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OCCASIONAL PAPERS

LIBRARY

of the

MUSEUM OF NATURAL HISTORY
The University of Kansas
Lawrence, Kansas

number 155, pages 1-33 11 march 1993

New Species of Centrolenid Frogs
from Northern Peru

William E. Duellman and Rainer Schulte

Museum of Natural History and Department of Systematica and Eeologx, The

University of Kansas, Lawrence, Kansas 66045-2454, USA (WED); Proyecto de

Investigation de la Biologia de las Cordilleras Orientates, Apartado 258,

Tarapoto, Peru (RS)

ABSTRACT Eight new species of centrolenid frogs are described from the eastern
slopes of the Cordillera Central and ridges extending eastward and southward from
the main cordillera of the Andes in Departamento San Martin in northern
Peru. Three of these species are described in the genus Centrolene; they have dark
dorsal pigmentation in preservative, and the males have humeral spines. Four
species are placed in the genus Cochranella; they have dark dorsal pigmentation in
preservative, but males lack humeral spines. One of the species of Cochranella is
known only from females. One species is white in preservative and is a member of
the genus Hyalinobatrachium. Three species recognized in the genus Cochranella
by Ruiz-Carranza and Lynch ( 1991 ) are placed in the genus Centrolene.

Key words: Anura, Centrolenidae, New species, Peru.

RESUMEN Ocho especies nuevas de ranas centrolenidas se describen de las
laderas orientales de la Cordillera Central y de las montanas que se extienden al este
y al sur de la cordillera principal de los Andes en el Departamento de San Martin en
el norte de Peru. Tres de estas especies se describen en el genero Centrolene: tienen
pigmentacion oscura en el dorso en preservative, y los machos tienen espinas
humerales. Tres especies se ponen en el genero Cochranella; tienen pigmentacion
oscura en el dorso en preservativo, pero los machos no tienen espinas humerales.
Solo las hembras se conocen de una de las especies de Cochranella. Una especie es
blanca en preservativo y es un miembro del genero Hyalinobatrachium. Tres
especies reconcidas en el genero Cochranella por Rufz-Carranza and Lynch ( 1991 )
se ponen en el genero Centrolene.

Palabras claves: Anura, Centrolenidae, Especies nuevas, Peru.

© Museum of Natural History. The University of Kansas. Lawrence. 1SSN:009 1-7958

2 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

In recent years, six species of frogs have been named in the genus
Centrolenella from the cloud forests on the eastern slopes of the Andes or
isolated mountain ranges in central and southern Amazonian Peru — C.
spiculata and C. truebae from Departamento Ayacucho (Duellman, 1976),
C. mariae from Departamento Huanuco (Duellman and Toft, 1979), C.
phenax and C. pluvialis from Departamento Cuzco (Cannatella and
Duellman, 1982), and C. azulae from Departamento Huanuco (Flores and
McDiarmid, 1989). Even with the descriptions of these new species (all
except the holotype of C. azulae were collected for the first time in the
1970s), it seemed reasonable that many additional species would be
discovered as new areas were explored, especially in northern Peru. This
supposition was verified with the discovery of C. euhystrix and C. hesperia
on the Pacific versant of the Andes in northern Peru in 1987 (Cadle and
McDiarmid, 1990) and by our discovery of eight undescribed species on
the eastern Andean slopes in Departamento San Martin in northern Peru in
1989.

In the Departamento San Martin, a complex series of ridges reaching
elevations of more than 1000 m extend eastward and southward from the
Cordillera Central of the Andes. These steep, rocky ridges support lower
montane rainforest and cloud forest, whereas the intervening valleys, most
of which are extensively cultivated, contain remnants of dry tropical
forest. Many small streams cascade down these slopes; major drainage
systems flow south (e.g., Rio Mayo) or east (e.g., Rio Cainarache) into the
Rio Huallaga. which separates the Cordillera Central from the Cordillera
Oriental. In northern Peru, the Cordillera Oriental terminates in a series of
ridges with elevations of 1000-1200 m. The Rfo Huallaga cuts a canyon
through these front ranges between Shapaja and Chazuta, to the south of
which is the northern end of the Cordillera Azul (Fig. 1 ).

Adult centrolenids were collected on the eastern slopes of the Cordillera
Central and on two associated ridges. One site is on the west slope of a
pass, Abra Tangarana, to the west of the valley of the Rio Mayo. The other
sites are on a ridge north of Tarapoto and east of the Rio Mayo.

METHODS AND MATERIALS

In the following descriptions, species are assigned to genera and species
groups according to the definitions provided by Rufz-Carranza and Lynch
(1991), and the numbered characters in the diagnoses follow the standard
established by Lynch and Duellman ( 1973) as modified by Cannatella and
Duellman (1982) and Flores ( 1985). None of the species of centrolenids in
Central America, Chocoan South America, Venezuela, and southeastern
Brazil is shared with the upper Amazon Basin and the Amazonian slopes of
the Andes. Consequently, comparisons of the new taxa are made only with

NEW CENTROLENID FROGS FROM NORTHERN I'ERl

Fig. 1 . Map of part of Departamento San Martfn, Peru, showing localities
mentioned in text; circled numbers are: 1 = Abra Tangarana, 2 = Cataratas
Ahuashiyacu. 3 = northern end of Cordillera Azul. Based on Mapa Fisico Politico
Departamento de San Martin, Ed. 2, 1985, Instituto Geografico Nacional. Lima,
Peru.

those species from the upper Amazon Basin and the Amazonian slopes of
the Andes.

All measurements were taken to the nearest 0.1 mm with dial calipers.
Abbreviations used in the descriptions are: ED (eye diameter) = greatest
length of eye visible externally; EW (eyelid width) = greatest diagonal
width of upper eyelid; FL (foot length) = distance from proximal base of
inner metatarsal tubercle to tip of fourth toe; HL (head length) = distance
between tip of snout and posterior edge of angle of jaws; HnL (hand length)
= distance between proximal base of palmar tubercle and tip of third finger;
HW (head width) = greatest width of head at lateral margins of jaws; IOD
(interorbital distance) = width of top of head between nearest medial
margins of upper eyelids; SL (snout length) = distance from tip of snout to
anterior corner of orbit; SVL (snout-vent length) = distance from tip of
snout to posterior end of body; T-E (tympanum-eye) = distance between
anterior border of tympanic annulus and posterior corner of eyelid; TL
(tibia length) = distance from knee to heel; TYM (tympanum) = horizontal

4 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

length of tympanum including annulus. Relative lengths of digits were
determined by adpressing adjacent digits. Webbing formulae follow the
scheme of Savage and Heyer ( 1967) as modified by Myers and Duellman
(1982). Nuptial pad terminology follows Flores (1985).

Museum collections are abbreviated as follows: KU = Museum of
Natural History, The University of Kansas; LSUMZ = Museum of Zoology,
Louisiana State University; MHNSM = Museo de Historia Natural,
Universidad Nacional Mayor de San Marcos, Lima, Peru. Comparisons of
the new taxa were made with KU specimens of the other species.

DESCRIPTIONS OF NEW SPECIES

Centrolene fernandoi new species

Figure 2

Holotype. — KU 211770, an adult male, from the west slope of Abra
Tangarana, 7 km (by road) northeast of San Juan de Pacaysapa (06°12'S,
76°44'W, 1080 m), Provincia Lamas, Departamento San Martin, Peru, one
of a series collected on 5 February 1989 by William E. Duellman and
Rainer Schulte.

Paratopotypes — KU 211771-75 and MHNSM 6176-78; same date
and collectors.

Diagnosis. — A species in the Centrolene prosoblepon group
characterized by: (1) vomerine teeth present; (2) bones green; (3) parietal
peritoneum white; visceral peritoneum clear; (4) color in life, lime-green
with bluish white flecks dorsally; in preservative, pale lavender; (5) modal
webbing between outer fingers III 2 — 2 IV; (6) modal webbing on foot I
1 — 1+II 1—2 III 1—2 IV 2—1 V; (7) snout rounded in dorsal view, bluntly
rounded in profile: (8) dorsal skin shagreened with scattered minute
spicules; (9) arms and legs lacking tubercles and dermal folds; (10) humeral
spine present; (11) tympanum distinct, oriented posterolaterally with slight
dorsal inclination; (12) prepollex enlarged; unpigmented nuptial
excrescences (Type I) present; prepollical spine absent; (13) pair of enlarged
tubercles below vent; (14) first finger longer than second.

Of the other Centrolene in the region with vomerine teeth, C. audax
differs from C . fernandoi (characters in parentheses) by having the snout
truncate in profile (bluntly rounded), skin on dorsum shagreened without
spicules (shagreened with scattered small spicules), and in life, golden
(bluish-white) flecks on dorsum, digits pale yellow (pale green), and iris
pale bronze (silvery-green) with black reticulations. Centrolene marine
differs by having large, pale spots on the dorsum, truncate snout in profile,
and only basal webbing between Fingers III and IV; C. azulae and C .
puyoensis differ by having ulnar folds, snout truncate in profile in C. azulae

NEW CENTROLENID FROGS FROM NORTHERN PERU

Fig. 2. Upper left — paratype of Cochranella croceopodes, KU 211804, female,
24.7 mm SVL. Upper right. — holotype of Centrolene fernandoi, KU 21 1770, male,
24.0 mm SVL. Middle left — holotype of Centrolene lemniscatum, KU 217300,
male. 27.0 mm SVL. Middle right — holotype of Hyalinobatrachium lemur, KU
211768, male, 20.4 mm SVL. Lower left — holotype of Cochranella saxiscandens,
KU 211779, male, 21.7 mm SVL. Lower right — holotype of Cochranella
tangarana, KU 211776, male, 23.3 mm SVL.

