Two New Species of Centrolenella, with a Brief Review of the Genus in Perú and Bolivia David C. Cannatella; William E. Duellman Herpetologica, Vol. 38, No. 3. (Sep., 1982), pp. 380-388. Stable URL: http://links.jstor.org/sici?sici=0018-0831%28198209%2938%3A3%3C380%3ATNSOCW%3E2.0.CO%3B2-8 Herpetologica is currently published by Herpetologists' League.

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Acknowledgments.-The Instituto Nacional Forestal (INAFOR) graciously issued permits for collecting in Guatemala. We thank Gretchen Bracher who skillfully prepared Figs. 1 , 2 , 4 and 5, and John Simmons for his darkroom expertise. Finally, we are grateful to Ron Savage for his enthusiastic field assistance that led to the discovery of Ptychohyla panchoi.

DUELLMAN, W. E. 1961. Descriptions of two species of frogs, genus Ptychohyla. Studies of American hylid frogs, V. Univ. Kansas Publ. Mus. Nat. Hist. 13:349-357. . 1963. A review of the Middle American tree frogs of the genus Ptychohyla. Univ. Kansas Publ. Mus. Nat. Hist. 15:297349. . 1970. The hylid frogs of Middle America. Univ. Kansas Mus. Nat. Hist. Monogr. 1:l-753.

GOSNER,K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica 16:183-190. HOLDRIDGE,L. R. 1964. Life Zone Ecology. Tropical Science Center, San Jose, Costa Rica. WASSERSUG,R. 1980. Internal oral features of larvae from eight anuran families: functional, systematic, evolutionary and ecological considerations. Univ. Kansas Misc. Publ. 68:l-146. Accepted: 25 April 1982 Associate Editor: Stephen Tilley

Museum of Natural History and Department of Systematics and Ecology, The University of Kansas, Lawrence, KS 66045, USA

Herpetologica, 38(3), 1982, 380-388 @ 1982 by The Herpetologists' League, Inc.

TWO NEW SPECIES OF CENTROLENELLA, WITH A BRIEF REVIEW OF THE GENUS IN PERU AND BOLIVIA DAVIDC. CANNATELLA AND WILLIAME. DUELLMAN ABSTRACT: Ten species of centrolenid frogs of the genus Centrolenella are recorded from Peni and Bolivia. Two species are named as new--C. pluuialis and C. phenax, both from cloud forests on the Amazonian slopes of the Andes in central and southern Peni. Limited material indicates the presence of five undescribed species in Peni. A key to the identification of named species is provided. Key words:

Anura; Centrolenidae; Centrolenella; Peni; Bolivia

IT is becoming apparent that centrolenid frogs of the genus Centrolenella are highly diverse in the American tropics, especially in the cloud forests on the slopes of the Andes. Duellman (1977) listed 55 s ~ e c i e sof Centrolenella. Ten additional sbecies have been named since that list (taxa named through 1974), and we are aware of many other new species awaiting descri~tion. The purposeLof this paper is to summarize our existing knowledge of this genus in the southwestern part of its range

in Peni and Bolivia. We describe two new species, review the status of the species already described, and provide a key for their identification. The definition of characters and arrangement of diagnoses follow Lynch and Duellman (1973); the webbing formula follows Savage and Heyer (1967). Natural history and color data are from the field notes of William E. Duellman. Acronyms for museum collections are: AMNH, American Museum of Natural History; BMNH, British Museum (Natural History); KU, University

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FIG. 1.-Top: Centrolenella phenax, KU 162263, holotype, adult male, SVL 21.8 mm. Bottom: Centrolenelk pluuialis, KU 173224, holotype, adult male, SVL 25.6 mm.

of Kansas Museum of Natural History; LSU, Louisiana State University Museum of Zoology. Centrolenella pluvialis sp. nov. Ho1otype.-KU 173224 (Fig. l), adult male, 25.6 mm SVL, from Pistipata, Rio Umasbamba (= Rio Santa Maria), 12 km SE Huyro, 1820 m, Departamento Cuzco, Peni (72'30' W, 13'03' S), obtained on 10 January 1977, by David C. Cannatella and William E. Duellman. Paratypes.-KU 173225-27, same data as the holotype. Referred specimens.-KU 173488 (cleared and stained skeleton), LSU 32464, Peni: Cuzco: Huyro. Diagnosis.-41) Prevomerine teeth absent; (2) color of bones in life unknown; (3) anterior half of parietal peritoneum white, elsewhere clear; visceral perito-

