HERP QL 668 E257 D84 1999 Scientific 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 Museum and Biodiversity Research Center, and Department of Ecology and Evolutionary Biology, The University of Kansas, Lawrence, Kansas 66045-2454, USA. CONTENTS ANTBISTIERVAC TT ceanbececcasebons tan eceeegsE BES AEE SAGE CREE: EEC BLOEE AE CER ErARE RAREST EAE orn Pree EE Eee earners 2 IRIE TUIMIEBIN] . sccoccodsencécae Seana cna cEAD or SSEE Sr BASSE EA? 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GROUP ssscescy caves tases seseessesssveccvadevonsvescesvasvasesedecessecdsssdensesasvectvets 35 REUTER A GIN MUIUINTSTRIGATLIS| GROUP osc: scsascesedsisscsverssesddastncassits «aiissesvarssvocssssvesdseesoseaveees 4] BLO GEO GIRUN IPI RING as cenonh deco Os aa ck One RES HEE EEE nS Sct BS er Eee 69 TLITTEUPANTTTUSRES (COUT EY DD ss Sa eee WP AN PXPIBINIDUT CIES) can scanancbeceo ce Boon GEO HERRERA Bare PASE er ECE ee 75 © Natural History Museum, The University of Kansas ISSN No. 1094-0782 pualyan, William & . [444 z i) SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS ABSTRACT We recognize 45 species in the leptodactylid frog genus Eleutherodactylus in the Andes of northern Ecuador. Twenty-one of these species have been known previously from Peru, but three of 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 species are members of the large Eleutherodactylus unistrigatus group, and nine species belong to the Eleutherodactylus conspicillatus group; one species belongs to the Eleutherodactylus nigrovittatus group, and four Peruvian species are in the Eleutherodactylus 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 Eleutherodactylus conspicillatus group occur at elevations of less than 1500 m, but two species extend to elevations in excess of 2000 m. Members of the Eleutherodactylus nigrovittatus and orestes groups mostly are confined to elevations above 3000 m, whereas species in the Eleutherodactylus unistrigatus group range through- out the elevations in the Andes to 3200 m. Among the isolated mountain ranges in northern Peru, the Cordillera del Condor has the largest number of species of Eleutherodactylus (19); of these, 11 are shared with Cordillera Oriental in Ecuador, nine with the Cordillera de Cutuct in Ecuador, and only five are among the 16 species known from the northern part of the Cordillera Central in Peru. The Cordillera Colan has five species of Eleutherodactylus; one is shared with the Cordillera Oriental in Ecuador and another with the Cordillera Central in Peru. Of the 11 species inhabiting the Cordillera de Huancabamba, three are shared with the Cordillera Occidental in Ecuador, two with the Cordillera Oriental in Ecuador, and two with the Cordillera Occidental in Peru. The absence of collections of Eleutherodactylus from many mountain ranges in northern Peru suggests that the number of species far exceeds that reported here. Key Words: Leptodactylidae, Eleutherodactylus, Andes of northern Peru, Taxonomy, Biogeography. RESUMEN Reconocemos 45 especies en el género leptodactilido Eleutherodactylus 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 Eleutherodactylus unistrigatus, y nueve especies pertenecen al grupo Eleutherodactylus conspcillatus; una especie pertenece al grupo Eleutherdactylus nigrovittatus, y cuatro especies peruanas estan en el grupo Eleutherodactylus orestes, el cual también 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, descripcién o referencia a una descripcion, habitat y distribucion geografica. Se proveen claves en ingles y espanol 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 Pert, todos los miembros del grupo Eleutherodactylus conspicillatus se encuentran en elevaciones menores a 1500 m, pero dos especies se extienden hasta elevaciones mayores a 2000 m. Miembros de los grupos Eleutherodactylus nigrovittatus y Eleutherodactylus 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 Eleutherodactylus (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 Eleutherodactylus; una ocurre también en la Cordillera Oriental en Ecuador y otra en la Cordillera Central en Pert. 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 Pert. La ausencia de colecciones de Eleutherodactylus 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, Eleutherodactylus, Andes del norte de Peru, Taxonomia, Biogeografia. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 3 INTRODUCTION Frogs of the genus Eleutherodactylus are speciose in the cloud forests on the slopes of the Andes in Ecuador and Colombia (Lynch et al., 1997; Lynch and Duellman, 1980, 1997). Several species of Eleutherodactylus have been de- scribed from cloud forests on the Amazonian slopes of the Andes in southern Peru (Duellman, 1978a, b), but until recently, few species have been recorded from northern Peru. The first were E. cajamarcensis and E. lymani, which were collected by G. K. Noble in 1916 and described by Barbour and Noble (1920); both species are now known to be rather widely distributed in northwestern Peru and southwestern Ecuador (Lynch and Duellman, 1997). More than half a century passed until more collections were made in the Andes in northern Peru. Field parties from The University of Kansas collected amphibians and reptiles in the Andes of northern Peru in 1979, 1989, and 1991, and field parties from Louisiana State University obtained specimens from the Cordillera de Huancabamba in 1974 and the Cordillera Colan in 1978. Likewise field parties from the University of Florida ob- tained specimens in 1970 and 1972. Rainer Schulte, a resi- dent of Tarapoto, Peru, and his field companions collected several species of amphibians from the eastern slopes of the Andes and outlying ranges, as well as the southern slopes of the Cordillera del Condor. Alfonso Miranda of the Universidad Nacional Cajamarca, Peru, obtained speci- mens in the northern part of the Cordillera Occidental, and Javier Icochea and Robert P. Reynolds collected several species on the eastern slopes of the Cordillera del Condor in 1994. Material resulting from these field expeditions led