6 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

and truncate in dorsal view in C. puyoensis, and less extensive webbing
between Fingers III and IV in C. puyoensis.

Description. — Nine adult males; SVL 22.5-26.4 mm. Body slender,
elongate. Head distinct, about as wide as long, round in outline as viewed
from below; HW 31.3-38.1% (3c = 33.2) of SVL; HL 30.0-36.0% (3c =
33.7) of SVL; snout rounded in dorsal view, bluntly rounded with slight
anteroventral inclination in profile: SL 34.2^11.0% (x = 37.3) of HL;
canthus rostralis straight, indistinct, round in section; loreal region concave;
lips barely flared anterior to orbit (Fig. 3C). Nostril ovoid, directed
anteriorly on protuberance; internarial area slightly depressed. Eye large,
ED 34.3^1.1% (x = 37.3) of HL, directed anterolateral^ at 37^0° (x =
39.2) from midline; greatest diagonal EW 57.1-67.0% (x = 60.0) of
IOD. Tympanum distinct, lower four-fifths visible, oriented posterolaterally
with slight dorsal inclination, separated from eye by distance 1.64-1.90 (x
= 1.71) times TYM, which is 31.2-41.4% (x = 35.7) of ED; tympanic
annulus slightly elevated anteriorly and ventrally; supratympanic fold
moderate, obscuring tympanic annulus dorsally.

Skin on dorsal surfaces of head, body, and limbs shagreened with small
spicules on side of head ventral and posterior to orbit and on dorsolateral
surfaces of body. Flanks smooth dorsally, granular ventrally. Skin on belly
and proximal four fifths of ventral surfaces of thighs coarsely granular;
other ventral surfaces smooth. Vent directed posteroventrally at upper level
of thighs; cloacal sheath short, unmodified; para- and subcloacal folds
absent; one pair of large, round, flat tubercles on ventral surfaces of thighs
below vent.

Breadth of upper arm about one half that of forearm; humeral spine
present; ulnar fold and tubercles absent; hand moderately large; HnL 88.0-
95.2% (x = 91.0) of HL; digits broad, bearing narrow lateral fringes,
including outer edge of Finger IV but not inner edge of Finger I; length of
digits II < I < IV < III; webbing basal between Fingers I and II; webbing
formula for other fingers II 2—2 III 2— (l-l1/:) IV (Fig. 5 C). Terminal
discs elliptical: width of disc on Finger III about 1.3 times TYM; width of
disc on Finger I 64.3-69.2% (x = 66.7) of that on Finger III. Subarticular
tubercles as wide as digits, round, elevated; penultimate tubercle on Finger
IV largest; supernumerary tubercles present on proximal segments (absent
on Fingers III and IV in some specimens); palmar tubercle large, broadly
ovoid with diagonal orientation, elevated, its length 15.5-20.5% (x = 17.7)
of HnL. Thenar tubercle elliptical, diffuse proximally, barely elevated,
about as long as palmar tubercle; prepollex slightly enlarged; Type I nuptial
excrescence present; prepollical spine absent.

Hind limbs long, slender; TL 54.3-59.2% (x = 56.7) of SVL; FL 43.0-
48.2% (x = 45.2) of SVL; dermal fringes, tarsal tubercles, and tarsal fold
absent; inner metatarsal tubercle elongately elliptical, barely elevated, not

NEW CENTROLENID FROGS FROM NORTHERN PERU 7

visible from above; outer metatarsal tubercle absent: toes moderately
slender, bearing lateral fringes, including outer edge of Toe V and inner
edge of Toe I; length of toes I < II < III < V < IV; toes about three-fourths
webbed; webbing formula I 1—2 II ( 1-1 '/:) — (2"-2+) III ( l-l1/:)— 2 IV 2 —
( 1-1 '/:) V. Terminal discs subtruncate; width of digit on Toe I 45.5-55.5%
(x = 50.3) of that on Toe IV, which is 69.2-73.3% (x = 71.3) of that on
Finger III; subarticular tubercles not as wide as digits, round, elevated.

Dentigerous processes of vomers ovoid, between nearly quadrangular
choanae, narrowly separated medially, each bearing 2-5 (x = 3.3)
teeth. Tongue nearly round, barely free posteriorly. Vocal slit elongate,
extending from posterolateral margin of tongue toward angle of jaw; vocal
sac single, median, subgular.

Color in life: Dorsum lime-green with small bluish-white flecks; digits
pale green; parietal peritoneum white; visceral peritoneum clear; heart not
visible; bones green; iris pale silvery-green with fine black reticulations.

Color in preservative: General appearance lavender with small, elevated,
cream flecks dorsally, cream ventrally. Under 25x magnification, dense
melanophores in skin of dorsal surfaces of head and body with scattered,
small, unpigmented elevations; equally dense melanophores in loreal
region; less dense melanophores on upper lip, in tympanic region, and on
dorsal surfaces of forelimbs. hind limbs, third and fourth fingers, tarsi, and
fourth and fifth toes. Edge of upper eyelid pigmented. Flanks, anterior and
posterior surfaces of thighs, inner two fingers and inner three toes, and all
ventral surfaces cream with no melanophores.

Measurements (mm with means in parentheses): SVL 22.5-26.4 (23.7),
TL 12.9-14.3 (13.4), FL 10.2-11.5 (10.7), HW 7.3-8.9 (7.9), HL 7.6-8.4
(8.0), SL 2.8-3.2 (3.0). ED 2.8-3.2 (3.0). IOD 2.8-3.1 (3.0). T-E 1.7-1.9
(1.8), TYM 1.0-1.2 (1.1), HnL 7.2-7.4 (7.3).

Distribution and ecology. — All individuals were on the upper surfaces
of leaves of trees 1.5-2.0 m above a small stream in a narrow ravine at
night. Although centrolenids were calling, no call was associated with this
species. The type locality is on the road between Moyobamba and
Tarapoto. The stream flows southward at a point about 0.5 km above
(northeast of) Somos Libres (a road maintenance camp) and about 1 km
below (southwest of) the crest of the ridge.

Etymology. — The specific name is a patronym for Fernando M. Cuadros
V. of the Universidad Nacional Mayor de San Marcos in Lima in recognition
of his cheerful companionship and dedication to collecting frogs in northern
Peru in 1989.

Remarks. — The most similar species to C. fernandoi is C. audax,
which is known from elevations of 1660-1700 in on the Amazonian slopes
of the Andes in northern Ecuador.

UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

A LathRlM* **'

Fig. 3. Heads of centrolenids: A. Cochranella chancas, KU 211778. B.
Cochranella croceopodes, KU 211804. C. Centrolene fernandoi, KU 211770. D.
Centrolene lemniscatum, KU 217300. Scale = 5 mm.

NEW CENTROLENID FROGS FROM NORTHERN PERU

J

Fig. 4. Heads of centrolenids: A. Hyalinobatrachium lemur, KU 211768. B.
Centrolene muelleri, KU 217301. C. Cochranella saxiscandens. KU 211779. D.
Cochranella tangarana, KU 211777. Scale = 5 mm.

10

UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

-I L 1 |

mmm

Ml » m
■ m ... %&
Mm

Fig. 5. Hands of centrolenids: A. Cochranella chancas, KU 211778. B.
Cochranella croceopodes, KU 211804. C. Centrolene fernandoi, KU 211770. D.
Centrolene lemniscatum, KU 217300. Scale = 5 mm.

NEW CENTROLENID FROGS FROM NORTHERN PERU

11

Fig. 6. Hands of centrolenids: A. Hyalinobatrachium lemur, KU 211768. B.
Centrolene muelleri, KU 217301. C. Cochranella saxiscandens, KU 211779. D.
Cochranella tangarana, KU 211777. Scale = 5 mm.

12 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

Centrolene lemniscatum new species

Figure 2

Holotype. — KU 217300. an adult male, from 14 km (by road) west of
Venceremos (05°44'S, 77°32'W, 2000 m). Provincia Rioja, Departamento
San Martin, Peru, obtained on 9 July 1989 by Rainer Schulte.

Diagnosis. — A species in the Centrolene prosoblepon group
characterized by: (1) vomerine teeth absent; (2) bones green; (3) parietal
peritoneum white; visceral peritoneum clear; (4) color in life, dorsum pale
green with white labial stripe continuous with broad lateral stripe on body;
in preservative, pale lavender with white lateral stripe; (5) webbing between
outer fingers III 21/:— 21/: IV; (6) webbing on foot I Vh— 2+II 1— 2+III 1+—
2Vi IV 21/: — 2"V; (7) snout bluntly rounded in dorsal view and in profile; (8)
dorsal skin spiculate; (9) arms and legs lacking tubercles and dermal folds;
(10) humeral spine present; (11) tympanum indistinct, oriented
posterolaterally; (12) prepollex enlarged; nuptial excrescense absent;
prepollical spine absent; (13) two pairs of enlarged tubercles below vent;
(14) first finger shorter than second.

Two other species in the region have white labial stripes. Of these,
Cochranella croceopodes differs by having a diffuse yellow line on the
flanks, first finger longer than the second, vomerine teeth, a distinct
tympanum, more webbing on the hand, and presumably no humeral spines
in males. Centrolene lemniscatum is like C. hesperium in having a distinct
lateral white stripe that is continuous with the labial stripe, no vomerine
teeth, and humeral spines. Centrolene lemniscatum differs from C.
hesperium (characters in parentheses) by having the snout rounded in
profile (sloping anteroventrally), and an indistinct tympanum (concealed),
and in lacking dermal folds on the limbs (present).

