previously known Ecuadorian species of Centrolenella, two species
heretofore unknown from Ecuador, and eleven new species named
herein. We have examined the type specimens of all Ecuadorian
taxa. Our investigations show the presence of 19 species of centro-
lenids in Ecuador. We present data on all the Centrolenella but
include Centrolene geckoideum only in the key. Additionally, we
treat the Colombian Centrolenella medemi, although the species has
yet to be found in Ecuador.

Most areas of Ecuador have been explored in the course of our
field work. Duellman worked mostly in the Amazonian lowlands
and slopes, whereas Lynch spent most of his time at high elevations
in the Andes and on the Pacific slopes of the Andes. Both of us
spent limited time on the mesic Pacific lowlands. Based on our ex-
perience, we have noted a high degree of endemism in the faunas
on the Pacific and Amazonian slopes and in some semi-isolated
mountain ranges in Ecuador. We suspect that the centrolenid fauna
is much larger than the 19 species now known from Ecuador."^ Ac-
cordingly, in this paper we examine the current state of our knowl-
edge of Ecuadorian centrolenids.

Acknowledgments

For the loan of specimens or for provision of working space in
their respective institutions, we are grateful to James R. Dixon, Alice
G. C. Grandison, M. S. Hoogmoed, Konrad Klemmer, Charles W.
Myers, Gi.inther Peters, Douglas A. Rossman, Charles F. Walker,
and George R. Zug. We are grateful to the following persons whose
efforts in the field enhanced our collections of centrolenid frogs:
Joseph T. Collins, Martha L. Crump, Stephen R. Edwards, Thomas
H. Fritts, Marsha Lynch, Bruce MacBryde, John E. Simmons, Ger-
ald R. Smith, Linda Trueb, and Charles F. Walker. Field work in
Ecuador was partially supported by Watkins Museum of Natural

" Subsequent to the completion of this paper, we received a specimen of an
apparently unnamed species of Centrolenella, obtained at the Estacion Biologica
Rio Palenque, Provincia Los Rios. The specimen would be identified in our
key as C. cochranae.

CENTROLENID FROGS OF ECUADOR 3

History Grants, University of Kansas (Duellman and Lynch), So-
ciety of Sigma Xi (Lynch), Penrose Fund (Grant no. 5549) of the
American Philosophical Society (Lynch), and the Committee on
Systematics and Evolutionary Biology, University of Kansas (Lynch).
Duellman's study of specimens in European museums was made
possible by a grant (no. 5063) from the Penrose Fund of the Amer-
ican Philosophical Society.

Abbreviations for collections used throughout the text are:

AMNH American Museum of Natural History

BMNH British Museum (Natural History)

CAS California Academy of Sciences

KU University of Kansas Museum of Natural History

LSU Louisiana State University

RMNH Rijksmuseum van Natuurlijke Historic

SMF Senckenbergische Museum Frankfurt

TCWC Texas Cooperatixe Wildlife Collection

UMMZ Uni\ersity of Michigan Museiun of Zoolog>'

USNM United States National Museum ( National Museum of Natural

History )

ZMB Zoologisches Museum Berlin

TAXONOMIC GHARACTERS

With the exception of Centrolene geckoideum, living centro-
lenids usually are small, slender-limbed, green frogs. Most preserved
examples seem to offer a limited suite of characters useful in their
classification. The apparent paucity of characters is in part illusory,
for there has been a multiplicitx^ of names applied to comparatively
few species in some areas; furthermore, there has been a limited
amount of well-preserved material with detailed records of color-
ation of living frogs. In the following paragraphs we discuss those
taxonomic characters found to be useful by us and contemporary
investigators (Savage, 1967; Savage and Starrett, 1967).

Prevomerine teeth. — Prevomerine teeth and dentigerous proc-
esses are absent in 9 species: anomalo, Inickleiji, fieischmanni, grif-
fithsi, megachemi, munozomm, peUucida, peristicta, and pipilata.
Judgment on the absence of teeth should be held in abeyance for
anomala and pcUucida, inasmuch as we have a single specimen of
each. Centrolenella peUucida is a member of the fieischniajini
group, and if it does have prevomerine teeth, it would be the only
species of the group to have them. Centrolenella anomala seems to
be allied most closely to C. cochranae, some specimens of which
lack prevomerine teeth. Prevomerine teeth usually are absent in
grandisonae. The teeth usually are present in cochranae, flavopunc-
tata, ocelUjera, prosohJepon, and siren, and are found in all ex-

4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

amples examined of audax (4), medemi (1), rnidas (11), and
resplendens (2).

Color of bones. — In living and recently preserved specimens of
many centrolenids, the bones are green. The color is seen most
easily on the ventral surface of the shank. We have seen living ex-
amples of all species discussed here except medemi. Color of the
bones was recorded for most specimens after field work in 1967; our
only material of two species was collected in 1967 (oceUifera and
resplendens ) , and no notations of bone color were made.

Based on color notes of living frogs, the following taxa have
green bones: audax, buckleyi, cochranae, favopunctata, grandisonae
(pale), griffithsi (pale), megacheira, midas, peristicta (pale), pipi-
lata, prosoblepon, and siren; white bones were observed in anomala,
fleischmanni, munozorum, and pelJiicido. Some specimens of green-
boned species collected in 1967 still retain green bones. Dissection
of preserved specimens of medemi, oceUifera, and resplendens, re-
vealed no green pigment in the bones. We are tempted to argue
that the bones of these species were white in life; however, we must
point out that the pale green bones of griffithsi quickly fade to white
in preservative.

Humeral spines. — On some centrolenids, a hook-like process ex-
tends anteriorly from the deltoid ridge of the humerus. This fea-
ture was used initially to distinguish Centrolenella and Cochranella
but is no longer regarded as adequate grounds for generic distinc-
tion (Coin, 1964). Humeral spines usually are present only in males,
and the spines appear to become more pronounced in larger, and
presumably, older individuals. The spine is usually absent in fe-
males (Eaton, 1958, reported small spines in female prosoblepon).
The spines are present in audax, buckleyi, grandisonae, medemi
(small), peristicta, pipilata, and prosoblepon. Normally griffithsi
lacks humeral spines, but one individual (KU 142649) has small
humeral spines. The distal end of the humerus of males is pro-
vided with expanded lateral flanges in some of our examples of
grandisonae, peristicta, and pipilata; sexual dimorphism of this sort
is well-known in some Eupsophus and some Leptodactylus (Lynch,
1971).

Snout shape. — The shape of the snout in dorsal view (Fig. 1)
varies from subacuminate (fieischmanni) to round {audax, buckleyi,
fiavopunctata, grandisonae, munozorum, oceUifera, peUucida, peri-
sticta, prosoblepon, and resplendens) or truncate (anomala, coch-
ranae, griffithsi, medemi, megacheira, midas, and siren). Truncate
snouts ( in dorsal view ) result from the protuberant nostrils lending
an angular appearance to the top of the snout. Protuberant nostrils

CENTROLEMD FROGS OF ECUADOR

Fig. 1. Heads of Centrolenella: Dorsal views. A. C. fleischmanni, KU
146606. B. C. audax, KU 143290. C. C. anomala, KU 143299. Lateral views.
D. C. respJcmlcns, KU 118053. E. C. munozorum, KU 118054. F. C. priffithsi,
KU 118040.

are also found among some of those species with round snouts ( dor-
sal view) but the nostrils do not extend as far anteriad.

In lateral view (Fig. 1), the snout profile varies from strongly
sloping (resplendens) to a weakly sloping or rounded (htickleyi), to
rounded flavopunctata, <i,mncUsonae, munozoritm, pellucida, peri-
sticta, and in some female prosohlepon), to truncate (anomala,
audax, cochranae, weakly in fleischmanni, <iriffithsi, medemi, mega-
cheira, midas, ocellifera, pipikita, most prosuhlepon, and siren). The
laterally truncate snouts again reflect the presence of protuberant
nostrils (more so than any differences in shape of the underlying
skeleton ) .

An additional feature of the snout is the angularit\' of the canthus
and loreal region. The canthus rostralis is scarcely evident in fleisch-
manni, munozorum, and pellucida, whereas it is much more distinct
in those frogs ha\'ing more vertical loreal regions ( all other species,
except grandisonae, peristicta, and pipilata).

Tympanum. — Tympana are present in all centrolenids we ex-
amined. The tympana are concealed beneath the skin in medemi,
munozorum, and pellucida, but at least partially \'isible in others
(lower one-fourth visible in buckleyi to entirely visible in some
fleischmanni, peristicta, and pipilata). The upper edge of the tym-
panum usual!)' is obscured by a supratympanic fold or may be
simply covered by a thickening of the skin in the absence of a dis-
tinct supratympanic fold. The extent of the exposed portion of the
tympanum is probabK' not a reliable feature for species recognition.

6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

The tympanic region is directed strongly dorsolaterally (as op-
posed to being nearly vertical in most frogs) in fleischmanni, muno-
zoriim, and peUucida; this feature, in combination with the weakly
marked canthus, seems to give these frogs a flatter head than that
seen in the other Ecuadorian species. These species also have a
strong dorsolateral orientation to the tympanum. The tympana are
strongly oriented laterally ( little or no dorsal or posterior inflection )
in grandisonae, oceUifera, and peristicta. The tympana are oriented
posterolaterally with little or no dorsal inflection in anomala, buck-
leiji, midas, pipilata, and siren. The tympana are oriented dorso-
laterally, but much less so than in fleischmanni and its allies, in
audax, cochranoe, griffithsi, megacheira, prosohlepon, and resplen-
dens; in these frogs there may be little to moderate posterior in-
flection of the tympana.

Skin texture. — The skin of the venter is coarsely areolate in all
species examined. The skin of the dorsal surfaces varies in texture
and is useful in species recognition. The dorsal skin of medemi is
smooth, unlike any other Ecuadorian centrolenid. Most species have
finely shagreened skin on the dorsum without warts or spinules
(audax, flavopunctata, fieischmanni, grandisonae, griffithsi, midas,
munozorum, oceUifera, peristicta, peUucida, prosohlepon, and siren).
Shagreened skin with scattered enameled^ warts is found in pipilata
and resplendens; resplcndens has many more warts (Plate 2C).
Centrolenella cochranae has scattered warts on a shagreened dor-
sum; C. anomala has a shagreened dorsum with more numerous
warts, and C. megacheira has pustular skin on the dorsum. The skin
on the dorsum of buckleyi is shagreened with spinules, which are
most numerous laterally.

Dernud ornamentation. — Few species of the family have what
Taylor and Cochran ( 1953 ) termed "decoration" or tarsal and ulnar
folds. We find that many of the folds and tubercles are enameled
and thus appear to be distinctive. Centrolenella grandisonae has
low tubercles along the outer edge of the forearm and tarsus; the
tubercles are present distally along the lateral edge of the hand and
foot to the digital pads. A similar arrangement of more pronounced
tubercles is found in C. peristicta (Fig. 3) and C. pipilata. Centro-
lenella resplendens has a scalloped fringe along the outer edge of
the hand and arm (Fig. 3), and foot and tarsus (Fig. 4); the fringe
continues around the heel. A thin fringe, without warts or scallop-
ing, occurs along the hand and forearm and the foot and tarsus of
C. peUucida.

' The term "enameled" is used throughout to denote the shiny white ele-
vations found in the skin of some centrolenids.

CENTROLENID FROGS OF ECUADOR

3S-

^Jr..-'.

Fig. 2. Anal ornamentation in Ccntrolenella. A. Ventral view of posterior
ventrum of C. siren, KU 146621. B. Posterior view of anal region of C pellu-
cida, KU 143298. C. Dorsal view of C. resplendens, KU 118053.

Most of the Ecuadorian frogs have two to four large, round
subanal tubercles (Fig. 2). Five species have more extensive anal
ornamentation. Six Ecuadorian species do not have subanal tuber-
cles (anomaJa, feiscJimanni. medemi, mtinozoruin, ocellifera, and
pelhicida).

The small tubercles lateral and posteroventral to the vent in
grandisonae, peristicta, and pipilata are enameled and seem to rep-
resent a rudimentary character-state for the "anal decoration" char-
acter-state reported by Taylor and Cochran ( 1953 ) ; none of these
species has enlarged para- or postanal warts or folds. CentroIeneUa
pelhicida has a transverse fold below the vent (Fig. 2); the fold is
enameled. The "anal decoration" is most extensive and unique in
C. resplendens (Fig. 2). A similar feature appears to be present in
Taylor and Cochran's (1953) surda of southeastern Brasil. In re-
splendens, a pair of thick folds extends laterally from the vent onto
the posterior surface of the thighs; the folds are connected at their
lateral extent by a semicircular fold extending medio-ventrally be-
neath the anus. These folds and the area between them are orna-
mented with enameled warts and short ridges.

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 3. Palmar views of hands of Centrolenella. A. C. griffithsi, KU 118040.
B. C. grandisonae, KU 118047. C. C. flavopunctata, KU 121050. D. C. muno-
zorum, KU 118054. E. C. resplendem, KU 118053. F. C. pcristicta, KU
118051.

Hands and feet. — The first finger is as long as or longer than the
second in all Ecuadorian centrolenids. In two Peruvian species we
have examined, the first finger is shorter than the second. The pal-
mar and subarticular tubercles are difficult to see on most preserved
examples and are not accorded significance here. The fingers bear
discs that are wider than long but vary between being more nearly
round or truncate. The species with more rounded finger discs are
])uckle\i'h fleischnianni, <i,riffithsi munozorum, pelliicida, pipilata,
resplendens; me^acheira is intermediate between rounded and trun-
cate discs include anomala, audax, cochranae, flavopunctata, gran-

CENTROLENID FROGS OF ECUADOR 9

disonae, medemi, midas, ocelJifera, peristicta, prosoblepon, and
siren. The finger discs are larger than those of the toes in all species
examined, but in several species (anomaki, aiidax, fieischmanni, me-
demi, megacheira, munozonim, peUiicida, pipilata, prosoblepon,
and resplendens) the discs are more nearly equal in size than in
cochranae, flavopunctata, grandisonae, griffithsi, midas, ocellifera,
peristicta, and siren. The fingers bear narrow lateral fringes in ano-
mala, audax, buckleiji, fieischmanni, grandisoyiae, megacheira, muno-
zonim, peUucida, peristicta, pipilata, prosoblepon, and resplendens
but not in cochranae, flavopunctata, griffithsi, medemi, midas, ocelli-
fera, or siren.

The extent of webbing of the hands and feet has been used
commonly in order to distinguish species of centrolenids. In record-
ing the extent of webbing we have followed Savage and Heyer
(1967). The least finger webbing is seen in anomala, cochranae,
griffithsi, megacheira, and siren (Fig. 3); buckleyi grandisonae, and
pipilata have only slighth' greater webbing of the fingers (Fig. 3).
In these eight taxa the webbing does not enclose the distal sub-
articular tubercle. The most extensive webbing seen in Ecuadorian
Centrolenella was found in the holot\pe of petersi and an example
refen-ed to fieischmanni ( KU 121052). In these specimens the web
reaches the disc of the fourth finger and nearly to the disc of the
third (IIIIJ^-OIV); the modal webbing for fieischmanni is III1M-2IV.
Tlie webbing completely encloses the distal subarticular tubercle of
fingers III and I\' in flavopunctata, medemi, peristicta, and resplen-
dens (Fig. 3) and is only slightly less extensive in fieischmanni,
midas, ocellifera, and prosoblepon (Fig. 3). The distal subarticular
tubercle of IV, but not III, is free of webbing in audax and pellucida.

Most species studied have a small outer metatarsal tubercle;
none was found in audax, flavomaculata, fieischmanni, medemi,
midas, munozorum, ocellifera, pellucida, resplendens, or siren.

The extent of webbing on the feet parallels that of the fingers,
but species group less well. The least webbing is seen in cochranae,
griffithsi, and siren, and with onh' slightly more webbing in anomala,
buckleyi, megacheira, and pipilata (Fig. 4). The webbing formulae
for these seven species range from 12-2,^nilM-3TII2--3IV3-2V of
cochranae and siren to the comparatively well-webbed pipilata
(Il-2^IIl-2TIIl-2MIV2)3-lV).Nine species have more webbing than
pipilata but less than the extensive webbing of medemi and re-
splendens (audax, flavopunctata, fieischmanni, grandisonae, midas,
ocellifera, pellucida, peristicta, and prosoblepon). Toe webbing is
most extensive in medemi where the web reaches the discs on all
but the fourth toe.

10

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 4. Plantar views of feet of Centrolenella. A. C. griffithsi, KU 118040.
B. C. grandisonae, KU 118047. C. C. resplendens, KU 1180653. D. C. jiavo-
punctata, KU 121048.

Peritoneal coloration. — The parietal peritoneum is white (heart
not visible in life) in all Ecuadorian Centrolenella except feisch-
tnanni, rnunozorum, and pellucida; in these three species the heart
is visible through the skin of the chest. The visceral peritoneum is
white in fleischmanni, munozorum, pellucida, and resplendens. The
latter is the only Ecuadorian species with both white parietal and
visceral peritonea.

In living frogs, if the heart is visible, the parietal peritoneum is
clear; if the heart is not visible, the parietal peritoneum is opaque

CENTROLENID FROGS OF ECUADOR 11

and white. The character-states of the visceral peritoneum are more
difficult to observe in living frogs. If the intestine is dark (as seen
through the skin of the venter), the visceral peritoneum is clear; if
the intestine is yellow or white, the visceral peritoneum is opaque
and white.

In preserved examples, inspection of peritoneal pigmentation is
best made by cutting into the abdomen and recording whether the
intestine is white or not (visceral peritoneum) and whether the in-
side of the belly beneath the liver is white or not (parietal peri-
toneum ) .

Three peritoneal pigmentation patterns are presently known: 1)
clear parietal, white visceral {jieischmanni and pulverata groups and
at least C. oJbotunica, etinj^natha, and vanzoUnii of southeastern
Brasil); 2) white parietal, clear visceral { pwsoblepon group); and
3) white parietal, white visceral (antioquiensis and resplendem) .

Ground color in preservative. — The ground color of preserved
frogs is cream to creamy-white in jieischmanni (including the holo-
type of petersi), munozorum, and pellucida. The ground color of
anonmla is pale brown. The ground color of flavopunctata, medemi,
and pipilata is gray to slate-gray. That of <i,randisonae, griffithsi,
ocellifera, peristicta, and resplendens is pale to dull lavender and
contrasts with the darker lavender ground color of audax, hiickleyi,
cochranac, megacheira, midas, prosohlcpon, and siren.

Color patterns. — The most simple color patterns are those of
audax, flavopunctata, niidas, and siren (white flecks on ground
color), griffithsi (lavender to black flecks on ground color), buck-
leiji (moderately distinct white labial stripe continuing onto flanks
separating lavender or purple dorsum from cream venter), and
jieischmanni and munozorum (faint peppered reticulation of dorsal
surfaces). No color pattern is seen in pellucida.

The color pattern of resplendens consists of many smaU white
spots (enameled warts) on a dull lavender ground color. Centro-
lenella medemi has a simple pattern of large cream spots on the
dorsal surfaces, unlike the pattern of any other centrolenid.

The other centrolenids with non-ocellated color patterns include
grandisonae, peristicta, and pipilata with small, diffuse black (or
lavender) spots and equally diffuse white spots on the dorsal sur-
faces and megacheira and prosohlepon with discrete dark lavender
spots on the dorsum and limbs (spotting reduced or absent in
prosoblepon) .

