A NEW SPECIES OF ARBOREAL ELEUTHERODACTYLUS (ANURA: LEPTODACTYLIDAE) FROM THE AMAZONIAN LOWLANDS OF CENTRAL PERU

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1 Herpetologica, 63(1), 2007, E 2007 by The Herpetologists League, Inc. A NEW SPECIES OF ARBOREAL ELEUTHERODACTYLUS (ANURA: LEPTODACTYLIDAE) FROM THE AMAZONIAN LOWLANDS OF CENTRAL PERU EDGAR LEHR 1,3,4,CLAUDIA TORRES 2, AND JUANA SUÁREZ 2 1 Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Königsbrücker Landstrasse 159, D Dresden, Germany 2 Museo de Historia Natural, Departamento de Herpetología, Universidad Nacional Mayor de San Marcos, Av. Arenales 1256, Jesús María, Ap , Lima, Peru ABSTRACT: A new species of arboreal Eleutherodactylus is described and assigned to the Eleutherodactylus unistrigatus Group from the Amazonian lowlands of northern Departamento de Cusco, Peru. The new species was found on vegetation cm above ground, has a SVL up to 21.9 mm, and is distinguished from other species of the genus by having a green dorsum with white spots; a pointed, brown snout; and by lacking a tympanic membrane and annulus. Males lack vocal slits and nuptial pads; females are unknown. Key words: Amazonian lowlands; Anura; Eleutherodactylus; Leptodactylidae; New species; Peru OVER 25 species of Eleutherodactylus have been discribed during recent years in montane forests of the Andes in northern (Duellman and Pramuk, 1999) and central Peru (e.g., Duellman and Hedges, 2005; Lehr, 2005; Lehr et al., 2004; Lehr et al., in press), increasing the number of species of Eleutherodactylus known from Peru to 87. Only five species have been described in the last decade from the Amazonian lowlands of Peru; these are E. buccinator Rodríguez 1994, E. delius Duellman and Mendelson 1995, E. luscombei Duellman and Mendelson 1995, E. metabates Duellman and Pramuk 1999, and E. skydmainos Flores and Rodríguez In October 2005, the junior authors surveyed the herpetofauna in the Amazonian lowlands in Departamento de Cusco. Among the anurans collected are two specimens of Eleutherodactylus that are remarkably distinct from all known species in coloration pattern and morphological characters. Herein, we describe the new species. MATERIAL AND METHODS The format for the description follows that of Lynch and Duellman (1997), with inclusion 3 PRESENT ADDRESS: Natural History Museum and Biodiversity Research Center, Division of Herpetology, The University of Kansas, Jayhawk Boulevard 1345, Lawrence, KS , USA. 4 CORRESPONDENCE: , elehr@ku.edu of a description of a holotype. Anuran taxonomy is currently in flux and familial placement of Eleutherodactylus may be subject to change, therefore we do not follow Frost et al., (2006). Field notes on the new species were recorded by C. Torres and J. Suárez on October For descriptions of coloration in life, photographs also were used. Specimens were preserved in 4% formalin and stored in 70% ethanol. Specimens were dissected to determine the nature of the gonads. Measurements taken with digital calipers and rounded to the nearest 0.1 mm were: SVL (snout vent length), TL (tibia length), FL (foot length, distance from proximal margin of inner metatarsal tubercle to tip of Toe IV), HL (head length, from angle of jaw to tip of snout), HW (head width, at level of angle of jaw), ED (eye diameter), IOD (interorbital distance), EW (upper eyelid width), IND (internarial distance), E N (eye nostril distance, straight line distance between anterior corner of orbit and posterior margin of external nares). Lengths of Toes III and V were estimated when both were adpressed against Toe IV; lengths of Fingers I and II were estimated when adpressed against each other. The otic region was dissected in order to evaluate the condition of the tympanic annulus. All drawings were made by the senior author using a stereomicroscope with a drawing tube attachment. Codes for museum collections follow those of 94

March 2007] HERPETOLOGICA 95 FIG. 1. Eleutherodactylus tantanti (MHNSM 23942, holotype, SVL 21.9 mm) in lateral (A) and ventral (B) views, photos by C. Torres. Leviton et al., (1985) and Frost (2006).