381

neum clear; (4) color in life green with pale cream dorsal flecks; in preservative, lavender with white spots; (5) modal webbing formula of fingers I1 2 + - 3 + I11 2.5--2.5 IV; (6) modal webbing formula of toes I 1.&2+ I1 1.5--2.5 I11 1.5--3IV 3--2- V; (7) snout weakly truncate in dorsal and lateral views; (8) dorsal skin spiculate; (9) limbs lacking dermal folds; (10) humeral spine absent; (11) lower three-fourths of tympanum visible, dorsolaterally oriented with weak posterior inclination. Ten species of Andean Centrolenella also have spiculate, lavender dorsal surfaces: Of these, C. buckleyi, C. lynchi, C. johnelsi and C. grandisonae differ from C. pluvialis by having humeral spines present in the males. Centrolenella truebae and C. megacheira can be distinguished from the new species by the dark flecks on the dorsum. Centrolenella spiculata differs by having prevomerine teeth and more extensive webbing; C. ocellata has large dorsal ocelli and a snout that is round in lateral profile. Centrolenella siren can be distinguished from C. pluvialis by a more truncate snout and yellow dorsal spots. Centrolenella bejaranoi differs from C. pluvialis in that the first finger is shorter than the second and the white dorsal spots in C. pluvialis are much larger than those of C. bejaranoi. Furthermore, the distribution of the white pigment of the parietal peritoneum is much more extensive in C. plvvialis. Description.-Adult males of moderate size for centrolenid frogs; SVL 24.9-26.5 mm (X = 25.4, n = 5); females unknown. Head slightly wider than body; snout short, weakly truncate in dorsal and lateral views; canthus round; loreal region concave; lips not flared; nostrils nearly terminal on snout, not flared, directed laterally; internarial region slightly concave. Eye moderately large, directed anterolaterall~.Supratympanic fold distinct; lower three-fourths of tympanum visible, dorsolaterally oriented with weak posterior inclination. Prevomerine teeth absent; choanae round; tongue circular, with

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slight indentation posteriorly; vocal slits present in males, extending from angle of jaw to midlateral edge of tongue. Humeral spine absent; dermal folds lacking; order of fingers from shortest to longest 1=24-3; fingers barely webbed, with slight fringe and truncate discs; webbing formula I1 (2-2+)--3+ I11 (2.53)-2.5 IV; subarticular tubercles small, round, single; supernumerary tubercles absent; palmar tubercle round, single; thenar tubercle elliptical. Hind limbs slender; tibia length 58.159.6% of SVL; tarsal fold absent; inner metatarsal tubercle small, flattened, elliptical; outer metatarsal tubercle absent; subarticular tubercles small, round; supernumerary tubercles absent; toes about one-half webbed; webbing formula I (1.52')-2' I1 (I+-1.5)-(2+-2.5) I11 l . S (2.5-3-) IV (3--3+)-2- V; discs rounded to weakly truncate, smaller than those on fingers. Skin on dorsal surfaces of body and limbs bearing spicules of heterogenous sizes; spicules also present on upper eyelids, dorsal surface of snout, upper lip below the eye, and on the heels and anal region; skin of belly and ventral thighs granular; pair of large distinct subanal tubercles present; anal opening directed posteriorly at upper level of thighs; anal tubercles present, spiculate. Color in preservative.-Dorsal surfaces lavender with white spots. Upper labial stripe with distinct white pigment. Ventral surfaces white or translucent. Color in life.-Dorsal surfaces green with pale cream flecks; labial stripe and discs cream; anterior half of parietal peritoneum white, elsewhere clear; visceral peritoneum clear; color of bones unknown. Iris pale bronze with black flecks. Distribution.-The species is known only from the type locality on the eastern slopes of the Andes in southern Peni (Fig. 2, locality 13). Etymology.-The specific epithet (Latin) is an adjective meaning "of rain", and alludes to the rainy conditions coincident with centrolenid calling activity.