to the descriptions of nine new species of Eleutherodactylus—E. schultei (Duellman, 1990a) and E. lirellus (Dwyer, 1995) from the Cordillera Central, E. petrobardus (Duellman, 1991a) from the Cordillera Occiden- tal, E. bearsei (Duellman, 1992a) and E. citriogaster (Duellman, 1992b) from eastern outliers of the Andes in Departamento San Martin, and E. ceuthospilus, rhodoplichus, sternothylax, and wiensi from the Cordillera de Huancabamba (Duellman and Wild, 1993). The latter au- thors also provided the first Peruvian records for three species (E. colodactylus, cryptomelas, and phoxocephalus) pre- viously known only from Ecuador. Reynolds and Icochea (1997) reported the first Peruvian locality for E. condor, pre- viously only known from Ecuador, and provided the first locality for E. peruvianus from the Andes of northern Peru. Flores and Rodriguez (1997) described E. karcharias from the northern part of the Cordillera Central. Thus, 17 spe- cies have been reported from the Andes of northern Peru. Among the collections of Eleutherodactylus from north- ern Peru are specimens of 18 more unnamed species, as well as examples of six species known from Ecuador but previously not reported from Peru. The purposes of this paper are to present descriptions of the new species and to provide a review of the species of Eleutherodactylus known to occur in the Andes and associated mountain ranges in northern Peru—departamentos Amazonas, Cajamarca, Piura, and San Martin. Despite the large num- ber of species now known from the region, our knowledge of anurans in the Andes of northern Peru is rudimentary. More thorough collecting surely will expand the presently known ranges of many species, and the exploration of pre- viously uncollected, isolated mountain ranges certainly will reveal additional species. We would not be surprised if the present number represents no more than half of the eleutherodactyline fauna of northern Peru. ACKNOWLEDGMENTS Duellman is indebted to his field companions—Tho- mas J. Berger, David C. Cannatella, Fernando M. Cuadros V, Michael E. Morrison, Rainer Schulte, and John J. Wiens— whose efforts contributed greatly to the amount of mate- rial available for study, to many residents of northern Peru who provided shelter, food, and assistance to the field par- ties, and to B. Anthony Luscombe of Lima, and Rainer Schulte of Tarapoto for logistical support. For the loan of specimens, we are grateful to David L. Auth of the Florida State Museum; Frank T. Burbrink, David A. Good, and Douglas A. Rossman of Louisiana State University; and Robert P. Reynolds of the National Museum of Natural History. We are grateful to Rainer Schulte for providing photographs of one species, Erik R. Wild for measuring many of the specimens and to Analia Pugener for consid- erable help with the Resumen and key in Spanish. The manuscript benefited from critical review by S. Blair Hedges and John D. Lynch and careful editing by Linda Trueb; we thank them profusely for their efforts. Permits for the collection and exportation of specimens were is- sued by Luis J. Cueto Aragon, Armando Pimental Bustamento, Gonzalo Bravo Mejia Munoz, and José Purisaca, Direccion General Forestal y de Fauna, Ministerio de Agricultura, Lima, Peru. The research reported herein is part of a study on patterns of speciation and biogeogra- phy of Andean anurans supported by a grant (BSR 8805920) from the National Science Foundation (W. E. Duellman, P.I.). MATERIALS AND METHODS Specimens in museum collections are identified by their catalogue numbers preceded by the following codes: ANSP = Academy of Natural Sciences of Philadelphia; BM = Brit- ish Museum (Natural History); KU = Natural History 4 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Museum, University of Kansas; LSUMZ = Museum of Zoology, Louisiana State University; MCZ = Museum of Comparative Zoology, Harvard University, MHNSM = Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru; MNCN = Museo Nacional de Ciencias Naturales, Madrid, Spain; NHMG = Naturhistoriska Museet Goteborg, Sweden; NHRM = Naturhistoriska Riksmuseet, Stockholm, Sweden; MSNT = Museo e Instituto di Zoologia Sistematica, Universita di Torino, Italy; USNM = National Museum of Natural His- tory; UF = Florida State Museum, University of Florida, Gainesville. All specimens from northern Peru and Ecua- dor that have been studied are listed in Appendix 1. Lo- calities from which specimens have been examined are listed with their geographic coordinates and elevations in Appendix 2. Measurements, definitions of structural characters, and numbered characteristics in diagnoses follow the meth- ods of Lynch and Duellman (1997). Diagnoses of E. bromeliaceus, colodactylus, cryptomelas, proserpens, and ver- sicolor were modified from those of Lynch (1979), and those of E. cajamarcensis, lymant, and phoxocephalus were taken from Lynch and Duellman (1997), whereas those of E. incomptus, condor, galdi, ockendeni, and peruvianus were modified from those of Lynch and Duellman (1980) and that of E. pecki from Duellman and Lynch (1988). A major problem in providing adequate diagnoses of Eleutherodactylus is the comparison of new taxa with the plethora of existing species. In order to be able to ascertain character states among the many species that might be confused with the species described or discussed herein, we tabulated character states for all diagnostic characters used by Lynch and Duellman (1997). For the purposes of this study, we included all described species of Eleutherodactylus from Peru and Ecuador, as well as the new taxa; it is extremely unlikely that species from any other region occur in northern Peru. To the nonspecialist, it may seem that we have elaborated unnecessarily the diagnoses of all of the taxa and the descriptions of the new taxa. However, the distinction between specimens of many species of Eleutherodactylus is difficult even for the special- ist. We present detailed diagnoses and descriptions in an endeavor to facilitate comparisons with other taxa that certainly will be discovered in