Description. — One adult male; SVL 27.0 mm. Body robust. Head
distinct, about as wide as long, round in outline as viewed from below; HW
30.3% of SVL; HL 30.7% of SVL; snout rounded with upper margin
truncate in dorsal view; bluntly rounded in profile; SL 41.0% of HL;
canthus rostralis straight, barely evident, round in section; loreal region
slightly concave; lips slightly flared anterior to orbit (Fig. 3D). Nostril
ovoid, directed anterolaterally on protuberance; internarial area barely
depressed. Eye moderately large, ED 34.9% of HL, directed anterolaterally
at 42° from midline; greatest diagonal EW 70.0% of IOD. Tympanum
indistinct, oriented laterally with slight posterior inclination; tympanic
annulus not evident; supratympanic fold moderately heavy, obscuring upper
part of tympanum.

Skin on dorsal surfaces of head and body spiculate, with numerous
small spicules below orbit; skin on limbs smooth. Flanks smooth dorsally.

NEW CENTROLENID FROGS FROM NORTHERN PERU 13

granular ventrally. Skin on belly and ventral surfaces of thighs coarsely
granular; other ventral surfaces smooth. Vent directed posteroventrally at
upper level of thighs; cloacal sheath short, unmodified; para- and subcloacal
folds absent; two pairs of moderately large, round, elevated tubercles on
ventral surfaces of thighs below vent.

Breadth of upper arm about one half that of forearm; humeral spine
present; ulnar fold and tubercles absent; hand large: HnL 106% of HL;
digits broad, bearing narrow lateral fringes, including outer edge of Finger
IV and inner edge of Finger I; length of digits I < II < IV < III; webbing
absent between Fingers I, II, and III; webbing formula for other fingers III
2 — 21/: IV (Fig. 5D). Terminal discs subtruncate; width of disc on Finger
III 65.5% of ED; width of disc on Finger I 57.9% of that on Finger III.
Subarticular tubercles nearly as wide as digits, round, elevated; distal
tubercle on Finger IV largest; supernumerary tubercles prominent on
proximal segments; palmar tubercle moderately large, ovoid with diagonal
orientation, elevated, its length 13.6% of HnL. Thenar tubercle ovoid,
diffuse proximally, barely elevated, much longer than palmar tubercle;
prepollex slightly enlarged; Type I nuptial excrescence present; prepollical
spine absent.

Hind limbs long, slender; TL 56.3% of SVL; FL 5 1 .9% of SVL; dermal
fringes, tarsal tubercles, and tarsal fold absent; inner metatarsal tubercle
elongately elliptical, elevated, visible from above; outer metatarsal tubercle
absent; toes moderately slender, bearing lateral fringes, including outer
edge of Toe V and inner edge of Toe I; length of toes I < II < III < V < IV;
toes about three-fifths webbed; webbing formula I l1/: — 2+II 1 — 2+III 1+ —
2Vi IV 21/: — 2 V Terminal discs subtruncate; width of digit on Toe I 56% of
that on Toe IV. which is 93.1% of that on Finger III; subarticular tubercles
as wide as digits, round, elevated.

Vomerine teeth absent; choanae widely separated, quadrangular. Tongue
shallowly cordiform, free posteriorly for about one third of its length. Vocal
slit elongate, extending from posterolateral margin of tongue toward angle
of jaw; vocal sac single, median, subgular.

Color in life: Dorsum medium green with irregular patches of yellowish-
green: white labial stripe continuous with distinct lateral white stripe above
insertion of arm extending to groin; flanks below lateral stripe cream;
narrow white stripe on outer edge of forearm and foot; digits pale yellowish-
green; parietal peritoneum white; visceral peritoneum clear; heart not
visible; bones green; iris silvery-bronze with black flecks.

Color in preservative: General appearance grayish lavender on dorsal
surfaces of head and body, cream on limbs and venter; white lateral stripe
distinct. Under 25x magnification, dense melanophores in skin of dorsal
surfaces of head and body; tips of spicules unpigmented: less dense
melanophores in loreal and tympanic regions and on dorsal surfaces of

14 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

forelimbs, thighs, shanks, tarsi, and fourth and fifth toes. Edge of upper
eyelid pigmented. Edge of upper lip and lateral stripe white; flanks, anterior
and posterior surfaces of thighs, fingers and and inner three toes, and all
ventral surfaces cream with no melanophores.

Measurements (in mm): SVL 27.0. TL 15.2. FL 14.0, HW 8.2, HL 8.3,
SL 3.4, E-N 1.8, ED 2.9, EW 2.1, IOD 3.0, HnL 8.8.

Distribution and ecology. — The holotype was on a bush in a stream in
cloud forest at night. The type locality is on the road between Pomacochas
(= Florida) and Rioja; Veneceremos is 89 km (by road) northwest of Rioja.

Etymology. — The specific name is a Latin adjective meaning adorned
with ribbons; it alludes to the distinctive pale dorsolateral stripes.

Remarks. — Centrolenella lemniscatum exhibits only slight differences
from C. hesperium, which is known only from forested slopes at elevations
of 1500-1800 m on the Pacific versant in northern Peru (Cadle and
McDiarmid, 1990). Conceivably, the single specimen of C. lemniscatum
represents geographic variation in C. hesperium, but no species of frog is
known to occur in the vicinity of the type locality of C. hesperium on the
Pacific slopes of Peru and on the eastern slopes of the Cordillera Central in
Peru.

Centrolene muelleri new species

Figure 7

Holotype. — KU 217301, an adult male from 14 km (by road) west of
Venceremos (05°44'S, 77°32'W, 2000 m), Provincia Rioja, Departamento
San Martin, Peru, obtained on 9 July 1989 by Rainer Schulte.

Diagnosis. — A species in the Centrolene prosoblepon group
characterized by: (1) vomerine teeth absent; (2) bones green; (3) parietal
peritoneum white; visceral peritoneum clear; (4) color in life, green with
dark greenish-black spots and pale yellow tubercles; in preservative, brown
with black spots and cream tubercles; (5) webbing on outer fingers III 2+ —
2+ IV; (6) webbing on foot I Vh— 2 II 1+— 2 III 2— 21/: IV 21/:— l1/: V; (7)
snout truncate in dorsal view, inclined anteroventrally in profile; (8) dorsal
skin smooth with dorsolateral rows of tubercles; (9) low, scalloped dermal
fold on outer edge of forearm and tarsus; ( 10) humeral spine present; (11)
tympanum distinct, oriented posterolaterally with slight dorsal inclination;
(12) prepollex enlarged; unpigmented nuptial excrescences (Type I)
present; prepollical spine absent; (13) pair of enlarged tubercles below
vent; (14) first finger longer than second.

Centrolene muelleri differs from all other members of the genus by
having large, conical tubercles on the dorsal surfaces of the head, body, and
limbs. Other species that have dermal fringes on the limbs include C.
gemmatum and C. hesperium, both of which have the first finger shorter

NEW CENTROLENID FROGS FROM NORTHERN PERU 15

than the second (first longer than second in C. muelleri), and C. pipilatum,
which has minute black flecks instead of large dark spots on the dorsum.

Description. — One adult male; SVL 23.5 mm. Body slender, elongate.
Head distinct, slightly wider than long, round in outline as viewed from
below; HW 32.3% of SVL; HL 31.1% of SVL; snout slightly protruding,
truncate in dorsal view; slightly inclined anteroventrally in profile; SL
37.0% of HL; canthus rostralis straight, indistinct, round in section; loreal
region concave; lips not flared anterior to orbit (Fig. 4B). Nostril ovoid,
directed dorsolaterally on protuberance; internarial area depressed. Eye
moderately large, ED 35.6% of HL, directed anterolateral^ at 42° from
midline; greatest diagonal EW 46.7% of IOD. Tympanum distinct,
completely visible, oriented posterolaterally with slight dorsal inclination,
separated from eye by distance 1.20 times TYM, which is 42.3% of ED;
tympanic annulus slightly elevated; supratympanic fold weak, barely
obscuring tympanic annulus dorsally.

Skin on dorsal surfaces of head, body, and limbs smooth with
dorsolateral row of tubercles extending from occiput to insertion of hind
limb; row of small tubercles on upper lip; one tubercle on each side of head
at anterior corner of orbit and one medial to posterior corner of orbit:
tubercles on dorsal surfaces of forelimbs. thighs, shanks, and tarsi. Flanks
smooth dorsally, granular ventrally. Skin on belly and ventral surfaces of
thighs coarsely granular; other ventral surfaces smooth. Vent directed
posteroventrally at upper level of thighs; cloacal sheath short; tubercular
para- and subcloacal folds present; one pair of large, subcorneal tubercles
on ventral surfaces of thighs below vent.

Breadth of upper ami about one half that of forearm: humeral spine
present: scalloped dermal folds on inner and outer edges of forearm; hand
large: HnL 105% of HL; digits broad, bearing broad lateral fringes,
including outer edge of Finger IV and inner edge of Finger I; length of
digits II < I < IV < III; webbing basal between Fingers I and II; webbing
formula for other fingers II 2+ — 3 III 2+ — 2+ IV (Fig. 6B). Terminal discs
broadly elliptical; width of disc on Finger III 1.45 times TYM; width of
disc on Finger I 50% of that on Finger III. Subarticular tubercles as wide as
digits, round, elevated: distal tubercle on Finger I conical; all tubercles
about same size; supernumerary tubercles present on proximal segments;
palmar tubercle moderately large, broadly ovoid with diagonal orientation,
elevated, its length 14.3% of HnL. Thenar tubercle ovoid, elevated, slightly
longer than palmar tubercle; prepollex enlarged; Type I nuptial excrescence
present; prepollical spine absent.

