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University of Kansas Publications MUSEUM OF NATURAL HISTORY

The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Pub- lications, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subject to critical review by intra- and extramural specialists; final acceptance is at the discretion of the publications committee.

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MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas

NUMBER 52, PAGES 1-11 APRIL 15, 1976

CENTROLENID FROGS FROM PERtJ

By William E. Duellman1

Although numerous species of the small frogs of the genus Centrolenella are known from Colombia and Ecuador, only one species has been reported from Peru. Boulenger (1918) named Hi/lella ocellata from Huancabamba, Departamento Pasco. Taylor and Cochran (1953) suggested that Boulenger's species was a cen- trolenid, and Goin (1964) placed ocellata in Centrolenella, an assignment verified by Lynch and Duellman (1973), who listed ad- ditional specimens from Departamento Ayacucho.

Field work in Peru in February 1975 resulted in the acquisition of an additional specimen of Centrolenella ocellata, in addition to series of two species previously known only from Ecuador, and two other unnamed species. These specimens, together with eight indi- viduals collected earlier, form the basis of this report. The termi- nology and methodology used herein is that of Lynch and Duell- man (1973).

Field work was made possible by grants from the National Geo- graphic Society and the National Science Foundation (GB 42481); I am grateful to my field companions Dana K. Duellman, John E. Simmons, and Linda Trueb for their patience in searching for elusive centrolenids. Thanks are due to Alice G. C. Grandison, British Museum (Natural History) (BMNH) and Douglas A. Ross- man, Louisiana State University Museum of Zoology (LSU) for the privilege of examining specimens in their care. Specimens in the Museum of Natural History at the University of Kansas are desig- nated KU. Representatives of all species are illustrated in Figure 1.

i Curator, Division of Herpteology, Museum of Natural History, and Professor, Depart- ment of Systematics and Ecology, University of Kansas, Lawrence, Kansas bt>U4t>.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Accounts of Species Centrolenella munozorum Lynch and Duellman

Centrolenella munozorum Lynch and Duellman, 1973:40 [Holotype. — KU 118054 from Santa Cecilia, Provincia Napo, Ecuador].

Diagnosis.— (I) Prevomerine teeth absent; (2) bones white; (3) parietal peritoneum clear; visceral peritoneum white; (4) color in life pale green with diffuse yellow spots; in preservative creamy white with minute reddish Drown flecks; (5) webbing between outer fingers III 2+-2 IV; (6) webbing on foot I 1-1X II VA-YA III 1/2-2M IV 2K-1/2 V; (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.

Centrolenella munozorum differs from other Peruvian species by having the heart visible through the ventral body wall and by being white in preservative. The only other species on the eastern slopes of the Andes having these characteristics, C. pellucida, differs by having ulnar, tarsal, and anal folds.

Distribution, — This species is known from three localities in Peru, two in cloud forest on the eastern front of the Andes and one in lowland tropical rainforest. The species has a known elevational range of 200-1840 m. The southernmost locality (Rio Cosnipata, Departamento Cuzco) is approximately 1600 kilometers (airline) south-southeast of the Ecuadorian localities on the Rio Aguarico.

Remarks. — The specimens from the Rio Piene and Rio Cosnipata have slightly less webbing on the hands and feet than do those from Ecuador. Also, the Peruvian frogs are slightly larger than those from Ecuador. The snout-vent length in eight Peruvian males is 22.7-25.7 (x = 24.1) mm and in five Ecuadorian males, 18.8-20.5 (x : 19.7) mm. One female from Ecuador has a snout-vent length of 20.7 mm, whereas two Peruvian females have snout-vent lengths of 23.3 and 24.4 mm. The coloration in life is the same in Ecua- dorian and Peruvian frogs. Although the differences in size and in the amount of webbing between the two samples are of a magni- tude usually exhibited by different species of Centrolenella, the enormous distance between the samples and the absence of other structural or color differences are suggestive of possible clinal varia- tion in size and amount of webbing.

The specimens from the Rio Piene and Rio Cosnipata were found in February 1975. Males were calling from the upper and lower surfaces of herbaceous leaves up to 2 m above small rivulets. At the Rio Piene one male was calling adjacent to a clutch of eggs on the underside of a leaf; the eggs had clear jelly and pale green yolks.