The other three species known from Ecuador (anomala, coch-
ranac, and ocellifera) and the Peruvian ocellata have a dorsal pat-
tern of ocelli. The ocelli are large in ocellata but small in the three

12

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 5. Dorsal patterns of ocellated Centrolenella. A. C. anomala, KU
143299. B. C. cochranae, KU 121035. C. C. oceUata, LSU 25990. D. C.
oceUifera, KU 118046.

Ecuadorian ocellated Centrolenella (Fig. 5). The ocelli of anomala
are black with white centers; those of cochranae are deep lavender
with white centers, and those of oceUifera faint lavender without
white centers. The pattern of oceUifera also includes small non-
ocellated white spots (Fig. 5).

KEY TO ECUADORIAN CENTROLENID FROGS

1. Discs on fingers as large as eye Centrolene geckoideum

Discs on fingers less than half size of eye 2

2. Dorsum tan with black ocelli enclosing cream (orange in life)

spots CentroJeneUa anomala

Dorsum white, cream, gray, or lavender ( green in life ) witii or
without spots or ocelli 3

3. Heart visible in life; bones white; dorsum cream or white in

preservative ocelli absent 4

Heart not visible in life, bones green (unknown in medemi,
oceUifera, and resplendens); dorsum lavender or slate gray in
preservative; ocelli and spots present or absent 6

4. Tympanum visible CentroleneUa fleischtnanni

Tympanum concealed 5

5. Forearm and tarsus each bearing a dermal fold on ventrolateral

edge; transverse dermal fold below anus ..

CentroleneUa peUucida

CENTROLENID FROGS OF ECUADOR 13

Forearm and tarsus lacking dermal folds; no transverse dermal
fold below anus Centrolenella munozorum

6. Webbing extending no more than midway between basal and

distal subarticular tubercles on fourth finger 7

Webbing extending at least to base of distal subarticular tuber-
cle on fourth finger 10

7. Dorsal skin pustular Centrolenella megacheira

Dorsal skin shagreened 8

8. Dorsum with distinct small black ocelli enclosing pale spots
Centrolenella cochranae

Dorsum without ocelli 9

9. Dorsum with white (yellow in life) flecks ___ Centrolenella siren

Dorsum with or without diff^use black flecks

Centrolenella griffithsi

10. Broad, scalloped dermal fringe on ventrolateral edges of fore-
arm and foot; U-shaped anal fold; snout sloping anteroventrally;
dorsum with white warts Centrolenella resplendens

Not nearly so fancy 11

11. Dorsum of body and limbs slate gray with large cream spots;

webbing extending to discs on all but fourth toe

Centrolenella medemi

Dorsum of body and limbs unicolor, ocellated, or flecked; web-
bing less extensive on feet — - 12

12. Dorsum with dark ocelli Centrolenella ocellifera

Dorsum without ocelli 13

13. Webbing extending beyond distal subarticular tubercle of

fourth finger 14

Webbing not extending beyond distal subarticular tubercle of
fourth finger 16

14. Dorsum unicolor or with distinct dark flecks; no pale flecks
Centrolenella prosoblepon

Dorsum with pale flecks; diffuse dark flecks present or not .. 15

15. Dorsum of body slate gray (dark green in life) with white

(yellow in life) flecks Centrolenella flavopunctata

Dorsum of body pale lavender (pale green in life) with dark
lavender spots and few white flecks Centrolenella peristicta

16. Dorsum with pale flecks, no dark flecks; prevomerine teeth
present 17

14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Dorsum unicolor with dark and pale flecks; prevomerine teeth
absent 18

17. Humeral spine present in males; pale flecks numerous on dor-
sum CentroleneUa audax

Humeral spine absent in males; pale flecks few, principally
laterally CentroleneUa midas

18. Dorsum unicolor CentroleneUa buckleyi

Dorsum with dark and pale flecks 19

19. Snout truncate in dorsal and lateral profiles

CentroleneUa pipUata

Snout round in dorsal and lateral profiles.

CentroleneUa urandisonae

to"

ACCOUNTS OF SPECIES

In the following accounts we attempt to make comparable state-
ments in the diagnoses and descriptions. In order to facilitate com-
parisons of primary character states among species, we have num-
bered what we consider to be the primaiy character in the diagnoses.
The webbing formula in the diagnosis of each species is the modal
formula; variation is given in the description. Measurements and
proportions are for Ecuadorian specimens only, unless othenvise
noted.

Specimens designated as paratypes and paratopotypes (in addi-
tion to holotypes) are those on which descriptions of new species
were based. In some cases measurements of referred specimens have
been incorporated into statements of ranges of variation.

Colored photographs of 15 species are reproduced on Plates 1
and 2 ( following pages 16 and 26 ) .

Taxonomic changes and their justifications, comments on type
specimens and literature, and observations on behavior and ecology
are given in the "Remarks" section of each account. All specimens
examined are listed in a terminal section of the paper.

CentroleneUa anomala new species
Plate 2H

Holotype.—KU 143299, an adult male, 24.1 mm, from the Rio
Azuela, 1740 m, Quito-Lago Agrio road, Provincia Napo, Ecuador,
obtained on 23 October 1971, by William E. Duellman.

D'uifinosis. — 1) prevomerine teeth absent; 2) bones white; 3)
parietal peritoneum white; visceral peritoneum clear; 4) color in

CENTROLENID FROGS OF ECUADOR 15

life tan with black ocelli with orange-tan centers; in preservative,
brown with black ocelli with white spots; 5) webbing between
outer fingers III3^-2MV; 6) webbing on foot 12--2}ai 1-2)^11 1-2^1 V2?^-
IMV; 7) snout truncate in dorsal and lateral profiles; 8) dorsal skin
shagreened with minute spicules and elevated warts corresponding
to ocelli; 9) arms and legs lacking dermal fringes; 10) humeral spine
absent in male; 11) lower two-thirds of tympanum visible, directed
posterolaterally with dorsal inclination.

Centrolenella anomaJa difi^ers from all other centrolenids by be-
ing tan instead of green; it further difi^ers from other species having
ocellated patterns (cochranae, ocellata, and ocellifera) by having
more ocelli and scattered black flecks between ocelli. Moreover,
anomala differs from ocellifera by having less webbing between the
outer fingers, from cochranae by lacking prevomerine teeth, and
from ocellata by ha\'ing the snout truncate and lower two-thirds of
the tympanum visible, whereas in ocellata the snout is round and
the tympanum is concealed.

Description. — Adult male moderate-sized, 24.1 mm in snout- vent
length; females not known. Head slightly wider than body; width
of head 33.2 percent of snout- vent length; snout short, truncate in
dorsal and lateral profiles (Fig. 1); canthus round; loreal region
concave; lips not flared; nostrils nearly terminal on snout, protuber-
ant, directed dorsolaterally; internarial area depressed. Eye large,
protuberant, directed anterolaterally. Supratympanic fold very
weak, barely covering upper edge of t>'mpanum; tympanum directed
posterolaterally with dorsal inclination. Prevomerine dentigerous
processes and teeth absent; choanae small, ovoid, near margin of
mouth; tongue not notched posteriorly, barely free behind; vocal
slits extending from lateral base of tongue to angles of jaws.

Humeral spine absent; ulnar fold and tubercles absent; first
finger longer than second; fourth finger nearly as long as third; lat-
eral fringes present on fingers; webbing absent between first and
second fingers, vestigial between second and third; webbing formula
for outer fingers III3+-2/-3IV; discs broad, truncate; subarticular tu-
bercles large, round, simple; supernumerary tubercles absent; pal-
mar single, ovoid; nuptial excrescences absent. Hind limbs slender;
length of tibia 54.4 percent of snout-vent length, tarsal folds and
tubercles absent; inner metatarsal tubercle large, elongate; outer
metatarsal tubercle small, round; subarticular tubercles large, round;
supernumerary tubercles absent; feet about two-thirds webbed;
webbing formula 12 2}nil-2)nil 1-2^1 V2?3-1W; discs on toes slightly
smaller and more nearly round than those on fingers.

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Skin on dorsal surfaces of head, body, forearms, and shanks with
many minute spicules and larger spicules corresponding to ocelli;
skin of belly and ventral surfaces of thighs granular; other surfaces
smooth; anal opening directed posteroventrally at midlevel of
thighs; anal folds and tubercles absent.

Color in preservative: dorsal surfaces of head, body, forearms,
fourth fingers, thighs, shanks, feet, and fourth and fifth toes brown
with dark brown flecks and head, body, forearms, and shanks with
black ocelli enclosing white spots (Fig. 5); other surfaces creamy
tan.

Color in life: dorsum tan with small black ocelli enclosing
orange-tan spots; chest white; heart not visible; throat, visceral peri-
toneum, and ventral surfaces of limbs lacking pigment; bones white;
iris bronze with black reticulations.

Distribution. — This distinctive species is known only from the
type locality, a rivulet flowing into the Rio Azuela on the east slope
of \^olcan Reventador at an elevation of 1740 m (Fig. 6). The small
stream is about 50 m north of the bridge over the Rio Azuela on the
Quito-Lago Agrio road. In this area there is a narrow relatively
gentle slope between the steep slopes of Volcan Reventador to the
west and the chasm of the Rio Coca to the east.

Remarks. — The single specimen was kept alive for two days and
two nights, during which time it was observed frequently. At no
time did it take on any greenish color; it remained tan with no
noticeable change in pigmentation. After 18 months in alcohol the
dorsum has a faint lavender tint.

The holotype was on a mossy limb of a bush about 1.5 m above
a cascading rivulet at night. Hylo pJiyUo^natha and foiu- other spe-
cies of Centrolenella (megacheira, pellucida, pipilata, and siren)
were found in the same stream and in other small streams nearby in
the cloud forest, but no other individuals of C. anomala were
observed.

Etymology. — The specific name is from the Greek anomalos
meaning unusual, deviating from the general rule; the name is used
in allusion to the distinctive tan color of this species.

Centrolenella audax new species
Holotype.— KV 146624, an adult male, 23.0 mm, from Salto de

Plate 1. A. Centrolenella fleisehnianni, KU 146607; B. C. numozonim, KU
123225; C. C. pellucida, KU 143298 ( holotvpe); D. C. midas, KU 123219
(holotype); E. C. siren, KU 143293; F. C. fiavopunctata, KU 123224; G. C.
cochranae, KU 123216; H. C. anomala, KU 143299 (holotype). All X 2.

PLATE 1

CENTROLENID FROGS OF ECUADOR

17

5 -

600 M.

I

CONTOUR

f 1

L . 1

OVER

3000 M

0

he

K

OVER
40

5000 M
100

ILOMETERS

76

Fig. 6. Distribution of CcntrolencIIa anomala (square), C. cochranae
(circles), C. occllifcra (triangles), C. resplendens (hexagons).

Agua, 2.5 km NNE of Rio Reventador on Quito-Lago Agrio road,
1660 m, Provincia Napo, Ecuador, obtained on 7 April 1972, by
William E. Duellman.

Paratypes.—KV 143290 and 143292, adult males, from 16.5 km
NNE of Santa Rosa on Quito-Lago Agrio road, 1700 m, Provincia
Napo, Ecuador, obtained on 18 October 1971, by Joseph T. Collins
and William E. Duellman.

Diag,nosis. — 1) prevomerine teeth 2-4; 2) bones green; 3) pa-
rietal peritoneum white; visceral peritoneum clear; 4) color in life
pale green with small yellow flecks; in preservative, lavender with
white flecks; 5) webbing between outer fingers III2"-2I\'; 6) web-
bing on foot 11-2 II1-2 III1-2^IV2^-1V; 7) snout round in dorsal
view, truncate in lateral profile; 8) dorsal skin shagreened; 9) arms

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

and legs lacking dermal folds; 10) humeral spine present in males;
11) lower four-fifths of tympanum visible, directed dorsolaterally
with slight posterior inclination.

The coloration of audax is like that of flavopunctata, midas, and
siren, but the males of these species lack humeral spines. No other
species having humeral spines has a green dorsum with yellow flecks.

Description. — Adult males moderate-sized, 23.0-23-6 mm
(x=23.3, N=3) in snout- vent length; females not known. Head
much wider than body, width of head 34.8-43.5 percent (x=37.8,
N=3) of snout-vent length; snout short, round in dorsal view,
truncate in lateral profile (Fig. 1); canthus round; loreal region
barely concave; lips not flared; nostrils four-fifths distance from eyes
to tip of snout, slightly protuberant dorsolaterally; internarial area
depressed. Eye moderately large, directed anterolaterally. Supra-
tympanic fold absent; lower four-fifths of tympanum visible, di-
rected dorsolaterally witli slight posterior inclination. Prevomerine
dentigerous processes posteromedially inclined, narrowly separated
medially, between longitudinally elliptical choanae, bearing 2-4
teeth; tongue nearly ronnd, barely free behind; vocal slits extending
from midlateral base of tongue to angles of jaws.

Humeral spine large, parallel to humerus; ulnar f(jld and tuber-
cles absent; first finger longer than second; third finger slightly
shorter than fourth; lateral fringes present on fingers; webbing ab-
sent between first and second fingers, vestigial between second and
third; webbing formula for outer fingers III(2^-2/2)-(2--2'^)IV; discs
broad, truncate; subarticular tubercles small, round, simple; super-
numerary tubercles absent; palmar tubercles rectangular, simple;
nuptial excrescences absent. Hind limbs moderately slender; length
of tibia 51.3-55.6 percent (x^54.0, N=3) of snout-vent length; tar-
sal folds and tubercles absent; inner metatarsal tubercle small, ellip-
tical, outer metatarsal tubercle absent; subarticular tubercles small,
round; supernumerary tubercles absent; feet about two-thirds
webbed; webbing formula Il-(2--2 )IIl-(2^-2M)IIIl-2 IV2 -IV; discs
on toes slightly smaller than those on fingers.

Skin on dorsal surfaces shagreened; skin on belly and proximal
ventral surfaces of thighs granular; other surfaces smooth; anal open-
ing directed posteroventrally at upper level of thighs; pair of large
tubercles ventral to anus.

Color in preservative: dorsal surfaces of head, body, forearms,
thighs, and shanks lavender with small white dots; hands, feet, and
ventral surfaces cream.

Color in life: dorsum green with gold flecks; fingers and toes
pale yellow; chest white; heart not visible; visceral peritoneum and

CENTROLENID FROGS OF ECUADOR

19

Fig. 7. Distribution of Centrolendla audax (triangles), C. grandisonae
(square), C. medcmi (hexagon), and C. prosoblepon (circles).

ventral surfaces of limbs iinpigmcnted; bones green; iris pale bronze
with black reticulations.

Distribution. — This species presently is known from only two
localities at elevations of 1660 and 1700 m on the Amazonian slopes
of the Cordillera Oriental of the Andes (Fig. 7). The type of locality
is a deeply recessed waterfall in a nearly vertical cliff at a point 2.5
km north-northeast of the bridge over the Rio Reventador on the
Quito-Lago Agrio road.

Remarks. — The holotype was found in a bromeliad on a cliff
below and in the spray zone of the waterfall. In another bromeliad
a metamoiphosing young having a snout-vent length of 13.5 mm and
a tail 19.5 mm in length was found. Two individuals were found at
night on vegetation over a stream 16.5 km north-northeast of Santa

20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Rosa; three other species of Centrolenella were found there — mega-
cheira, pipilata, and siren.

Etymology. — The specific name is Latin, meaning daring, and is
used in allusion to the precipitous regions inhabited by the species.

Centrolenella buckleyi (Boulenger)
Plate 2E

Hylella buckleyi Boulenger, 1882:420 [Syntypes.— BMNH 78.1.25.16 from In-
tac, Provincia Imbabura Ecuador; BMNH 80.12.5.201 from "Paitanga"
( =Pallatanga), Provincia Chimborazo, Ecuador].

Hyla purpurea Nieden, 1923:267 [Substitute name for Hyllela buckleyi Bou-
lenger].

Cochranella buckleyi — Taylor, 1951:35.

Centrolenella buckleyi — Coin, 1964:6.

Diagnosis. — 1) prevomerine teeth absent; 2) bones green; 3) pa-
rietal peritoneum white; visceral peritoneum clear; 4) color in life
dark green; in preservative, purple; 5) webbing between outer fin-
gers III2]^-2^IV; 6) webbing on foot 1 1J^-2II 1-2^111 B^-2?nV3^-lW;
7) snout round in dorsal view, in profile round above and slightly
sloping anteriorly; 8) dorsal skin shagreened with minute spinules;
9) arms and legs lacking dermal folds; 10) humeral spine present in
males; 11) lower one-fourth to one-half of tympanum visible, di-
rected posterolaterally with slight dorsal inclination.

Centrolenella buckleyi is like many specimens of griffithsi and
some prosoblepon in being uniform lavender above in preservative,
but it differs from both of these by having an inclined snout in lat-
eral profile, spinules in the dorsal skin, and only the lower one-fourth
to one-half of the tympanum visible. It further differs from griffithsi
by having more webbing on the hand and humeral spines in males,
and from prosoblepon by having less webbing on the hand and lack-
ing prevomerine teeth.

Description. — Adults large, snout-vent length 28.4-29.5 mm
(x=29.0, N=2) in males, 29.8-34.4 mm (x=31.7, N=5) in females.
Head not as wide as body; width of head 32.5-38.6 percent (x=34.7,
N=7) of snout- vent length; snout short, round in dorsal view, round
above and slightly sloping anteriorly in profile; canthus round; loreal
region concave; lips flared; nostrils two-thirds distance from eyes to
tip of snout, barely protuberant laterally; internarial area flat. Eyes
moderately large, directed anterolaterally. Supratympanic fold mod-
erately heavy; lower one-fourth to one-half of tympanum visible,
directed posterolaterally with slight dorsal inclination. Prevomerine
dentigerous processes and teeth absent; choanae small, ovoid;
tongue cordiform, shallowly notched behind; free posteriorly for
about one-fourth of its length; vocal slits extending posterolaterally
from midlateral base of tongue to angles of jaws.

CENTROLENID FROGS OF ECUADOR 21

Humeral spine short, pointed, nearly parallel to humerus; ulnar
fold and tubercles absent; first finger equal in length to second;
fourth finger slightly shorter than third; lateral fringes present on
fingers; webbing absent between first, second, and third fingers;
webbing formula for outer fingers III(2"-2)4)-(2/4-2;3)IV; discs broadly
rounded; subarticular tubercles moderately large, conical, simple;
distal tubercle on fourth finger weakly bifid; supernumerary tuber-
cles small, numerous on basal segments; palmar tubercle large,
ovoid; nuptial excrescences absent. Hind limbs moderately robust;
length of tibia 49.4-56.4 percent (x=51.2, N=7) of snout- vent length;
tarsal folds and tubercles absent; inner metatarsal tubercle large,
ovoid; outer metatarsal tubercle small, round; subarticular small,
round; supernumerary tubercles small, present basally; toes about
three-fourths webbed; webbing formula I( l/3-l73)-(2-2 )n( l-^)-
2"^I(l-l73)-(2+-3')IV(2•:^-3 )-(lM-2 )V; discs on toes slightly smaller
than those on fingers.

Skin on dorsum shagreened with minute spinules; skin on belly
and proximal posteroventral surfaces of thighs granular; other sur-
faces smooth; anal opening directed posteroventrally at midlevel of
thighs; several small and four large tubercles below anus.

Color in preservative: dorsal surfaces, except fingers and toes,
purple; other surfaces cream; narrow cream stripe on edge of upper
lip in some specimens.