Hind limbs long, slender; TL 6 1 .2% of SVL; FL 44.4% of SVL; pair of
flaplike tubercles on heel; scalloped dermal fold on outer edge of tarsus;
inner tarsal fold absent; inner metatarsal tubercle elongate, elliptical, barely
elevated, visible from above; outer metatarsal tubercle absent; toes

16 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

moderately slender, bearing lateral fringes, including outer edge of Toe V
and inner edge of Toe I; length of toes I < II < III < V < IV; toes about three-
fourths webbed; webbing formula I 1 Vi— 2 II T— 2 III 2— 2Vi IV 21/: —1 Vi
V Terminal discs subtruncate; width of digit on Toe I 53.8% of that on Toe
IV, which is 81.3% that on Finger III; subarticular tubercles about as wide
as digits, round, subconical.

Vomerine teeth absent; choanae round, widely separated. Tongue
narrowly cordiform. barely free posteriorly. Vocal slit elongate, extending
from posterolateral margin of tongue toward angle of jaw; vocal sac single,
median, subgular.

Color in life: Dorsum green with small black spots and cream tubercles;
parietal peritoneum white; visceral peritoneum clear; heart not visible;
bones green.

Color in preservative: General appearance grayish brown with small
dark spots and cream spots corresponding to tubercles on head, body, and
limbs dorsally; cream ventrally. Under 25x magnification, dark spots on
head and body formed by aggregations of melanophores; elsewhere on
body melanophores not so dense; narrow band of melanophores on dorsal
surfaces of thighs and upper arms; melanophores present on outer two toes,
absent on fingers. Edge of upper eyelid pigmented. Flanks, anterior and
posterior surfaces of thighs, fingers, inner three toes, and all ventral surfaces
cream without melanophores.

Measurements (in mm): SVL 23.5. TL 14.4. FL 10.4. HW 7.6. HL 7.3.
SL 2.7, ED 2.6, EW 1 .4, IOD 3.0, T-E 1 .2, TYM 1.1. HnL 7.7.

Distribution and ecology. — The holotype was on a low bush at the
edge of a stream in cloud forest at night. See the account of Centrolene
lemniscatum for a description of the type locality.

Etymology. — The specific name is a patronym for Paul S. Muller, of the
Universitiit des Saarlandes, who originally supported Schulte's field studies
in Peru.

Remarks. — The holotype is slightly desiccated; however, all of the
characters are readily discernable.

Cochranclla chancas new species

Figure 8

Holotype. — KU 211778, an adult male, from the west slope of Abra
Tangarana, 7 km (by road) northeast of San Juan de Pacaysapa (06°12'S,
76°44'W, 1080 m), Provincia Lamas, Departamento San Martin. Peru,
obtained on 5 February 1989 by William E. Duellman.

Diagnosis. — A species in the Cochranella ocellata group characterized
by: (1) vomerine teeth absent; (2) bones green; (3) parietal peritoneum
white; visceral peritoneum clear; (4) color in life, dull yellowish-green with

NEW CENTROLENID FROGS FROM NORTHERN PERU 17

minute yellow flecks; in preservative, dull gray with cream flecks; (5)
webbing basal between outer fingers; (6) webbing on foot I 2+ — 3 II 2+ — 3
III 2 — 3 IV 3 — 2 V; (7) snout bluntly rounded in dorsal view, truncate in
profile; (8) dorsal skin smooth with minute spicules on head and in thoracic
region; (9) arms and legs lacking tubercles and dermal folds; (10) humeral
spine absent in males; (11) tympanum weakly defined, oriented
posterolaterally with slight dorsal inclination; (12) prepollex not enlarged,
Type I nuptial excrescence present; prepollical spine absent; (13) one pair
of large tubercles below vent; ( 14) first and second fingers equal in length.

Cochranella chancas is compared with other species having a white
parietal peritoneum (heart not visible) and green bones that lack vomerine
teeth and humeral spines. Of these species, C. megacheira and C. truebae
differ from C. chancas by having dark dorsal markings (black spots in C.
megacheira and many black flecks in C. truebae). Cochranella cochranae
and C. ocellata differ from C. chancas by having well-defined ocelli on the
dorsum. Cochranella phenax, C. pluvialis, C. siren, and C. euhystrix have
spicules over the entire dorsum, whereas in C. chancas the dorsum is
smooth except for minute spicules on the head and in the thoracic region.
Moreover, C. phenax (male SVL IQ.l-ll.X mm) and C. siren (male SVL
19.8-22.0 mm) are smaller than C. chancas (male SVL 24.9 mm) and have
proportionately shorter snouts that are truncate in dorsal view (bluntly
rounded in C. chancas), whereas C. euhystrix (male SVL 28.5-3 1 .3 mm) is
larger. Two other species are similar to C. chancas; C. bejaranoi is like C.
chancas in coloration and in having most of Finger IV free of webbing, but
it differs by having spicules over the entire dorsum of the head and body, in
having the first finger shorter than the second, and in lacking enlarged
tubercles below the vent; C. fiavopunctata also is like C. chancas in
coloration, but it differs by having all skin on the dorsum smooth to
shagreened without spicules, only 1 '/: phalanges of Finger IV free of
webbing, and first finger longer than second. Some individuals of C.
fiavopunctata and C. siren have vomerine teeth.

Description. — One adult male; SVL 24.9 mm. Body slender, elongate.
Head distinct, about as wide as long, round in outline as viewed from
below; HW 29.7% of SVL; HL 30.5% of SVL; snout slightly protruding
with upper margin truncate in dorsal view, truncate in profile: SL 36.8% of
HL; canthus rostralis straight, indistinct, round in section; loreal region
concave; lips noticeably flared anterior to orbit (Fig. 3A). Nostril ovoid,
directed anterolaterally on protuberance; internarial area depressed. Eye
large, ED 42.1% of HL. directed anterolaterally at 35° from midline;
greatest diagonal EW 88.9% of IOD. Tympanum barely distinct, completely
visible, oriented posterolaterally with slight dorsal inclination, separated
from eye by distance 1.36 times TYM, which is 34.4% of ED; tympanic
annulus slightly elevated anteriorly and ventrally; supratympanic fold weak.

18 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

barely obscuring tympanic annulus dorsally.

Skin on dorsal surfaces of head, body, and limbs smooth except for
small, round elevations corresponding to pale flecks and small spicules on
head and in thoracic region. Flanks smooth dorsally. finely granular
ventrally. Skin on belly and proximal three fourths of ventral surfaces of
thighs coarsely granular; other ventral surfaces smooth. Vent directed
posteroventrally at upper level of thighs; cloacal sheath short, unmodified;
para- and subcloacal folds absent; one pair of large, round, flat tubercles on
ventral surfaces of thighs below vent.

Breadth of upper arm about one half that of forearm; humeral spine
absent; ulnar fold and tubercles absent; hand moderately large; HnL 90.8%
of HL; digits broad, bearing narrow lateral fringes, including outer edge of
Finger IV and inner edge of Finger I; length of digits I = II < IV < III;
webbing basal between Fingers III and IV absent between other fingers
(Fig. 5A). Terminal discs elliptical; width of disc on Finger III equals
TYM; width of disc on Finger I 68% of that on Finger III. Subarticular
tubercles about as wide as digits, round, elevated; distal tubercle on Finger
IV largest; supernumerary tubercles low, diffuse, present only on proximal
segments; palmar tubercle large, nearly rectangular with diagonal
orientation, elevated, its length 18.8% of HnL. Thenar tubercle elliptical,
diffuse distally, barely elevated, nearly as long as palmar tubercle; prepollex
not enlarged; Type I nuptial excrescence present; prepollical spine absent.

Hind limbs long, slender; TL 59.0% of SVL; FL 43.4% of SVL; dermal
fringes, tarsal tubercles, and tarsal fold absent; inner metatarsal tubercle
ovoid, elevated, visible from above; outer metatarsal tubercle absent; toes
moderately slender, bearing lateral fringes, including outer edge of Toe V
and inner edge of Toe I; length of toes I < II < III < V < IV; toes about three-
fifths webbed; webbing formula I 2+ — 3 II 2+— 3 III 2—3 IV 3-2 V.
Terminal discs subtruncate; width of digit on Toe I 739£ of that on Toe IV,
which equals that on Finger III; subarticular tubercles not as wide as digits,
round, elevated.

Vomerine teeth absent; choanae widely separated, subtriangular with
apex anteromedially and rounded posterolaterally. Tongue shallowly
cordiform. barely free posteriorly. Vocal slit elongate, extending from
posterolateral margin of tongue toward angle of jaw; vocal sac single,
median, subgular.

Color in life: Dorsum dull yellowish-green with minute yellow flecks
(elevated); parietal peritoneum white; visceral peritoneum clear; heart not
visible; bones green; iris pale yellowish-bronze.

Color in preservative: General appearance grayish-brown with minute
cream flecks dorsally, cream ventrally. Under 25x magnification, dense
melanophores in skin of dorsal surfaces of head. body, forearms, and
shanks with scattered, small, unpigmented elevations; equally dense

NEW CENTROLENID FROGS FROM NORTHERN PERU 19

melanophores in loreal and tympanic regions; less dense melanophores on
dorsal surfaces of fourth fingers, thighs, tarsi, and fourth and fifth toes. Edge
of upper eyelid unpigmented. Upper lip. flanks, upper arms, anterior and
posterior surfaces of thighs, inner three fingers and toes, and all ventral
surfaces, except chest, cream with no melanophores; chest white.