the future. Measurements were taken with dial calipers to the near- est 0.1 mm. If sex and reproductive condition were not evident externally (nuptial pads or vocal sacs in males and eggs visible through the body wall in females), sex and reproductive condition were determined by dissection. The following abbreviations are used: E-N = eye-nostril dis- tance; HL = head length; HW = head width; IOD = inter- orbital distance; SVL = snout-vent length. Photographs noted as (ERW) were taken by Erik R. Wild; all others were taken by Duellman. Except for the sites visited by Reynolds and Icochea, who used GPS devices, geographical coordinates were obtained from maps, principally the Mapa Fisico Politico, 1:1,000,000 (1973) but also the Carta Nacional del Peru, 1:100,000 (1986) for those regions so mapped; both sets of maps were produced by the Instituto Geografico Militar del Peru. Coordinates for Ecuadorian localities were ob- tained from the 1974 edition of the Mapa de Ecuador, 1:1:000,000, produced by the Instituto Geografico Militar, Quito. Elevations were obtained by altimeters or in some cases from maps. ANDES OF NORTHERN PERU Equaled in heights and expanse only by the Himalayas, the majestic Andean mountain chain was described by Enock (1907) as: “Heavenward thrown, crumpled, folded, ridged and fractured, with gleaming ‘porcelain’ gnomons pointing to the sun; shattered strata and shear crevasse; far terraces and grim escarpments, hung over with filmy mist-veils, and robed with the white clothing of crystalised rains and mists; the birthplace of the winds and hails; the father of rivers whose floods are borne a thousand leagues away—the mighty cordillera is!” This description is espe- cially appropriate for the Andes of northern Peru (departamentos Amazonas, Cajamarca, Piura, and San Martin), where north-south mountain ranges are separated by deep valleys. GEOLOGICAL History In northern Peru and southern Ecuador, a major struc- tural and physiographic break exists in the Andes (Figs. 1, 2); this is the Huancabamba Depression, a complex sys- tem of relatively low ridges, basins, and valleys. Therein is the lowest pass in the Andes between Colombia and southern Chile—Abra de Porculla at 2145 m. In the Huancabamba Depression, there is a structural deflection of the Andean faults that corresponds to two major tec- tonic segments of the Andes (Sillitoe, 1974). South of the depression the axis of the Andes is northwest to south- east; north of the depression the axis is north-northeast— south-southwest. Geological evidence points to extensive marine trans- gressions in the region of the Huancabamba Depression during the Cretaceous (Ham and Herrera, 1963). Orogenic events associated with plate tectonics along the west coast of South America in the Late Cretaceous resulted in the uplift of the Andes to elevations probably not exceeding 1000 m above sea level (Zeil, 1979). The major uplift of the ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU > Andes south of the depression was in the Miocene (Harrington, 1962; Aubodin et al., 1973; Sempere et al., 1990); the final major uplift was in the Pliocene with some additional orogeny in the Pleistocene (James, 1973; Gansser, 1973; Noble et al., 1990). In contrast, the uplifted terrain resulting from the Late Cretaceous orogeny subsequently was eroded to low hills before the major uplift of the Andes north of the Huancabamba Depression was initiated in the Pliocene; this uplift continued into the Quaternary, as evi- denced by many active volcanoes in Ecuador and Colom- bia (Herd and Naeser, 1974; Sauer, 1971; Shagam, 1975; Simpson, 1979). Thus, the Huancabamba Depression is not only a physiographic anomaly in the Andes and a tectonic border, but it bridges tectonic segments that were uplifted at different times. The present elevations and drainage patterns in the Andes of northern Peru and the Huancabamba Depres- sion probably were achieved in the Pleistocene (Gansser, 1973; Harrington, 1956). During the Pleistocene, climatic fluctuations included cooler, drier conditions during gla- cial phases, and warmer, more moist conditions during interglacial phases. According to Sauer (1971), during in- terglacials, climates were depressed 1500-2000 m in the Ecuadorian Andes, whereas in the Peruvian Andes they were depressed 1000-1500 m on the eastern slopes and 500— 1000 m on the western slopes (Hastenrath, 1967; Dollfus, 1976). These postulated changes are in accord with the paleoecological work in Colombia by van der Hammer (1974), van der Hammen and Cleef (1986), and Clapperton (1987), and in the Ecuadorian Andes by Colinvaux (1988). These climatic fluctuations presumably resulted in alter- nating isolation and interconnection of montane environ- ments, thereby providing corridors for, and barriers to, dispersal in the Andes (Simpson, 1979; Colinvaux, 1993). PHYSIOGRAPHY A casual glance at a physiographic map of South America reveals the long Andean mountain chain border- ing the west side of the continent; as such, one might ex- pect major drainages to be east and west. Although there are many rivers originating in the Andes, some of the most significant drainages are north and south. This is especially evident in Colombia where the Rio Magdalena and Rio Cauca form broad valleys between three major north-south cordilleras. Between the Nudo de Pasto (Macizo Colombiano) in southern Colombia, the Andes are formed by two major ranges in Ecuador—the western Cordillera Occidental and the eastern Cordillera Oriental. Between the eastern and western cordilleras are 10 basins that are completely or partially separated by transverse ridges that in most cases connect the cordilleras. South of the dry valley of the Rio Jubones, the Cordillera Occidental diminishes in isolated ranges, such as the Estribuciones de Celica. On the other hand, the Cordillera Oriental is continuous with the Cor- dillera de Huancabamba and Cordillera de Tabacones in northern Peru, both of which terminate north of the Rio Chamaya, a tributary of the Rio Maranon (Fig. 2). In Peru, the