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Remarks.-The t y p e locality, t h e Cooperativa Pistipata, consists of coffee and tea plantations with remnants of cloud forest. The frogs were taken near our campsite at a bridge over the Rio Umasbamba (Rio Santa Maria). They were calling on a rainy night, on herbs on a dripping cliff and from the upper sides of leaves over water, all adjacent to the river. The call is a shrill "bree-ee-ee-eeeep". The egg clutches were on the tips of the upper surfaces of the leaves. This area is also the type locality of Eleutherodactylus mendax (Duellman, 1978). Centrolenella phenax sp. nov. Ho1otype.-KU 162263 (Fig. l ) , adult male, 21.8 mm SVL, from Tutumbaro, Rio Piene, 1840 m, Departamento Ayacucho, Peni (73'55' W, 12'42' S), obtained on 20 February 1975, by William E. Duellman and John E. Simmons. Paratypes.-KU 162264, 162266-67, obtained on 20-21 February 1975, same locality and collectors as the holotype. Diagnosis.-(1) Prevomerine teeth absent; (2) bones green in life; (3) parietal and visceral peritoneum clear; (4) in life, dorsal surfaces dark green with creamy white flecks; in preservative, lavender with white spots; (5) very scant webbing between fingers III-IV; (6) modal webbing formula of toes I 2--2.5 I1 1.5-4I11 1.5-3+ IV 3 1 2 - V; (7) snout weakly truncate in dorsal and lateral views; (8) dorsal skin spiculate (see Discussion); (9) limbs lacking dermal folds; (10) humeral spine absent; (11) lower four-fifths of tympanum visible, dorsolaterally inclined with weak posterior orientation. As noted in the account of C. pluvialis, several other species possess lavender spiculate dorsa: Centrolenella buckleyi, C. lynchi, C. johnelsi and C. grandisonae differ from C. phenax in having humeral spines in the males. Dark dorsal flecks distinguish C. truebae and C. megacheira from this new species. Centrolenella ocellata has large dorsal ocelli, a snout that is round in lateral profile, a longer snout, more vertically oriented

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383

I

I

FIG.2.-Map of Ecuador, Peni and Bolivia showing localities of Centrolenella (complete citations in text): 1, Santa Cecilia; 2, Rio Yasuni; 3, Cordillera Col6n; 4, Tocache; 5, Utiquinia-Tapiche; 6, Serrania de Sira; 7, Finca Panguana; 8, Huancabamba; 9, Valle de Perene; 10, Tutumbaro; 11, Huanhuachayocc; 12, San Josk; 13, Pistipata; 14, Huyro; 15, Rio Cosiiipata; 16, Quincemil, 40 km E; 17, Villa Tunari, 58.1 km SW.

tympanum, and relatively longer tibia; tibia length/SVL = 63.3 and 65.6%in the two specimens of ocellata, in contrast to 56.0-57.9% in phenax. Also, the combi-

nation of clear visceral and parietal peritoneum distinguishes C. phenax from these other centrolenids, except for some specimens of C. spiculata (see Discus-

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FIG. 3.-Lateral and ventral views of the head of C. siren (KU 146610, holotype?, left, and C. phenax (KU 162264, paratype), right, illustrating differences in the shape of the snout. Scale equals 1 mm.

sion). However, C. spiculata possesses prevomerine teeth and more extensive webbing than C. phenax. Centrolenella bejaranoi has only a small portion of the parietal peritoneum pigmented, and it also differs from C. phenax in that the first finger is shorter than the second. Furthermore, in C. phenax the dorsal spots encompass a spicule and the surrounding flat area. In C. bejaranoi, the dorsal spots are much smaller and associated only with the upraised portion of the spicule proper. Centrolenella phenax differs from C. pluvialis in being a much smaller frog, with less webbing and no white peritoneal pigments or white band on the upper lip. Centrolenella phenax was tentatively referred to C. siren by Duellman (1976), who noted that the Ecuadorian specimens had yellow spots, whereas the Peruvian specimens had white spots. Comparison of recently collected series of C. siren from Ecuador reveals the following consistent differences between the two