Color in life: dorsal surfaces bright to dark green, sharply de-
marcated laterally from white lower flanks; chin and most of venter
pale green; parietal peritoneum yellow; heart not visible; edge of
upper lip, outer edge of tarsus, and anal stripe white; bones green;
iris pale copper flecked with black.

Distribution. — Centrolenella huckleiji occurs from the Merida
Andes in western Venezuela southward through the Andes and in-
terandean valleys to southern Ecuador (Fig. 8). Although the spe-
cies has been recorded at elevations as low as 1500 m, authenticated
records in Colombia and Ecuador are all above 2000 m; we have
found the species at elevations up to 3000 m.

Remarks. — We have found individuals in terrestrial bromeliads
and in vegetation in cienegas, the quaking meadows in many high
Andean localities. Francisco Leon of the Universidad Catolica in
Quito collected two specimens from bromeliads in a Eucalyptus
grove near Machachi. Thomas H. Fritts obtained one from a bro-
meliad on an island in Laguna Cuicocha. Centrolenella buckleyi
may breed in the same situations as other centrolenids — rapid,
mountain streams — but buckleyi frequents non-stream situations
(Laguna Cuicocha, the bog-like cienegas) where trees, if present,

22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

600 M. CONTOUR;

r~l OVER 3000 M.

.'~ol OVER 5000 M
0^0 100

KILOMETERS I

76

Fig. 8. Distribution of Centrolenella huckleiji (circles), C. peristicta
(square), and C. pipilata (triangles); open symbols are literature records.

are small and rarely overhang water. Because of the absence of
streams, we suspect that the tadpoles may develop in bromeliads or
in cienegas, a radical departture from the adaptive zone of centro-
lenid frogs.

As noted by Goin (1961:101), one of the syntypes (BMNH
80.12.5.201) is now represented by disarticulated bones and the
other (BMNH 78.1.25.16) is missing.

Rivero (1968) named Centrolenella hiickleyi venezuelensis from
the Merida Andes of western Venezuela. We have not been con-
cerned with northern populations which may or may not be conspe-
cific with biickleyi; thus, we use the binomial for the populations in
Ecuador.

CENTROLENID FROGS OF ECUADOR 23

Centrolenella cochranae (Goin)

Plate IG

Cochranelle cochranae Goin, 1961:97 [Holotype.— BMNH 1912.11.1.68 from

El Topo, Rio Pastaza, Provincia Tungurahua, Ecuador].
Centrolenella cochranae Goin, 1964:6.

Diagnosis. — 1) prevomerine teeth 0-3; 2) bones green; 3) pari-
etal peritoneum white; visceral peritoneum clear; 4) color in life
pale green with minute puiple ocelli with red centers; in preserva-
tive, lavender with purple ocelli with white centers; 5) webbing be-
tween fingers III3-2?1IV; 6) webbing on foot I2-2MIIlM-2Min2-3IV3+-
1%V; 7) snout truncate in dorsal and lateral profiles; 8) dorsal skin
shagreened with elevated warts corresponding to ocelli; 9) arms and
legs lacking dermal folds; 10) humeral spine absent in males; 11)
lower three-fourths of tympanum visible, directed dorsolaterally.

The first impression of tlie dorsal pattern of cochranae is of black
flecks on a green ground color; however, upon closer inspection it is
evident that the markings actually are small ocelli (Fig. 5). Centro-
lenella cochranae differs from other species having ocellated dorsal
patterns (anomaJa, ocellafa, and ocellifera) by having smaller ocelli
and prevomerine teeth (absent in some cochranae). Furthermore,
the species differs from anomala by having a green instead of brown
dorsum, from ocellifera by haxing less webbing between the outer
fingers, and from oceUata by liaving the snout trimcate and the lower
three-fourths of the tympanimi \isible, whereas in oceUata the snout
is round and the tympanum concealed.

Description. — Adults moderately large; snout-vent length 23.8-
26.7 mm (x=25.0, N^6) in males, 30.0 mm in single female. Head
noticeably wider than body; width of head 33.0-35.3 percent (x=34.0,
N=7) of snout-vent length; snout short, truncate in dorsal and lateral
profiles; canthus round, loreal region concave; lips moderately flared;
nostrils nearly terminal, directed anterolaterally; internarial area
depressed. Eye large, protuberant, directed anterolaterally. Supra-
tympanic fold absent; lower three-fourths of tympanum visible, di-
rected dorsolaterally. Prevomerine dentigerous processes absent in
one specimen, present in six; processes transverse between choanae,
bearing 1-3 teeth; choanae small, ovoid, near edge of mouth; tongue
cordiform, free posteriorly; vocal slits extending from midlateral
base of tongue to angles of jaws.

Humeral spine absent; ulnar fold and tubercles absent; first fin-
ger longer than second; fourth finger slightly shorter than third;
lateral fringes absent on fingers; webbing absent between first and
second fingers, vestigial between second and third; webbing formula
for outer fingers 111(3 -3+)-(2?i-3)IV; discs broader than deep.

24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

rounded; subarticular tubercles small, round, simple; supernumerary
tubercles absent; palmar tubercle large, elliptical; nuptial excres-
cences absent. Hind limbs slender; length of tibia 55.7-61.8 percent
(x=58.7, N-7) of snout- vent length; tarsal folds and tubercles absent;
inner metatarsal tubercle large, flat, elongate; outer metatarsal tu-
bercle small, conical; subarticular tubercles small, round; super-
numerary tubercles absent; feet about two-thirds webbed; webbing
formula 12- ( 2^-2)^ ) II ( ]Ji- !%)-{ 2--23i ) 1112"- ( 3--3- ) IV ( 3--3^ ) - ( 2-2" ) V;
discs on toes smaller and more nearly round than those on fingers.

Skin on dorsal surfaces smooth with scattered spicules; skin on
belly and ventral surfaces of thighs granular; other surfaces smooth;
anal opening directed posteroventrally at midlevel of thighs; pair of
large tubercles below anus.

Color in preservative: dorsal surfaces of head and body lavender
with small black ocelli enclosing minute white dots or some solid
black flecks; other dorsal surfaces creamy tan with black flecks on
thighs and shanks of some specimens.

Color in life: pale green with black or purple ocelli enclosing
pink or red dots; chest white; heart not visible; visceral peritoneum
and ventral and concealed surfaces of limbs unpigmented; bones
green; iris pale gray with black flecks.

Distribution. — Most localities from which CentroJenella coch-
ranae is known are in the lower valley (1100-1300 m) of the Rio
Pastaza on the eastern face of the Andes (Fig. 6); additionally the
species is known from an elevation of 1150 m in the Cordillera del
Due above the Rio Coca ( ± 180 km NNE of the Rio Pastaza val-
ley). On the basis of these few localities it seems that cochranae,
along with favoptnictata, inhabits intermediate elevations between
the ranges of the lowland species (medemi, midas, miinozorum, and
resplendens) and the numerous species in the higher cloud forests
(anomala, aiidax, megacheira, peUiicida, pipilata, and siren).

Remarks. — Coin (1961:97) described cochranae as lacking pre-
vomerine teeth, but he (1964:2) noted the presence of teeth in six
other specimens. Six of our seven specimens have prevomerine teeth,
and one lacks not only teeth but dentigerous processes.

We compared two of our specimens (KU 121033, 123217) with
the holotypes of cochranae (BMNH 1912.11.1.68) and C. ocellata
(Boulenger, 1918) (BMNH 1912.11.1.19) from Huancabamba, De-
partamento Pasco, Peru. Our specimens compare favorably with the
holotype of cochranae but are different from ocellata in several
characters. Centrolenella ocellata dift'ers from C. cochranae as fol-
lows: 1) prevomerine teeth absent; 2) ocelli much larger (Fig. 5);
3) snout truncate; 4) first finger shorter than second.

CENTROLENID FROGS OF ECUADOR 25

All of our specimens were obtained at night from vegetation
overhanging small mountain streams. The call consists of a single,
high-pitched note. Other than the calls, no evidence of reproduction
was noted while we were collecting in the range of Cochrane (April,
June, July, August). Individuals seemed to be sparsely distributed
in the habitat.

Centrolenella flavopunctata new species
Plate IF

HoJotype. — KU 121048, an adult male, 21.6 mm, from Mera,
Provincia Pastaza, Ecuador, obtained on 14 July 196(S, by John D.
Lynch.

Paratopofypes.—KU 121041, 28 June 1968, John D. Lynch;
121043-46, 2 July 1968, John D. Lynch and Gerald R. Smith; 121049,
14 July 1968, John D. Lynch; 121050-51, 24 July 1968, John D.
Lynch.

Diagnosis. — 1) prevomerine teeth 0-3 on low processes; 2) bones
green; 3) parietal peritoneum white; visceral peritoneum clear; 4)
color in life green with pale yellow flecks; in preservative, slate gray
with white flecks; 5) webbing between outer fingers III2 -IMIV; 6)
webbing on foot I1-1MI0-2TII1-2-IV2-1V; 7) snout round in dorsal
and lateral profiles; 8) dorsal skin shagreened; 9) arms and legs
lacking dermal folds; 10) humeral spine absent in males; 11) lower
three-fourths of tympanum visible, directed dorsolaterally with
slight posterior inclination.

CentrolcncUa flavopunctata is like inidas and siren in having a
lavender dorsum with white flecks (in life, green with gold flecks)
and no black flecks, but it differs from both species by having a
rounded, instead of truncate, snout and slightly more webbing on
the hands and feet. Centrolenella siren further differs from flavo-
punctata by having nearly the entire tympanum visible and strongly
inclined posteriorly, whereas onh' about three-fourths of the tym-
panum is visible and directed dorsolaterally in flavopunctata.

Description. — Adults rather small; snout-vent length 20.6-23.2
mm (x=21.6, N=:7) in males, 24.1-25.7 mm (x=24.9, N=3) in females.
Head slightly wider than body; width of head 34.6-38.3 percent
(x=36.2, N=10) of snout- vent length; snout short, round in dorsal and
lateral profiles; canthus round; loreal region barely concave; Hps not
flared; nostrils nearh' terminal on snout, not protuberant, directed
antero-dorsolaterally; internarial area barely depressed. Eye moder-
atel)' large, directed anterolaterally. Supratympanic fold absent;
lower three-fourths of tympanum visible, directed dorsolaterally
with slight posterior inclination. Prevomerine dentigerous processes

26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

small, low, widely separated between longitudinally rectangular
choanae, bearing 0-3 teeth; tongue broadly cordiform, barely free
posteriorly; vocal slits extending from midlateral base of tongue to
angles of jaws.

Humeral spine absent; ulnar fold and tubercles absent; first fin-
ger longer than second; fourth finger much shorter than third; fringes
absent on fingers; webbing absent between first and second fingers;
webbing formula for other fingers IIl-(2?i-3^)III(lJ^-2)-(l-l?^)IV; discs
truncate, subarticular tubercles small, round, simple; supernumerary
tubercles absent; palmar tubercle large, elliptical; nuptial excres-
cences absent (Fig. 3). Hind limbs moderately robust; length of
tibia 55.3-62.0 percent (x=58.0, N=10) of snout- vent length; tarsal
folds and tubercles absent; inner metatarsal tubercle small, flat,
elliptical; outer metatarsal tubercle absent; subarticular tubercles
small, round; supernumerary tubercles absent; feet about three-
fourths webbed; webbing formula I(0-l)-(l-l?i)n(0-l)-(0-2)ni(0-l)-
(l/4-2"^)IV(2"-2^)-(0-l)V; discs on toes smaller and more nearly round
than those on fingers ( Fig. 4 ) .

Skin on dorsal surfaces shagreened; skin on belly and proximal
posteroventral surfaces of thighs granular; other surfaces smooth;
anal opening directed posteriorly at upper level of thighs; pair of
large tubercles below anus.

Color in preservative: dorsal surfaces of head, body, and limbs
lavender or slate gray with many minute white flecks; other surfaces
dull creamy tan.

Color in life: dorsal surfaces of head, body, and limbs pale green
with numerous minute yellow flecks on body and limbs; edge of
upper lip pale yellow; fingers and toes yellow; chest white; visceral
peritoneum unpigmented; throat pale bluish green; bones green;
iris pale grayish white, with or without golden tint, with dark gray
or brown flecks or fine reticulations.

Distribution. — Most specimens are from elevations of 1000-1800
m in the Pastaza VaHey in the eastern slope of the Andes; the species
is also known from an elevation of 720 m in the Serrania de Um-
baqui, 200 km NNE of the former locality (Fig. 9). One faded
specimen with no visible yellow flecks from San Jose Abajo, Provin-
cia Napo (AMNH 22187) tentatively is referred to this species. The
locality is between 700 and 1000 m on the eastern slope of Volcan
Sumaco, about 130 km NE of the Pastaza Valley. Centrolenella

Plate 2. A. CentroJcneUa me^acheira, KU 143245 (holotype); B. C. grif-
fithsi, KU 121039; C. C. rcsplcndens, KU 118053 (holotype); D. C. pipilata,
KU 143278 (holotype); E. C. buckleyi, KU 144131; F. C. prosohlepon, KU
146609; G. C. pensticta, KU 121053. All x 2.

PLATE 2

CENTROLExNID FROGS OF ECUADOR

27

\

600 M.

CONTOUFfl

f 1

OVER

3000 M.

lo\

OVER

5000 M.

0
K

40

100

ILOMETERS

Fig. 9. Distribution of Centrolenella flavopunctata (circles), C griffithsi
(hexagons), C. megacheira (solid squares), C. niidas (triangles), and C. siren
(open squares); half closed squares are localities for both C. megacheira and
C. siren.

flavopunctata, togetlicr with C. cochranae, inhabits intermediate
elevations between tlie ranges of the lowhmd species (medemi, mi-
das, munozonnn, and respJendens) and the cloud forest inhabitants
at higher ele\ations (anomala, audax, megacheira, pellucida, pipilata,
and siren). Although cochranae and favopunctata occur at about
the same elevations on the eastern slopes of the Andes, the two
species have not been found in sympatr).

Remarks. — One was found on a bush in forest by day; most were
collected in a deep ravine. Males were perched on small herbs in
the spray-zone of a small waterfall. No eggs were obser\'ed although
one gravid female was found. At several small streams between
Mera and the Rio Alpayacu, males were calling on 2, 14, and 24 July

28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

196(S. Recently metamorphosed young having snout-vent lengths of
14.5 and 16.0 mm were found on 28 June and 6 July 1968.

Etymology. — The specific name is a combination of the Latin
flaviis, meaning golden yellow, and the Latin punctattis, meaning
dotted, and is used in reference to the dorsal coloration.

Centrolenella fleischmanni (Boettger)
Plate lA

Hijlella fleischynanni Boettger, 1893:251 [Holotype.— SMF .3760 from San Jose,

Provincia San Jose, Costa Rica].
Hylella cappcUei Lidth de Jeude, 1904:94 [Holotype.— RMNH 4463 from

Saramacca, Surinam; synonymy ^de Coin, 1964:1].
Centrolenella fleischmanni — Noble, 1924:67.
Centrolenella cappellei — Noble, 1926:18.
Cochranella fleischmanni — Taylor, 1951:34.
Cochranella petersi Coin, 1961:96 [Holotype.— BMNH 1902.5.27.24 from Rio

Durango, Provincia Esmeraldas, Ecuador]. New synonym.
Centrolenella fleischmanni — Coin, 1964:1.
Centrolenella petersi — Coin, 1964.6.

Diagnosis. — 1) prevomerine teeth absent, 2) bones white; 3)
parietal peritoneum clear; visceral peritoneum white; 4) color in life
pale green with pale yellow spots; in preservative, cream with faint
dark flecks; 5) webbing between outer lingers III2-1/2IV; 6) web-
bing on foot 11-2110-21111-2 IV2 -IV; 7) snout subacuminate in
dorsal view, round in lateral profile; 8) dorsal skin shagreened; 9)
arms and legs lacking dermal folds; 10) humeral spine absent in
males; 11) tympanum almost entirely visible, directed dorsolaterally
with slight posterior inclination.

Centrolenella fleischmanni differs from other Ecuadorian centro-
lenids that have the heart visible in life and a white dorsum in
preservative (munozoriim and peUucida) by having the snout sub-
acuminate in dorsal view and tiomcate in lateral profile, and the
tympanum distinct. The other species have round snouts and the
tympanum concealed. Furthermore, pelliicida differs from fleisch-
manni by having ulnar, tarsal, and anal folds. Centrolenella oro-
costalis from Venezuela differs from fleischmanni by having minute
enamel (yellow in life) flecks on the dorsum.

Description. — Adult males small; snout-vent length 19.2-21.2 mm
(x=20.4, N=5). Head wider than body; width of head 38.3-40.6 per-
cent (x=39.2, N-4) of snout- vent length; snout short, shallow,
roundly subacuminate in dorsal view, round in lateral profile (Fig.
1); canthus round; loreal region shallowly concave; lips not flared;
nostrils three-fourths distance from eye to tip of snout, barely pro-
tuberant, directed dorsolaterally; intemarial area slightly depressed.
Eye large, protuberant, strongly oriented anteriorly. Supratympanic

CENTROLENID FROGS OF ECUADOR

29

600 M. CONTOUR

r~n OVER 3000 M.

[o~] OVER 5000 M
040^100

KILOMETERS

- 5

Fig. 10. Distribution of Centrolenella fleischmanni (circles), C. munozorum
(triangles), and C. pellttcida (square); open symbol is a literature record.

fold absent; t\'mpanum small, about one-fourth diameter of eye,
nearly entirely visible, directed dorsolaterally with slight posterior
inclination. Prevomerine dentigerous processes and teeth absent;
choanae small, round, near margin of mouth; tongue ovoid, free pos-
teriorly for about one-fourth of its length; vocal slits extending from
midlateral base of tongue to angles of jaws.

Humeral spine absent; ulnar folds and tubercles absent; first
finger longer than second; fourth finger nearly as long as third; lateral
fringes present on fingers; webbing vestigial between first and second
and second and third fingers; webbing formula for outer fingers 111(0-
2)-(l)3-l/2)IV; discs moderate, rounded, subarticular tubercles small,
round, simple; supernumerar\' tubercles absent; palmar tubercle
small, ovoid; nuptial excrescences absent. Hind limbs slender;

30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

length of tibia 51.9-57.3 percent (x=55.1, N=4) of snout-vent length;
tarsal folds and tubercles absent; inner metatarsal tubercle low, Hat,
elliptical; outer metatarsal tubercle absent; subarticular tubercles
small, round; supranumerary tubercles absent; feet about three-
fourths webbed; webbing fonmila I(0-l)-( l}^-2)II(0-l)-(2-2^)III(l-
2+)-(2-20IV(2-2^)-lV; discs of toes round, slightly smaller than
those on fingers.

Skin on dorsal surfaces of head and body shagreened; skin on
belly and ventral surfaces of thighs granular; other surfaces smooth;
anal opening directed posteriorly at upper level of thighs; anal folds
and tubercles absent.

Color in preservative: dorsum creamy white with scattered black
flecks visible under microscope; eyelids and hepatic peritoneum
white; skin elsewhere transparent.

Color in life: dorsum pale green with pale yellow or yellowish
green spots, so large in some individuals so as to give appearance of
a pale frog with darker green reticulations; heart visible; belly white;
tips of digits yellow; other surfaces unpigmented; bones white; iris
white to pale yellow.