Cordillera Occidental forms the backbone of the Andes; it is confluent with the Cordillera Oriental at the Nudo de Pasco. To the north of the Nudo de Pasco, the valley of the Rio Maranon separates the Cordillera Cen- tral from the Cordillera Occidental. The northern part of the Cordillera Oriental is separated from the Cordillera Central by the valley of the Rio Huallaga, which, as it curves eastward, forms the northern boundary of the Cor- dillera Oriental. Thus the easternmost range of the Andes in Peru north of about 7°S Lat. is the Cordillera Central. In southern Ecuador and northern Peru, the complex topography is associated with north and south drainages (Fig. 2). For example, the Cordillera del Condor is bordered on the west by the Rio Narangaritza (flowing northward) and on the east by the Rio Cenepa (flowing southward). The major drainage system in the region is the Rio Maranon, which at its confluence with the Rio Huallaga forms the Rio Amazonas. With the exception of the Rio Chamaya, the major tributaries of the Rio Maranon flow southward (e.g., Rio Huancabamba, Rio Cenepa, Rio Chinchipe) or northward (e.g., Rio Chotano, Rio Utcubamba, Rio Chiriaco). These rivers separate distinct north-south mountain ranges (e.g., Cordillera de Huancabamba, Cordillera Colan; Fig. 3). The change in di- rection (from north to east) of the Rio Maranon forms the northern boundary of the Cordillera Central. The north- ern part of the Cordillera Central is dissected by rivers that flow northward (Rio Chiriaco and Rio Utcubamba) and southward (Rio Mayo). The resulting complex topography contains major isolated highland areas (e.g., Cordillera Colan) and the high elevations to the east of the Rio Mayo; many ridges more than 1000 m above sea level extend southward almost to the Rio Huallaga. Unlike other moun- tain ranges in the region, the highest parts of the Cordil- lera del Condor consist of sandstone table mountains (Fos- ter and Beltran, 1997). CLIMATE Temperature is dependent primarily on elevation. In the lower valley of the Rio Maranon, temperatures may exceed 40°C. At high elevations (> 3000 m), mean monthly temperatures are relatively constant throughout the year, but the daily fluctuation can be as much as 20°C with noc- turnal lows below freezing (Schwerdtfeger, 1976). Rainfall patterns are determined by wind patterns and topography. The prevailing winds from the Amazon Ba- 6 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS ; | Méndez e | @ Cities and towns o©— Collecting localities Major roads ---— International boundary —-— Departmental and provincial boundaries % : Rio Comainas <. (gc =e x Alfonso Ugarte / 4 Abra de Zamora | &P. V. Comainas ) 5 oy f —S | RO = ) @y ¥ | ; t=) Ss BS 2B 1 Zamora oS ( Se ' 06p\\¥ (2 OA &a5? A AZ On a oly e Palambla : . =e Huancabamba AN 2 ING | ElTambo /\ } j ; \ \ { Loreto Canchaque ya p= [ \ | La Peca Te Abra Pardo-de Miguel p< Venceremos— - y Rio Coren ayaa qeouenH \ 8 Bagua eque \ xe \ ___Moyobamba Yurimaguas SS Rioja x See Chachapoyas re WN, ~ ¥— Abra Tangarana Levanto AS ° Rio > San Anares \ c a Ae es Cainarachi de Cutervo \? j Cataratas 9 NS Leimebamba wie Ahaushiyacu By) . 63 Ingenio Rio 9 P) Santa Cruz ; C2 DY ae NS. Zapatero—o Tarapoto San Martin Pacific Ocean Fig. 1. Political map of the Andes of northern Peru and southern Ecuador showing major rivers and localities mentioned in the text. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 7 < 1000 m 1000-2000 m 2000-3000 m 3000-4000 m > 4000 m 50 Kilometers ee BS oly =) = ls ic} Pacific Ocean Fig. 2. Physiographic map of the Andes of northern Peru and southern Ecuador. SCIENTIFIC PAPERS, Meters 81° 4000 79° Cordillera Oriental 3000 Cerros de Amotape - 1000 Meters 4000 Cordillera de pF Tabacones Cordillera de Huancabamba = 3000 Ss <— Rio Maranon Meters — 4000 Cordillera Occidental 1000 =< Rio Nangartitza Rio Maranon NATURAL History Museum, THE UNIVERSITY OF KANSAS 78° 76° Cordillera del Condor 4°10'S x Rio Cenepa < Rio Santiago <— Rio Morona <— Rio Pastaza Cordillera Cordillera Central < Rio Huallaga Cordillera Central 6°10'S <— Rio Sonche <— Rio Mayo <— Rio Huallaga Fig. 3. Profiles of the Andes in southern Ecuador and northern Peru. Vertical exaggeration 35X. sin bring moisture to the eastern slopes of the mountain ranges throughout the region. Generally rainfall is high- est (+ 2500 mm annually) at elevations of 500-1500 m on the windward slopes and lower at higher elevations and on the leeward slopes. Deep valleys, such as those of the Rio Maranon and its tributaries in the Huancabamba De- pression are in rain shadows and receive only 250-500 mm of rain annually; most of this falls in October—May. Influenced by the cold Humboldt Current, the winds blowing off of the Pacific Ocean have relatively little mois- ture; some of this moisture is dissipated as fog on the dry western flanks of the Cordillera Occidental, where annual precipitation may be less than 150 mm. However, as noted by Foster and Beltran (1997), the relatively low passes in the western cordilleras permit the westerlies to provide rainfall on the higher mountains to the east (e.g., Cordil- lera del Céndor and Cordillera de Huancabamba). Of course, the former receives its greatest rainfall from the winds off the Amazon Basin. Sporadically, rainfall is in- creased along the Pacific lowlands and on the western slopes of the Andes by the southward movement of the warm waters of the counter-equatorial current, El] Nino. VEGETATION The complex topography and associated climatic pat- terns result in a great variety of vegetation formations. Herein we use the terminology for vegetation formations (and associated maps) in Peru proposed by Tosi (1960, based on Holdridge’s [1967] classification) and extend these into Ecuador, as classified and mapped by Canadas (1983). As noted by Savage (1975) and Lynch and Duellman (1997), the Holdridge system seems to be too sophisticated ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 9 for determining broad patterns of animal distribution. Therefore, we have simplified the terminology by com- bining some of the categories. A transect from west to east across northern Peru at approximately 05°30' S. Lat. begins in coastal desert, as- cends dry, rocky, slopes to humid highlands, drops to an arid valley and repeats this sequence until descending into the Amazonian lowlands. In making this transect, one passes through dry scrub forest, dry montane forest, hu- mid montane forest, and lowland rainforest. Each of the major vegetation formations in northern Peru is discussed briefly below; Tosi’s (1960) terminology is in parentheses. Desert scrub.