At the Rio Cosnipata C. munozorum was found with C. ocellata, C. spiculata, and C. truebae, and at the Rio Piene C. munozorum occurred with C. tiren,

CENTROLENID FROGS FROM PERU

Fig. 1. — Upper left: Centrolenella munozorum, KU 162248, 9, 24.4 mm. Upper right: C. ocdlata, KU 162262, 6, 25.4 mm. Center: C. siren, KU 162263, $, 22.8 mm. Lower left: C. spicuhta, KU 162283, 3, 21.7 mm. Lower right: C. truebae, KU 162268, <3 , 23.6 mm.

Centrolenella ocellata (Boulenger)

Hylella ocellata Boulenger, 1918:433 [Holotype.— BMNH 1912.11.1.19 from

Huancabamba, Departamento Pasco, Peru]. CoclirancUa ocellata — Taylor and Cochran, 1953:1628. Centrolenella ocellata — Goin, 1964:6.

Diagnosis. — (1) Prevomerine teeth absent; (2) bones green; (3) parietal peritoneum white; visceral peritoneum clear; (4) color in life dark green with pale bluish white ocelli; in preservative, lavender with cream ocelli; (5) webbing between outer fingers III 3-2% IV; (6) webbing on foot I 2-2M II 2-1M III 1^-3" IV 3-2 V; (7) snout round in dorsal and lateral profiles; (8) dorsal skin sha- greened; (9) arms and legs lacking dermal folds; (10) humeral spine absent in males; (11) tympanum entirely visible, directed dorsolaterally with slight posterior inclination.

4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Centrolenella ocellata differs from other Peruvian centrolenids by having moderately large dark-edged pale spots and no dark flecks on the dorsum and only basal webbing between the outer fingers. It superficially resembles the Ecuadorian C. cochranae, which has less webbing between the outer fingers, a truncate snout and much smaller ocelli on the dorsum.

Distribution. — This species is known from three localities at ele- vations of 1630-1700 m along the Cordillera Oriental in central and southern Peru. Two localities, Rio Cosnipata and Huanhuachayocc are in cloud forest on the eastern front of the Andes, whereas Huancabamba is in a north-south valley (Amazonian drainage).

Remarks.— The holotype of C. ocellata (BMNH 1912.11.19) is a female having a snout-vent length of 29 mm; the outer finger is nearly one-half webbed. Three additional specimens are males having snout- vent lengths of 21.0, 24.5, and 25.1 mm. In each the outer finger is about one-fourth webbed. The specimen from the Rio Cosnipata was green with small bluish white flecks. The margin of the upper lip was creamy white. The heart was not visible, and the iris was grayish white with fine black reticulations.

At the Rio Cosnipata one male was perched on a herb in cloud forest on the night of 10 February 1975. Three other species of centrolenids (C. munozorum, C. spiculata, and C. truebae) were found at the same locality.

Centrolenella siren Lynch and Duellman

Centrolenella siren Lynch and Duellman, 1973:54 [Holotype. — KU 146610 from the Rio Salado, Provincia Napo, Ecuador].

Diagnosis. — (1) Prevomerine teeth absent; (2) bones green; (3) parietal peritoneum white; visceral peritoneum clear; (4) color in life green with white flecks; in preservative, lavender with creamy white flecks; (5) webbing between outer fingers III 3-2/2 IV; (5) webbing on foot I 2-28 II 2-28 III 18-3 IV 3-2 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 two-thirds of tympanum visible, directed posterolaterally, inclined dorsally.

This small species with minute pale flecks differs from all other Peruvian species principally in coloration. The only other Peruvian species with discrete pale dorsal spots is C. ocellata, which has much larger spots, broader discs on the fingers, and a round snout. Two species, C. flavomaculata and C. midas, occurring on the Ama- zonian slopes of Ecuador are like C. siren in coloration, but both have much more webbing between the fingers.

Distribution. — Centrolenella siren is known from several locali- ties at elevations of 1400-1740 m on the Amazonian slopes of Ecuador and from the Rio Piene at an elevation of 1840 m on the Amazonian slopes of Departamento Ayacucho, Peru.