Distribution. — This is the most widespread species of Centro-
lenella, ranging from Veracruz and Guerrero, Mexico to Ecuador
and Surinam (Coin, 1964; Savage, 1967). In Ecuador it occurs on
the Pacific lowlands and to an elevation of 1460 m at Tandapi on
the Pacific slopes of the Cordillera Occidental of the Andes (Fig. 10).

Remarks. — Numerous males were calling over small streams in
April at the Estacion Biologica Rio Palenque north of Quevedo.
The single specimen from Tandapi was on a leaf of an herb about
2 m above the ground at night.

Coin (1961:96) diagnosed petersi as differing from fleischmanni
by having more extensive webbing on the hand. One specimen ( KU
121052) is like the holotype in having a webbing formula for the
outer fingers IIIO-I/3IV, whereas other Ecuadorian specimens have
III(l/2-2)-l/2lV. Examination of series of specimens from Costa Rica
and Panama reveals that most fall within the range of variation in
the webbing of the hand as displayed by the three Ecuadorian speci-
mens, but three have only one free digit on the third finger and
about one and one-half digits free on the fourth finger. Due to the
lateral fringes on the fingers, the determination of the point of de-
parture of the web is highly subjective in some specimens. We have
observed living fteischmanni in Mexico, Guatemala, Costa Rica, and
Panama, as well as in Ecuador. Comparison of colored photographs
of living individuals from throughout this range reveals that nortii-

CENTROLENID FROGS OF ECUADOR 31

ern frogs tend to be less distinctively marked than southern ones,
but the reticulate pattern is evident in some Panamanian specimens.

Despite the wide geographic range of the species as outlined by
Goin (1964:4), he was reluctant to accept the occurrence of fleisch-
manni in Ecuador; he stated: "I have examined the specimen
(USNM 60520) that Noble (1924:67) recorded as fieischmanni from
Guevedo [=Quevedo, 56 km south of a locality where we obtained
four specimens], Ecuador, and agree with him that this individual
looks like typical fieischmanni from along the north coast of South
America. It seems improbable to me, however, that this species
actually occurs in Ecuador."

We have examined the type specimens of all of the nominal
species included in the foregoing synonymy and have compared our
findings with data obtained from series of living and preserved
frogs. We conclude that the holotypc of Cochranella petersi and
KU 121052 represent the extreme in variation of webbing in Cen-
trolenella fieischmanni and that Cochranella petersi Goin, 1961, is a
junior synonym of Centrolenella fieischmanni (Boettger, 1893).

Centrolenella grandisonae Cochran and Goin

Centrolenella grandisonae Cochran and Goin, 1970:513 [Holotype. — BMNH
1910.7.11.68 from Pueblo Rico, Departamento Caldas, Colombia].

Diagnosis. — l)prevomerinc teeth usually absent; 2) bones pale
green; 3) parietal peritoneum white; visceral peritoneum clear; 4)
color in life green with minute yellow, white, and black flecks; in
preservative, pale lavender with small gray spots and white flecks;
5) webbing between outer fingers 1112^2-2+1 V; 6) webbing on foot
I1-2-II1-2-I1I1-2IV2+1MV; 7) snout round in dorsal and lateral pro-
files; 8) dorsal skin shagreened; 9) row of low, indistinct tubercles
on ventrolateral edges of forearm and tarsus; 10) humeral spine
present in males; 11) tympanum entirely visible, directed laterally
with posterodorsal inclination.

Two other Ecuadorian species are lavender (in preservative)
with dark spots and white flecks. Of these, pipilata differs from
grandisonae by having ulnar and tarsal folds, truncate snout, and
incised webbing. Centrolenella peristicta is nearly identical to
grandisonae in coloration and structural features, except that peri-
sticta has more webbing on the hand ( V/z phalanges free on fourth
finger; 2^ free in grandisonae) and is smaller (mean snout-vent length
19.7 mm; 24.8 in grandisonae ) .

Description. — Adults moderately large; snout-vent length in
males 23.7-25.8 mm (x=24.8, N=4); females not known. Head about
as wide as bodv; width of head 24.6-33.3 percent (x=31.1, N=4) of

32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

snout-vent length; snout short, rounded in dorsal view, in lateral
view inclined above, truncate below, giving a rounded appearance;
canthus rounded; loreal region barely concave; lips not flared; nos-
trils nearly terminal on snout, slightly protuberant dorsolaterally;
internarial area slightly depressed. Eye large, directed anterolater-
ally. Supratympanic fold weak; entire tympanum visible, directed
laterally with posterodorsal inclination. Prevomerine dentigerous
processes and teeth absent; choanae small, round; tongue cordiform,
notched behind, free posteriorly for about one-fourth of its length;
vocal slits extending from midlateral base of tongue to angles of
jaws.

Humeral spine blunt, perpendicular to humerus; row of low
tubercles on ventrolateral edge of forearm; first finger longer than
second; fourth finger much shorter than third; lateral fringes present
on fingers; webbing absent between first and second fingers, vestigial
between second and third fingers; webbing formula for outer fingers
III(2)4-2M)-(2--2')IV; discs truncate; subarticular tubercles small,
round, simple; supernumerary tubercles absent; palmar tubercle
ovoid, simple; nuptial excrescences absent (Fig. 3). Hind limbs
slender; length of tibia 54.9-57.3 percent (x=55.8, N=4) of snout- vent
length; row of low, indistinct tubercles on ventrolateral edge of
tarsus; inner metatarsal tubercle small, elliptical; outer metatarsal
tubercle small, ovoid; subarticular tubercles small, round; super-
numerary tubercles absent; toes about two-thirds webbed; webbing
formula I(0-l?0-l'3-2^)II(0-l)-( lM-20^I(l-lK)-(2-20IV(2-2}i)-(l-
l/2)V; discs smaller and more nearlv round than those on fingers
(Fig. 4).

Skin on dorsal surfaces shagreened; skin on belly and proximal
posteroventral surfaces of thighs granular, other surfaces smooth;
anal opening directed posteriorly at upper level of thighs; pair of
large tubercles below anus.

Color in preservative: dorsal surfaces lavender with large dark
smudges and small white flecks; other surfaces cream.

Color in life: green with minute vellow and white flecks and
dark green and black spots on body and limbs; tips of digits pale
yellow; humeral spine bluish green, vocal sac green; chest white;
heart not visible; visceral peritoneum transparent; bones green; iris
pale golden bronze flecked with black.

Distribution. — Centrolenella grandisonae occurs at moderate ele-
x^ations on the Pacific slopes of the Cordillera Occidental of the
Andes from southwestern Colombia to northwestern Ecuador (Fig.
7). In Colombia it is known from Pueblo Rico, 1540 m and Santa

CENTROLENID FROGS OF ECUADOR 33

Leticia, 2000 m, Departamento Caldas, and in Ecuador from Tan-
dapi, 1460 m, Provincia Pichincha.

Remarks. — Three calling males were found along the small
stream in cloud forest at Tandapi in July 1967. With the exception
of one metamorphosing young found along the Rio Tandapi, gran-
disonae was observed only along the one stream, where males were
calling from leaves of bushes and trees by a waterfall. None of the
other four species of CentroleneUa known from Tandapi was found
along this stream. The metamorphosing young has a snout- vent
length of 13.5 mm and a tail stub of 2 mm.

The holotype of grandisonae is the largest known specimen of
the species ( 27.4 mm ) and is the only one having prevomerine teeth.

CentroleneUa griffithsi (Coin)

Plate 2C

Cochranella griffithsi Coin, 1961:99 [Holotyi^e.— BMNH 1940.2.20.4 from Rio
Saloya, Provincia Pichincha, Ecuador].

CentroleneUa griffithsi Goin, 1964:6.

Diagnosis. — 1) prevomerine teeth absent; 2) bones pale green;
3) parietal peritoneum clear; visceral peritoneum white; 4) color in
life yellowish green with or without dark flecks; in presei"vative, dull
lavender; 5) webbing between outer fingers III3-2/3IV; 6) webbing
on foot I2-2KII 1-2^111 1-2MIV233-1V; 7) snout truncate in dorsal and
lateral profiles; 8) dorsal skin shagreened; 9) arms and legs lacking
dermal folds; 10) humeral spine absent in males; 11) lower three-
fourths of tympanum visible, directed dorsolaterally with posterior
inclination.

The diagnostic characters of griffithsi resemble those of three
other CentroleneUa — huckleiji, megacheira, and prosoblepon. Of
these, megacheira is much larger and has pustular dorsal skin. Cen-
troleneUa huckleiji and prosoblepon have more webbing, and the
males have humeral spines.

Description. — Adults moderately large; snout- vent length 19.7-
26.1 mm (x=24.1, N=14) in males^ 21.6-24.8 (x=23.4, N=3) in fe-
males. Head slightly wider than body; width of head 30.6-34.7 per-
cent (x=32.3, N=17) of snout-vent length; snout short, truncate in
dorsal and lateral profiles ( Fig. 1 ) ; canthus round; loreal region
concave; lips not flared; nostrils nearly terminal on snout, slight
protuberant dorsolaterally; internarial area barely depressed. Eye
moderately large, directed anterolaterally. Supraty^mpanic fold
barely evident; lower three fourths of tympanum visible, directed
dorsolaterally with posterior inclination. Prevomerine dentigerous
processes and teeth absent; choanae large, quadrangular; tongue

34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

cordiform, distinctly notched behind, barely free posteriorly; vocal
slits extending from midlateral base of tongue to angles of jaws.

Humeral spine absent; ulnar fold and tubercles absent; first fin-
ger longer than second; fourth finger noticeably shorter than third;
lateral fringes absent on fingers; webbing absent between first, sec-
ond, and third fingers; webbing formula for outer fingers III3-2/3IV;
discs broad, rounded; subarticular tubercles small, round, simple;
supernumerary tubercles absent; palmar tubercle large, elliptical,
simple; nuptial excrescences absent (Fig. 3). Hind limbs slender;
length of tibia 52.4-62.3 percent (x=55.2, N=17) of snout- vent length;
tarsal folds and tubercles absent; inner metatarsal tubercle small,
ovoid; outer metatarsal tubercle absent; subarticular tubercles small,
round; supernumerary tubercles absent; feet about one-half webbed;
webbing formula I(2-2-)-(2--2)i)II(l^-l?^)-(2>i-3-)nilJ^(2)^-3-)IV(2M-
3")-2~V; discs on toes smaller and more nearly round than those on
fingers ( Fig. 4 ) .

Skin on dorsal surfaces shagreened; skin on belly and postero-
ventral surfaces of thighs granular; other surfaces smooth; anal
opening directed posteriorly at upper level of thighs; pair of large
tubercles below anus.

Color in preservative: dorsal surfaces of head, body, foreanus,
shanks, and feet dull lavender with or without minute black flecks;
other surfaces cream.

Color in life: dorsal surfaces pale yellowish green with or with-
out dark green flecks; tips of digits pale yellow; chest white; heart
not visible; visceral peritoneum transparent; bones pale green; iris
whitish bronze.

Distribution. — CentroIeneUa griffithsi occurs on the Pacific slopes
of the Cordillera Occidental of the Andes from southwestern Colum-
bia to northwestern Ecuador (Fig. 9); the species inhabits cloud
forests at elevations of 1200-2170 m.

Remarks. — Most of our specimens were obtained from low vege-
tation in cloud forest at Tandapi. Males were calling from leaves of
herbs and bushes over cascading streams. One was found in the axil
of an elephant-ear plant by day. Calling males were obsen'ed by
Lynch on each of his five trips to Tandapi ( March 196S, June 1968,
July 1967, 1968, and 1970). Sparsely distributed and sporadically
calling males were heard every evening at Tandapi, but the breed-
ing season clearly is in March, when most available sites were occu-
pied by calling males, and vegetation overhanging streams was
festooned with egg masses. Eggs were not observed in June or July.

In July 1968, while we were collecting astroblepid catfishes in
the gravel at the bottom of a small stream, two centrolenid tadpoles

CENTROLENID FROGS OF ECUADOR 35

were found. These may be the tadpoles of griffithsi, the most abun-
dant of the five species of CentroleneUa known from Tandapi; only
griffithsi adults were found along the stream. Structurally identical
tadpoles were collected at Pilalo, Provincia Cotopaxi, 2500 m.

Goin (1969:99) gave the type locality as "Rio Saloya, Ecuador,
4000 feet." This locality probably is the point on the Rio Saloya
where it is crossed by the road from Chillogallo to Santo Domingo
de los Colorados, the only road west from Quito in 1940, when the
specimens were collected. The point where the road crosses the Rio
Saloyo is about 1200 m. The species is abundant in the next valley
south, that of the Rio Pilaton.

CentroleneUa medemi Cochran and Goin

CentroleneUa medemi Cochran and Goin [Holot\pe. — USNM 152277 from
Puerto Asis, Comisaria Putuniayo, Colombia].

Diagnonis. — l)prevomerine teeth 2-3; 2) bone color unknown;
3) heart apparently not visible; 4) color in life unknown; in pre-
servative, dark gray with many cream spots; 5) webbing between
outer fingers III2 -i^^IV; 6) webbing on foot lO-OIIO-OIIIO-lIVl-OV;
7) snout truncate in dorsal and lateral profiles; 8) dorsal skin
smooth; 9) arms and legs lacking dermal folds; 10) small humeral
spine present in males; 11) tympanum concealed.

CentroleneUa medemi is unique among known centrolenids by
having a dark gray dorsum with many large, round spots on all dor-
sal surfaces. The only other Amazonian species having humeral
spines, audax and pipilota, have distinct t\inpana, less webbing, and
a pale green dorsum with small )'ellow flecks (audax) or dark green
with black spots and yellow flecks (pipilata).

Description. — Adult male moderate in size, 25.3 mm in snout-
vent length; head much wider than body; width of head 39.5 percent
of snout- vent length; snout moderately short, truncate in dorsal and
lateral profiles; canthus round; loreal region barely concave; lips not
flared; nostrils nearly terminal, not protuberant, directed dorso-
laterally; internarial area flat. Eye moderately large, directed antero-
laterally. Supratympanic fold absent; t\mpanum concealed. Pre-
vomerine dentigerous processes short, trans\erse between choanae,
bearing 2-3 teeth; choanae small, ovoid, near margin of mouth;
tongue ovoid, barely free posteriorly; vocal slits extending from
midlateral base of tongue towards angles of jaws.

Humeral spine curved, not projecting; ulnar folds and tubercles
absent; lateral fringes absent on fingers; webbing formula I2-2/2II2"-
3TII2 -I/2IV; discs large, truncate; subarticular tubercles large,
round, simple; supernumerary tubercles absent; palmar tubercle

36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

large, simple, ovoid; nuptial excrescences absent. Hind limbs robust;
length of tibia 68.8 percent of snout-vent length; tarsal folds and
tubercles absent; inner metatarsal tubercle low, elongate; outer meta-
tarsal absent; subarticular tubercles small, round; supernumerary
tubercles absent; feet essentially fully webbed; webbing formula
lO-OIIO-OIIIO-lIVl-OV; discs on toes slightly smaller and more nearly
round than those on fingers.

Skin on belly granular; skin on other surfaces smooth; anal open-
ing directed posteriorly at upper level of thighs; anal folds and
tubercles absent.

Color in preservative: all dorsal surfaces slate gray with many
round cream spots; ventral surfaces cream. Color in life unknown.

Distri])Ution. — Known only from the type locality at an elevation
of about 280 m on the Rio Putumayo in Amazonian Colombia
(Fig. 7).

Remarks. — This distinctive species is included here in the antic-
ipation that it will be found in Ecuador; the type locality is only
20 km north of the Ecuadorian border.

Centiolenella megacheira new species

Plate 2A

Holotype. — KU 143245, an adult male, 27.1 mm, from a stream
16.5 km NNE of Santa Rosa, 1700 m. on Quito-Lago Agrio road,
Provincia Napo, Ecuador, one of a series obtained on 17 October
1971, by Joseph T. Collins and William E. Duellman.

Parotopotypes.—KU 143246-72, BMNH 1971.1854-55, CAS
135498-9, UMMZ 131668 (3) obtained on 17-19 October 1971, by
Joseph T. Collins and William E. Duellman.

Diagnosis. — 1) prevomerine teeth absent; 2) bones green; 3)
parietal peritoneum white; visceral peritoneum clear; 4) color in life
green with black spots; in preservative, lavender with black spots;
5) webbing between outer fingers III3-273IV; 6) webbing on foot
I2--2^IIl,'2-2MIIIlM-2:^nV3 -I73V; 7) snout truncate in dorsal and lat-
eral profiles; 8) dorsal skin pustular; 9) arms and legs lacking dermal
folds; 10) humeral spine absent in males; 11) lower three-fourths of
tympanum visible, directed dorsolaterally with slight posterior in-
clination.

CentrolencUa megacheira is distinguished from all other Andean
species by its large size, pustular dorsal skin, and small amount of
webbing on the hands and feet. The presence of black flecks on the
dorsum also is characteristic of prosoblepon, males of which have
humeral spines.

CENTROLENID FROGS OF ECUADOR 37

Descriptions. — Adults large; snout- vent length 27.1-32.8 mm
(x=28.2, N=20) in males, 32.1-32.8 mm (x=32.5, N=3) in females.
Head much wider than body; width of head 32.8-37.4 percent
(x=34.3, N=23) of snout-vent length; snout short, truncate in dorsal
and lateral profiles; canthus rounded; loreal region slightly concave;
lips barely flared; nostrils four-fifths distance from eye to tip of
snout, slightly protuberant dorsolaterally; internarial area depressed.
Eye large, directed more anteriorly than laterally. Supratympanic
fold weak; lower three-fourths of tympanum visible, directed dorso-
laterally with slight posterior inclination. Prevomerine dentigerous
processes and teeth absent; choanae small, ovoid; tongue cordiform,
barely free posterif)rly; vocal slits extending from midlateral base of
tongue to angles of jaws.

Humeral spine absent; ulnar fold and tubercles absent; hand
large; first finger equal in length to second; fourth finger slightly
shorter tlian third; lateral fringes present on fingers; webbing absent
between first and second fingers; wel)bing formula for other fingers
n(2+-2M)-(33^-3?4)III(2^^-3-)-(2J^-3)IV; discs broader than deep,
rounded; subarticular tubercles large, ovoid, simple; supernumerary
tubercles absent; palmar tubercle large, ovoid, simple; nuptial ex-
crescences absent. Hind limbs slender; length of tibia 53.7-62.3 per-
cent (x=56.9, N=23) of snout- vent length; tarsal folds and tubercles
absent; inner metatarsal tubercle large, elliptical; outer metatarsal
tubercle small, ovoid; subarticular tubercles small, round; super-
numerary tubercles absent; feet about one-half webbed; webbing
formula ' I ( 2-2 ) - ( 2^-2)^ ) II ( l-H^ - ( 2^^-2;:i ) III ( VA-m ) - ( 2]^-3- ) IV ( 2%-
31 )-( 1/2-2 )V; discs on toes slightly smaller and more nearly round
than those on fingers.

Skin on dorsal surfaces pustular; skin on belly and ventral sur-
faces of thighs granular; other surfaces smooth; anal opening di-
rected posteriorly at upper level of thighs; pair of large tubercles
below anus.

Color in preservative: dorsal surfaces of head, body, hind limbs,
and forearms lavender with small, round, black dots; hands, feet,
upper arms, and ventral surfaces cream.

Color in life: dorsum green with black dots; flanks cream; chest
white; heart not visible; visceral peritoneum unpigmented; other
ventral surfaces green; iris pale grayish bronze; bones green.