—(Desierto subtropical y tropical; Maleza desértica subtropical y tropical). Essentially, the entire Pa- cific lowlands of Peru is extremely dry; with annual rain- fall less than 150 mm, only a few xeric-adapted plants are present. In the drier regions, vegetation may be absent or consist only of a terrestrial gray Tillandsia. Farther inland and close to the base of the Andes, vegetation becomes more diverse with legumes (e.g., Prosopis juliflora and Capparis sp.) and cacti (e.g., Cererus, Lemairocereus, and Opuntia. Thorn forest.—(Bosque espinoza subtropical y tropi- cal). In regions receiving up to 500 mm of rainfall annu- ally, this type of forest develops on the western slopes of the Cordillera Occidental to elevations of about 1200 m. It also is prevalent in the interior valleys of the rios Chamaya, Chinchipe, Huancabamba, Maranon, and Utcubamba, as well as in the Catamayo Basin in Ecuador. Trees consist principally of legumes (Prosopis and Acacia) with other drought-resistant trees especially near streams—Bombax, Bursera, Jacaranda, and Pithecolobium. Cacti (Cereus, Opun- tia, and Lemairocereus) are numerous, especially in the middle Maranon Valley, where dense cactus forest pre- dominates. Both terrestrial and arboreal bromeliads (e.g., Tillandsia) are locally abundant. In many areas, especially near rivers, there are irrigated fields of rice or sugar cane. Dry forest.—(Bosque seco tropical; Bosque muy seco tropical; bosque seco subtropical). Occurring peripheral to the thorn forests in the interior river valleys and in the rain shadow in the lower Rio Mayo Valley, there is a forest that develops in areas receiving 500-1000 mm of rain an- nually. This forest has no closed canopy and is made up of moderate-sized trees of diverse genera (e.g., Bauhinia, Bombax, Bursera, Cordia, Centrolobium, Curatella, Inga, Pithecolobium, Tabebuia). The epiphytic Spanish moss (Tillandsia usneoides) is prevalent locally, and both terres- trial and arboreal bromeliads (e.g., Pitcairnia and Tilland- sia) are abundant locally. In the lower Rio Mayo Valley and extending to the middle part of the Rio Huallaga, the dry forest receives more rainfall (1000-2000 mm annually). In this region, the forest contains several genera of trees (e.g., Brosimum, Cedrela, Myroxylon, Swietenia) characteristic of humid tropical forest. Montane dry forest.—(Bosque seco montano bajo). Iso- lated patches of this forest occur at elevations of 2000-2500 m in the upper Rio Maranon Valley, in the Cordillera Occi- dental in the vicinity of Cajamarca and southward, in the vicinity of Chachapoyas in the Cordillera Central, and in the mountains south and west of Loja in Ecuador. Natural vegetation consists of trees such as Jacaranda acutifolia, Caesalpinia tinctoria, and various Acacia and Mimosa. Gen- erally, these areas are heavily cultivated and have plantings of Agave americana and Eucalytus globulus. Humid Montane Forest.— (Bosque humedo montano). This is the dominant type of forest on the lower slopes (up to about 2500 m) of the cordilleras Huancabamba, Tabacones, Condor, and Colan, where it is continuous to the Cordillera Central. Principal trees include Berberis, Polylepis, and Eugenia. At higher elevations (>3000 m) in the Cordillera Occidental, this vegetation is reduced to few trees, bushes (Baccharis) and bunch grasses. Much of this forest has been cleared and cultivated; at higher elevations Eucalyptus has been planted. Very humid montane forest.—(Bosque muy humedo montano). This is the “ceja de la montana” or “cloud for- est” characteristic mostly of windward slopes at elevations of 2500-3000 m and receiving 1000-2000 mm of rainfall annually. This forest exists on the higher ridges of the cor- dilleras Huancabamba, Tabacones, Condor, and Colan, as well as on the slopes (especially eastern) of the Cordillera Central in Peru and the Cordillera Occidental in Ecuador. A variety of trees, often stunted and covered with lichens and mosses, includes Polylepis and Podocarpus. Also charac- teristic are bushes (Baccharis) and the spiny bamboo (Chusquea spicata). Arboreal bromeliads are locally abundant. Humid subtropical forest—(Bosque humedo y muy humedo subtropical). Covering the lower (500-1900 m) eastern slopes of the Cordillera Central and middle Rio Mayo Valley, and lower slopes of the Cordillera Oriental in Ecuador, this type of forest develops in areas receiving as little as 1000 mm of rainfall annually to others that re- ceive rainfall in excess of 3000 mm. The forest consists of a variety of moderate to large trees including Juglans neotropica, Cedrela fissipes, Tabebuia, and genera common in the Amazonian lowlands—Brosimum, Cordia, Inga, Piper, Swietenia. In some areas, the forest has been cleared for citrus and coffee plantations. Subtropical pluvial forest—(Bosque pluvial subtropi- cal). In northern Peru, this type of forest is known only on the highest ridges of the Cordillera Central east of the Rio Mayo Valley, where rainfall is expected to be in excess of 4000 mm annually. This type of forest consists of stunted trees with emergent palms, such as Euterpe. 10 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Humid tropical forest—(Bosque humedo tropical). This is the Amazonian rainforest, which receives rainfall in excess of 2000 mm annually and has a great richness of tree species. The trees form a continuous, or nearly so, canopy 30-40 m above the ground, but there are canopy emergents, such as species of Cedrela, Ceiba, and Ficus. This diverse forest contains many kinds of palms (e.g., Bactris, Triartea, Scheelea), but under certain edaphic and hydrologi- cal conditions, the forest is clearly dominated by one spe- cies of palm, Mauritia reflexa. Other formations.