CENTROLENID FROGS FROM PERO 5

Remarks. — The Peruvian specimens include four adult males and one juvenile. Ecuadorian males have snout-vent lengths of 19.8-22.0 mm (x = 20.8, N = 14); those from Peru are 20.8-23.5 mm (x = 22.2, N = 4) in length. The only other noticeable dif- ference is that Ecuadorian specimens have yellow flecks instead of white, as in the Peruvian frogs. The specimens from the Rio Piene had a white margin to the upper lip; the vocal sac was green, and the iris was silvery white with black reticulations.

At the Rio Piene males of C. siren were calling at night from dense herbaceous growth above a low-gradient stream. The short call consists of three quickly repeated notes and is distinct from the higher-pitched single note produced by C. munozorum, which was calling at the same time.

Centrolenella spiculata new species

Holotype.—KU 162284, an adult male, 21.8 mm, from the Rio Cosiiipata, 4 kilometers southwest of Santa Isabel, Departamento Cuzco, Peru, 1700 m, obtained on 10 February 1975 by Linda Trueb.

Parotopotype. — KU 162283, collected with the holotype.

Diagnosis. — (1) Prevomerine teeth 1-4; (2) bones green; (3) parietal peritoneum white; visceral peritoneum clear; (4) color in life dark green; in preservative, dark lavender; (5) webbing be- tween outer fingers III 2-1 IV; (6) webbing on foot I 1-1M II 1-1& III 1-2 IV 2-1 V; (7) snout round in dorsal and lateral profiles; (8) dorsal skin bearing many minute spicules; (9) arms and legs lacking dermal folds; (10) humeral spine absent in males; (11) lower three-fourths of tympanum visible, directed dorsolaterally, slightly inclined posteriorly.

The combination of a uniformly dark green, finely spiculate dorsum, extensively webbed fingers with large discs, and the ab- sence of humeral spines in males readily distinguishes C. spiculata from all other Andean species of Centrolenella. The Ecuadorian C. megacheira also has spiculate dorsal skin but differs from C. spiculata in being much larger and by having little webbing be- tween the fingers and black spots on the dorsum.

Description. — Adults moderately small; snout-vent length 21.7- 22.8 mm (x = 22.2, N = 7) in males; females unknown. Head slightly wider than body; width of head 35.5-36.9% (x 36.2, N = 7) of snout- vent length; snout very short, round in dorsal and lateral profiles, canthus round; loreal region barely concave; lips not flared; nostrils nearly terminal on snout, not protuberant; inter- narial area not depressed. Eye moderately large, directed antero- laterally. Supratympanic fold barely evident; lower three-fourths of tympanum visible, directed dorsolaterally, slightiy inclined pos- teriorly. Prevomerine dentigerous processes short, transverse be- tween choanae, each bearing 0-2 teeth; choanae small, ovoid;

6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

tongue broadly cordiform, shallowly notched posteriorly, barely free behind; vocal slits extending from midlateral base of tongue towards angles of jaws.

Humeral spine absent, forearm robust; ulnar fold and tubercles absent; fingers short with lateral fringes; first finger slightly longer than second; fourth finger slightly shorter than third; webbing vestigial between first and second fingers; webbing formula for other fingers II lX-(2X-3) III (lK-2)-(l-H) IV; discs large, broad, truncate; subarticular tubercles large, subcorneal, simple; super- numerary tubercles absent; palmar tubercle low, ovoid; nuptial excrescences absent. Hind limbs moderately robust; length of tibia 55.2-59.9% (x = 57.0, JV::7) of snout-vent length; tarsal folds and tubercles absent; inner metatarsal tubercle small, elongate; outer metatarsal tubercle absent; subarticular tubercles small, round; supernumerary tubercles absent; feet about three-fourths webbed; webbing formula I 1-18 II l-(lM-2) III (l-13t)-(l-2) IV (lX-2)-l V; discs on toes smaller than those on fingers, truncate.

Skin on dorsal surfaces smooth, with minute white spicules on dorsum of body and dorsal and lateral surfaces of head; skin on belly and proximal surfaces of thighs areolate; other surfaces smooth; anal opening directed posteriorly at upper level of thighs; anal tubercles and folds absent.

