— HERP QL 668 .E257 D84 1999 bcientific Papers Natural History Museum The University of Kansas 30 July 1999 Number 13:1-78 Frogs of the Genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of Northern Peru By William E. Duellman and Jennifer B. Pramuk Natural History Miiseiiui ami Biodiversity Research Center, and Department of Ecology and Evolutionary Biology, The University of Kansas, Lawrence, Kansas 66045-2454, USA. CONTENTS ABSTRACT 2 RESUMEN 2 INTRODUCTION 3 Acknowledgments 3 MATERIALS AND METHODS 3 ANDES OF NORTHERN PERU 4 Geological History 4 Physiography 5 Climate 5 Vegetation 8 GENERA OF ELEUTHERODACTYLIINE FROGS 10 JT SUMMARY OF TAXONOMIC CHARACTERS 11 ^ IDENTinCATION OF SPECIES 17 Key to the Species 17 Clave de las especies 18 r^' ACCOUNTS OF SPECIES 20 "^ Eleutherodactylus conspicillatus Group 20 ^ Eleutherod/ictyll/s N/GROvnr/iTus Group 33 ^ Eleutherodactylus orestes Group 35 S:::. Eleutherodactylus unistricatus Grovf 41 r: BIOGEOGRAPHY 69 2> LITERATURE CITED 72 APPENDICES 75 ^ Specimens Examined 75 ^ Gazetteer 77 ^ © Natural History Museum, The University of Kansas ISSN No. 1094-0782 a ^C/A 2 Scientific Papers, Natural History Museum, The University of Kansas leptodactylid frog Ekuthewdnch/his in the Andes of ^ 7,S'*1 ABSTRACT We recognize 45 species in the genus northern Ecuador. Twenty-one of these species have been known previously from Peru, but three of 'T^'^0^ them are reported for the first time from the Andes. Six other species known previously from Ecuador -^ are recorded for the first time in Peru, and 18 species are described as new. The majority (31) of the 45 ! /^*7£/ species are members of the large EleiithewdiKtylus iinistrigntus group, and nine species belong to the Eleutherodactylns conspicillatus group; one species belongs to the Eleutherodactyhis nigrovittahis group, and four Peruvian species are in the EleutJierodnctylus orestes group, which otherwise is known from three species in the Andes of southern Ecuador. Each species is treated in an account that includes a diagnosis, description or reference to a description, habitat, and distribution. Keys in English and Spanish to the species in the Andes of northern Peru are provided; 16 species are illustrated in color. In the Andes of southern Ecuador and northern Peru, all members of the Eleutherodactyhis conspicillatus group occur at elevations of less than 1500 m, but two species extend to elevations in excess of 2000 m. Members of the Eleutherodactyhis nigrovittatus and orestes groups mostly are confined Eleutherodactyhis unistrigatus group range through- St, to elevations above 3000 m, whereas species in the o' out the elevations in the Andes to 3200 m. Among the isolated mountain ranges in northern Peru, the o* Cordillera del Condor has the largest number of species of Eleutherodactyhis (19); of these, 11 are shared «? N with Cordillera Oriental in Ecuador, nine with the Cordillera de Cutucu in Ecuador, and only five are •• ^ <D ' among the 16 species known from the northern part of the Cordillera Central in Peru. The Cordillera ^ -< '' Colan has five species of Eleutherodactyhis; one is shared with the Cordillera Oriental in Ecuador and Cordillera Central in Peru. Of the 11 species inhabiting the Cordillera de Huancabamba, •Q S w another with the o S --^ three are shared with the Cordillera Occidental in Ecuador, two with the Cordillera Oriental in Ecuador, £/fHf/!erai(7cfi//"s (0 § - and two with the CordiUera Occidental in Peru. The absence of collections of from many ^ '-^ ^ mountain ranges in northern Pem suggests that the number of species far exceeds that reported here. ' ° in Key Words: Leptodactylidae, Eleutherodactyhis, Andes of northern Peru, Taxonomy, Biogeography. S] RESUMEN Reconocemos 45 especies en el genero leptodactilido Eleutherodactyhis en los Andes del ;§ norte Peru. Vientiuna de estas especies eran conocidas previamente para Peru, pero tres de ellas se reportano por primera vez para los Andes. Otras seis especies conocidos para Ecuador son reportadas por primera vez para Peru, y 18 especies son descriptas como nuevas. La mayoria (31) de las 45 especies son miembros del grupo Eleutherodactyhis unistrigatus, y nueve especies pertenecen al grupo Eleutherodactyhis conspcillatus; una especie pertenece al grupo Eleutherdactyhis nigrovittatus, y cuatro especies peruanas estan en el grupo Eleutherodactyhis orestes, el cual tambien es conocido por tres especies en los Andes del sur de Ecuador Para cada una de las especies se realiza un resumen que uncluye diagnosis, descripcion o referenda a una descripcion, habitat y ciistribucion geografica. Se proveen claves en ingles y espaiiol para la identificacion de las especies en los Andes del norte de Peru; 16 especies se ilustradan a color. En los Andes del sur de Ecuador y el norte de Peru, todos los miembros del grupo Eleutherodactyhis conspicillatus se encuentran en elevaciones menores a 1500 m, pero dos especies se extienden hasta elevaciones mayores a 2000 m. Miembros de los grupos Eleutherodactyhis nigrovittatus y Eleutherodactyhis orestes estan restringidos a elevaciones