2 March 2007] HERPETOLOGICA 95 FIG. 1. Eleutherodactylus tantanti (MHNSM 23942, holotype, SVL 21.9 mm) in lateral (A) and ventral (B) views, photos by C. Torres. Leviton et al., (1985) and Frost (2006). Specimens examined are listed in Appendix I. Eleutherodactylus tantanti sp. nov. Holotype. MHNSM (Fig. 1A,B), an adult male obtained from Myaria (11u S, 73u W) at 321 m on 17 October 2005, Distrito de Echarate, Provincia de La Convención, Departamento de Cusco, Peru, by V. Villalobos. Paratype. MHNSM 23943, an adult male obtained from Myaria (11u S, 73u W) at 351 m on 16 October 2005, Distrito de Echarate, Provincia de La Convención, Departamento de Cusco, Peru, by J. Suárez and C. Torres. Diagnosis. A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus group having the following combination of characters. (1) Skin on dorsum shagreen; that on venter areolate; discoidal fold not evident; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus absent; (3) snout short, triangular, pointed in dorsal view, protruding well anterior to margin of lower jaw in lateral view; (4) upper eyelid without tubercles, its width narrower than IOD; cranial crests absent; (5) vomerine teeth minute, embedded in mucosa of mouth; (6) males lacking vocal sac, vocal slits, and nuptial pads; (7) Finger I shorter than Finger II; discs on outer fingers broadly expanded, rounded; (8) fingers with lateral fringes; (9) ulnar and tarsal tubercles present, coalescing into a fold; (10) heel without tubercles; inner tarsal fold absent; (11) inner metatarsal tubercle, moderate, ovoid, as large as outer; outer metatarsal tubercle small, ovoid, contacting tarsal fold; few, round supernumerary plantar tubercles; (12) toes with lateral fringes; basal webbing between Toes III and IV and IV and V; Toe V much longer than Toe III; discs slightly smaller than those on fingers; (13) dorsum green with white spots; venter greenish yellow; groin and anterior surfaces of thighs green with white spots; iris pale beige, with fine, brown reticulation; (14) SVL in males mm (n 5 2), females unknown. Eleutherodactylus tantanti is readily distinguished from other species of the genus by its lack of a tympanum, pointed snout, and green dorsum with white spots. Seven other species of Eleutherodactylus from the eastern Andes and Amazonian lowlands of Peru lack a tympanum, but none of them is green: E. colodactylus, E. cruciocularis, E. flavobracatus, E. imitatrix, E. lirellus, E. martiae, ande. ventrimarmoratus. With respect to habitus and coloration, E. tantanti resembles species of the E. lacrimosus Group; these are E. aureolineatus, E. bromeliaceus, E. lacrimosus, E. mendax, E. olivaceus, E. pardalinus, and E. schultei. All of the latter species have a tympanum and none is green with white spots. Eleutherodactylus tantanti is most similar to E. acuminatus Shreve In both species, the dorsum is green and the snout acuminate in dorsal view, but E. tantanti lacks a tympanum (present in E. acuminatus), possesses fingers and toes with lateral fringes (fringes absent in E. acuminatus), and lacks black canthal, interorbital, and supratympanic stripes (present in E. acuminatus). Both E. tantanti and E. galdi have a green dorsum with white spots, but E. tantanti lacks cranial crests (present in E. galdi), lacks a tympanum (present in E. galdi), and is a lowland species