Color in preservative: dorsal surfaces, exclusive of fingers and toes, uniformly dark lavender; other surfaces cream.

Color in life: dorsum dark green; tips of digits yellowish green; margin of upper lip greenish white; parietal peritoneum white; visceral peritoneum clear; heart not visible; bones green; iris dull bronze with fine black reticulations.

Distribution. — CentroleneUa spiculata is known from the type locality at 1700 m and from San Jose, Rio Santa Rosa, 1000 m, Departamento Ayacucho, Peru.

Remarks. — At the Rio Cosiiipata, two individuals were calling at night from the upper sides of leaves of herbs adjacent to a small stream. The call is a short "peep." The type locality is between kilometer markings 150 and 151 on the road from Paucartambo to Pilcopata. In this area, which supports cloud forest, many streams cascade down steep slopes on the north side of the Rio Cosiiipata.

Etymology. — The specific name is derived from the Latin spicu- lum, meaning a small, sharp structure.

CentroleneUa truebae new species

Holotype.—KU 16226S, an adult male, 23.6 mm, from Rio Cosiiipata, 4 kilometers southwest of Santa Isabel, Departamento Cuzco, Peru, 1700 m, one of a series collected on 10-11 February 1975 by William E. Duellman, John E. Simmons, and Linda Trueb.

Paratopotypes.—KU 162269-S1, collected with the holotype.

Diagnosis. — (1) Prevomerine teeth absent; (2) bones green;

CENTROLENID FROGS FROM PERU 7

(3) parietal peritoneum white; visceral peritoneum clear; (4) color in life green with greenish black flecks and creamy white spots; in preservative, lavender with black flecks and small white spots; (5) webbing between outer fingers III 3-3 IV; (6) webbing on foot I 2-2£ II lfc-3 III 2-3 IV 3-2 V; (7) snout round in dorsal view, truncate in lateral profile; (8) dorsal skin shagreened with small spinules; (9) arms and legs lacking dermal folds; (10) humeral spine absent in male; (11) lower three-fourths of tympanum visi- ble, directed posterolaterally with slight dorsal inclination.

Ccntrolenella truebae differs from other species on the Ama- zonian slopes, except C. pipilata and ocellata, by having dark flecks and pale spots on the dorsum. Centrolenelhi ocellata has cream- centered ocelli on the dorsum. Ccntrolenella pipilata has dermal fringes on the outer edges of the hands, forearms, and feet, and males have a humeral spine. Other species on the Amazonian slopes are either uniform green or are green with yellow flecks or diffuse spots.

Description. — Adults moderately large; snout- vent length 22.6- 24.8 mm (x = 23.3, N = 13) in males, 24.8 mm in one female. Head noticeably wider than body; width of head 35.8-37.2% (x = 36.3, N = 14) of snout-vent length; snout short, round in dorsal view, truncate in lateral profile; canthus round; loreal region barely concave; lips not flared; nostrils three-fourths distance from eye to tip of snout, protuberant laterally; internarial area slightly de- pressed. Eye large, protuberant, directed more anteriorly than laterally. Supratympanic fold weak; lower three-fourths of tym- panum visible, directed posterodorsally with a slight dorsal inclina- tion. Prevomerine dentigerous processes and teeth absent; choanae moderately large, ovoid; tongue broadly cordiform, shallowly 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 and second fingers equal in length; fourth finger somewhat shorter than third; lateral fringes present on fingers; webbing absent be- tween first, second, and third fingers; webbing formula for outer fingers III 3-3 IV; discs broad, rounded; subarticular tubercles moderately large, flattened, simple; supernumerary tubercles minute; present on proximal segments of digits; palmar tubercle moderately large, ovoid; nuptial excrescences absent. Hind limbs slender; length of tibia 55.1-60.0% (x = = 57.7, N - ■ 14) of snout-vent length; tarsal folds and tubercles absent; inner metatarsal tubercle small, flattened, elliptical; outer metatarsal tubercle small, low ovoid, sub- articular tubercles small, round; supernumerary tubercles absent; toes about one-half webbed; webbing formula I 2-(2/4-3) II (lf-2)- (2X-3) III (lK-2)-(3--3) IV (2K-3)-(lX-2) V; discs round, slightly smaller than those on fingers.