Measurements (in mm): SVL 24.9. TL 14.7. FL 10.8, HW 7.4, HL 7.6.
SL 2.8, ED 3.2. EW 2.4. IOD 2.7. T-E 1 .5, TYM 1.1, HnL 6.9.

Distribution and ecology. — The only known specimen was perched on
a leaf over a stream in a narrow ravine at night. See the account of
Centrolene fernandoi for a description of the type locality, the only site
from which this species is known.

Etymology. — The specific name is a noun in apposition and is the name
of the local indigenous people, Chancas, who were nearly exterminated by
the Incas; the survivors are centered in the town of Lamas.

Remarks. — Loathful though we are to describe a new species on the
basis of a single individual, the holotype of C. chancas has a combination
of characters unknown in any other member of the genus.

Cochranella croceopodes new species

Figure 2

Holotype— KU 211804, an adult female, from 23.2 km (by road)
northeast of Tarapoto (06°27'S, 76°48'W. 800 m), Provincia San Martin.
Departamento San Martin, Peru, obtained on 15 February 1989 by William
E. Duellman.

Paratype. — KU 21 1799, an adult female, from Cataratas Ahuashiyacu,
730 m, 14 km (by road) northeast of Tarapoto, Provincia San Martin,
Departamento San Martin. Peru, obtained on 11 February 1989 by John
J.Wiens.

Diagnosis. — A species in the Cochranella granulosa group
characterized by: (1) vomerine teeth present; (2) bones green; (3) parietal
and visceral peritonea white; (4) color in life, uniform dull green dorsally
with white margin of upper lip, diffuse yellow line on flanks, and digits
yellow; in preservative, dark lavender; (5) modal webbing between outer
fingers III 2— LIV; (6) modal webbing on foot I 1— LII 1+— 2 III L— 2
IV 2 — 1 V; (7) snout bluntly rounded in dorsal view and nearly truncate in
profile; (8) dorsal skin shagreened; (9) arms and legs lacking tubercles and
dermal folds; ( 10) state of humeral spine unknown; (11) tympanum distinct,
oriented laterally with slight dorsal inclination; (12)prepollex not enlarged;
prepollical spine absent; state of nuptial excrescences unknown; (13) pair
of enlarged tubercles below vent; ( 14) first finger longer than second.

Cochranella croceopodes is compared with other species having lateral
stripes — Centrolene hesperium and C. lemniscatum. Both of these species

20

UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

Fig. 7. Holotype of Centrolene muelleri, KU 217301. male, 23.5 mm SVL.

lack vomerine teeth, have broader and more distinct lateral stripes, and the
first finger is shorter than the second. Furthermore, the males have humeral
spines.

Description. — Two adult females; SVL 24.7-24.8 mm. Body robust.
Head distinct, about as wide as long, round in outline as viewed from
below; HW 33.9-34.0% of SVL; HL 33.9-34.8% of SVL; snout slightly
protruding, bluntly rounded in dorsal view and nearly truncate with slight
anteroventral inclination in profile; SL 38.1% of HL; canthus rostralis
straight, indistinct, round in section; loreal region barely concave; lips not
flared anterior to orbit (Fig. 3B). Nostril ovoid, directed anterolaterally on
protuberance; internarial area depressed. Eye moderately large, ED 38.1%
of HL, directed anterolaterally at 45° from midline; greatest diagonal EW
67.7-71.0% of IOD. Tympanum distinct, almost entirely visible, oriented
dorsolateral^ with slight posterior inclination, separated from eye by
distance 1.38-1.58 times TYM, which is 37.5-40.6% of ED; tympanic
annulus slightly elevated anteriorly, posteriorly, and ventrally;
supratympanic fold moderately heavy, obscuring tympanic annulus
dorsally.

Skin on dorsal surfaces of head, body, and limbs shagreened. Flanks
smooth dorsally, finely granular ventrally. Skin on belly and ventral surfaces
of thighs coarsely granular; other ventral surfaces smooth. Vent directed
posteroventrally at upper level of thighs; cloacal sheath short, unmodified;

NEW CENTROLENID FROGS FROM NORTHERN PERU 2 1

para- and subcloaca! folds absent; one pair of large, round, slightly elevated
tubercles on ventral surfaces of thighs below vent.

Breadth of upper arm about one half that of forearm; humeral spine
absent (males unknown); ulnar fold and tubercles absent; hand moderately
large; HnL 88.4-90.5% of HL; digits broad, bearing narrow lateral fringes,
including outer edge of Finger IV but excluding inner edge of Finger I;
length of digits II < I < IV < III; webbing absent between Fingers I and II;
webbing formula for other fingers II 2~ — 3 III 2 — 1 + IV (Fig. 5B). Terminal
discs elliptical; that on fourth finger slightly indented medially; width of
disc on Finger III 1.33-1.38 times TYM; width of disc on Finger I 62.5-
66.7% of that on Finger III. Subarticular tubercles round, subconical;
penultimate tubercle on Finger IV largest, as wide as digit; supernumerary
tubercles low, present on proximal segments of Fingers I — I II; palmar
tubercle large, broadly ovoid with diagonal orientation, elevated with
median depression, its length 17.1-19.0% of HnL. Thenar tubercle
elongately elliptical, diffuse proximally, barely elevated, longer than palmar
tubercle; prepollex not enlarged. Condition of nuptial excrescence and
prepollical spine unknown.

Hind limbs long, slender; TL 53.8-59.7% of SVL; FL 43.7-50.8% of
SVL; dermal fringes, tarsal tubercles, and tarsal fold absent; inner
metatarsal tubercle elliptical, elevated distally, visible from above; outer
metatarsal tubercle absent; toes moderately slender, bearing lateral fringes,
including outer edge of Toe V and inner edge of Toe I; length of toes I < II
< III < V < IV; toes about three-fourths webbed; webbing formula I 1 — ( 1+-
2) II 1+ — 2 III (T-l1/: )— 2 IV (2-2) — 1 V Terminal discs elliptical;
width of digit on Toe I 6 1 .5% of that on Toe IV. which equals that on Finger
III; subarticular tubercles as wide as digits, round, elevated.

Dentigerous processes of vomers elliptical, slightly inclined
posteromedially, narrowly separated medially, each with four or five (x =
4.8) teeth; choanae widely separated, subtriangular with anteromedial apex
and lateral rounded base. Tongue slightly longer than wide, barely free
posteriorly. Condition of vocal slit and sac unknown.

Color in life: Dorsum dull green; margin of upper lip white; diffuse
yellow line on flanks from axilla to groin; fingers and toes yellow; throat
and ventral surfaces of limbs pale green; parietal and visceral peritonea
white; heart not visible; bones green; iris dull bronze with back flecks and
brownish suffusion anteriorly and posteriorly.

Color in preservative: General appearance dark lavender dorsally, cream
ventrally. Under 25x magnification, lavender with underlying
melanophores in skin of dorsal surfaces of head and body, and in loreal and
tympanic regions; less dense melanophores on dorsal surfaces of limbs,
third and fourth fingers, and fourth and fifth toes. Edge of upper eyelid
pigmented. Edge of upper eyelid pigmented. Upper lip, lower one half of

22 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

tympanum, flanks, posterior surfaces of thighs, inner two fingers, inner
three toes, and all ventral surfaces cream with no melanophores.

Measurements (in mm; holotype followed by paratype): SVL 24.7,
24.8; TL 13.3, 14.8; FL 10.8, 12.6; HW 8.4, 8.4; HL 8.6, 8.4; SL 3.2, 3.2;
ED 3.2, 3.2; EW 2.1, 2.2; IOD 3.1, 3.1; T-E 1.9, 1.8; TYM 1.2, 1.3; HnL
7.6, 7.9.

Distribution and ecology. — The species is known only from two
localities on a northwest-southeast ridge north of Tarapoto. The type
locality is on the north slope and in the drainage of the Rio Cainarachi. The
locality is at the edge of the Tarapoto- Yurimaguas road, 3.2 km north of the
tunnel through the upper part of the ridge. The second locality is on the
south slope of the same ridge (for description see account of Cochranella
sa.xiscandens). The holotype was found at night on the upper surface of an
aroid on a vertical rock face about 3 m above the ground in an area of
dripping water. The paratype was on a low herb next to a cascading stream
at night.

Etymology. — The specific name is an adjective derived from the Greek
croceos meaning yellow-orange and the Greek podos meaning foot; the
name refers to the bright yellow hands and feet.

Remarks. — The definitive placement of this species in Cochranella
must await the discovery of males in order to determine the presence or
absence of humeral spines.

Cochranella sa.xiscandens new species
Figure 2

Holotype. — KU 211779, an adult male, from Cataratas Ahaushiyacu
(06°30'S, 76°20'W, 730 m). 14 km (by road) northeast of Tarapoto,
Provincia San Martin, Departamento San Martin. Peru, one of a series
collected on 8 February 1989 by William E. Duellman, Rainer Schulte, and
John J. Wiens.

Paratypes.— KU 211780-88 and MHNSM 6166-71 collected with the
holotype; KU 211789-98, 211800-01 from the type locality collected on
1 1 February 1989 by John J. Wiens: KU 21 1802-03 from 15 km northeast
of Tarapoto, 800 m, collected on 1 1 February 1989 by Rainer Schulte; KU
217299 from the type locality obtained on 3 June 1989 by Rainer Schulte.