—Local climatic and edaphic condi- tions may result in different local vegetation formations. For example, the Abra de Zamora at 2800 m in the Cordil- lera Oriental of Ecuador is windswept and has vegetation consisting of thick growths of low bushes supporting mosses and large bromeliads. Although Tosi (1960) mapped the highest reaches of the Cordillera Colan as very humid montane forest, data on a field tag of an Eleutherodactylus collected by Thomas S. Schulenberg at 3300 m in that cordillera noted “in a grassy bog above treeline.” In the poorly explored Cordillera del Condor, the flat-topped sandstone mountains at 2000-2300 m are mostly are covered by sclerophyllous shrublands—shrubs and small trees 2-5 m high; according to Foster and Beltran (1997), these shrublands are composed mostly of species of Ilex, Weinmannia, Clusia, and Persea. GENERA OF ELEUTHERODACTYLIINE FROGS Four genera within the tribe Eleutherodactylini, as defined by Lynch (1971) occur in the Andes of northern Peru. Only species in the genus Eleutherodactylus are treated herein. However, we provide brief diagnoses of external features of all four genera presently recognized in the Andes of northern Peru, so that specimens can be allocated to the proper genus. There is no substantial evidence that any of the genera are monophyletic, and Eleutherodactylus may be paraphyletic with respect to the other genera. For example, Lynch (1986) questioned the allocation of spe- cies to Phyllonastes or Phrynopus, and Harvey and Keck (1995) noted the occurrence of “diagnostic” characters of Ischnocnema in some species of Elettherodactylus. Eleutherodactylus Duméril and Bibron, 1841 Diagnosis.—Terrestrial or arboreal frogs characterized by (1) skin on venter smooth or areolate; (2) relative lengths of Fingers I and II variable; (3) relative lengths of Toes III and V variable; (4) digits bearing terminal discs and pads; (5) tips of digits rounded, elliptical, or truncate; (6) tarsal tubercle present or absent; (7) palmar and subarticular tu- bercles not greatly enlarged; (8) adult size 10-120 mm SVL. Content.—About 600 species in five subgenera, only one of which (Eleutherodactylus) includes the 45 species known from the Andes of northern Ecuador. Distribution.—Tropical and subtropical America (ex- clusive of arid regions), through the West Indies and into southwestern United States. Remarks.—The 45 species known from the Andes of northern Peru are members of four species groups, which are defined in the following Accounts of Species. Ischnocnema Reinhardt and Lutken, 1862 Diagnosis.—Terrestrial frogs characterized by (1) skin on venter smooth; (2) Finger I > II; (3) Toe V > III; (4) digits bearing small terminal discs; pads present or absent; (5) tips of digits rounded; (6) tarsal tubercle absent; (7) pal- mar and subarticular tubercles greatly enlarged; (8) adult size 30-55 mm SVL. Content.—Five species, of which only Ischnocnema saxatilis and I. simmonsi are in the Andes of northern Ecua- dor (Duellman, 1990b; Lynch, 1974a). Distribution.—Upper Amazon Basin, Andean slopes in southern Ecuador, northern Peru, and Bolivia. Remarks.—Harvey and Keck (1995) questioned the validity of Ischnocnema.' Phrynopus Peters, 1874 Diagnosis.— Terrestrial frogs characterized by (1) skin on venter smooth or areolate; (2) relative lengths of Fin- gers I and II variable; (3) Toe III longer than, or equal in length to, Toe V; (4) digits not bearing terminal discs or pads; (5) tips of digits rounded; (6) tarsal tubercle absent; (7) palmar and subarticular tubercles not greatly enlarged; (8) adult size 18-45 mm SVL. Content.—More than 20 species, of which three have been reported from the Andes of northern Peru; descrip- tions of three others are being written (Duellman, in prep.). Distribution.—Humid habitats at moderate to high elevations in the Andes from Colombia to Bolivia. In the Andes of northern Peru, Phrynopus nebulanastes and P. parkeri are endemic to the Cordillera de Huancabamba, and P. simonsii is widespread in the northern part of the Cor- dillera Occidental (Cannatella, 1984; Lynch, 1975a); one of the undescribed species is from the northern part of the Cordillera Central, and two are from the Cordillera Occi- dental. ‘It is doubtful that the Amazonian Ischnocnema quixensis and the three Andean species are related to the type species, /. verrucosus, in southeast- ern Brazil. If that should be shown to be true, Oreobates Jiménez de la Espada, 1872 (type species by monotypy O. quixensis) is an available ge- neric name. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 11 Remarks.—In reviewing the osteology of Phrynopus, Phyllonastes, and other genera, Lynch (1986) noted that the Phrynopus peruvianus group and Phyllonastes may be re- lated. Phyllonastes Heyer, 1977 Diagnosis.—Terrestrial frogs characterized by (1) skin on venter smooth; (2) Finger I shorter than or equal in length to Finger II; (3) Toe III shorter than Toe V; (4) digits not bearing terminal discs or pads; (5) tips of at least Toes III and IV pointed; (6) prominent tarsal tubercle present; (7) palmar and subarticular tubercles not greatly enlarged; (8) adult size 13-20 mm SVL. Content.—Four species, two of which occur in the Andes of northern Peru. Distribution.—Phyllonastes myrmecoides inhabits the upper Amazon Basin, whereas three montane species have been reported from only a few localities—P. heyeri from Alamor, Ecuador, and the Cordillera de Huancabamba in Peru (Lynch, 1986)”; P. lochites from the Cordillera del Con- dor in Ecuador (Lynch, 1976); and P. lynchi from the north- ern part of the Cordillera Central in Peru (Duellman, 1991b). Remarks.