Skin on dorsal surfaces of head and body smooth with small

8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

spicules most numerous on side of head posterior to eye; skin on belly and posteroventral surfaces of thighs granular; other surfaces smooth; anal opening directed posteriorly at upper level of thighs; anal tubercles and folds absent.

Color in preservative: dorsal surfaces of head, body, forearms, thighs, shanks, and feet, exclusive of toes 1-3 and distal parts of toes 4-5 pale lavender with many dark lavender flecks and a few small cream spots; other surfaces, including margin of upper lip and sharply demarcated flanks, cream.

Color in life: dorsal surfaces green with greenish black flecks and small creamy white spots; tips of digits yellowish green; margin of upper lip and axilla cream; chest white; heart not visible; bones green; iris pale gold with black reticulations.

Distribution. — This species is known only from the type locality in cloud forest on the Amazonian front of the Andes in southern Peru.

Remarks. — Individuals were perched on and calling from the upper sides of ferns along a dripping cliff at night. The call is a series of short cricket-like chirps. See the account of C. spiculata for a description of the type locality.

Etymology. — The species is named for Linda Trueb who first traced the call to this Centrolenella and whose efforts in Peru and elsewhere in South America have resulted in many specimens of centrolenid frogs.

Discussion

Lynch and Duellman (1973) commented on the association of Ecuadorian species of Centrolenella with the species groups defined by Savage (1967). Two general groups of Centrolenella are readily recognized. The fleischmanni group, characterized by white bones, colorless parietal peritoneum, and a white ground color in pre- servative, is represented in Peru solely by C. munozorum. The other four species known from Peru belong to the prosoblepon group, characterized by green bones, white parietal perito- neum, and a lavender ground color in preservative. Lynch and Duellman (1973) associated C. siren with C. auclax, cochranae, flavovunctata, miclas, and prosol)lepon, and defined another as- semblage consisting of C. bucklcyi, granclisonae, giffithsi, mega- cheira, peristicta, and pipilata. Three Peruvian species, C. ocellata, spiculata, and truebae, fit into the latter assemblage. As defined by Lynch and Duellman, the assemblages are only loose phenetic groupings, because knowledge of the systematics of South American centrolenids is still too fragmentary to permit a phylogenetic syn- thesis.

With the exception of the specimen of C. munozorum from Finca Panagua at 200 m, all specimens of Peruvian Centrolenella come from localities between elevations of 1000-1840 m on the

CENTROLENID FROGS FROM PERtj

()

Kilometers Contours in Meters

Fig. 2. — Map of east-central Peru showing localities mentioned in text: 1. Finca Panagua, 2. Huancabamba, 3 Tutumbaro, 4. San Jose, 5. Huanhua- chayocc, 6. Tambo, 7. Ayacucho, 8. Santa Isabel, 9. Pilcopata. With the ex- ception of those pertinent to the text, only the major rivers are shown. Based on Mapa Fisico Politico del Peru, 1:1,000,000, Instituto Geografico Militar, Lima, 1973.

Amazonian slopes of the Cordillera Oriental of the Andes (Fig. 2). According to Tosi's (1960) application of Holdridge's Life Zone classification, the montane localities where centrolenids were found in Ayacucho and Cuzco are located in very humid lower montane forest and very humid subtropical forest, respectively. Huanca- bamba is located in humid subtropical forest, and Finca Panguana is in humid tropical forest.

The present summary of Peruvian centrolenids can be consid- ered as nothing more than a first step on a long stairway winding through the extensive lowland and montane forests of Amazonian Peru before ascending to a plateau of knowledge comparable with that of Central American centrolenids. I suspect that several species now known from Ecuador will be found in Peru and that many species await discovery.