Diagnosis. — A species in the Cochranella ocellata group characterized
by: (1) vomerine teeth present; (2) bones green; (3) parietal peritoneum
white; visceral peritoneum clear; (4) color in life, dull dark green to black
dorsally and paler green laterally; in preservative, dark olive4orown to
black; (5) modal webbing between outer fingers III 2 — 2 IV; (6) modal
webbing on foot I 1+— l1/: II 1—2" III 1—2 IV 2—1 V; (7) snout bluntly
rounded in dorsal view and in profile; (8) dorsal skin spiculate; (9) arms

NEW CENTROLENID FROGS FROM NORTHERN PERU

23

Fig. 8. Holotype of Cochranella chancas, KU 211778. male. 24.9 mm SVL.

and legs lacking tubercles and dermal folds; inner tarsal fold present; ( 10)
humeral spine absent; (11) tympanum distinct, oriented dorsolaterally with
slight posterior inclination; (12) prepollex enlarged; unpigmented nuptial
excrescences (Type I) present; prepollical spine absent; (13) no enlarged
tubercles below vent; (14) first finger longer than second.

Of the nine other species of Cochranella in the region. C. resplendens
differs from C. saxiscandens by having dermal folds on the limbs (as do
Centrolene azulae and puyoensis); C. ametarsia and C. ritae differ by
having black flecks on the dorsum. Four of the other species (C. cochranae,
flavopunctata, midas, and siren) differ from C. saxiscandens by having
pale spots or flecks on the dorsum. Only C. spiculata and C. tangarana are
like C. saxiscandens in having a uniformly darkly pigmented dorsum that
is finely spiculate; both of these species differ from C. saxiscandens
(characters in parentheses) by having a lavender dorsum (dark gray to
black) in preservative, melanophores absent on the ventral surfaces of
shanks and tarsi (present), and inner tarsal fold absent (present).

Description.— Thirty adult males; SVL 20.8-23.2 mm (x = 22.0). Body
moderately robust. Head distinct, about as wide as long, round in outline as
viewed from below; HW 31.2-37.3% (x = 35.1) of SVL; HL 32.0-36.2%
(x = 34.3) of SVL; snout rounded with upper margin truncate in dorsal
view; bluntly rounded in profile; SL 34.1-43.8% (x = 38.6) of HL: canthus
rostralis slightly curved, indistinct, round in section; loreal region barely
concave; lips not flared anterior to orbit (Fig. 4C). Nostril round, directed
dorsolaterally on slight protuberance; internarial area not depressed. Eye

24 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

moderately large, ED 32.4-45.6% (x - 37.2) of HL, directed anterolaterally
at 40-42° from midline; greatest diagonal EW 55.4-63.8% (x = 57.7) of
IOD. Tympanum distinct, lower four-fifths visible, oriented dorsolateral^
with slight posterior inclination, separated from eye by distance 1.20-1.43
(x = 1.33) times TYM, which is 26.4-43.7% (x = 35.3) of ED; tympanic
annulus slightly elevated; supratympanic fold heavy, obscuring dorsal part
of tympanum.

Skin on dorsal surfaces of head and body finely spiculate; spicules also
present on upper lip and in tympanic region. Dorsal surfaces of limbs
smooth except for a few low spicules on proximal part of forearm in some
specimens. Flanks smooth. Skin on belly and ventral surfaces of thighs
coarsely granular; other ventral surfaces smooth. Vent directed
posteroventrally at upper level of thighs; cloacal sheath short, unmodified;
para- and subcloacal folds absent; no conspicuous tubercles on ventral
surfaces of thighs below vent.

Breadth of upper arm about one half that of forearm; humeral spine
absent; ulnar fold and tubercles absent; hand large; HnL 92-111% (x =
101) of HL; digits broad, bearing narrow lateral fringes, including outer
edge of Finger IV but not inner edge of Finger I; length of digits II < I < I V
< III; webbing basal between Fingers I and II; webbing formula for other
fingers II (VA-l-)— (2-3) III (2"-2)— (2~-2+) IV (Fig. 6C). Terminal discs
subtruncate; width of disc on Finger III about twice TYM; width of disc on
Finger I 62.5-81.3% (x = 72.4) of that on Finger III. Subarticular tubercles
not as wide as digits, round, elevated; all tubercles about same size, except
distal tubercle on Finger IV largest; small supernumerary tubercles
variously evident on proximal segments; palmar tubercle large, ovoid with
diagonal orientation, elevated, its length 15.8-18.8%- (x = 16.9) of
HnL. Thenar tubercle elliptical, elevated, much longer than palmar tubercle;
prepollex slightly enlarged; Type I nuptial excrescence present; prepollical
spine absent.

Hind limbs long, slender; TL 56.2-6 1.1% (x = 58.4) of SVL; FL46.4-
52.8% (x = 49.3) of SVL; dermal fringes and tarsal tubercles absent; inner
tarsal fold present; inner metatarsal tubercle elliptical, slightly elevated,
barely visible from above; outer metatarsal tubercle absent; toes moderately
slender, bearing lateral fringes, including outer edge of Toe V and inner
edge of Toe I; length of toes I < II < III < V < IV; toes about three-fourths
webbed; webbing formula I (1 -1 )— ( 1-1 + ) II ( 1-L)— 1 III 1 — (2-2) IV
(2~— 2) — ( 1-1+) V Terminal discs subtruncate: width of digit on Toe I 53.8-
61.5% (x = 58.7) of that on Toe IV. which is 75.0-92.8% (x = 83.2) that on
Finger III; subarticular tubercles not as wide as digits, round, elevated.

Vomerine tooth rows transverse to slightly inclined posteromedially,
narrowly separated medially, between ovoid choanae, each with one to five
(x = 2.9) teeth. Tongue broadly cordiform, barely free posteriorly. Vocal

NEW CENTROLENID FROGS FROM NORTHERN PERU 25

slit elongate, extending from posterolateral margin of tongue toward angle
of jaw; vocal sac single, median, subgular.

Color in life: Dorsum dull dark green to black; flanks paler green; tinge
of yellow on digits; parietal peritoneum white; visceral peritoneum clear;
heart not visible; bones green; iris dull gray with black reticulations, tinge
of red dorsal ly.

Color in preservative: General appearance dark olive-brown to black
dorsally, cream ventrally. Under 25x magnification, dense melanophores in
skin of dorsal surfaces of head and body; less dense melanophores on
dorsal surfaces of limbs, including digits and webbing. Edge of upper
eyelid pigmented. Venter cream with melanophores on margin of chin and
on ventral surfaces of forearms, shanks, and tarsi.

Measurements (in mm; mean and 1 SD in parentheses): SVL 20.8-23.2
(22.0 ± 1.1), TL 12.3-13.7 (12.9 ± 0.4), FL 9.9-1 1 .5 (10.9 + 0.4), HW 6.9-
8.4 (7.7 ± 0.4), HL 6.9-8.0 (7.5 ± 0.3), SL 2.5-3.4 (2.9 ± 0.2). ED 2.4-3.3
(2.8 ±0.2). EW 1.5-2.3 ( 1.9 ± 0.2). IOD 2.3-3.1 (2.6 ± 0.2), T-E 1.1-1.4
(1.2+1.1), TYM 0.9-1 .2 (1.0 + 0.1), HnL 6.8-8.4 (7.6 ± 0.4).

Distribution and ecology. — The species is known from only two sites
on the south slope of the northwest-southeast range of mountains
immediately to the north of Tarapoto. Both sites are on the road between
Tarapoto and Yurimaguas. The type locality is a rocky gorge into which
two streams plunge. The sides of the gorge are nearly vertical rocky walls,
and the floor of the gorge is littered with huge boulders. About 50 m beyond
the bases of the waterfalls, the two streams merge, and the resulting wider
stream flows over, under, and around these boulders, many of which
support a thin layer of moss. Small ferns and herbs also grow on some of
the boulders. Small trees, ferns, and a variety of herbs grow in patches of
soil on the floor of the gorge.

A few of the frogs were perched on ferns on boulders, but most were on
the boulders within 10 cm of the waterline at night. Several individuals
were on the bare, wet undersides of boulders over the stream. When
disturbed, the frogs ran rapidly across the boulders instead of leaping
away. The call is a soft chiip.

Etymology. — The specific name is an adjective derived from the Latin
saxum meaning rock and the Latin scandens meaning climbing: the name
alludes to the habit of these frogs of scampering across bare rocks.

Remarks. — Most centrolenids are active on vegetation above streams
at night. However, some species are known to inhabit rocks in streams.
Cochranella orejuela were found at night on rocks along a steep stream
bank or on rocks within the stream (Duellman and Burrowes, 1989).
Centrolene medemi were found on rock cliffs in the spray zone of waterfalls
(WED, pers. observ.), as were Cochranella euhystrix (Cadle and
McDiarmid (1990). Centrolene geckoideum and C. paezorum inhabit

26 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

similar rocky areas and apparently do not perch on vegetation (Lynch et al.,
1983: Ruiz-Carranza et al., 1986; WED, pers. observ.).

Cochranella tangarana new species

Figure 2

Holotype. — KU 211776, an adult male, from the west slope of Abra
Tangarana, 7 km (by road) northeast of San Juan de Pacaysapa (06°12'S,
76°44'W, 1080 m), Provincia Lamas, Departamento San Martin, Peru, one
of a series collected on 5 February 1989 by William E. Duellman, Rainer
Schulte, and John J. Wiens.