species: Centrolenella siren has yellow spots (white in C. phenax), a sharply truncate snout (subtruncate and slightly longer, Fig. 3), and white parietal peritoneum (clear). The spots in C. siren are seen under close examination to be small masses of white pigment in preserved specimens. In C. phenax, the spots are formed instead by the simple absence of the purplish melanophores from the area, rather than the presence of discrete spots of pigment. Description.-Adult males of small size for centrolenid frogs; SVL 20.2-22.1 mm (Z = 21.3, n = 4). Head slightly wider than body; snout short, weakly truncate in dorsal and lateral views; canthus round; loreal region slightly depressed; lips not flared; nostrils almost terminal on snout, slightly protuberant, directed anterolaterally; internarial region depressed. Eye moderately large, directed anterolaterally. Supratympanic fold not distinct; lower four-fifths of tympanum visible, dorsolaterally inclined and with weak posterior

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orientation. Prevomerine teeth absent, dentigerous processes questionably present in some; choanae round; tongue round with slight posterior indentation; vocal slits present in males, extending from angle of jaw to midlateral margin of tongue. Humeral spine absent; dermal folds absent on forelimbs; order of fingers from shortest to longest 1=24-3; only scant webbing between fingers I11 and IV; fringes on fingers absent; finger discs broad, subtruncate; subarticular tubercles small, round, single; supernunlerary tubercles absent; palmar tubercle round, single; thenar tubercle elliptical. Hind limbs slender; tibia length 56.057.9% of SVL; dermal folds absent on hindlimbs; inner metatarsal tubercle elliptical, flattened; outer metatarsal tubercle absent; subarticular tubercles small, round; supernumerary tubercles absent; toes about one-third to one-half webbed; webbing formula I (2-2+)-2.5 I1 (1.51.75)-(2.75-3) 111 (I+-1.5)--(3-3") IV (3--3+)-(2--2) V; discs rounded, smaller than those on fingers. Skin on dorsal surfaces finely spiculate; spicules also on limbs, eyelids, tympanic region, and posterior portion of upper lip. Skin of belly and ventral thighs granular; flanks smooth. Anal opening directed posteriorly at upper level of thighs; a few small spicules in anal region; pair of large, indistinct subanal tubercles present. Color in preservative.-Dorsal surfaces lavender with white spots; ventral surfaces translucent. Color in life.-Dorsal surfaces dark green with minute creamy white flecks. Vocal sac green; bones green. Iris silvery white with black reticulations. Distribution.-The species is known from the type locality (Fig. 2, locality 10). Etymology.-The specific name phenax (Greek) is a noun in apposition, and signifies an imposter, in reference to the superficial resemblance of this species to C. siren. Remarks.-On the two days when the specimens were taken, the range of tem-

385

perature was 14.0-28.5 C, and total rainfall was 13.5 mm. Tutumbaro is a tiny village about 5 km south of Ayna. T h e locality is in disturbed cloud forest in a river valley with bamboo, ferns and elephant ear plants (Xanthosoma), but no tree ferns and few bromeliads. The frogs were calling at night from the upper sides of leaves of herbs and elephant ear plants within 1 m of a slow stream. Centrolenella bergeri was collected syntopically with C. phenax at this site. DISCUSSION The possibility always exists that isolated populations that are named as species are really nothing more than geographic variants. However, we note that the diagnostic characters used herein, with the exception of the peritoneal pigmentation, are invariant within species, assuming that one examines adults of the same sex. This observation holds true even for widespread species such as C. fleischmanni, C. prosoblepon and C. buckleyi. We have found that the spiculate dorsal skin and the extent of the parietal pigmentation-two of the diagnostic characters-may b e variable within some species. Examination of KU holdings of C. buckleyi, C. grandisonae, C, lynchi, C. megacheira and C. truebae demonstrates that in each of these species, the males are spiculate but the females possess no spicules, or only a few in the tympanic and postocular regions. Also, juveniles of t h e species have few or no spicules. An example of a spiculate dorsum can be seen in Fig. 1. Many, but not all, of the males of C. buckleyi are extensively spiculate. The non-spiculate males lack greatly distended vocal sacs, indicating that they were either not calling or not in breeding condition at the time of capture. This suggests that the degree of spiculation in males is related to breeding condition. These observations do not exclude skin texture as a useful taxonomic character. Rather, one must exercise