3 96 HERPETOLOGICA [Vol. 63, No. 1 FIG. 2. Lateral (A) and dorsal (B) views of head, and ventral views of hand (C) and foot (D) of Eleutherodactylus tantanti (MHNSM 23942). Folds and coalescing tubercles on outer surfaces of arm and foot in gray. known from elevations between m, whereas E. galdi is an Andean species known from elevations of m (Lynch and Duellman, 1980). Because of its green dorsum, Eleutherodactylus tantanti might be confused with the species of the genera Sphaenorhynchus (Hylidae) or Cochranella (Centrolenidae) that inhabit the Amazonian lowlands. Although species of Sphaenorhynchus have a green dorsum and pointed snout, they have a tympanum and extensively webbed hands and feet. Some Cochranella have a green dorsum with white spots, but they have a tympanum and broadly rounded snout. Description of the holotype. Head smaller than body, nearly as wide as long; head width 34.2% SVL; head length 35.2% SVL; snout short, acuminate in dorsal view, protruding well anteriad to margin of lower jaw in lateral view (Fig. 2A,B); eye diameter slightly greater than eye nostril distance (eye nostril distance 85.2% eye diameter); nostrils slightly protuberant, directed laterally; canthus rostralis rounded in dorsal view; loreal region nearly vertical; lips rounded; upper eyelid lacking tubercles; upper eyelid width 58.8% IOD; supratympanic fold weak, barely evident; tympanic annulus and tympanic membrane absent; conical postrictal tubercles present on each side, coalesced into short ridge. Choanae ovoid, not concealed by palatal shelf of maxillary arch; vomerine teeth minute, embedded in mucosa of mouth roof, barely visible, situated posteromedial to choanae; tongue as long as wide (length 3.9 mm, width at middle of tongue 3.6 mm), slightly notched posteriorly, posterior fourth free. Skin on dorsum shagreen; dorsolateral folds absent; skin on flanks on its upper half as on dorsum, and venterolaterally, coarsely areolate; skin on thighs, belly and chest coarsely areolate, skin on other ventral surfaces smooth; discoidal fold not evident; cloacal sheath short; large tubercles in cloacal region absent. Outer venterolateral surface of left forearm bearing a row of four, low, elongate tubercles coalescing to a fold, outer venterolateral surface of right forearm bearing two, low, elongate tubercles; palmar tubercles slightly elevated, outer palmar tubercle bifid, approximately size of inner, narrow, ovoid palmar tubercle; no supernumerary tubercles; subarticular tubercles round in dorsal and lateral views; fingers with broad lateral fringes, outer fringe of Finger IV continuing to proximal edge of palm with its shape slightly undulated, contacting fold or tubercles on venterolateral surface of forearm; Finger I shorter than Finger II; discs on fingers broadly expanded, most prominent of Finger III and IV, rounded terminally; all fingers having ventral pads well defined by circumferential grooves (Fig. 2C). Hind limbs slender, tibia length 50.2% of SVL; foot length 45.7% of SVL; upper and lateral surfaces of hind limbs shagreen; ventral surface of thigh coarsely areolate; heel without tubercles; row of low, elongate tubercles coalesced to a tarsal fold with an undulated shape on outer ventral surface of tarsus; inner metatarsal tubercle elevated, elongate, about subconical outer metatarsal tubercle; few, low plantar supernumerary tubercles; subarticular tubercles well defined, round in dorsal view and subconical in lateral view; toes with broad lateral fringes, outer fringe of Toe

4 March 2007] HERPETOLOGICA 97 V contacting outer row of fold of plantar surface with its shape undulated, contacting outer tarsal fold (Fig. 2D); basal webbing between Toes III and IV, and IV and V; discs on toes slightly smaller than those on fingers, most prominent of Toes IV and V; well defined by circumferential grooves; relative lengths of toes: 1, 2, 3, 5, 4 (Fig. 2D); Toe III much shorter than Toe V (disc of Toe III not reaching level of disc of Toe V, when both adpressed against Toe IV); Toe III not extending to level of penultimate subarticular tubercle on Toe IV. Measurements (in mm) of holotype. SVL: 21.9; TL: 11.0; FL: 10.0; HL: 7.7; HW: 7.5; ED: 2.7; IOD: 3.4; EW: 2.0; IND: 1.6; E N: 2.3. Coloration of holotype in preservative. Entirely cream with pattern of pale brown blotches and white spots as described below, but less distinct. Coloration of holotype in life. Dorsum green with small white spots (fewer on head and shoulder), snout pale brown with white spots; brown spots on upper lip below eye, on knees, on heel, and on Fingers III IV and Toes IV V; flanks green with white spots, venterolateral areas white; groin green with white spots; chest and anterior half of belly yellowish white, throat, posterior half of belly and ventral surface of shanks greenish yellow; ventral surface of lower jaw white; outer margins of arms and legs white; other ventral surfaces greenish yellow; iris grayish tan with fine, pale brown reticulations. Variation. The second known specimen is a male with following measurements (mm): SVL: 19.6; TL: 11.2; FL: 9.1; HL: 6.3; HW: 6.5; ED: 2.1; IOD: 3.0; EW: 1.3; IND: 1.6; E N: 2.3. The specimen has a similar coloration pattern as described for the holotype, except for a less distinct brown coloration. Etymology. The specific name tantanti is associated with the call of this or a similar anuran species in the language of the Asháninka. The Asháninka are an indigenous tribe inhabiting the Amazonian lowlands of Peru (including the type locality of the new species) and western Brazil. Distribution and ecology. Eleutherodactylus tantanti is known only from the type locality (Fig. 3) supporting primary lowland forest. One specimen (MHNSM 23943) was FIG. 3. Location of type locality (star) of Eleutherodactylus tantanti in Peru. found on a leaf 150 cm above the ground at 8.51 pm; the second individual (MHNSM 23942) was found sleeping on a leaf of a tree 800 cm above ground at 10 am. Other species of Eleutherodactylus collected at Myaria are E. diadematus, E. fenestratus, E. ockendeni, and E. peruvianus. REMARKS Although arboreal frogs often are heard, they frequently are out of reach and remain uncollected. Thus the canopy of tropical forests is probably one of the least known strata with respect to species diversity. Guayasamin et al., (2006) described E. aureolineatus (lacrimosus Assemblage) from specimens collected up to 40 m in the canopy in the western Amazon Basin of northern Ecuador. We suspect that Eleutherodactylus tantani is a canopy inhabitant because one individual was obtained opportunistically when botanists were collecting leaves from a 8-m high branch, that had been cut from a tree. One of the characteristics of the new species is the lack of external ears. The absence of a tympanum in eleutherodactyline frogs has been positively correlated with increased elevation in central Peruvian eleutherodactyline frogs (Lehr et al., in press). RESUMEN Describimos una nueva especie arbórea perteneciente al grupo Eleutherodactylus