Paratopotypes — KU 21 1777 (male) and MHNSM 6179 (female); same
date and collectors.

Diagnosis. — A species in the Cochranella ocellata group characterized
by: (1) vomerine teeth present; (2) bones green; (3) parietal peritoneum
white; visceral peritoneum clear; (4) color in life, uniform dark green
dorsally; in preservative, dark lavender; (5) modal webbing on outer fingers
III 2— 1!/2 V; (6) modal webbing on foot I 1 — V/i II 1—2 III 1—2 VI 2-
1 V; (7) snout truncate in dorsal view and in profile; (8) dorsal skin finely
spiculate; (9) arms and legs lacking tubercles and dermal folds; (10)
humeral spine absent; (11) tympanum distinct, oriented dorsolaterally with
slight posterior inclination; (12) prepollex enlarged; nuptial excrescences
(Type I) present: prepollical spine absent; (13) pair of enlarged tubercles
below vent; ( 14) first finger equal to, or barely longer than, second.

Of the nine other species of Cochranella in the region, C. resplendens
differs from C. tangarana by having dermal folds on the limbs (as do
Centrolene azulae and puyoensis); C. ametarsia and C. ritae differ by
having black flecks on the dorsum. Four of the other species (C. cochranae,
flavopunctata, midas, and siren) differ from C. tangarana by having pale
spots or flecks on the dorsum. Only C. sa.xiscandens and C. spiculata
resemble C. tangarana in having a uniformly darkly pigmented dorsum
that is finely spiculate. Centrolenella tangarana differs from C.
sa.xiscandens (characters in parentheses) by having a lavender dorsum
(dark gray to black) in preservative, melanophores absent on ventral
surfaces of shanks and tarsi (present), and inner tarsal fold absent (present).
Centrolenella tangarana is most similar to C. spiculata which differs by
having a shorter snout that is round in profile, more conspicuous tympanum,
more conspicuous spicules on the head and body, and spicules on the dorsal
surfaces of the forearms and shanks.

Description. — Two adult males and one adult female; SVL 23.3, 22.5,
and 22.5 mm, respectively. Body moderately robust. Head distinct, about
as wide as long, round in outline as viewed from below; HW 31.5-35.5%
(x = 33.3) of SVL; HL 33.5-36.0% (x = 34.4) of SVL; snout truncate in

NEW CENTROLENID FROGS FROM NORTHERN PERU 27

dorsal view and in profile; SL 37.0-40.7% (x = 38.7) of HL: canthus
rostralis curved, indistinct, round in section; loreal region concave; lips
slightly flared anterior to orbit (Fig. 4D). Nostril ovoid, directed
dorsolaterally on protuberance; internarial area depressed. Eye moderately
large. ED 30.7-35.5% (x = 33.6) of HL, directed anterolateral^ at 37-38°
from midline; greatest diagonal EW 63.3-65.5% (x = 64.7) of
IOD. Tympanum distinct, completely visible, oriented dorsolaterally with
slight posterior inclination, separated from eye by distance 1.7-1.9 (x =
1.8) times TYM, which is 37.0-45.8% (x =40.7) of ED; tympanic annulus
slightly elevated; supratympanic fold weak, barely obscuring tympanic
annulus dorsally.

Skin on dorsal surfaces of head and body finely spiculate (less so in
female); dorsal surfaces of limbs smooth; small spicules on side of head
below orbit and in tympanic region. Flanks smooth. Skin on belly and
ventral surfaces of thighs coarsely granular; other ventral surfaces
smooth. Vent directed posteroventrally at upper level of thighs; cloacal
sheath short, unmodified: para- and subcloacal folds absent; one pair of
large, round, slightly elevated tubercles on ventral surfaces of thighs below
vent.

Breadth of upper arm about one half that of forearm; humeral spine
absent; ulnar fold and tubercles absent; hand moderately large; HnL 95-
101%- (x = 97.3) of HL; digits broad, bearing narrow lateral fringes,
including outer edge of Finger IV but not inner edge of Finger I; length of
digits I > II < IV < III; webbing basal between Fingers I and II; webbing
formula for other fingers II ( 1—2") — 3 III (2"-2)— (L-2) IV (Fig.
6D). Terminal discs subtruncate; width of disc on Finger III about 1.5
TYM; width of disc on Finger I 71.4-78.5% (x = 75.6) of that on Finger
III. Subarticular tubercles as wide as digits, round, elevated; all tubercles
about same size; supernumerary tubercles present on proximal segments;
palmar tubercle large, nearly quadrangular with diagonal orientation,
elevated, its length 15.8-16.4% (x = 16.2) of HnL. Thenar tubercle broadly
elliptical, barely elevated, longer than palmar tubercle; prepollex enlarged;
Type I nuptial excrescence present; prepollical spine absent.

Hind limbs long, slender; TL 57. 1-59.6% (x = 58.6) of SVL; FL 46.7-
48.9% (x = 47.7) of SVL; dermal fringes, tarsal tubercles, and tarsal fold
absent; inner metatarsal tubercle elliptical, slightly elevated, barely visible
from above; outer metatarsal tubercle absent; toes moderately slender,
bearing lateral fringes, including outer edge of Toe V and inner edge of Toe
I: length of toes I < II < III < V < IV; toes about three-fourths webbed;
webbing formula I (1-1 + )— (l'/:-2) II (1-L)— 2 III 1— (2"-2) IV (2-2)—
1 V Terminal discs subtruncate; width of digit on Toe I 61.5-66.7% (x =
63.9) of that on Toe IV which is 84.6-92.9% (x = 87.7) of that on Finger
III; subarticular tubercles as wide as digits, round, elevated.

28 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

Vomerine tooth rows elongate, posteromedially inclined, narrowly
separated medially, between ovoid choanae, each with two to five (x = 3.5)
teeth. Tongue narrowly cordiform, barely free posteriorly. Vocal slit
elongate, extending from posterolateral margin of tongue toward angle of
jaw; vocal sac single, median, subgular.

Color in life: Dorsum uniform dark green; anterior and posterior surfaces
of thighs pale green; digits pale yellow; vocal sac pale green; parietal
peritoneum white; visceral peritoneum clear; heart not visible; bones green;
iris bronze with black reticulations.

Color in preservative: General appearance dark lavender dorsally, cream
ventrally. Under 25x magnification, dorsal surfaces of head and body dark
lavender except for tips of spicules, which are unpigmented; moderately
dense melanophores in loreal, labial, and tympanic regions, and on dorsal
surfaces of limbs; less dense melanophores on flanks, dorsal surfaces of
Fingers III and IV and Toes III— V, and anterior and posterior surfaces of
thighs. Edge of upper eyelid pigmented. Venter cream; scattered
melanophores apically and posterolaterally on chin in one male (KU
211777).

Measurements (in mm; male holotype and male paratype, followed by
female paratype): SVL 23.3, 22.5, 22.5; TL 13.3, 13.4, 13.3; FL 11.1, 10.5,
1 1 .0; HW 7.6, 7.4, 8.0; HL 7.8, 7.6, 8. 1 ; SL 3.0, 3. 1 , 3.0; ED 2.4, 2.7, 2.8;
EW 1.7, 1.9, 1.9; IOD 2.6, 3.0, 2.9; T-E 1.2, 1.0, 1.1; TYM 1.1, 1.0. 1.1;
HnL 7.9, 7.3, 7.6.

Distribution and ecology. — The species is known only from the type
locality and another ravine about 0.5 km to the northeast. See the account
of Centrolene fernandoi for a description of the type locality.

All individuals were found at night perched on the dorsal surfaces of
ferns, leaves of trees, aroids, and other herbaceous plants at the edge of, or
overhanging, a small cascading stream in a narrow rocky ravine. All were
less than 2 m above the ground or water. The call is a harsh, high-pitched
"tzeeet."

Etymology. — The specific name is a noun in apposition and refers to the
type locality. Tangarana is the local name for the tree, Triplaris poeppigiana
(Polygonaceae), colonized by small red ants that have a painful sting.

Remarks.— A male 22.0 mm SVL (LSUMZ 37104) from the Cordillera
Colon, Departamento Amazonas, Peru, superficially is like Cochranella
tangarana, but it lacks vomerine teeth and has large pale spots on the
dorsum.

Hyalinobatrachium lemur new species
Figure 2

Holotype. — KU 211768, an adult male, from the west slope of Abra
Tangarana, 7 km (by road) northeast of San Juan de Pacaysapa (06°12'S,

NEW CENTROLENID FROGS FROM NORTHERN PERU 29

76°44'W. 1080 m), Provincia Lamas, Departamento San Martin, Peru,
obtained on 5 February 1989 by William E. Duellman.

Paratopotype. — KU 211769, collected with the holotype.

Diagnosis. — A species in the Hyalinobatrachium fleischmanni group
characterized by: (1) vomerine teeth absent; (2) bones white; (3) parietal
peritoneum clear; visceral peritoneum white; (4) color in life, pale green
with diffuse, dull yellow spots; in preservative, creamy white with scattered
melanophores: (5) modal webbing on outer fingers III 2 — 1 IV; (6) modal
webbing on foot I 1 — l1/: II 1 — 2 III 1 — 2 IV 2 — 1 V; (7) snout truncate in
dorsal view and in profile; (8) dorsal skin smooth; (9) arms and legs lacking
tubercles and dermal folds; (10) humeral spine absent; (11) tympanum
indistinct: (12) prepollex not enlarged: nuptial excrescences absent;
prepollical spine absent; (13) no enlarged tubercles below vent; (14) first
finger longer than second.