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caution in comparing specimens of different sexes or breeding condition. We have found also that the extent of white parietal peritoneal pigment is intra- and interpopulationally variable in two species. Two of five specimens of C. spiculata (LSU 25978-82) from the same population lack any white pigment on the parietal peritoneum or behind the eye. T h e other three specimens have the opaque white layer covering about half of the parietal peritoneum, and also behind the eyes. Other specimens of the same species (KU 162283-84) from another locality lack white pigment. At least two populations of C. buckleyi lack white pigment entirely, but among other populations the white pigment may be present immediately ventral to the heart only, or may cover up to half of the parietal peritoneum. In contrast, there is no variation with some species for which large sample sizes are available (e.g., C. siren, more than 60 specimens examined). The variation noted above suggests that while the white pigment and the skin texture can be useful in diagnosing species, the diagnosis should not be based solely on these characters; furthermore, when large samples are available, the variation should be carefully assessed. The two species described herein can be properly diagnosed without reference to the extent of the peritoneal pigments. Savage (1967) distinguished three groups of Centrolenella in Central America. Lynch and Duellman (1973) pointed out that several Ecuadorian centrolenids did not fit comfortably into Savage's groups. Likewise, some of the Peruvian frogs are not referable to these groups. At least three Peruvian species (C. phenax, C. truebae and C. mariae) differ trenchantly from other centrolenids in the lack of both parietal and visceral peritoneal pigments. (C. truebae and C, mariae were originally described as having white parietal peritoneum.) Furthermore, some specimens of C. spiculata and C. buckleyi likewise lack these pigments, and C.

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bejaranoi has white pigment only on the portion of the ~ a r i e t a lperitoneum immediately ventral to the heart (Cannatella, 1980). In view of the difficulty of assigning with confidence many of the South American Centrolenella to any of Savage's groups, it seems best to refrain from speculating on relationships of these two new species. The following eight species have been recorded also from Peru and Bolivia. These brief accounts summarize the distribution (Fig. 2) and the specimens examined. Centrolenella bejaranoi.-Known only from the type locality in the Yungas region of Bolivia. For description see Cannatella (1980). We have examined the type series, KU 182369-71, from 58.1 km (by road) SW Villa Tunari, 1980 m, Depto. Cochabamba, Bolivia 17"ll' S, 65'50' W. Centrolenella bergeri.-Known only from three localities in the cloud forests of Bolivia and southern Peru. We have examined the type series (KU 182363-68) and an egg clutch (KU 182372), all from the same locality as C. bejaranoi. I n addition we have seen the following specimens from Peru: KU 162248-50, Cuzco: Rio Cosiiipata, 4 km SW Santa Isabel, 1700 m; KU 162251-58,162259-60 (eggs), Ayacucho: Tutumbaro, Rio Piene, 1840 m. For description see Cannatella (1980). Centrolenella mariae.-We examined the only known specimen, the holotype, KU 174713 from Peru: Huanuco: Serrania de Sira, +I550 m. A description is given in Duellman and Toft (1979). Centrolenella midas.-Toft and Duellman (1979) noted without comment the occurrence of this species in Peru. The specimens (KU 172165-66, Peru: Huanuco: Finca Panguana, Rio Llullapichis, 4-5 km upstream from Rio Pachitea, 200 m) are the first reported from Peru and represent a significant range extension from the type locality in northern Ecuador. For a description see Lynch and Duellman (1973). Centrolenella munoxorum.-The range of this species of Centrolenella is sur-