mayores de 3000 m, mientras que especies en el grupo Eleutherodactylus unistrigatus sew extiendenpor todas las elevaciones de los Andes hasta los 3200 m. Entre las Cordilleras aisladas en el norte del Peru, la Cordillera del Condor tiene el numero mas grande de especies de Eleutherodactyhis (19); de estas, 11 se encuentran tambien en la Cordillera Oriental en Ecuador, nueve en la Cordillera de Cutucu en Ecuador, y solamente cinco estan entre las 16 especies conocidas en el parte norte de la Cordillera Central en Peru. La Cordillera Colan tiene cinco especies de Eleutherodactyhis; una ocurre tambien en la Cordillera Oriental en Ecuador y otra en la Cordillera Central en Peru. De las 11 especies que habitan la Cordillera de Huancabamba, tres ocurren tambien en la Cordillera Occidental en Ecuador, dos en la Cordillera Oriental en Ecuador, y dos en la Cordillera Occidental en Peru. La ausencia de colecciones de Eleutherodactyhis de muchas Cordilleras en el norte de Peru sugiere que el numero de especies excede en mucho al que aqui se reporta. Palabras claves: Leptodactylidae, Eleutherodactyhis, Andes del norte de Peru, Taxonomia, Biogeografia. — Eleutherodactylus in the Andes of Northern Peru INTRODUCTION Frogs of the genus Eleutherodactylus are speciose in the cies have been reported from the Andes of northern Peru. cloud forests on the slopes of the Andes in Ecuador and Among the collections of Eleutherodactylus from north- Colombia (Lynch et al., 1997; Lynch and Duellman, 1980, ern Peru are specimens of 18 more unnamed species, as 1997). Several species of Eleuthewdacti/his have been de- well as examples of six species known from Ecuador but scribed from cloud forests on the Amazonian slopes of the previously not reported from Peru. The purposes of this Andes in southern Peru (Duellman, 1978a, b), but until paper are to present descriptions of the new species and recently, few species have been recorded from northern to provide a review of the species of Eleutherodactylus Peru. The first were £. cajauwrceusis and £. h/uuvii, which known to occur in the Andes and associated mountain described by were collected by G. K. Noble in 1916 and ranges in northern Peru—departamentos Amazonas, Barbour and Noble (1920); both species are now known to Cajamarca, Piura, and San Martin. Despite the large num- be rather widelv distributed in northwestern Peru and ber of species now known from the region, our knowledge 1997). southwestern Ecuador (Lynch and Duellman, More of anurans in the Andes of northern Peru is rudimentary. than half a century passed until more collections were More thorough collecting surely will expand the presently made in the Andes in northern Peru. known ranges of many species, and the exploration of pre- Field parties from The University of Kansas collected viously uncollected, isolated mountain ranges certainly amphibians and reptiles in the Andes of northern Peru in will reveal additional species. We would not be surprised 1979, 1989, and 1991, and field parties from Louisiana State if the present number represents no more than half of the University obtained specimens from the Cordillera de eleutherodactyline fauna of northern Peru. Huancabamba in 1974 and the Cordillera Colan in 1978. Acknowledgments Likewise field parties from the University of Florida ob- tained specimens in 1970 and 1972. Rainer Schulte, a resi- Duellman is indebted to his field companions—Tho- dent of Tarapoto, Peru, and his field companions collected mas J. Berger, David C. Cannatella, Fernando M. Cuadros several species of amphibians from the eastern slopes of V, Michael E. Morrison, Rainer Schulte, and John J. Wiens the Andes and outlying ranges, as well as the southern whose efforts contributed greatly to the amount of mate- slopes of the Cordillera del Condor. Alfonso Miranda of rial available for study, to many residents of northern Peru the Universidad Nacional Cajamarca, Peru, obtained speci- who provided shelter, food, and assistance to the field par- mens in the northern part of the Cordillera Occidental, and ties, and to B. Anthony Luscombe of Lima, and Rainer Javier Icochea and Robert P. Reynolds collected several Schulte of Tarapoto for logistical support. For the loan of species on the eastern slopes of the Cordillera del Condor specimens, we are grateful to David L. Auth of the Florida in 1994. Material resulting from these field expeditions led State Museum; Frank T. Burbrink, David A. Good, and to the descriptions of nine new species of Douglas A. Rossman of Louisiana State University; and Eleutherodactylus—E. schultei (Duellman, 1990a) and E. Robert P. Reynolds of the National Museum of Natural lirellus (Dwyer, 1995) from the Cordillera Central, E. History. We are grateful to Rainer Schulte for providing petrobardus (Duellman, 1991a) from the Cordillera Occiden- photographs of one species, Erik R. Wild for measuring tal, E. bearsei (Duellman, 1992a) and E. citriogaster many of the specimens and to Analia Piigener for consid- (Duellman, 1992b) from eastern outliers of the Andes in erable help with the Resumen and key in Spanish.