Distribution. — The species currently is known at two localities at
elevations of 1700 and 1740 m on the Amazonian slopes of the Cor-
dillera Oriental in Ecuador (Fig. 9). The type locality is a small
stream which drains into a tributary of the Rio Salado. This stream

38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

is crossed by the Quito-Lago Agrio road at a point 16.5 km north-
northeast of the village of Santa Rosa.

Remarks. — Individuals were found at night on leaves and stems
of bushes and trees overhanging streams in cloud forest. Males were
calling in October 1971. At the type locality, axidax, pipilata, and
sire7i were found with megacheira; at the Rio Azuela, anomala, pel-
liicida, pipilata, and siren occurred along the same streams with
megacheira.

Etymology. — The specific name is from the Greek megas, mean-
ing large, and the Greek cheiros, meaning hand; the name is used in
reference to the exceedingly large hands of this species.

Centrolenella midas new species
Plate ID

Holotype. — KU 123219, an adult male, 19.2 mm, from Santa
Cecilia, 340 m, Provincia Napo, Ecuador, obtained on 22 June 1968,
by Linda Trueb.

Paratypes.—¥A] 107026, 23 November 1966, William E. Duell-
man; KU 146625, 2 April, Martha L. Crump; KU 150622, 25 June
1971, Philip S. Humphrey; KU 150623, 28 August 1971, Martha L.
Crump, all from the type locality. KU 125334, 23 May 1969, Thomas
H. Fritts, and UMMZ 129314, 6 May 1969; Charles F. Walker from
Lago Agrio, 330 m, Provincia Napo, Ecuador.

Diagnosis. — 1) prevomerine teeth 1-3; 2) bones green; 3) pari-
etal peritoneum white; visceral peritoneum clear; 4) color in life
dark green with yellow flecks; in preservative, lavender with white
flecks; 5) webbing between outer fingers III2-2 IV; 6) webbing on
foot I1-2II1-2III1-2^IV2^-1V; 7) snout truncate in dorsal and lateral
profiles; 8) dorsal skin shagreened; 9) arms and legs lacking dermal
folds; 10) humeral spine absent in males; 11) lower two-thirds of
tympanum visible, directed dorsolaterally with strong posterior in-
clination.

Centrolenella midas is like favopiinctata and siren in having a
lavender dorsum with white flecks ( in life, green with gold flecks)
and no black flecks; it differs from favopiinctata by having a trun-
cate, instead of round snout and slightly less webbing on the hands
and feet. Centrolenella siren differs from midas by having less web-
bing and a more prominent tympanum oriented posterolateral ly,
instead of dorsolaterally.

Description. — Adults small; snout- vent length 17.4-19.2 mm
(x=18.4, N=3) in males, 20.6-25.6 mm (x=22.7, N=7) in females. Head
wider than body; width of head 33.3-39.1 percent (x=36.0, N=10) of
snout-vent length; snout short, truncate in dorsal and lateral profiles;

CENTROLENID FROGS OF ECUADOR 39

canthus round; loreal region concave; lips rounded; nostrils nearly
terminal on snout, slighth' protuberant dorsolaterally; intemarial
area depressed. Eye moderately large, directed anterolaterally.
Supratympanic fold absent; lower two-thirds of tympanum visible,
directed dorsolaterally with strong posterior inclination. Dentigerous
processes of prevomers small, low, widely separated between mod-
erately large, round choanae, bearing 1-3 teeth; tongue broadly
cordiform, barely free posteriorly; vocal slits extending from mid-
lateral base of tongue to angles of jaws.

Humeral spine absent; ulnar fold and tubercles absent; first fin-
ger longer than second; fourth finger noticeably shorter than third;
webbing vestigial between first and second fingers; webbing formula
for other fingers 11(2-2 )-( 3 -3M) III (2--2^)-( U^-2^)IV; discs truncate;
subarticular tubercles small, round, simple; supernumerary tubercles
absent; palmar tubercle large, elliptical; nuptial excrescences absent.
Hind limbs moderately slender; lengtli of tibia 53.1-62.0 percent
(x=57.7, N=10) of snout- vent length; tarsal folds and tubercles ab-
sent; inner metatarsal tubercle elongate, rounded in section; outer
metatarsal tubercle absent; subarticular tubercles small, round;
supernumerary tubercles absent; feet about two-thirds webbed; web-
bing formula I(0-13^)-(2--2^)n(.^M)-(2-20mi-(2-2K)IV(2-2^)-IV;
discs on toes more nearly round than those on fingers.

Skin on dorsal surfaces shagreened; skin on belly and proximal
postero ventral surfaces of thighs granular; other surfaces smooth;
anal opening directed posteriorly at upper level of thighs; pair of
large tubercles below anus.

Color in preservative: dorsal surfaces of head, body, and limbs
lavender with few small white flecks on body; other surfaces creamy
white.

Color in life: dorsum of head, body, and limbs dark green with
a few small yellow flecks dorsolaterally on body; hands and feet dull
greenish yellow; chest white; heart not visible; visceral peritoneum
unpigmented; iris silvery bronze with black reticulations; bones
green.

Distribution. — This species is known from three localities at ele-
vations of 330-570 m along the Rio Aguarico in the upper Amazon
Basin in Ecuador (Fig. 9). In this area it occurs in sympatry with
munozorum and resplendens.

Remarks. — Individuals have been found throughout the year on
leaves of herbs and trees along small rivulets in rainforest. The call
consists of three short notes.

Etymology. — The specific name is that of a king in Greek my-
thology, at whose touch everything tm-ned to gold. The name is

40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

associated with this frog known along the Rio Aguarico, meaning
rich water, in reference to gold found in the river, and in allusion to
the gold flecks on the frogs.

Centrolenella munozorum new species
Plate IB

Holotype. — KU 118054, an adult male, 20.2 mm, from Santa
CeciHa, 340 m, Provincia Napo, Ecuador, obtained on 18 June 1967,
by John D. Lynch.

Paratypes.—KV 105251, 13 July 1966, Charles M. Fugler; KU
123225, 13 July 1968, William E. Duellman; KU 150620, 13 August
1971, Martha L. Crump; KU 150621, 10 October 1971, Martha L.
Crump, all from Santa Cecilia, and UMMZ 129313 from Lago Agrio,
330 m, 14 km E Santa Cecilia, Provincia Napo, Ecuador, 6 May
1969, Charles F. Walker.

Dia(^nosis. — 1) prevomerine teeth absent; 2) bones white; 3)
parietal peritoneum clear; visceral peritoneum white; 4) color in
life pale green with pale greenish yellow spots; in preservative,
creamy white with pale gray reticulations; 5) webbing between
outer fingers IIIIJ^-IJIIV; 6) webbing on foot I0-1II0-L^^III1-2IV2--
IV; 7) snout round in dorsal and lateral profiles; 8) dorsal skin
shagreened; 9) arms and legs lacking dermal folds; 10) humeral
spine absent in males; 11) tympanum concealed; strongly directed
dorsolaterally.

Two other Ecuadorian species have the heart visible in life and
a white dorsum in preservative. Centwlenella peUiicida differs from
munozorum by having ulnar, tarsal, and anal folds, and fieischmanni
differs by having the snout subacuminate in dorsal view and truncate
in lateral profile and the tympanum is distinct; in munozorum and
pellucida the snout is round, and the tympanum is concealed. The
Venezuelan orocostalis has enamel white (yellow in life) flecks on
the dorsum.

Description. — Adults small; snout- vent length in males 18.8-20.5
mm (x=19.7, N-5), in one female 20.7 mm. Head wider than body
width of head 37.7-40.7 percent (x=38.4, N=6) of snout-vent length
snout short, shallow, round in dorsal and lateral profiles (Fig. 1)
canthus round; loreal region shallowly concave; lips slightly flared
nostrils about four-fifths distance from eye to tip of snout, barely
protuberant, directed dorsolaterally; internarial area slightly de-
pressed. Eye moderately large, protuberant, directed anterolaterally.
Supratympanic fold absent; tympanum concealed, strongly directed
dorsolaterally. Prevomerine dentigerous processes and teeth absent;
choanae small, ovoid, near margin of mouth; tongue ovoid, barely

CENTROLENID FROGS OF ECUADOR 41

free posteriorly; vocal slits extending from posterolateral base of
tongue to angles of jaws.

Humeral spine absent; ulnar fold and tubercles absent; first finger
longer than second; fourth finger nearly as long as third; lateral fringes
present on fingers; webbing vestigial between first and second and
second and third fingers; webbing fomiula for outer fingers 111(1/2-
2")-(l-l/2)IV; discs small, rounded, subarticular tubercles small, round,
simple; supernumerary tubercles present on proximal segments of
Erst and second fingers; palmar tubercle small, round; nuptial ex-
crescences absent (Fig. 3). Hind limbs slender; length of tibia 57.5-
59.6 percent (x=5S.4, N=5) of snout-vent length in males, 54.1 per-
cent in one female; tarsal folds and tubercles absent; inner meta-
tarsal tubercle small, elongate; outer metatarsal tubercle absent;
subarticular tubercles small, round; supernumerary tubercles absent;
fringe on inner edge of first toe; toes about three-fourths webbed;
webbing formula 10- (1-^0 II (0-1) -(l^-2-)III( 0-1)- (2--20IV(2--2) -
(0-1 )V; discs on toes round, slightly smaller than those on fingers.

Skin on dorsal surfaces of head and body shagreened; skin on
belly and proximal ventral surfaces of thighs weakly granular; other
surfaces smooth; anal opening directed posteriorly at upper level of
thighs; anal folds and tubercles absent.

Color in preservative: dorsal surfaces of head, body, forearms,
thighs, and shanks creamy white with many minute black flecks
giving appearance of gray with unpigmented spots; eyelid and he-
patic peritoneum white; other surfaces unpigmented.

Color in life: dorsum pale green with pale yellow or yellowish
green spots; limbs pale green, with slightK- darker crossbars in one;
thighs unpigmented; iris pale gold.

Distribution. — Currently this species is known from only two
localities along the Rio Aguarico in the Amazonian lowlands of
Ecuador (Fig. 10).

Remarks. — All individuals were found in lowland rainforest. At
Santa Cecilia the frogs were found on leaves of bushes and trees at
night: one over a pond, one away from water in primary forest, one
on a palm frond 2 m above a stream, and one on an herbaceous leaf
more than 2 m above a stream. The specimen from Lago Agrio was
obtained from the foliage of a large tree that was felled during the
clearing of primary forest.

Etymology. — The specific name is a patronym for Ing. Ildefonso
Muiioz B. and Sra. Blanca Munoz, our congenial hosts at Santa
Cecilia.

42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Centrolenella ocellifera (Boulenger)

Hijla oceUifera Boulenger, 1899:277 [Holotype.— BMNH 98.5.19.3 from Pa-

raniba, Provincia Imbaljura, Ecuador].
Cochranella oceUifera — Taylor, 1951:35.
Centrolenella oceUifera — Coin, 1964:6.

Diagnosis. — 1) prevomerine teeth absent; 2) bones white (?);
3) parietal peritoneum white; visceral peritoneum clear; 4) color in
life green with yellow spots; in preservative, pale lavender with faint
purple ocelli enclosing white spots; 5) webbing between outer fin-
gers III2-UnV; 6) webbing on foot Il-2IIl-2-IIIl-2IV2^-lV; 7 snout
round in dorsal view, truncate in lateral profile; 8) skin shagreened;
9) arms and legs lacking dermal folds; 10) humeral spine absent in
males; 11) lower two-thirds of tympanum visible, directed laterally
with slight posterodorsal inclination.

Centrolenella ocellifera has shagreened skin and few large ocelli
on the dorsum. Other species having ocellated dorsal patterns (ano-
mala, cochranae, and ocellata) have minute spicules in the dorsal
skin and less webbing between the outer fingers. Furthermore,
ocellifera difl^ers from anomala by having a green instead of brown
dorsum, from ocellata by having the lower two-thirds of the tym-
panum visible (concealed in ocellata), and from cochranae by hav-
ing larger ocelli, snout round in dorsal view, and prevomerine teeth
absent (cochranae has small ocelli, snout truncate in dorsal view,
and prevomerine teeth usually present ) .

Description. — Adults moderate-sized; snout-vent length 20.0 mm
in male, 26.7 mm in female. Head as wide as body; width of head
32.9-37.0 percent (x=35.0, N=2) of snout- vent length; snout short,
round in dorsal view, truncate in lateral profiles; canthus round;
loreal region concave; lips slightly flared; nostrils four-fifths distance
from eye to tip of snout, not protuberant, directed laterally; inter-
narial area flat. Eye large, protuberant, directed anterolaterally.
Supratympanic fold not evident; lower two-thirds of tympanum vis-
ible, directed laterally with slight posterodorsal inclination. Prevo-
merine teeth absent in one male, 2-3 on transverse processes between
choanae in one female; choanae small, rectangular; tongue ovoid,
slightly free posteriorly.

Humeral spine absent; ulnar folds and tubercles absent; first fin-
ger larger than the second; fourth finger nearly as long as third;
fingers extensively webbed; webbing formula I2/4-2II2-3III2-(l/2-
ri ) IV; fringe on outer edge of fourth finger; discs moderately broad,
truncate; subarticular tubercles small, round, simple (distal tubercle
on fourth fingers bifid in female); supernumerary tubercles absent;
palmar tubercle single, ovoid; nuptial excrescences absent. Hind

CENTROLENID FROGS OF ECUADOR 43

limbs long, slender; tibia length 52.4-59.5 percent (x=55.9, N=2) of
snout-vent length; tarsal folds and tubercles absent; inner metatarsal
tubercle moderately small, elliptical; outer metatarsal tubercle ab-
sent; subarticular tubercles small, round; supernumerary tubercles
absent; feet about three-fourths webbed; webbing formula I1-2II1-
2 III(1"-1)-2IV2'^-1V; fringe on inner edge of first toe; discs on toes
smaller and more nearly round than those on fingers.

Skin on dorsal surfaces shagreened, that on belly and ventral
surfaces of thighs weakly granular; other surfaces smooth; anal
opening directed posteriorly at upper level of thighs; anal folds and
tubercles absent.

Color in preservative: dorsal surfaces of head, body, and shanks
pale lavender, scattered dark brown flecks; three or four thin purple
ocelli with cream centers in scapular region ( Fig. 5); chest and belly
cream; other surfaces transparent.

Color in life: body green with pale yellow spots and dark green
flecks on body; limbs paler green; chest white; heart not visible;
visceral peritoneum transparent; iris gray-bronze.

Distribution. — Centrolenella ocellifera inhabits the Pacific slopes
of the Andes (Fig. 6). In addition to the holotype from Paramba
(777 m), we have a specimen from Tandapi (1460 m) and have
seen a specimen in the Gustavo Orces-V. collection in the United
States National Museum from Pilalo (2320 m).

Renuirks. — Our specimen was found at night on a fern in cloud
forest. The fern was at the edge of a cliff about 10 m above the Rio
Pilaton.

Centrolenella pelliicida new species
Plate IC

Holotype. — KU 14329S, a gravid female, 22.0 mm from the Rio
Azuela, 1740 m, Quito-Lago Agrio road, Provincia Napo, Ecuador,
obtained on 20 October 1971, by William E. Duellman.

Diapwsis. — 1) prevomcrine teeth absent; 2) bones white; 3)
parietal peritoneum clear; visceral peritoneum white; 4 ) color in life
pale green with diftuse yellow spots; in preservative, uniform cream;
5) webbing between outer fingers III2 -2IV; 6) webbing on foot
Il-2^IIl-l)2lIIl-2'IV2 -IV; 7) snout round in dorsal and lateral pro-
files; 8) dorsal skin shagreened; 9) unscalloped dermal fold on outer
edge of hand and forearm and on foot and tarsus; transverse dermal
fold below anus; 10) humeral spine absent in males; 11) tympanum
concealed.

Among the Ecuadorian species (flei.schmonni, peUucida, and
munozonim) having the heart visible in life and the dorsum white

44 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

in preservative, C. peUucida is unique in possessing ulnar, tarsal, and
anal folds. Moreover, in C. jleischmanni the snout is subacuminate
in dorsal view and truncate in lateral profile, and the tympanum is
distinct; in C. peUucida and C. rnunozorum the snout is round, and
the tympanum is concealed. The Venezuelan orocostalis, a member
of the fleischmoimi group, also lacks dermal folds.

Description. — Males unknown; adult female small, 22.0 mm in
snout-vent length; head slightly wider than body; width of head 36.4
percent of snout-vent length; snout short, round in dorsal and lateral
profiles; canthus round; loreal region barely concave; lips not flared;
nostrils nearly terminal, directed laterally, not protuberant; inter-
narial area flat. Eye large, protuberant, strongly oriented anteriorly.
Supratympanic fold absent; tympanum concealed. Prevomerine
dentigerous processes and teeth absent; choanae large, oval; tongue
ovoid, barely free posteriorly.

Humeral spine absent; narrow, unscalloped dermal fold on ven-
trolateral edge of forearm and outer edge of hand; fringes absent on
fingers; first finger longer than second; fourth finger slightly shorter
than third; fingers extensively webbed; webbing formula I2-2+II2"-
3III2 -2IV; discs moderately large, rounded; subarticular tubercles
small, low; supernumerary tubercles absent; palmar tubercle small,
indistinct. Hind limbs slender; tibia length 56.4 percent of snout-
vent length; narrow, unscalloped dermal fold along outer edge of
tarsus and fifth toe; inner metatarsal tubercle small, ovoid; outer
metatarsal tubercle absent; subarticular tubercles small, low; super-
numerary tubercles absent; feet about three-fourths webbed; web-
bing formula Il-2ni-l)2nil-2 IV2^-IV; discs on toes round, slighdy
smaller than those on fingers.

Skin on dorsum of head and body shagreened; skin on other sur-
faces smooth; anal opening directed posteriorly at upper level of
thighs; transverse dermal fold below anus at posteroventral edge of
thighs (Fig. 2).

Color in preservative: dorsum creamy white with minute purple
flecks visible under magnification; hepatic peritoneum white; skin on
ventral surfaces transparent.

Color in life: dorsum pale green with diffuse yellow spots; venter
and hidden surfaces of limbs lacking pigment; fingers and toes yel-
low; parietal peritoneum clear; heart visible; bones white; iris pale
silvery bronze.

Distribution. — This small species is known only from the type
locality on the east slope of Volcan Reventador on the Amazonian
slopes of the Andes (Fig. 10).

Remarks. — The holotype was on the leaf of an herb over a small

CENTROLENID FROGS OF ECUADOR 45

stream at night. See the account of anomala for a detailed descrip-
tion of the type locaUty and comments on associated species.

Etymology. — The specific name is Latin meaning transparent and
is appUed to this species having a transparent parietal peritoneum.

Centrolenella peristicta new species
Plate 2G

Holotype. — KU 11(S051, an adult male, 20.6 mm, from Tandapi,
1460 m, Provincia Pichincha, Ecuador, obtained on 23 July 1967, by
John D. Lynch.

Paratopotypes.—KV 118052, 24 July 1967, Marsha Lynch;
121053, 28 July 1968, Gerald R. Smith.