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passed only by that of C.fleischmanni. It is found in the Amazonian lowlands from northern Ecuador to southern Peru. We have examined the following in addition to the Ecuadorian specimens listed by Lynch and Duellman (1973): KU 154749, 172167-69, Peru: Huanuco: Finca Panguana, Rio Llullapichis, 4-5 km upstream from Rio Pachitea, 2200 m; KU 15248889, 155493-97, 155524 (larvae), 175504, Ecuador: Napo: Santa Cecilia, 340 m; KU 175215, Ecuador: Napo: Rio Yasuni (200 km upstream from Rio Napo); LSU 3246873, Peni: Cuzco: 40 km E Quincemil. The latter specimens represent a significant southward range extension. Lynch and Duellman (1973) included a description of the species. Some of the specimens listed by Duellman (1976) have been referred to C. bergeri by Cannatella (1980). Centrolenella oce1lata.-The following Peruvian specimens were examined: LSU 25989-90, Ayacucho: Huanhuachayocc on Tambo-Valle de Apurimac Trail, 1630 m; BMNH 1912.11.1.19, Pasco: Huancabamba, 1700 m; AMNH 95216, Valle de Perene, 1200 m. Duellman (1976) provided a diagnosis based on the holotype. A female (KU 162262) with a SVL of 24.5 mm was referred to C. ocellata by Duellman (1976). This specimen, and KU 162265 (a juvenile), resemble C, ocellata in having many white spots on the shank and a hint of spots (although not really ocelli) on the dorsum. The specimens superficially resemble C. phenax in having a weakly truncate snout (round in ocellata) and clear parietal peritoneum (white in ocellata); however, the specimens differ from C. phenax in the shape of the head and several other rather subjective features; for this reason, we refer them tentatively to a possibly undescribed species. C e n t r o l e n e l l a spicu1ata.-AMNH 172250-51 from Peru: Valle de Perene, 1200 m represent the third record of this species, known from southern and central montane forests in Peru. Other specimens examined are LSU 25978-82, Ayacucho: San Jose, Rio Santa Rosa, 1000 m,

and KU 162283-84, Cuzco: Rio Cosiiipata, 4 km SW Santa Isabel, 1700 m. For a description see Duellman (1976). Centrolenella truebae.-Known only from the type locality in southern Peruthe Rio Cosfiipata, 4 km SW Santa Isabel, 1700 m, 13'05' S, 71'18' W, KU 16226882. Duellman (1976) provided a description of the species. I n addition to the one undescribed species listed under ocellata, we are aware of four undescribed species from Peru represented by four poorly preserved individuals: AMNH 42630 and 42631 from Tocache, 457-762 m; AMNH 43577 from the Peru-Brasil frontier, Utiquinia-Tapiche; and LSU 37104, from Depto. Amazonas, Prov. Bagua, Cordillera C o b , SE La Peca, 1799-1867 m. Owing to the poor state of preservation of these specimens, we defer from describing them until better material becomes available. Sympatry of species of Centrolenella occurs at the following localities (by number, Fig. 2): 1, 7 (munozorum, midas), 4 (two undescribed species), 9 (ocellata, spiculata), 10 (bergeri, phenax), 15 (bergeri, spiculata, truebae, one undescribed species), 17 (bejaranoi, bergeri). KEY TO CENTROLENELLA OF PERUAND BOLIVIA 1. Dorsum in life pale green with diffuse yellow spots, in preservative creamy white; visceral peritoneum white, parietal peritoneum clear ............................................ 2 Dorsum in life green, with or without pale or dark flecks; in preservative lavender; visceral peritoneum clear .................................. 3 2. Snout subtruncate in lateral view; webbing formula of fingers 111 2.25-2+ IV; webbing formula of toes I 1 . 5 2 - I1 1 . 5 2 111 1 . 2 5 2.5 IV 2.75-1.5 V; SVL of males 22.7-26.5 mm .................. C. bergeri Snout round in lateral view; webbing formula of fingers III 1.5-

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discs only slightly wider than fin1.25 IV; webbing formula of toes I ger .................................... C. midas &1 I1 G 1 . 5 I11 1-2 IV 2--1 V; SVL of males 18.8-20.5 mm -.-___-. Acknowledgments.-We are grateful to Alice G. .................................. C. munozorum C. Grandison, Charles W. Myers and Douglas A. 3. Dorsal pattern of many greenish Rossman for the loan of specimens and to Nelly Camllo de Espinoza of the Museo de Historia Natblack (dark purple in preservative) ural Javier Prado in Lima, Peni for logistic support spots and a few cream flecks ........ and many courtesies. We thank Thomas J. Berger ......................................... C. truebae and Jonathan A. Campbell for checking the key. Dorsum uniform green or marked Permits for collecting were generously provided by with pale spots or flecks .............. 4 Ing. Carlos Ponce del Prado, Direccibn de Conservacibn, Lima Peni, and Prof. Gaston Bejarano B., 4. Large pale dorsal spots (about one Departamento Nacional de Vida Silvestre, La Paz, mm diameter), generally bordered Bolivia. Field work was supported by a grant (DEB by ring of dark pigment .............. 5 76-09986) from the National Science Foundation to Dorsum uniform green or with William E. Duellman and by a NSF Graduate Fellowship to David C. Cannatella. small (much less than 1 mm) pale spots ............................................ 6 5. Prevomerine teeth present; first CANNATELLA, D. C. 1980. Two new species of finger about as long as second; paCentrolenella from Bolivia (Anura: Centrolenirietal peritoneum clear .... C. mariae dae). Proc. Biol. Soc. Wash. 93:714-724. Prevomerine teeth absent; first finDUELLMAN, W. E. 1976. Centrolenid frogs from ger distinctly shorter than second; Peni. Occ. Pap. Mus. Nat. Hist. Univ. Kans. 52: 1-11. parietal peritoneum white .._._.___. 1977. Liste der rezenten Amphibien und ..........................................C. ocellata