5 98 HERPETOLOGICA [Vol. 63, No. 1 unistrigatus. La nueva especie tiene una longitud rostro-cloacal de hasta 21.9 mm y se diferencia de otras especies del género por presentar un dorso verde con puntos blancos y un rostro café con forma punteaguda, y por carecer de tímpano y membrana timpánica. Los machos no tienen hendiduras vocales ni almohadillas nupciales; las hembras son desconocidas. La nueva especie ha sido encontrada en vegetación entre m sobre el nivel del suelo en el bosque Amazónico del norte del Departamento de Cusco, Perú. Acknowledgments. For comments on the manuscript we are grateful to A. Crawford, W. E. Duellman, L. Trueb, and two anonymous reviewers. Loan of material was kindly provided by J. H. Córdova and R. Heyer. C.T. and J.S. thank the Environmental Resources Management Peru and Repsol YPF Peru for logistic support, for financing fieldwork, and for providing collecting permits issued by the Ministerio de Agricultura (INRENA), Lima, Peru. The research was supported by a postdoctoral grant to E. Lehr by the Alexander von Humboldt-Foundation. LITERATURE CITED DUELLMAN, W. E., AND S. B. HEDGES Eleutherodactyline frogs (Anura: Leptodactylidae) from the Cordillera de Yanachaga in Central Peru. Copeia 2005: DUELLMAN, W. E., AND J. R. MENDELSON, III Amphibians and reptiles from northern Departamento Loreto, Peru: Taxonomy and Biogeography. University of Kansas Science Bulletin 55: DUELLMAN, W. E., AND J. B. PRAMUK Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of northern Peru. Scientific Papers Natural History Museum University of Kansas 13:1 78. FLORES, G., AND L. O. RODRÍGUEZ Two new species of the Eleutherodactylus conspicillatus group (Anura: Leptodactylidae) from Perú. Copeia 1997: FROST, D. R Amphibian Species of the World: an Online Reference. Version 4 (17 August 2006). Electronic Database accessible at org/herpetology/amphibia/index.php. American Museum of Natural History, New York, New York, U.S.A. FROST, D. R., T. GRANT, J.FAIVOVICH, R.H.BAIN, A.HAAS, C. F. B. HADDAD, R. O. DE SÁ, A. CHANNING, M. WILKINSON, S.C.DONNELLAN, C.J.RAXWORTH, J.A. CAMPBELL,B.L.BLOTTO,P.MOLER,R.C.DREWES,R.A. NUSSBAUM, J. D. LYNCH, D. M. GREEN, AND W. C. WHEELER The amphibian tree of life. Bulletin American Museum of Natural History 297: GUAYASAMIN, J. M., S. R. RON, D. F. CISNEROS-HEREDIA, W. LAMAR, AND S. F. MCCRACKEN A new species of frog of the Eleutherodactylus lacrimosus assemblage (Leptodactylidae) from the western Amazon Basin, with comments on the utility of canopy surveys in lowland rainforest. Herpetologica 62: LEHR, E A new species of the Eleutherodactylus nigrovittatus group (Anura: Leptodactylidae) from Andean Peru. Herpetologica 61: LEHR, E., C. AGUILAR, AND W. E. DUELLMAN A striking new species of Eleutherodactylus from Andean Peru (Anura: Leptodactylidae). Herpetologica 60: LEHR, E., M. LUNDBERG, C. AGUILAR, AND R. von MAY New species of Eleutherodactylus (Anura: Leptodactylidae) from the eastern Andes of central Peru with comments on central Peruvian Eleutherodactylus. Herpetological Monographs 20: LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. DAWSON Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985: LYNCH, J. D., AND W. E. DUELLMAN The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). The University of Kansas, Museum of Natural History Miscellaneous Publication 69:1 86. Lynch, J. D., AND W. E. DUELLMAN Frogs of the genus Eleutherodactylus in western Ecuador. Systematics, ecology, and biogeography. The University of Kansas, Natural History Museum Special Publication 23: RODRÍQUEZ, L. O A new species of the Eleutherodactylus conspicillatus group (Leptodactylidae) from Peru. Alytes 12: Accepted: 20 September 2006.Associate Editor: Paul Chippindale APPENDIX I Comparative Specimens Examined Eleutherodactylus acuminatus: PERU: SAN MARTIN: Rio Cainarache, 33 km NE Tarapoto, on road to Yurimaguas: KU ; Rio Shilayo, Tarapoto, 500 m: KU ; Lamas: 15.4 km SW Zapatero, 950 m: KU ; LORETO: San Jacinto, 175 m: KU ; CUSCO: San Martin-3, ca. 5 km N of the Camisea River: UNSNM , Pagoreni on the Camisea River: UNSNM Eleutherodactylus bromeliaceus: ECUADOR: MORONA- SANTIAGO: Mirador: KU (paratype); El Cruzado: KU (paratype); PERU: SAN MARTÍN: Rioja: E slope Abra Pardo de Miguel, 2180 m: KU ; PASCO: 2.9 km N, 5.5 km E (airline) Oxapampa: KU , ; PASCO: 2.5 km N, 11 km E (airline) Oxapampa: KU ; PASCO: ca. 8 km SW La Suiza on E slope: KU ; PASCO: Oxapampa: Antenna de Pajonal, 10u S, 75u W, 2780 m: MTD ; PASCO: Oxapampa: San Alberto, ca. 10u S, 75u W, 2200 m: MHNSM 18660, , MTD , Eleutherodactylus colodactylus: PERU: PIURA: summit cordillera between Chanchaque and Huancabamba, 3100 m: KU , ; 31 km W Huancabamba, 3080 m: KU ; 33 km W Huancabamba, ca m: KU Eleutherodactylus cruciocularis: PERU: JUNÍN: Pampa Hermosa: 10u S, 75u W, 1540 m: MHNSM (holotype), MHNSM , MTD 45634, (all paratypes).