Diagnosis. — 1) prevomerine teeth absent; 2) bones pale green;
3) parietal peritoneum white; visceral peritoneum clear; 4) color in
life pale green with minute \ellow, white, and black Hecks; in pre-
servative, pale lavender with dark lavener spots and few white
flecks; 5) webbing between outer fingers III2 -IMIV; 6) webbing on
foot 11-2 II1-2III1-2^IV2-1V; 7) snout round in dorsal and lateral
profiles; 8) dorsal skin shagreened; 9) row of low tubercles on ven-
trolateral edges of forearm and tarsus; 10) humeral spine present in
males; 11) tympanum entirely visible, directed laterally with shght
dorsal inclination.

Two other Ecuadorian species are lavender (in preser\'ative )
with dark spots and white flecks. Of these pipilota differs from
peristicta by haxing ulnar and tarsal folds, truncate snout, and in-
cised webbing. Centrolenella grandisonae is nearly identical to
peristicta in coloration and structural features, except that grandi-
sonae has less webbing on the hand (2* phalanges free on fourth
finger; VA free in peristicta) and is larger (mean snout- vent length
24.8 mm; 19.7 mm in peristicta) .

Description. — Adults small; snout- vent length 18.7-20.6 mm
(x=19.7, N=2) in males, 20.5 mm in one female. Head no wider than
body; width of head 32.5-32.7 percent (x=32.6, N^3); snout short,
round in dorsal and lateral profiles; canthus round; loreal region
concave; hps rounded; nostrils nearly terminal on snout, slightly
protuberant dorsolaterally; internarial area depressed. Eye moderate
sized, directed more laterally than anteriorly. Supratympanic fold
weak; tympanum entirely visible, directed laterally with slight dor-
sal inclination. Prevomerine dentigerous processes and teeth absent;
choanae small, ovoid; tongue cordiform, shallowly notched behind,
barely free posteriorly; vocal slits extending from midlateral base of
tongue to angles of jaws.

Humeral spine curv^ed, terminus parallel to humerus; row of low

46 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

tubercles on ventrolateral edge of forearm; first finger longer than
second; fourth finger much shorter than third; lateral fringes present
on fingers; webbing vestigial between first and second fingers; web-
bing formula for other fingers II ( 2-2/2 )-3'^III2--l)2lV; discs truncate;
subarticular tubercles small, round; distal subarticular tubercle on
fourth finger bifid; supernumerary tubercles small, round, present on
proximal segments of digits 2-4; palmar tubercle large, ovoid; nup-
tial excrescences absent ( Fig. 3 ) . Hind limbs slender; length of tibia
52.2-58.8 percent (x=54.8, N=3) of snout-vent length; row of low
tubercles on ventrolateral edge of tarsus; inner metatarsal tubercle
elliptical; outer metatarsal tubercle small, ovoid; subarticular tu-
bercles small, round; supernumerary tubercles absent; feet about
two-thirds webbed; webbing formula Il-(l)2-2)III-(2--2+)IIIl-(2--2+)
IV(2'-2")-lV; discs smaller and more nearly round than those on
fingers.

Skin on dorsum shagreened; skin on belly and proximal postero-
ventral surfaces of thighs granular; other surfaces smooth; anal
opening directed posteriorly at upper level of thighs; pair of large
tubercles below anal opening.

Color in preservative: dorsal surfaces lavender with dark spots
and scattered cream flecks; other surfaces cream.

Color in life: dorsal surfaces pale to medium green with dark
green spots and white and yellow flecks; flanks yellow-green; belly
yellow; vocal sac green; tips of digits pale yellow; heart not visible;
bones green; iris grayish bronze with brown or copper ring around
pupil.

Distribution. — This species is known only from Tandapi (for-
merly Cornejo Astorga), a village at the point where the Quito-Santo
Domingo de los Colorados road crosses the Rio Pilaton at an eleva-
tion of 1460 m on the Pacific slopes of the Cordillera Occidental of
the Andes (Fig. 8).

Remarks. — All three individuals were obtained in July on vege-
tation in cloud forest at night. One was on an elephant-ear leaf; one
was on a fern in the spray zone of a waterfall, and one was on a
bush over a waterfall. None was calling.

Etymology. — The specific name is derived from the Greek
peristiktos, meaning dappled, and refers to the spotted color pattern.

Centrolenella pipilata new species
Plate 2D

Holotype. — KU 143278, an adult male, 22.9 mm, from a stream
16.5 km NNE of Santa Rosa, 1700 m, on Quito-Lago Agrio road,

CENTROLENID FROGS OF ECUADOR 47

Provincia Napo, Ecuador, obtained on 17 October 1971, by William
E. Duellman.

Paratopotypes. —KU 143279-83, 17-18 October 1971, William E.
Duellman and Joseph T. Collins.

Diagnosis. — 1) prevomerine teeth absent; 2) bones green; 3)
parietal peritoneum white; visceral peritoneum clear; 4) color in life
dark green with diffuse black spots and pale yellow flecks; in pre-
servative, gray with dark and pale flecks; 5) webbing between outer
fingers UI2K-2%IV; 6) webbing on foot indented 11-2 111-2^1111-
2)3lV2/3-IV; 7) snout truncate in dorsal and lateral profiles; 8) dorsal
skin shagreened with elevated warts corresponding to pale flecks;
9) unscalloped fringe on outer edge of hand, forearm, and foot;
fringe with low scallops on tarsus; 10) humeral spine present in
males; 11) tympanum entirely visible, directed posterolaterally with
no dorsal inclination.

CentroIeneJIa pipilata resembles two other Ecuadorian species in
coloration, but both grandisonae and peristicta differ in having
ulnar and tarsal tubercles instead of folds and in having round, in-
stead of truncate, snouts. Two other Ecuadorian species have ulnar
and tarsal folds; pellucida is white in preservative and in life lacks
markings and a white peritoneum, whereas resplendens is a much
larger, fringe-limbed frog. Males of both of those species lack hu-
meral spines.

Description. — Adults moderately small; snout-vent length 19.5-
22.9 mm (x=21.5, N=10) in males, 21.8-22.1 mm (x=21.9, N=2) in
females. Head noticeably wider than body; width of head 32.9-38.9
percent (x=35.2, N=12) of snout- vent length; snout extremely short,
truncate in dorsal and lateral profiles; canthus round; loreal region
barely concave; lips not flared; nostrils three-fourths distance from
eye to tip of snout, protuberant anterodorsolaterally; internarial area
slightly depressed. Eye large, protuberant, directed more anteriorly
than laterally. Supratympanic fold weak; tympanum entirely visible,
directed posterolaterally with no dorsal inclination. Prevomerine
dentigerous processes and teeth absent; choanae small, ovoid; tongue
cordiform, shallowly notched behind, barely free posteriorly; vocal
slits extending from midlateral base of tongue to angles of jaws.

Humeral spine blunt, oriented at about 30° from humerus; un-
scalloped dermal fringe on ventrolateral edge of forearm and outer
edge of hand; first finger longer than second; fourth finger notice-
ably shorter than third; lateral fringes on fingers; webbing absent
between first, second, and third fingers; webbing formula for outer
fingers ni(2"-2/2)-(2-2+)IV; webbing incised; discs broad, rounded;
subarticular tubercles moderately large, subconical, simple; super-

48 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

numerary tubercles small, present on proximal segments of all digits;
palmar tubercle large, ovoid; nuptial excrescences absent. Hind
limbs moderately slender; length of tibia 55.7-61.9 percent (x=59.7,
N=12) of snout- vent length; scalloped dermal fold on ventrolateral
edge of tarsus; unscalloped dermal fold on outer edge of foot; inner
metatarsal tubercle large, ovoid; outer metatarsal tubercle small,
round; subarticular tubercles small, round; supernumerary tubercles
minute, present on proximal segments of digits; toes about two-
thirds webbed; webbing formula l{l-VA)-{2-2')lll-i2-2ys)lU{l-l}i)-
(2"-2}2)IV(2M-2/'2)-( 1-1^2 )V; discs on toes slightly smaller than those
on fingers.

Skin on dorsal surfaces shagreened with elevated warts corre-
sponding to white spots; skin on belly and ventral surfaces of thighs
granular; other surfaces smooth; anal opening directed posteroven-
trally at midlevel of thighs; many small and two large tubercles
below anus.

Color in preservative: all dorsal surfaces lavender with dark
spots and white dots; ventral surfaces cream.

Color in life: dorsum dark green with diffuse black flecks and
pale yellow flecks; flecks on side of head silvery white; chest white;
heart not visible; other ventral surfaces pale green; discs pale yel-
low; bones green; iris pale bronze with black reticulations.

Distribution. — In Ecuador, C. pipilata is known from two locali-
ties ( 1700 and 1740 m ) on the Amazonian slopes of the Cordillera
Oriental of the Andes (Fig. S).

Remarks. — All individuals were found at night on vegetation
along cascading mountain streams in cloud forest. Both females
were in amplexus; one deposited a clutch of 18 eggs having clear
jefly and pale green yolks. At the Rio Azuela, pipilata was found in
sympatry with anomala, megacheira, peUucida, and siren, and at
16.5 km north-northeast of Santa Rosa, it was found with audax,
megacheira, and siren.

At first we thought our specimens might be Centrolenella johnelsi
Cochran and Coin, but the much larger size of johnelsi ( 6 29.3 mm)
and certain structural differences negated this assignment. Further-
more, johnelsi is known only from San Pedro, Departamento An-
tioquia, in northern Colombia.

See the account of megacheira for a description of the type
locality.

Etymology. — The specific name is an adjectival derivative of the
Latin verb pipila, meaning to peep, and refers to the characteristic
call of this and many other centrolenid frogs.

CENTROLENID FROGS OF ECUADOR 49

Centrolenella prosoblepon (Boettger)
Plate 2F

Hyla prosoblepon Boettger, 1892:45 [Syntypes.— SMF 3756 and ZMB 28019

from "Plantago Cairo" (La Junta), near Limon, Provincia Limon, Costa

Rica].
Hyella puncticrus Boulenger, 1896:431 [Syntypes.— BHNH 96.10.8.70-71 from

La Palma, Provincia San Jose, Costa Rica].
Hyla parahamhae Boulenger, 1898:125 [Holotype.— BMNH 98.4.28.163 from

Paramlja, Provincia Iml^abura, Ecuador]. New synomym.
Ccntrolene prosoblepon — Noble, 1924:66.
Centrolene paramhae (emendation) — Dunn, 1933:73.
Cochranellc panimbac — Taylor, 1951:35.
Cochranella parabambac — Taylor, 1951:35.
Centrolenella prosoblepon — Coin, 1964:5.
Centrolenella parahamhae — Coin, 1964:6.

Diagnosis. — 1) prevomerine teeth 0-4; 2) bones green; 3) pari-
etal peritoneum white; visceral peritoneum clear; 4) color in life
green, usually with black dots; in preservative, lavender, usually
with dark lavender dots; 5) webbing between outer fingers III2-
mV; 6) webbing on foot I1-2II1-2 III1-2IV2 -IV; 7) snout round
in dorsal view, truncate in lateral profile; 8) dorsal .skin shagreened;
9) arms and legs lacking dermal folds; 10) hum(>ral spine present in
males; 11) lower two-thirds of tympanum visible, directed dorso-
laterally with posterior inclination.

This species most closely resembles megacheira and griffitlisi,
both of which have less webbing and lack humeral spines in males.
Centrolenella megacheira further differs from prosoblepon by hav-
ing much larger hands and pustular, instead of shagreened, skin on
the dorsum. Unspotted prosoblepon are colored like buckletji, which
also has humeral spines in males, but buckleiji has less webbing, an
inclined snout, and only the lower one-fourth of the tympanum
visible.

Description. — Adults large, snout-vent length 21.7-25.6 mm
(x=24.1, N=5) in males, 25.4-27.2 mm (x=26.4, N=.3) in females.
Head slightly wider than body; width of head 33.1-35.6 percent
(x=34.1, N=8) of snout- vent length; snout moderately short, round
in dorsal view, truncate in lateral profile; canthus round; loreal re-
gion concave; lips slightly flared; nostrils three-fourths distance from
eyes to tip of snout, slightly protuberant dorsolaterally; internarial
area slightly deprc>sst>d. Eye moderately large, directed anterolater-
ally. Supratympanic fold weak; lower two-thirds of tympanum vis-
ible, directed dorsolaterally with posterior inclination. Prevomerine
dentigerous processes posteromedially inclined, narrowly separated
medially between moderately large, ovoid choanae, bearing 0-4
teeth; tongue broadly cordiform, shallowly notched behind, barely

50 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

free posteriorly; vocal slits extending from posterolateral edges of
tongue to angles of jaws.

Humeral spine pointed, oriented about 30° anterior to humerus;
ulnar fold and tubercles absent; first finger longer than second;
fourth finger slightly shorter than third; lateral fringes present on
fingers; webbing absent between first and second fingers, vestigial
between second and third; webbing formula for outer fingers III(2~-
2'^)-(l-l/3)IV; discs broad, truncate; subarticular tubercles small,
round, simple; supernumerary tubercles absent; palmar tubercle
large, ovoid, simple; nuptial excrescences absent. Hind limbs mod-
erately slender; length of tibia 51.4-57.6 percent (x=54.6, N=8) of
snout-vent length; tarsal folds and tubercles absent; inner metatarsal
tubercle small, flat, elliptical; outer metatarsal tubercle small, ovoid;
subarticular tubercles small, round; supernumerary tubercles absent;
feet about two-thirds webbed; webbing formula I( 1-1/2 )-( 2^-2^ )ni-
(2-2*)IIIl-(2 -2^)IV(2-2^)-lV; discs truncate, slighdy smaller than
those on fingers.

Skin on dorsal surfaces of head, body, forearms, and shanks
shagreened; skin on belly and posteroventral surfaces of thighs gran-
ular; other surfaces smooth; anal opening directed posteroventrally
at upper level of thighs; pair of large tubercles below anus.

Color in preservative: dorsal surfaces, exclusive of two inner
fingers and first three toes lavender, usually with numerous small
dark lavender dots; other surfaces cream.

Color in life: dorsal surfaces green with or without black flecks;
tips of digits pale yellow; chest white; heart not visible; throat
green; bones green; iris grayish white to pale bronze with brown or
gray flecks.

Distribution. — This species occurs to elevations of about 1200 m
from lower Central America ( Caribbean and Pacific slopes in Costa
Rica and Panama) southward on Pacific slopes and lowlands to
western Ecuador, where it has been taken at elevations of 220 to 800
m (Fig. 7). We have examined two specimens from the Pacific
versant of Colombia. Cochran and Coin (1970:508) listed C. para-
barnhae from Medellin, Rio Mecaya, and Serrania de Macarena,
Colombia. The last two localities are east of the Andes, so it is
doubtful if the specimens from there actually are prosoblepon.

Remarks. — The identity of Centrolenella parabambae has been
greatly confused in the literature. As noted by Savage (1967:330):
"Apparently the name [C. parabambae] has been applied to any
population of small, uniformly lavender (in preservative) centro-
lenids with vomerine teeth from Panama, Colombia, and Ecuador."
Savage noted that Panamanian frogs referred to '^parambae" by

CENTROLENID FROGS OF ECUADOR 51

Dunn (1933) are spinosa. Two characteristics of parabambae (as
given in the type description by Boulcnger, 1898) are responsible
for the confusion: 1) absence of a humeral spine; the holotype is a
female; 2) uniformly lavender dorsum, except for dark flecks on
hind limbs.

We compared a nearly uniformly lavender female (KU 121055)
from Santo Domingo de los Colorados, Ecuador, with the holotype
and noted that structurally the two frogs were identical; the holo-
type has small indefinite dark flecks on the thighs, shanks, and feet,
whereas KU 121055 has one fleck on one shank and two and three
flecks on the feet. Subsequent comparison of KU 121055 with two
females and five males of prosoblepon from Ecuador revealed that
the only difterences were in the numbers and disposition of dark
flecks on the dorsum. The numbers of flecks in the nine specimens
are (average value for limbs; means in parentheses after ranges):
head 0-38 (11.0), body 0-58 (23.1), forearm 0-11 (6.1), thigh 0-10
(5.2), shank 0-23 (9.4), foot 2-11 (5.1). In those specimens having
many flecks on the body ( 49 and 58 ) , the flecks are present over the
entire dorsum; in those having fewer flecks (19, 21, 24, 32), the
flecks are present only latc^rally. One individual has only five flecks
(all laterally), and two lack flecks on the body. Two individuals
have three and five small white flecks dorsally.

One specimen from Colombia has reduced flecking, whereas the
other is normal. Cursory examination of series of prosoblepon from
Costa Rica and Panama re\'ealed considerable variation in the num-
ber of flecks on the dorsum, but no specimens lacked flecks. Taylor
(1952:772) mentioned a specimen from Moravia, Costa Rica in which
the spotting was ". . . sparse and confined largely to limbs and pos-
terior part of dorsum." Tlie apparent continuum of variation in
dorsal pattern and the absence of structural features to distinguish
flecked and plain indi\'iduals necessitates the placement of Centro-
lenella parabambae (Boulenger, 1898) in the synonymy of Centro-
lenella prosoblepon (Boettger, 1892).

Three of our specimens were found on low vegetation in forest;
four others were calling from vegetation over a cascading mountain
stream at Balzapamba in July 1970.

Tadpoles agreeing with the description of prosoblepon by Star-
rett (1960:12) were collected late at night as they swam on the
bottom of silt-bottomed pools in streams at Balzapamba.

Centrolenella resplendens new species
Plate 2C
Holotype— KU 118053, an adult male, 27.3 mm, from Santa

52 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Cecilia, 340 m, Piovincia Napo, Ecuador, obtained on 14 June 1967,
by John D. Lynch.

Diagnosis. — 1) prevomerine teeth 2 on low processes; 2) bones
white (?); 3) parietal and visceral peritonea white; 4) color in life
dark green with white to bluish green flecks; in preservative, dull
lavender with white flecks; 5) webbing between outer fingers
III2--1IV; 6) webbing on foot Il-l73lI0-2^IIIl-2IV2--0V; 7) snout
round in dorsal view and gradually inclined anteroventrally in lat-
eral view; 8 ) dorsal skin shagreened with elevated warts correspond-
ing to white flecks; 9) scalloped dermal fold on outer edge of hand,
forearm, and elbow, on heel, tarsus, and foot; U-shaped anal fold;
10) humeral spine absent in males; 11) lower four-fifths of tym-
panum visible, directed dorsolaterally.

Centrolenella resplendens is unlike any other Centrolenella
known from South America; eiiknemos and pidverata from Central
America are like resplendens in having scalloped dermal folds on
the limbs. Centrolenella pidverata differs from the other two species
by having the heart visible in life, dorsum white in preservative,
snout short and slightly inclined anteroventrally, and dorsal skin
uniformly granular. In contrast, in euknemos and resplendens the
heart is not visible in life, the dorsum is lavender in preservative, the
snout is long and gradually inclined anteroventrally, and the dorsal
skin is shagreened with scattered elevations. Centrolenella re-
splendens differs from euknemos by having a deeper snout that is
round instead of subacuminate in dorsal view, more extensive anal
folds, and more extensive webbing. The modal webbing formulae
in euknemos are: hand— I3-3II3-3?niI2-2IV, foot I1-1}^II1-2MII1-
2IV2-1V (Savage and Starrett (1967:606).