Reptilien: Hylidae, Centrolenidae, Pseudidae. 6. Outer fingers moderately to extenDas Tierreich 95:1-225. sively webbed (I11 %2- IV); pre-. 1978. New species of leptodactylid frogs vomerine teeth present ................ 9 of the genus Eleutherodactylus from the Cosriipata Valley, Peni. Proc. Biol. Soc. Wash. 91: Outer fingers scantily webbed (I11 418430. 2.5--2+ IV or less); prevomerine W. E., AND C. A. TOFT. 1979. Anteeth absent ................................ 7 DUELLMAN, urans from Serrania de Sira, Amazonian Peni: tax7. Pale spots closely spaced (about 1 onomy and biogeography. Herpetologica 35:60mm or less apart), encompassing 70. LYNCH,J. D., AND W. E. DUELLMAN.1973. A respicule only; first finger shorter view of the centrolenid frogs of Ecuador, with than second ................ C. bejaranoi descriptions of new species. Occ. Pap. Mus. Nat. Pale spots widely spaced (about 2Hist. Univ. Kans. 16:1-66. 4 mm apart), encompassing spicSAVAGE, J. M. 1967. A new tree-frog (Centrolenidae) from Costa Rica. Copeia 1967:325331. ule and surrounding skin; first finger about equal to second ............ 8 SAVAGE,J. M., AND W. R. HEYER. 1967. Variation and distribution of the tree-frog genus Phyllo8. Parietal peritoneum clear; spicmedusa in Costa Rica, Central America. Beitr. ules on dorsum generally homogeNeotrop. Fauna 5:111-131. nous in size; dorsal spots disTOFT, C. A., AND W. E. DUELLMAN. 1979. Anurans of the lower Rio Llullapichis, Amazonian tinctive; SVL of males 20.2-22.1 Peni: a preliminary analysis of community strucmm .................................. C. phenax ture. Herpetologica 35:71-77. Parietal peritoneum white; spicules on dorsum distinctly heterogenous in size; dorsal spots not disAccepted: 3 May 1982 tinctive; SVL of males 24.9-26.5 Associate Editor: Stephen Tilley . . mm ................................ C. pluvzalzs 9: Dorsal skin spiculate-; finger disc Museum of Natural History and Deat least twice width of finger ...... partment of Systematics and Ecology, . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. spiculata The University of Kansas, Lawrence, KS Dorsal skin not spiculate; finger 66045, USA

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You have printed the following article: Two New Species of Centrolenella, with a Brief Review of the Genus in Perú and Bolivia David C. Cannatella; William E. Duellman Herpetologica, Vol. 38, No. 3. (Sep., 1982), pp. 380-388. Stable URL: http://links.jstor.org/sici?sici=0018-0831%28198209%2938%3A3%3C380%3ATNSOCW%3E2.0.CO%3B2-8

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Literature Cited A New Tree-Frog (Centrolenidae) from Costa Rica Jay M. Savage Copeia, Vol. 1967, No. 2. (Jun. 5, 1967), pp. 325-331. Stable URL: http://links.jstor.org/sici?sici=0045-8511%2819670605%293%3A1967%3A2%3C325%3AANT%28FC%3E2.0.CO%3B2-5

Two New Species of Centrolenella, with a Brief Review of the Genus ...

Mar 20, 2008 - http://www.jstor.org/journals/herpetologists.html. Each copy of ... Two species are named as new--C. pluuialis and C. phenax, both from cloud.

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