—See Remarks under Phrynopus. SUMMARY OF TAXONOMIC CHARACTERS Morphological characters, size, and color pattern of members of this genus in western Ecuador were described in detail and illustrated by Lynch and Duellman (1997). Consequently, we treat the characters rather briefly herein; 22 morphological characters and eight features of color pattern for species of Eleutherodactylus in the Andes of northern Peru are tabulated in Tables 1 and 2. As deemed necessary, we attempt to clarify some of these characters below. Skin texture —Although statements in the diagnoses and descriptions may suggest that various states are dis- crete, they are not in many cases. For example, the distinc- tion between the dorsal skin being shagreen with scattered tubercles versus finely tuberculate is not discrete, and cer- tainly different persons may describe the conditions in different ways. The ventral skin commonly has been de- scribed as granular; we use the term areolate. In contrast, fine granules on the dorsum give the skin a finely textured appearance that we term shagreen. If the granules are large and subconical or conical, we define the texture as tuber- culate. Dorsolateral folds on the body are evident in seven species of the Eleutherodactylus conspicillatus Group (E. avicuporum, citriogaster, condor, karcharias, lymani, and peruvianus) and in E. araiodactylus. Of these, E. avicuporum and E. karcharias also have interrupted, longitudinal folds on the flanks. In the Andes of northern Peru, E. avicuporum is unique in having a dermal interocular ridge. This fea- ture also is present in another member of the Eleutherodactylus conspicillatus Group, E. skydmainos (Flores and Rodriguez, 1997) and in some other species of the (e.g., E. quinquagesimus [Lynch and Duellman, 1997)). Tubercles.—The most noticeable and taxonomically useful tubercles are those on the upper eyelid and heels. We divege from Lynch and Duellman (1997) in the recog- nition of eyelid tubercles; they used the term only for the presence of enlarged conical or subconical tubercles on the eyelid. No species in the Andes of northern Peru possess such distinctive tubercles, but many species have a few pungent tubercles on the upper eyelid. If a tubercle is present on the heel, it usually is small, but four species (Eleutherodactylus galdi, lanthanites, muscosus, and quaquaversus) have large, conical tubercles on the heel. Tuberculation on the tarsus is highly variable. Tu- bercles are absent in several species (e.g., Eleutherodactylus citriogaster, colodactylus, proserpens, and rufioculis). Tubercles are present on the outer surface of the tarsus in many spe- cies (e.g., E. anemerus, bearsei, schultei, and versicolor); the outer tarsal tubercles are noticeably conical in E. galdi. The inner edge of the tarsus may be unadorned (e.g., E. condor, infraguttatus, percnopterus, and phoxocephalus), bear a low, elliptical tubercle distally (e.g., E. ceuthospilus, cuneirostris, nephophilus, and petrobardus), or have a low fold distally (e.g., E. atrabracus, avicuporum, melanogaster, and rhodoplichus). A row of tubercles may be present on the ventrolat- eral surface of the forearm. These ulnar tubercles are ab- sent in several species (e.g., Eleutherodactylus bromeliaceus, incomptus, lymani, and phoxocephalus). The tubercles are especially prominent in E. cryptomelas and E. infraguttatus, conical in E. galdi, and coalesced to form a short fold in E. pinguis. Two species (Eleutherodactylus anemerus and E. proserpens) have a noticeable tubercle (papilla) on the tip of the snout, and E. phoxocephalus has a low vertical keel on the snout. Eleutherodactylus karcharias is unique in hav- ing a finlike middorsal tubercle on the body. Tympanum.—As emphasized by Lynch and Duellman (1997:28): “The terms ‘tympanum’ and ‘external ear’ have been used to identify a combination of characters rather than a single character. The ‘tympanum’ is a combination 2A specimen (LSUMZ 39364) from the Cordillera Colan extends the known range of this species eastward into Departamento Amazonas, Peru. ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS NA _ JNOUS UO [aay [PITIAA, eP1oqn) [esiO Ppa oxTUNs UMOUAUN S}I][s [LOA JO UOIpUOD, juasqe S}I[s [RO0A, ATjeseq paqqam saoj. Apogq uo poy [e19}e] OsTy, PIO} [euep zepnd019}U], ynous jo dy uo apueqny, juasqe sa1oydojuopo aura, , 5) yes >I L4 ayeount, §=juessqy dOls a) papunoy ‘pi Ajaynoy juesqy yoours aye[Oa1y suai a juasqy >I 4 jeoudyq -‘said/sqy dors Vv pepunoy ‘“wmoeqns juasqy uaa13eys aye[Oaly lojoo1s4aa S) juesqy Il>I L4 ayeounly, = juesqy dOls Vv ‘pa Ajayndy = ud eqns juesqy uaaiseys a}e[Oo1y xvphyjousays q juasqy >I os jeoudyy wesqy dors Vv ‘pryunfg ‘umoeqns juesqy = ayepMosaqny, a}e[Oory snyjidipuasas 5) [jews >I La papunoy juasag GOs Vv ‘ad “pout summdeqns juesqy useiseys aye[Oa1y 1ajjnyas =) juasqy Il>I Il jeondyy wesqy dors q pepunoy papunoy juesqy Yyyoows aye[Oa1y ostjnoouna 5) juesqy Il>I La popunoy ywosag dOls Vv ‘ad ‘pul —- ayeurummdy juesqy uaaiseys a}e[Oa1y snyoijsopoys =) [jews Il>1 141 jeondyq yuesag Gols V ‘pa Ajandy =“ uM eqns juasqy uaei3eys aye[oa1y isnya1jdopoy. B) [eoru0> >I 14 jeondyq yueseig QOl> ») papunoy ~“uwmdeqns juasqy a}e[Od1V uaaiseys snsaaavnbonb =) juasqy Il>1 L4 pepunoy juasag QdOl> Vv pepunoy ,uIndeqns juesqy a}e[Ooly a}e[Ooly suadsasoid q juesqy Il>1 It [Tews juasqy COIs Vv ‘pi yunig “pr Ajanoy juesqy a}e[Oo1y ayejoary psinsuid 5 juasqy ll>I eed jeondyq wesqy dors Vv ‘ummdeqng = papunoy juesqy uaai3eys ayejoary —, smjujdao0xoyd D yews Il>I 14 jeoudyyq wesqy dolls V pepunoy papunoy juesqy iepnysng aye[Oory snpavgorjad q juasqy II _ ayeounty, ‘said/'sqy GOls Vv pepunoy ‘uwmoeqns juasqy Iepnod1eqny, aye[Oa1y (snajdoussad B) [Tews >I REL pepunoy jyuesag GOs V pepunoy —“wmnoeqns juasqy yoours aye[Oo1y 1yoad q juesqy ll>1 = yews juesqy dOIs q papunoy papunoy juasqy yoours aye[Oo1Vy ysoytoyod B) pepunoy [>] 14a jeoundyq jywosag dOl= Vv pepunoy “umdeqns juasqy a}e[Ooly uaaiseys 1uapuayoo 5) yTews >I La jeondyq jyuesaig Gols Vv papunoy papunoy yuasqy yjoows a}e[Ooly snypiydoydau S) Jeoruos Il>I Tel ayeounly, juesag dolls Vv ‘pi yung = - pa qunyg juesqy yyoows a}e[Ooly /Snsoosnul | juesqy I = yews juosqy dors q pepunoy papunoy juosqy a}e[Ooly aLOIY —_g, a] SUSOUvjaUL q juasqy I La jeoudyq jesag dors d