Description. — Adult male large, 27.3 mm in snout-vent length;
females not known. Head slightly wider than body; width of head
34. (S percent of snout-vent length; snout long, round in dorsal view,
gradually sloping from nostrils to tip in lateral profile ( Fig. 1 ) ;
canthus round; loreal region barely concave; lips slightly ffared; nos-
trils about two-thirds distance from eyes to tip of snout, not pro-
tuberant, directed dorsolaterally; internarial area barely depressed.
Eye large, directed more laterally than anteriorly. Supratympanic
fold absent; tympanum directed dorsolaterally. Prevomerine den-
tigerous processes low, short, transverse between choanae, each
bearing two teeth; choanae large; longitudinally rectangular; tongue
ovoid, barely free posteriorly; vocal slits extending from postero-
lateral base of tongue to angles of jaws.

Humeral spine absent; scalloped dermal fold around elbow,
along ventrolateral edge of forearm and outer edge of fourth finger;

CENTROLENID FROGS OF ECUADOR 53

first finger longer than second; fourth finger slightly shorter than
third; lateral fringes on fingers; webbing absent between first and
second fingers, vestigial between second and third; \\'ebbing formula
for outer fingers III2-1IV; discs large, nearly round; subarticular
tubercles small, round, simple; supernumerary tubercles absent;
palmar tubercle simple, large, ovoid; nuptial excrescences absent
(Fig. 3). Hind limbs moderately robust; length of tibia 53.5 percent
of snout-vent length; scalloped fold around heel, along ventrolateral
edge of foot and outer edge of fifth toe; inner metatarsal tubercle
large, triangular; outer metatarsal tubercle absent; subarticular tu-
bercles small, round; supernumerary tubercles absent; feet nearly
fully webbed; webbing formula Il-l73lI0-2-IIIl-2IV2 -OV; discs on
toes round, slightly smaller than those on fingers (Fig. 4).

Skin on dorsal surfaces of head, body, and limbs, and lateral
surface of head and flanks shagreened with small tubercles corre-
sponding to white spots; belly and proximal posteroventral surfaces
of thighs granular; other surfaces smooth; anal opening directed
posteriorly at upper level of thighs, bordered laterally and ventrally
by fleshy tuberculate U-shaped fold; two conical tubercles below
fold (Fig. 2).

Color in preservati\'(>: head, flanks, and dorsal surfaces of body
and limbs (except tluiml) and toes 1-3) dull lavender with many
small, white spots; venter and posterior surfaces of thighs cream.

Color in life: dorsum dark green with white to bluish green
spots; fringes on forearm and foot and fold around anus white;
venter yellow; iris gray.

Distribution. — Known only from the type locality and Santa
Maria de Sucumbios, Colombia, about 40 km NNW of Santa Cecilia
(Fig. 6).

Remarks. — The holotype was perched on a leaf of a bush about
30 cm ab()\'e the ground during light rain at night in primary rain-
forest. Subsequent work at Santa Cecilia from 1967 through 1972
has resulted in the collection of thousands of frogs, but no additional
specimens of resplendens have been found.

The Colombian specimen (AMNH 88083) is a juvenile having a
snout-vent length of 16.5 mm. The white dermal folds and flecks are
essentially the same as those in the holotype, but the webbing, as
topical of juveniles, is less extensive.

EtipnoJogy. — The specific name is derived from the Latin verb
resplendo meaning to glitter, and is used in allusion to the jewel-like
appearance of the li\'ing frog.

54 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Centrolenella siren new species
Plate IE

Holotype. — KU 146610, an adult male, 20.7 mm, from a small
tributary of the Rio Salado, about 1 km upstream from the Rio Coca,
1410 m, Provincia Napo, Ecuador, one of a series obtained on 7
April 1972, by William E. Duellman and John E. Simmons.

Paratopotypes. — KU 146611-23, same date and collectors.

Diagnosis. — 1) prevomerine teeth 0-2; 2) bones green; 3) pari-
etal peritoneum white; visceral peritoneum clear; 4) color in life
green with yellow flecks; in preservative, lavender with white flecks;
5) webbing between outer fingers 111273-2)21 V; 6) webbing on foot
I2-2)nil)^-2;3lIIlM-3 IV3-2 V; 7) snout truncate in dorsal and lateral
profiles; 8) dorsal skin shagreened; 9) arms and legs lacking dermal
folds; 10) humeral spine absent in males; 11) lower four-fifths of
tympanum visible, directed posterolaterally with slight dorsal in-
clination.

CentroJenella siren is like favopnnctata and miclas in having a
lavender dorsum with white flecks (in life, green with gold flecks)
with no black flecks, but it differs from both of those species by
having much less webbing on the hands and feet and a more promi-
nent tympanum oriented posterolaterally, instead of dorsolaterally.
Centrolenella fiavopunctata further differs from siren by having a
rounded, instead of truncate, snout.

Description. — Adults moderately small; snout-vent length 19.8-
22.0 mm (x=20.8, N=14) in males; females unknown. Head much
wider than body; width of head 30.0-37.0 percent (x=34.7, N=14) of
snout-vent length; snout extremely short, truncate in dorsal and lat-
eral views; canthus round; loreal region barely concave; lips not
flared; nostrils nearly terminal on snout, not protuberant, directed
laterally; internarial area flat. Eye moderately large, directed antero-
laterally. Supratympanic fold absent; lower four-fifths of tympanum
visible, directed posterolaterally with slight dorsal inclination. Pre-
vomerine dentigerous processes small, low, widely separated, be-
tween small round choanae, bearing 0-2 teeth; tongue round, barely
free posteriorly; vocal slits extending from midlateral base of tongue
to angles of jaws.

Humeral spine absent; ulnar fold and tubercles absent; first fin-
ger about equal in length to second; fourth finger noticeably shorter
than third; fringes absent on fingers; webbing absent between first
and second fingers, vestigial between second and third; webbing
formula for outer fingers III (232-273) -(2)4-2)2) IV; discs truncate; sub-
articular tubercles small, round, simple; supernumerary tubercles
absent; palmar tubercle large, elliptical; nuptial excrescences absent.

CENTROLENID FROGS OF ECUADOR 55

Hind limbs slender; length of tibia 54.8-60.1 percent (x=57.3, N=14)
of snout-vent length; tarsal folds and tubercles absent; inner meta-
tarsal tubercle small, ovoid; outer metatarsal tubercle absent; sub-
articular tubercles small, round; supernumerary tubercles absent;
feet about one-half v^^ebbed; webbing formula I2-(2+-2K)II(l?4-lJ2)-
{2}i-2%)UlVA-{2%-3)lV3-{l%-2)V; discs on toes smaller and more
nearly round than those on fingers.

Skin on dorsal surfaces shagreened; skin on belly and ventral
surfaces of thighs granular; other surfaces smooth; anal opening di-
rected posteriorly at upper level of thighs; pair of large tubercles
below anus ( Fig. 2 ) .

Color in preservative: dorsum of head, body, and limbs lavender
with many small, white flecks on head and body; other surfaces pale
cream.

Color in life: dorsum green with gold flecks; fingers and toes
pale yellow; chest white; heart not visible; other ventral surfaces
pale dull green; bones green; iris pale bronze with fine black re-
ticulations.

Distrihiition. — CentroIeneUa siren occurs in cloud forest on the
Amazonian slopes of the Andes, where it is known from elevations
of 1410-1740 m (Fig. 9). CentroIeneUa siren occurs in sympatr)^
with anomala, audax, megacheira, peUucida, and pi))ilata.

Remarks. — All individuals were found at night on low vegetation
along small mountain streams in cloud forest. The type locality is a
small stream at the south edge of the Papallacta-Lago Agrio road
about 300 m west of the bridge across the Rio Salado and approxi-
mately 1 km up the Rio Salado from its confluence with the Rio
Coca.

Etymo]o<i,y. — The specific name refers to the nymphs of Classical
mythology, who by their sweet singing enticed seafarers to destruc-
tion; the name is used here in allusion to the fact that the call of
these small frogs entices collectors to the nocturnal perils of streams.

DISCUSSION

Savage ( 1967 ) recognized three species groups of CentroIeneUa
in Central America. He characterized the feischmanni group as
having white bones in life, a colorless parietal peritoneum, a white
visceral peritoneum, a white ground color in preservative, and in
lacking prevomerine teeth and dentigerous processes. The pro-
soblepon group was characterized as having green bones in life, a
white (opaque) parietal peritoneum, a colorless visceral peritoneum,
lavender ground color in preservative, and prevomerine teeth and

56 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

dentigerous processes. He recognized a third group for the some-
what annectant piilverata.

Three Ecuadorian species (feischmanni, munozonwi, and pellu-
cida) clearly fit into Savage's fleiscJimanni group. With the excep-
tions of onomala, tnedcmi, oceUifera, and resplendens, the remaining
Ecuadorian CentroleneUa agree with Savage's definition of the pro-
soblepon group. The presence of prevomerine teeth and processes
is variable in the Ecuadorian representatives of the prosohlepon
group (sensii Savage) but is usually constant within species. How-
ever, a single specimen of cochranae and one of prosohlepon lack
teeth and processes, whereas all other specimens of these species
examined have them.

Except for the absence of prevomerine teeth and processes,
biickleyi, <i,rondisonae, griffithsi, me<i,acheira, peristicta, and pipilata
agree with the definition of the Central American prosohlepon
group. However, the agreement is weakened by the pale-green
(contrasted with green) bones of grandisonae, griffithsi, and peristicta
(a character-state of the pulverata group). Savage characterized the
pulverota group as having pale green bones, a colorless parietal
peritoneum (heart visible), opaque visceral peritoneum, yellowish-
white ground color in preservative, and prevomerine teeth and
processes. On examining the holotype of pulverata (ZMB 7842) in
1969, Ducllman noted the ground color was lavender. Aside from
having pale green bones, grandisonae, griffithsi, and peristicta do
not agree with the definition of the pulverata group. We know of
no Ecuadorian CentroleneUa assignable to the pulverata group. The
Venezuelan CentroleneUa antisthenesi may be allied with pulverata;
antisthenesi has a clear parietal peritoneum, opaque visceral peri-
toneum, prevomerine teeth, lavender dorsum, green bones, and sub-
anal tubercles and lacks a humeral spine.

Three Ecuadorian species ( and possibly the Colombian medemi)
depart from the fleischmanni-prosohlepon arrangement. Centro-
leneUa anomala is prosohlepon-Wke, except in lacking prevomerine
teeth, having white bones, and in lacking a lavender ground color.
The opaque parietal peritoneum and brown ground color exclude it
from the homogeneous fleiscJimanni group. CentroleneUa oceUifera
and resplendens presumably have Vv^hite bones (green bones were
not recorded in their color descriptions in life). Both have prevo-
merine teeth and low dentigerous processes, an opaque parietal
peritoneum, and lavender ground color. CentroleneUa resplendens
differs further in having white visceral peritoneum (a character-
state of the fleischmanni and pulverata groups). The combination

CENTROLENID FROGS OF ECUADOR 57

of characteristics cited above does not recommend them for any of
Savage's (1967) groups.

We noted the absence of hirge subanal tubercles in anomala,
fleischmanni, medemi, munozorum, ocellifera, and peUucida, and
their presence in the remaining Ecuadorian CentwJeneUa. Subanal
tubercles are absent in most Central American centrolenids; we
found them in ilex, prosoblepon, and pulverata and in antioqtiiemis
(Colombia) and antisthenesi (Venezuela). The absence of subanal
tubercles in the frogs of the fleischmanni group and in anomala,
medemi, and ocellifera (enigmatic in having white bones) thus may
not be significant. Tubercles are present in resplendens (also enig-
matic in presumably having white bones) but these may not be
homologous to those in the frogs lacking the elaborate post- and
para-anal ornamentation of resplendens.

Humeral spines are not known in any frog of the fleischmanni
group. Coin's (1964) and Savage's (1967) conclusion that presence
or absence of the spines cannot be used to support generic sepa-
ration of Centrolenella and Cochranella is substantiated here in that
an otherwise "normal" firiffithsi has humeral spines. This specimen
may prove to be distinct from <iriffithsi but at present no features
other than the humeral spine support that argument. Accordingh',
the presence of humeral spines possibly may be intraspecifically
variable, at least in iiriffithsi. Small spines occur on the humeri of
medemi but not in anomala, ocellifera, or resplendens (species with
presumed or known white bones). The presence of humeral spines
in aiidax, huckleyi, grandisonae, medemi, peristicta, pipilata, and
prosoblepon does not suggest to us that these frogs are more closely
related to one another than they may be to species lacking humeral
spines. This assemblage seems to be heterogeneous: three have pre-
vomerine teeth and processes, two have pale green as compared to
green bones, and one lacks subanal tubercles.

Centrolenella antioquiensis and resplendes differ from all other
known Centrolenella by having opaque parietal and visceral peri-
tonea. Otherwise they share few characters; antioquiensis has a
humeral spine and green bones (absent and white in resplendens)
and lacks dermal ornamentation and prevomerine teeth (both pres-
ent in resplendens) .

The Ecuadorian Centrolenella thus far described do not readily
fit into the arrangement of Central American forms proposed by
Savage (1967). Inspection of Taylor and Cochran's (1953) account
of southeastern Brasilian centrolenids reveals comparable lack of fit
with Savage's three groups in Central America. Because we antic-
ipate the discovery of many undescribed Centrolenella in north-

58 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

western South America, at this time we prefer not to alter Savage's
groupings or define new species groups based on our study of Ecua-
dorian species. We offer only the following comments on putative
relationships of the 19 species, as described below.

Centrolenella fleischtnanni, munozontm, and peUucida constitute
a well-circumscribed group of species having white bones and a
colorless parietal peritoneum ( heart visible ) in life, a white visceral
peritoneum, a white ground color in preservative, and in lacking
distinct canthi, subanal tubercles, humeral spines, and prevomerine
teeth and processes.

Centrolenella audax, cochranae, flavopunctata, midas, prosoble-
pon, and siren constitute a group having green bones and an opaque
parietal peritoneum ( heart not visible ) in life, a clear visceral peri-
toneum, a very pale to dark lavender ground color in preservative,
and in having distinct canthi, subanal tubercles, and prevomerine
teeth and processes. Two of these (audax and prosoblepon) have
humeral spines.

Centrolenella buckleyi, grandisonae, griffithsi, megacheira, peri-
sticta, and pipilata fall into a somewhat heterogeneous group agree-
ing in most respects with that listed above, except in the uniform
absence of prevomerine teeth and processes. The bones are pale
green in grandisonae, griffithsi, and peristicta; griffithsi and mega-
cheira lack humeral spines.

Centrolenella anomala differs markedly from all other centro-
lenids in the combination of white bones and an opaque parietal
peritoneum in life, clear visceral peritoneum, a brown ground color
in life and in preservative, distinct canthi, and in lacking prevo-
merine teeth and processes, subanal tubercles, and humeral spines.
Its ocellated color pattern on a shagreened dorsum with warts and
the reduced webbing of the fingers lend it a striking similarity to
cochranae; the similarity is out-weighed by the numerous differences
between the two species.

Centrolenella ocellifera also differs markedly from other centro-
lenids. The presumably white bones in life, opaque parietal peri-
toneum, clear visceral peritoneum, and absence of subanal tubercles,
coupled with the pale lavender ground color in preservative, distinct
canthi, prevomerine teeth and processes, ocellated color pattern, and
the absence of humeral spines suggests some relationship with ano-
mala or with the gn^up of species listed above, which included
audax and cochranae.

Centrolenella resplendens appears to be trenchantly different
from other Ecuadorian Centrolenella chiefly because of its dermal
ornamentation ( scalloped limb fringes, para- and postanal ornamen-

CENTROLENID FROGS OF ECUADOR

59

3000

2000

1000
M.

I

>ae

IS
5 -8

!-!v

PACIFIC

g <a i^ C3

.§

t)

I

m^

AMAZONIAN

Si 5

'V.'.'i

Paramo and
Subporamo

Cloud
Forest

Montane
Rainforest

Lowland
Rainforest

Fig. 11. Altitudinal distributions of Centrolenella in Ecuador; vertical line
= continental divide.

tation, and enameled warts ) . Also it differs from other known cen-
trolenids by the following combination of characters: white bones
in life, opaque parietal peritoneum, white visceral peritoneum, lav-
ender ground color in preservative, and prevomerine teeth.

Centrolenella medemi differs from all Ecuadorian centrolenids
in having smooth skin on the dorsum and the decidedly non-centro-
lenid color pattern. We do not know the color of the bones in living
individuals. Humeral spines are present, although small, and sub-
anal tubercles are lacking. The fully-webbed digits, elongate hind-
limbs, and completely concealed tympana, coupled with skin texture
and color pattern suggested the possibilit\' that the frog was not a
centrolenid but a hylid of the CnjptobatmcJms and Stephania group;
however, dissection of the tarsus revealed that the tarsal bones are
fused, as in all centrolenids but not in hvlids.

Our knowledge of the distribution of Centrolenella in Ecuador
is limited; even so, some patterns consistent with other groups of
frogs ( eleutherodactylines, dendobatids, and hylids) are apparent.
We can divide the Ecuadorian Centrolenella into six ecogeographic
assemblages. Each is associated with definite vegetation fonnations
and geographic regions (Fig. 11). These can be summarized as
follows :

Loiulaml Amazonian rainforest. — Four species (medemi, midas,
munozorum, and resplendens) occur only at elevations of less than
600 m in Amazonian rainforest, where they apparently are limited
to breeding along small, sometimes intermittent, streams having a
low gradient. The apparent restricted ranges of the species probably
is a reflection of inadequate collecting; most likely the species will
eventually be found in more distant areas in the upper Amazon
Basin.

60 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Montane rainforest, Amazonian slopes. — Two species (cochranae
and fJavopunctata) inhabit elevations of 700-1300 m on the lower
eastern slopes of the Andes. In this area they live along streams with
steep gradients. Warm temperatures and high amomits of rainfall
prevail throughout the year.

Cloud forest, Amazonian slopes. — This is the richest area in
Ecuador for centrolenids; six species (anomala, aiidax, megacheira,
pellucida, pipilata, and siren) occur at elevations of 1400-1800 m on
the Amazonian slopes of the Andes. This area of cool cloud forest
daily bathed in mist has luxuriant vegetatitm and many cascading
streams.

Paramo and subparamo. — This cool, windy region has been
greatly disturbed by man. Some areas are noticeably drier than
others. Throughout the paramo precipitation is lower than in cloud
forests, but the low temperatures result in low rates of evaporation.
A single species (])uckleyi) occurs in wet meadows and terrestrial
bromeliads in the interandean valleys and on high Amazonian and
Pacific slopes between 2500 and 3000 m. Records from lower ele-
vations on the Andean slopes are questionable.

Cloud forest. Pacific slopes. — Five species (feischmanni, gran-
disonae, griffithsi, ocellifera, and peristicta) occur at elevations of
1400-2200 m on the Pacific slopes of the Andes, where the habitat
is like that at similar elevations on the Amazonian slopes.

Pacific lowlands. — Two species (fleischmanni and prosoblepon)
are known from mesic forests on the Pacific lowlands, where rainfall
is far more seasonal than in the Amazon lowlands. Centrolenella
fleischmanni occurs up to 1460 m, and prosoblepon goes up to at
least 800 m.

Excluding the high Andean buckleyi, we find 12 species on the
Amazonian slopes and lowlands and only six species on the Pacific
slopes and lowlands. In addition to the fewer species on the Pacific
slopes, there is a difference in altitudinal assemblages. The two spe-
cies on the Pacific lowlands also inhabit the lower Andean slopes; in
the Amazonian region, two distinctive assemblages occur within the
same elevational range — one group in the lowlands, and one on the
slopes. No species is known from both Pacific and Amazonian slopes
or lowlands, except buckleyi which is at high elevations.