Key to Peruvian Species of Centrolenella

1. Dorsum in life pale green with diffuse yellow spots, in pre- servative creamy white; visceral peritoneum white; heart visible; bones white C. munozorum

10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Dorsum in life dark green with or without pale or dark flecks, in preservative lavender; visceral peritoneum clear; heart not visible; bones green 2

2. Dorsal pattern consisting of many greenish black (dark pur- ple in preservative) flecks and a few cream spots C. truebae Dorsum uniform green or marked with pale spots or flecks _ 3

3. Dorsum uniform green C. spiculata

Dorsum with pale flecks or spots 4

4. Dorsal flecks minute; snout truncate C. siren

Dorsal spots larger, bordered by dark pigment; snout round

C. ocellata

Resumen

La fauna de sapos centrolenidos del Peru conocida hasta ahora esta compuesta por cinco especies. Dos de ellas son descritas aqui como nuevas: Centrolenella spiculata y C. truebae, las cuales pro- vienen del Rio Cosnipata en el Departamento del Cuzco; la primera tambien se encuentra en el Rio Santa Rosa en el Departamento de Ayacucho.

De las otras especies, Centrolenella ocellata (Roulenger), des- crita para Huancabamba, Departamento de Pasco, tambien se la encuentra en las laderas Amazonicas, en su extension hasta el Rio Cosnipata. Otras dos especies, C. munozorum y C. siren, conocidas previamente solo para el Ecuador, extienden su rango hasta el Peru. Centrolenella munozorum se encuentra en el Rio Llullapichis (Huanuco), Rio Cosnipata (Cuzco), y Rio Piene (Ayacucho), mientras C. siren se encuentra en el Rio Piene.

Centrolenella munozorum es un miembro del grupo fleisch- manni; mientras que las restantes especies peruanas pertenecen al grupo prosoblepon, Todas las especies peruanas se encuentran a lo largo de arroyos en selvas humedas tropicales y subtropicales, ambientes que probablemente albergan a varias otras especies de Centrolenella que todavia permanecen sin ser descritas.

Specimens Examined

Only Peruvian specimens are listed; for Ecuadorian specimens see Lynch and Duellman ( 1973).

Centrolenella munozorum. — PERU: Ayacucho: Tutumbaro, Rio Piene, 1840 m, KU 162251-57, 162258 (cleared and stained), 162259-60 (eggs), 162261 (tadpoles). Cuzco: Rio Cosnipata, 4 km SW Santa Isabel, 1700 m, KU 162248-49. Huanuco: Finca Pangnana, Rio Llullapichis, 4-5 km upstream from Rio Pachitea, 200 m, KU 154749.

Centrolenella ocellata. — PERU: Ayacucho: Huanbuachayocc on Tambo- Valle de Apurimac trail, 1630 m, LSU 25989-90. Cuzco: Rio Cosnipata, 4 km SW Santa Isabel, 1700 m, KU 162262. Pasco: Huancabamba, 1700 m, RMNH 1912.11.1.19.

CENTROLENID FROGS FROM PERU 11

Ccntrolenella siren. — PERU: Ayacuclw: Tutumbaro, Rio Piene, 1840 m, KU 162263-67.

Centrolenella spiculata. — PERU: Ayacucho: San Jose, Rio Santa Rosa, 1000 m, LSU 25978-82. Cuzco: Rio Cosmpata, 4 km SW Santa Isabel, 1700 m, KU 162283-84.

Ccntrolenella truebae. — PERU: Cuzco: Rio Cosnipata, 4 km SW Santa Isabel, 1700 m, KU 162268-81, 162282 (cleared and stained).

Literature Cited

Boulenger, G. A. 1918. Descriptions of new South-American batrachians.

Ann. Mag. Nat. Hist, 9(2) :427-433. Goix, C. J. 1964. Distribution and synonymy of Ccntrolenella fleischmanni in

northern South America. Herpetologica, 20:1-8.

Lynch, J. D., Duellman, W. E. 1973. A review of the centrolenid frogs of

Ecuador, with descriptions of new species. Occas. Pap. Mus. Nat. Hist.

Univ. Kansas, 16:1-66. Savage, J. M. 1967. A new tree-frog ( Centrolenidae ) from Costa Rica.

Copeia, 1967(2) : 325-331. Taylor, E. H., Cochrax, D. M. 1953. Frogs of the family Centrolenidae from

Brasil. Univ. Kansas Sci. Bull., 35:1625-1656.

Tosi, J. A., Jr. 1960. Zonas de vida natural en el Peru. Inst. Interamer. Cien. Agricol. Bol. Tech., 5:1-271.

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