6 March 2007] HERPETOLOGICA 99 Eleutherodactylus flavobracatus: PERU: PASCO: obtained at km 34 on road from Oxapampa to Yaupi: 10u S, 75u W, 1770 m: MHNSM (holotype), MHNSM 19848, MTD , (all paratypes). Eleutherodactylus galdi: PERU: CAJAMARCA: Tabaconas-Namballe, 05u S, 79u W, MHNSM Eleutherodactylus imitatrix: PERU: MADRE DE DÍOS: Puerto Maldonado, 200 m: KU , , , Eleutherodactylus lirellus: PERU: SAN MARTÍN: Rioja, Río Cerranayacu, 76 km NW Roja, 1200 m: KU , (all paratypes). Eleutherodactylus mendax: PERU: PASCO: Auquimarca, Puqiopata forest, 10u 43,759 S, 75u 42,809 W, 3310 m: MTD 44329; Auquimarca, Puqiopata forest, 10u43,789S, 75u 42,839 W, 3325 m: MTD 44330; Auquimarca: Cillapata, 2900 m: MTD 45080; near road from Uchuhuerta to Auquimarca, 10u S, 75u W, 2850 m: MHNSM 19865; near road from Uchuhuerta to Auquimarca, 10u S, 75u W, 3000 m: MHNSM 19866, MTD ; Paucartambo, 2932 m: MTD , Eleutherodactylus pardalinus: PERU: JUNÍN: Huasahuasi: 11u S, 75u W, 2640 m: MHNSM (holotype), MHNSM , MTD (all paratypes). Eleutherodactylus schultei: PERU: AMAZONAS: 5 km N Levanto: 06u 179 S, 77u 519, 2850 m: KU (holotype), KU (paratypes), KU (paratypes); ca. 5 km NW Mendoza, ca m: KU (paratypes).

OCCASIONAL PAPERS OF THE MUSEUM OF ZOOLOGY UNIVERSITY OF MICHIGAN

OCCASIONAL PAPERS OF THE MUSEUM OF ZOOLOGY UNIVERSITY OF MICHIGAN A NEW SPECIES OF ELEUTHERODACTYLUS FROM THE CORDILLERA OCCIDENTAL OF COLOMBIA (AMPHIBIA : ANURA: LEPTODACTY LIDAE) Frogs of the fitzingeri