Greater species richness in cloud forest results in higher numbers
of sympatric species there. At Tandapi at an elevation of 1460 m
on the Pacific slopes, there are five species (fleischmanni, grandi-
sonae, griffithsi, ocellifera, and peristicta). At the Rio Azuela at an
elevation of 1740 m on the Pacific slopes, five species (anomala,
megacheira, pellucida, pipilata, and siren) were found along one

CENTROLENID FROGS OF ECUADOR 61

small rivulet. Three species (midos, munozortim, and resplendens)
have been found at one locality, Santa Cecilia, at an elevation of
340 m in the Amazon Basin. Two species (fleischmanni and pro-
soblepon) occur in sympatry at the Rio Palenque at an elevation of
220 m on the Pacific lowlands.

These limited data probably are only suggestive of the true eco-
logical and altitudinal distributions of these small frogs. Hopefully
this discussion will sen'e as a basis for future investigations and not
be destined to be the only commentary on them before they and
their habitat are eliminated bv human activitv'.

SUMMARY

Our collections from Ecuador contain many centrolenid frogs.
Our taxonomic conclusions are based on the examination of holo-
types of all of the Ecuadorian centrolenids, study of series of speci-
mens, and obscrv^ations of the frogs in the field.

We recognize 19 species in Ecuador; one of these, Centrolene
geckoideiim, is not treated in this paper. One Colombian species,
CentroJencUa medemi, is treated, because it probably occurs in east-
ern Ecuador. Eleven new species are named; se\'en (CentvoleneUa
anomala, audax, favopunctata, meiiacheira, pelhicida, pipikita, and
siren) are from the Amazonian slopes of the Andes; three (Centro-
leneUa midas, munozorum, resplendens) are from the Amazon Basin,
and one (Centrolenelhi peristicta) is from the Pacific slopes of the
Andes. Centrolenella parabambae (Boulenger, 1898) and Centro-
lenella pefersi (Coin, 1961) are placed in the synonymies of Cen-
trolenella prosoblepon ( Boettger, 1892) and Centrolenella fleisch-
tnanni (Boettger, 1893), respectively.

Savage (1967) placed the Central American species of Centro-
lenella in three groups: 1) fleischmanni, 2) prosoblepon, and 3)
pulverata. The distinction of these three groups in Ecuador is not
clear. Three species (fleischmanni, munozorum, peUucida) clearly
belong to the fleischmanni group. Centrolenella audax, buckleyi,
cochranae, flavopunctata, <i,randisonae, <iriffithsi, megacheira, midas,
peristicta, pipUata, and siren can be placed in the prosoblepon
group, redefined to include species with or without prevomerine
teeth. Centrolenella anomala, ocellifera, and resplendens cannot be
placed in any of the defined groups.

The maximum di\'ersity of Centrolenella in Ecuador is in the
cloud forests. Six species (five occur s\'mpatrically ) inhabit the
cloud forests on the Amazonian slopes of the Andes. Five sxmpatric
species occur in the cloud forests on the Pacific slopes of the Andes.
In other regions there are fewer species: Amazonian lowlands (4),

62 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

lower Amazonian slopes of the Andes (2), Pacific lowlands (2).
Only Centwlenella buckleiji occurs in the high Andes and inter-
andean valleys.

We provide the first records of four species from Colombia:
Centrolenella griffithsi, ilex, prosoblepon, and resplendens.

RESUMEN

Nuestros colecciones del Ecuador contienen muchas ranas de la
familia Ccntrolenidae. Nuestros conclusiones taxonomicos son ba-
sada en el examinacion de los holotipos de todos los centrolenidos
ecuatorianos, estudio de serie de ejemplares, y observaciones de las
ranas en sus ambientes naturales.

Se reconocen 19 especies de centrolenidos en Ecuador; una,
Centrolene geckoideiim, no esta tratado en esta publicacion. Un
especie colombiano, CentroJenelh medemi, esta tratado, porque
probablamente se occuren en el oriente del Ecuador. Once especies
nuevas se nombran; siete (Centroleiiella anomaJa, atidax, favopiinc-
tata, megacheira, peUucida, pipilota, siren) se encuentran en las
laderas amazonicas de los Andes, tres (Centrolenella midas, muno-
zoriini, resplendens) son de la cuenca amazonica, y una (Centro-
lenella peristicta) es de las laderas de Pacifico de los Andes. Cen-
trolenella parahamhae (Boulcnger, 189(S) y Centrolenella petersi
(Coin, 1961) son colocadas en las sinonimias de Centrolenella pro-
soblepon (Boettger, 1892) y Centrolenella fleischmanni (Boettger,
1S93), respectivamente.

Savage (1967) puse las especies centroamericanas de Centro-
lenella en tres grupos: 1) fleischmanni, 2) prosoblepon, and 3)
pulverata. La distincion de estos tres grupos en los centrolenidos
ecuatorianos no esta lucida. Tres especies (fleischmanni, muno-
zorum, pellucida) claramente pertenecen al grupo fleischmanni.
Centrolenella audax, biickleyi, cochranae, flavopiinctata, grandi-
sonae, griffithsi, megacheira, midas, peristicta, pipilata prosoblepon,
y siren se pueden colocar en un grupo prosoblepon redefinido para
incluir especies con los dientes prevomerianos presentes o ausentes.
Centrolenella anomala, ocellifera, y resplendens no se pueden colocar
en los grupos definidos.

La diversidad maxima en Centrolenella en Ecuador esta en el
bosque neblino. Seis especies (cinco occuren simpatricamente)
habitan los bysques neblinos en las laderas amazonicas de los Andes.
Cinco especies simpatricos occuren en los bosques neblinos en las
laderas de Pacifico de los Andes. En otras regiones hay menos
especies: Tierras bajas amazonicas (4), laderas bajas amazonicas

CENTROLENID FROGS OF ECUADOR 63

de los Andes (2), la costa (2). Solamente Centrolenella buckleyi
occure en los Andes altos v en los valles interandinos.

Suplimos las anotaciones primeras de cuatro especies de Colom-
bia: Centrolenella griffithsi, ilex, prosoblepon, y resplendens.

SPECIMENS EXAMINED

All specimens from Ecuador are listed first, alphabetically by
province. Specimens from other countries are listed after Ecua-
dorian ones.

Centrolenella alhotunica. — BRASIL: Sao Paulo: Paranapiacaba, KU 74310-
11.

Centrolenella anomala. — Napo: Rio Azuela, 1700 m, KU 143299.

Centrolenella antioquiensis. — COLOMBIA: Quindio: Salento, 1900 in, KU
133466.

Centrolenella anti.sthenesi. — VENEZUELA: Aragua: Rancho Grande, 1075
ni, KU 133467-80.

Centrolenella audax. — Napo: Salto de Agua, 2.5 km NNE Rio Reventador,
1660 m, KU 146624, 146831; 16.5 km NNE Santa Rosa, 1700 m, KU 143290,
14.3292.

Centrolenella fof/r/:/c[/j.— "Paramo," AMNH 20504. Aztiay: Sinicay, 2560
m, AMNH 17464. Carchi: Tulcan, 3000 m, Kl^ 118005-8. Chimhorazo: Palla-
tanga, 1520 m, BNLNH 80.12.5.201. Imhahura: Laguna Cnicocha, 10 km W
Quiroga, 3000 m, KU 138822. Pirhincha: Llave Pongo, AMNH 20141; Macha-
chi, 2950 m, KU 14829-30. Zannmi-Chinrhipc: 13.5 km E Loja, 2800 m, KU
142648; Sabanilla, AMNH 13530. COLOMBIA: Caiica: Coconuco, 3300 m,
KU 145087; road to Pacific coast from El Tambo, 2170 m, KU 144131-2; road
to Quintana, Qncbrada Santa Tereza, 2200 m, KU 144133-4.

Centrolenella cochranae. — Napo: south slope Cordillera del Due, 1150 m,
KU 1232] 6-8. Pastaza: Abitagua, 8 km NW Mera, 1300 m, KU 121033-35.
Tun'Aurahua: El Topo, 1220 m, BNLNH 1912.11.1.68; 11 km E Rio Negro,
1170 m, KU 146605.

Centrolenella eukncmos. — PANAMA: Darien: Laguna, 820 m, KU 77534;
Rio Jaque, 1.5 km aboxe Rio Imamado, 50 m, KU 116439-41. Panama: south
slope Cerro La Campana, 820 m, KU 116436. San Bias: Camp Summit, 300-
400 m, KU 116437-8.

Centrolenella eunjgnatha.—BEASlL: Guanahara: Tijuca, KU 93220-4.

Centrolenella flavopunctata. — Napo: Bermejo No. 4 (well site), 15 km
ENE Umbaqui, 720 m, KU 123224; San Jose Abajo, AMNH 22187. Pastaza:
Mera, 1140 m, KU 121041-6, 121048-51; Rio Alpayacu, 1 km E Mera, KU
121047; 13 km WSW Puyo, 1000 m, TCWC 24032.

Centrolenella fleischmanni. — Esmeraldas: Rio Durango, 110 m, BMNH
1902.5.27.24. Los Rios: Estacion Biologica Rio Palenque, 56 km N Que\edo, 220
m, KU 146606-8, 147580. Pichincha: Tandapi, 1460 m, KU 121052. COSTA
RICA: Caiiago: Tapanti, 1200 m, KU 65189-200. San Jose: La Palma, 1,520
m, KU 36885-95; San Jose, 1200 m, SMF 3760. MfiXICO: Oaxaca: 4.3-14.8
km N San Gabriel Mixtepec, 580-860 m, KU 137347-51; 9.1-12.7 km S Valle
Nacional, 610-790 m, KU 137352-5. PANAMA: Darien: Rio Jaque, 1.5 km
above Rio Imamado, 50 m, KU 116442-55. SURINAM: Saramacca, RMNH
4463.

Centrolenella pramlisonae. — Pichincha: Tandapi. 1460 m, KU 118036,
118047-50. COLOMBIA: Caldas: Pueblo Rico, BMNH 1910.7.11.68. Cauca:
Santa Leticia, 2000 m, KU 144129-30.

64 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Centrolenella griffithsi. — Pichincha: 4 km W Chiriboga, 2120 m, KU
142649; Las Maguinas, 2150 m, AMNH 20146; Rio Saloya, 1200 m, BMNH
1940.2.20.4; Tandapi, 1460 m, KU 118009-35, 118037-45, 118148, 121036-9,
138823-4. COLOMBIA: Caiica: road to Pacific coast from El Tambo, 2170 m,
KU 139497-8.

Centrolenella inedemi. — COLOMBIA: Putttmayo: Puerto Asis, USNM
152277.

Centrolenella megacheira. — Napo: Rio Azuela, 1740 m, KU 143273-7; 16.5
km NNE Santa Rosa, 1700 m, BMNH 1971.1854-55, CAS 135498-9, KU
143245-72, UMMZ 131668 (3).

Centrolenella midas.—Napo: Lago Agrio, 330 m, KU 125334, UMMZ
129314; Puerto Libre, 570 m, KU 123220-3; Santa Cecilia, 340 m, KU 107026,
123219, 146625, 150662-23.

Centrolenella miinozorutn. — Napo: Lago Agrio, UMMZ 129313; Santa Ce-
cilia, 340 m, KU 105251, 118054, 123225, 150620-21.

Centrolenella ocellata. — PERU: Arjacucho: Huanhauchayocc on Tambo-
Valle del Apurimac trail, LSU 25989-90. Pasco: Huancabamba, BMNH
1912.11.1.19.

Centrolenella oceUifera. — Imhahura: Paramba, 777 m, BMNH 98.5.19.3.
Pichincha: Tandapi, 1460 m, KU 118046.

Centrolenella orocostalis. — VENEZUELA: Aragua: Rancho Grande, KU
133481.

Centrolenella pellucida. — Napo: Rio Azuela, 1740 m, KU 143298.

Centrolenella peristicta. — Pichincha: Tandapi, 1460 m, KU 118051-2,
121053.

Centrolenella pipilata.—Napo: Rio Azuela, 1740 m, KU 143284-7, UMMZ
131669 (2); 16.5 km NNE Santa Rosa, 1700 m, KU 143278-83.

Centrolenella prosohlepon. — Bolivar: Balsapampa, 800 m, KU 132462-5.
Imhahura: Lita, 520 m, KU 133482-3; Paramba, 777 m, BMNH 98.4.28.163.
Los Rios: Estacion Biologica Rio Palenque, 56 km N Quevedo, 220 m, KU
146609. Pichincha: Santo Domingo de los Colorados, 600 m, KU 121054-5,
UMMZ 131671. COLOMBIA: Caiica: La Costa, El Tambo, 1200 m, KU
145085; Rio Michenque, El Tambo, 900 m, KU 145086. COSTA RICA:
Alajtiela: Cinchona, 1600 m, KU 65172-8. Heredia: La Concordia, 1900 m,
KU 65159-66. Linum: Plantage Cairo, near Limon, SMF 3756, ZMB 28019.
San Jose: La Palma, BMNH 96.10.8.70-71, KU 65180-3. PANAMA: Bocas
del Toro: Rio Claro near junction with Rio Changena, 910 m, KU 116469-78.
Chiriqui: Finca Ojo de Agua, southeast slope Cerro La Pelota, 1440 m, KU
96346-53; Finca Santa Clara, 1200 m, KU 116458-63.

Centrolenella ptdverata. — PANAMA, Chiriqui: ZMB 7842. Darien: Rio
Jaque, 1.5 km above Rio Imamado, 50 m, KU 116493.

Centrolenella resplendens. — Napo: Santa Cecilia, 340 m, KU 118053. CO-
LOMBIA: Putumaiio: Santa Maria de Sucumbios, AMNH 88083.

Centrolenella siren.— Napo: Rio Azuela, 1740 m, KU 143295-7, 143555,
UMMZ 131670 (3); Rio Salado, 1 km upstream from Rio Coca, KU 146610-23;
16.5 km NNE Santa Rosa, 1700 m, KU 143288-9, 143291, 143293-4.

Centrolenella vanzolinii.—BRASlh: Guanaharo: Tijuca, KU 93226-30.

LITERATURE CITED

BOETTGER, O.

1892. Katalog der Batrachier — Sammlung im Museum der Sencken-
bergischen Naturforschenden Gesellschaft im Frankfurt am Main.
Frankfurt, 73 pp.

1893. Fin neuer Laubfrosch aus Costa Rica. Ber. Naturf. Gesellsch.
Frankfurt, 1892-1893 ( 1893) :251-252.

CENTROLENID FROGS OF ECUADOR 65

BOULENGER, G. A.

1882. Catalogue of Batrachia Salientia s. Ecaudata in tlie collection of the
the British Museum, ed. 2. London, xvi + 503 pp.

1896. Descriptions of new batrachians collected by Mr. C. F. Under-
wood in Costa Rica. Ann. Mag. Nat. Hist. (6)18:340-342.

1898. An account of the reptiles and batrachians collected by Mr. W. F.
H. Rosenberg in western Ecuador. Proc. Zool. Soc. London, 1898
(1): 107-126.

1899. Descriptions of new batrachians in the collection of the British Mu-
seum (Natural History). Ann. Mag. Nat. Hist. (7)3:273-277.

1918. Descriptions of new South American batrachians. Ibid., 9(2):427-
433.
Cochran, D. M. and C. J. Goix

1970. Frogs of Colombia. Bull. U.S. Natl. Mus., 288:xii+655 pp.
DuELLMAN, W. E. and J- B. Tulecke

1960. The distribution, variation, and life history of the frog Cochranella
viriclissima in Mexico. Amer. Midi. Nat., 63:392-397.

Dunn, E. R.

1933. Amphibians and reptiles from El \'alle de Anton, Panama. Occ.
Papers Boston Soc. Nat. Hist., 8:65-79.
Eaton, T. H., Jr.

1958. An anatomical study of a NeotroiJical tree frog, Centrolene pro-
sohlepon (Salientia: Centrolenidae ) . Univ. Kansas Sci. Bull., 39:
459-472.
Coin, C. J.

1961. Three new centrolenid frogs from Ecuador. Zool. Anz., 166:95-104.
1964. Distril)ution and synonmy of Ceiitrolcnella jieischmanni in northern

South America. Herpetologica, 20:1-8.
1966. A new frog of the genus CentrolcncUa from Suriname. Stud. Fauna

Suriname and other Guyanas, 32:77-80.
1968. A new centrolenid frog from Guvana. Quart. Jour. Florida Acad.

Sci., 30:115-118.
Goodman, D. E., and C. J. Goin

1970. The habitat of Centrolene geckoidetini in Ecuador. Heipetologica,
26:276.

Jimenez de la Espada, M.

1872. Nuevos batracios americanos. An. Soc. Espana Hi.st. Nat., 1:85-88.
LiDTH de Jeude, T. W. van

1904. Reptiles and batrachians from Surinam. Notes Leyden Mus. 25:83-
94.
LuTz, B. and G. R. Kloss

1952. Anfibios anuros do alto Solimoes e Rio Negro. Mem. Inst. Oswaldo
Cruz, 50:625-678.
Lynch, J. D.

1971. Evolutionary relationships, osteology, and zoogeography of lepto-
dactyloid frogs. Misc. Publ. Mus. Nat. Hist. Univ. Kansas, 53:1-238.

Nieden, F.

1923. Das Tierreich. Amphibia. Anura I. Berlin, xxxii+584 pp.
Noble, G. K.

1920. Two new batrachians from Colombia. Bull. Amer. Mus. Nat. Hist.,
42:441-446.

1924. Some Neotropical batrachians preserved in the United States Na-
tional Museum with a note on secondary sexual characters of these
and otlier amphibians. Proc. Biol. Soc. Washington, 37:65-72.

1926. An analysis of the remarkable cases of distribution among the
Amphibia, with descriptions of new genera. Amer. Mus. Nox'itates,
212:1-24.

66 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

RiVERO, J. A.

1968. Los centrolenidos de Venezuela. Mem. Soc. Cien. Nat. La Salle,
28(8I):301-334.
Savage, J. M.

1967. A new tree-frog ( Centrolenidae ) from Costa Rica. Copeia,
1967(2) :325-331.
Savage, J. M. and W. R. Heyer

1967. Variation and distribution of tlie tree-frog genus PhtjUomedusa in
Costa Rica, Central America. Beitr. Neotrop. Fauna, 5:111-131.
Savage, J. M. and P. H. Starrett

1967. A new fringe-limbed tree-frog (family Centrolenidae) from lower
Central America. Copeia, 1967(3):604-609.
Starrett, P. H.

1960. Descriptions of tadpoles of Middle American frogs. Misc. Publ.
Mus. Zool. Univ. Michigan, 110:1-37.
Taylor, E. H.

1949. Costa Rican frogs of the genera Centwlene and CentroJenella.
Univ. Kansas Sci. Bull., 33:67-89.

1951. Two new genera and a new family of tropical American frogs. Proc.
Biol. Soc. Washington, 64:33-40.

1952. The frogs and toads of Costa Rica. Univ. Kansas Sci. Bull., 35:
577-942.

1958. Notes on Costa Rican Centrolenidae with descriptions of new fonns.
Zb/f/., .39:41-68.
Taylor, E. H. and D. M. Cochran

1953. Frogs of the family Centrolenidae from Brasil. Ibid., 35:1625-1656.

University of Kansas Publications

MUSEUM OF NATURAL HISTORY

The University of Kansas Publications, Museum of Natural
Histoiy, beginning with volume 1 in 1946, was discontinued
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Monographs of the Museum of Natural History were initiated
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