Hyalinobatrachium lemur differs from H. bergeri and H. pellucidum by
having smooth, instead of shagreened, skin on the dorsum and by having
more extensive webbing on Finger IV ( 1 free phalanx in H. lemur, as
opposed to 2 free phalanges in the others). Hyalinobatrachium munozorum
is like H. lemur in having smooth skin on the dorsum, but it differs by
having a round, instead of truncate, snout in profile, less webbing on Finger
IV (2 phalanges free), and smaller terminal discs on the fingers.

Description. — One adult male and one adult female: SVL 20.4 and 2 1 .3
mm, respectively. Body robust. Head not distinct from body, wider than
long, round in outline as viewed from below; HW 39.2, 37. 1 % of SVL; HL
30.9, 29.6% of SVL; snout slightly protruding, rounded with upper margin
truncate in dorsal view; truncate in profile; SL 44.4% of HL: canthus
rostralis slightly curved, indistinct, round in section; loreal region concave;
lips not flared anterior to orbit (Fig. 4A). Nostril ovoid, directed laterally
on low protuberance; internarial area barely depressed. Eye moderately
large, ED 36.5% of HL, directed anterolaterally at 35, 36° from midline;
greatest diagonal EW 60.0, 66.7% of IOD. Tympanum indistinct, oriented
dorsolaterally, separated from eye by distance 1.1, 1.3 times TYM, which is
30.4% of ED; tympanic annulus slightly elevated; supratympanic fold
weak, barely obscuring upper part of tympanum.

Skin on dorsal surfaces of head, body, and limbs smooth. Flanks smooth
dorsally. finely granular ventrally. Skin on belly and ventral surfaces of
thighs granular: other ventral surfaces smooth. Vent directed posteriorly at
upper level of thighs; cloacal sheath short, unmodified; para- and subcloacal
folds absent; one pair of elongate, elevated tubercles on ventral surfaces of
thighs below vent.

Breadth of upper arm about one half that of forearm; humeral spine
absent; ulnar fold and tubercles absent; hand large; HnL 96.8, 96.4% of
HL; digits broad, bearing broad lateral fringes; fringes narrow on outer
edge of Finger IV and inner edge of Finger I; length of digits II < I < IV <

30 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

III; fingers about one-half webbed; webbing formula I 2 — (2-3) II (l1/:-
2+)— 3 III (2—2) — ( 1 — 1 ) IV (Fig. 6A). Terminal discs round; width of disc
on Finger III equals twice TYM; width of disc on Finger I 80.8, 84.6% of
that on Finger III. Subarticular tubercles not as wide as digits, round, barely
elevated; all tubercles about same size; supernumerary tubercles absent;
palmar tubercle moderately large, round, indistinct, its length 13.2, 12.7%
of HnL. Thenar tubercle elliptical, diffuse, barely elevated, longer than
palmar tubercle; prepollex not enlarged; nuptial excrescence and prepollical
spine absent.

Hind limbs long, slender; TL 53.4, 49.5% of SVL; FL 53.5, 46.5% of
SVL; dermal fringes, tarsal tubercles, and tarsal fold absent; inner
metatarsal tubercle elliptical, barely elevated, visible from above; outer
metatarsal tubercle absent; toes moderately slender, bearing lateral fringes,
including outer edge of Toe V and inner edge of Toe I; length of toes I < II
< III < V < IV; toes about three-fourths webbed; webbing formula I (1-
D— (l1/: -2) II (1-1+)— (2-2) III 1—2 IV 2— (1-1+) V. Terminal discs
round; width of digit on Toe I 88.9, 77.8% of that on Toe IV. which is 64.3,
69.2% of that on Finger III; subarticular tubercles not as wide as digits,
round, barely elevated.

Vomerine teeth absent; choanae widely separated, ovoid. Tongue broadly
cordiform, barely free posteriorly. Vocal slit elongate, extending from
posterolateral margin of tongue toward angle of jaw; vocal sac single,
median, subgular, extending onto chest.

Color in life: Dorsum pale green with small, diffuse yellow spots; digits
pale yellow; parietal peritoneum clear; visceral peritoneum white; heart
visible; bones white; iris cream below, yellow above.

Color in preservative: General appearance white dorsally and cream
ventrally. Under 25x magnification, with exception of unpigmented areas
corresponding to yellow spots in life, scattered melanophores in skin of
dorsal surfaces of head. body, forelimbs. thighs, shanks, tarsi, fourth fingers,
and fourth and fifth toes. Other surfaces cream with no melanophores.

Measurements (in mm; holotype followed by paratype): SVL 20.4,
2 1 .3; TL 10.9, 1 1 .4; FL 10. 1 , 9.9; HW 8.0, 7.9; HL 6.3, 6.3; SL 2.8, 2.8; ED
2.3.2.3; EW 1.5. 1.5; IOD 2.3. 2.5; T-E 0.9, 0.6; TYM 0.7. 0.7: HnL 6.1,
6.6.

Distribution and ecology. — At the type locality, both individuals were
on the upper surfaces of leaves of trees 1.5-2.0 m above a stream in a
narrow ravine at night. The call is a soft "peep." See the account of
Centrolene fernandoi for a description of the type locality.

Etymology. — The specific name is a Latin noun in apposition meaning
ghost of the departed. The name refers to the characteristic loss of color in
preserved specimens of members of the genus Hyalinobatrachium.

NEW CENTROLENID FROGS FROM NORTHERN PERU 3 1

Remarks. — The paratype contains numerous, large, unpigmented ova.
Three specimens of Hyalinobatrachium (KU 217295-97 from Cataratas
Ahuashiyacu, 730 m, 14 km [by road] north of Tarapoto, Departamento
San Martin, Peru) are poorly preserved males, 21.5, 20.3, and 19.9 mm
SVL. They appear to have rounded snouts in profile, smooth dorsal skin,
two phalanges on Finger IV free of webbing, and comparatively small
terminal discs on the fingers. Thus, they are referred to H. munozorum, a
species that is widely distributed in the Amazonian lowlands and on the
lower slopes of the Andes from southern Colombia to central Peru.

DISCUSSION

Cadle and McDiarmid (1990) emphasized the frustrations of
understanding relationships among centrolenid frogs; this dilemma was
resolved partially by Ruiz-Carranza and Lynch (1991). who defined three
genera. According to their classification, frogs of the genus Centrolene
share the derived character state of a humeral spine in males. Those of the
genus Hyalinobatrachium are united by a single synapomorphy — a
bulbous, instead of the usual trilobate, liver. Centrolenids having the
plesiomorphic states of the absence of humeral spines in males and a
trilobate liver were relegated to the genus Cochranella.

The generic assignment of species known only from females can be a
problem. For example. Ruiz-Carranza and Lynch (1991) placed three
species known to them only from females — azulae, mariae, and
puyoensis — in Cochranella. Males of mariae have humeral spines (Martin
Hensel, pers. comm.); thus, the proper name for the species is Centrolene
mariae. Flores and McDiarmid ( 1989) considered azulae and puyoensis to
be closely related to mariae; if their suppositions are correct, those two
species also should be placed in Centrolene.

According to the classification of Ruiz-Carranza and Lynch (1991), as
amended herein, the centrolenid fauna of Peru consists of six species of
Centrolene (azulae, fernandoi, hesperium, lemniscatum, mariae, and
muelleri), 10 species of Cochranella (chancas, croceopedes, euhystri.x,
ocellata, phena.x, pluvialis, saxiscandens, spiculata, tangarana, and
truebae), and three species of Hyalinobatrachium (bergeri, lemur, and
munozorum).

All but three of the 19 currently recognized species of centrolenids in
Peru have been named in the past decade; one of us (WED) has seen
specimens of at least four additional unnamed species. As more biological
exploration is undertaken on the humid eastern slopes of the Andes and
extra- Andean ridges rising from the Amazonian lowlands, we expect many
more species of centrolenids to be discovered.

32 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155

Acknowledgments: Duellman's research on patterns of speciation and
biogeography of Andean anurans is supported by a grant (BSR 8805920)
from the National Science Foundation. We are grateful to our held
companions — Fernando M. Cuadros V., Michael E. Morrison, and John J.
Wiens — for enduring many wet nights in the quest for frogs. We are
grateful to Roy W. McDiarmid and John D. Lynch for helpful suggestions
and to the latter for providing us with a copy of the manuscript defining the
genera of centrolenid frogs. The illustrations were executed by Amy
Lathrop. and the photographic prints were made by John E. Simmons from
color slides. Erik R. Wild took most of the morphometric data. Permits for
collecting and exportation of specimens were provided by Ing. Gonzalo
Bravo Meji'a Muiioz and Biol. Jose Purisaca of the Direccion General
Forestal y de Fauna, Ministerio de Agricultura, Lima, Peru. We are grateful
to Gerardo Lamas and his staff at the Museo de Historia Natural,
Universidad Nacional Mayor de San Marcos, for many courtesies, and to
B. Anthony Luscombe, Asociacion de Ecologia y Conservacion, in Lima,
Peru, for logistic aid.

LITERATURE CITED

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(Anura: Centrolenidae) from northwestern Peru. Proc. Biol. Soc. Washington

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Cannatella, D. C, and W. E. Duellman. 1982. Two new species of Centrolenella,

with a brief review of the genus in Peru and Bolivia. Herpetologica 38:380-388.
Cannatella. D. C, and W. W. Lamar. 1986. Synonymy and distrihution of

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NEW CENTROLENID FROGS FROM NORTHERN PERU 33

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