pepunoy ‘“wmoeqns juesqy a}e[oa1y aye[OIIVy SN]]2A1] q Tews I WG aqyeouniy = jyuasealg GOs Vv pepunoy papunoy juasqy yyooursg a}e[Ooly snyoynsvifui B) juasqy ll>I I jeoudyg jyuesqy dors Vv pepunoy pi Ajaynoy juasqy yoours a}e[Ooly (snyduoout .) yeoruo> >I L4 ajyeounly, §=ywesag dOl> Vv ayeouniy == ayeuTUMD Vy juasqy a}e[OaIIV yyoows 1pjvs .) juesqy ll>I — jeoudyy juasarg dor Vv ‘pi yunig “pr Ajanoy juesqy o}yeounIL, aye[Oo1y snysi4oxa q juesqy WI — ayeounty, «= uasqy = =dOls Vv ‘pi Ajay ~—s papuNoy juasqy yoouws YyOowSs /Slqsou1gung 2) yTeus Il>1 La jeoudyq” juosaig dOl< Vv pepunoy ‘wmoeqns juasqy uaeiseys a}e[Oory sujawuojdhsg d juesqy » yews ll>I La papunoy jyuasag GOs d popunoy papunoy juasqy a}e [Oly a}ejoory 98M pAjavpojoo qd juasqy I La jeoudyq” jqesaig dors Vv ‘pa Ajay pa AjoyNDVy juesqy uaaiseys a}epoory snyiydoyjnaa 5) juesqy ll>1 i pepunoy jyuosag QOS Vv pepunoy papunoy juasqy uaarseys a}epoory sisuaoivuinlv9 » [Tews >I La jeondyq jyuesag dors V ‘pa Ajaynoy = uM eqng juesqy yoours aye[oaly snaouijaulo.1g =) juesqy >I Ua ajyeound, jussqy dOl< Vv papunoy papunoy juesqy —-ayeNosaqny, a}e[Ooly lasavaq Ret juesqy I< 4 ayeountLT = yuasag GOs Vv pepunoy ‘pi jyunyg Juesolg yoours aye[ooly unsodng1av q juesqy Tea La yews juesqy dOIs V ‘piyunig = “pr qunyg juesqy uaeiseYys a}e[Ooly (SNOvAQuqD 2) juesqy >I La pepunoy juesqy dOrls .) pepunoy papunoy juesqy yoours aye[Oaly snyaAuojppav Vv juasqy =" eel yews juasqy dOlI> Vv pepunoy papunoy juasaly yoous yoous snjAjovpoiwav 2) juasqy ll>I Tea ayeounty, = Juasqy dors Vv ayeounly, «= ,oyeuTUIND juesqy = ayeNOAaqny, a}epoory (snuaulauv D juesqy Tear — jeoudyq_ yuesqy dOl>> .) ‘ad pout = ayeurumm oy juesqy YyyOous a}ejooly snqouruinap uOHIpuoD sepieqn, [3] sesurny siosuy = Sapseqny, YpIA UoyTpuoD aYOLg [esiop SPIO} umsiop Io}UdA satads a0], ]2°H s1osuLy [eysIq uo sysiq pyeAq pyeAq wmnuedurdy ynous ynous [etoyZJOsIOg = ud UTS uo UTYS ‘TI 20], << A a0, = D ‘AT 20], JO aposaqny szepnoyreqns ayeury[Nued 07 Burpus}xe jou ynq |] AOL, < A POL = ‘A 90], = [I] POL = V :uoNtpuoDd doy, ‘sa0} = J ‘s1o8uy = J ‘sour [eySiq ‘yUasqe snfnuue pure ouviquiow = q ‘AT[eQUEA ATUO JUAPIAY SN_NuUe puke JUAsqe oULIqWO = D ‘UPYs ysno1yy AIISIA sn[NUUR Jo adURIAa;UMIID JO JSOW ynq JUasqe auLIquiaW = g ‘jUaUTUIOId snynUUR puUk oURIqUIEI = YW :UOyIpUOD LMUedWAL ‘ayeUTUMSegns = “UMdeNS ‘ATTRIJUAAOIE}SOd pourpout = ad -pouy ‘papunos Ayunyq = ‘pa yunqg ‘papunos Ajaynoe = “pa Ajaynoy :ynousg “Mag UsaYy}AOU Jo sapuy oy} WOIT sMpApovposayjna]y Ul S19}IeIeY [eINJON.YS JO $9}L}S *] A[GLL. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 13 of (1) a differentiated tympanic membrane and (2) a tym- panic annulus.” They recognized four states of tympanic annuli and membranes. In most Elewtherodactylus in the Andes of northern Peru, a prominent tympanic membrane and annulus are present. The membrane is not differenti- ated but most of the annulus is visible beneath the skin in E. melanogaster, pataikos, and rufioculis, whereas only the ventral part of the annulus is visible below the skin in E. acuminatus, ardalonychus, quaquaversus, and wiensi. There is no external evidence of an ear in E. colodactylus and E. lirellus. Fingers and toes.—In all members of the Eleutherodactylus orestes and unistrigatus groups in the Andes of northern Peru, the first finger (thumb) is shorter than the second. The first finger is longer than the second in most species in the Eleutherodactylus conspicillatus Group, but the two digits are about equal in length in E. cuneirostris and E. karcharias, as well as in E. araiodactylus in the Eleutherodactylus nigrovittatus Group. As shown by Lynch and Duellman (1979), the relative lengths of Toes III-IV is an important taxonomic character. Except for E. araiodactylus, in which Toes III and V are of equal length, Toe V is longer than Toe III in all species in the region. In members of the Eleutherodactylus conspicillatus and orestes groups, Toe V is only slightly longer than Toe III, and nei- ther toe extends to the level of the distal subarticular tu- bercle on Toe IV. In species in the Eleutherodactylus unistrigatus Group, Toe V is noticeably longer than Toe III and extends to the middle or distal border of the distal subarticular tubercle of Toe IV. With the exception of species in the Eleutherodactylus nigrovittatus and orestes groups, in which the discs on the fingers and toes are barely expanded, the discs on Fingers I and II and all toes are expanded. Their shapes are de- scribed arbitrarily as rounded, elliptical (much wider and long), or truncate. Most species have lateral fringes on the digits. We do not distinguish between fringes (thin) and keels (thick). Webbing between the toes is absent in all species, except three members of the E. conspicillatus group (E. avicuporum, karcharias, and metabates) in which basal webbing is present. Coloration.—The dorsal coloration can be used readily for identification of most species of frogs—not so for most species of Eleutherodactylus, in which distinctive coloration may be present only in the groin or on the anterior and/or posterior surfaces of the thighs (Table 2). In northern Peru, only two species are easily identified by their dorsal col- oration—uniform green (in life) in E. acuminatus and uni- form orange-red (in life) in E. anemerus. In preservative, the dorsal coloration of most species is varying shades of tan, brown, or gray, usually with darker markings. These may consist of distinct spots (e.g., E. cuneirostris), chev- rons (V-shaped marks with the apex anteriad) (e.g., E. lanthanites and E.lymani), or a W-, X-, or H-shaped mark in the scapular region (e.g., E. ardalonychus, exoristus, and sternothylax). Other dorsal patterns include dark longitu- dinal streaks or dashes (e.g., E. ceuthospilus, galdi, and percnopterus), a large middorsal blotch (e.g., some E. proserpens), and pale vermiculations (E. muscosus). Identi- fication is confused further by pattern polymorphism is several species. For example, some individuals of E. colodactylus, proserpens, rhodoplichus, and sternothylax have pale dorsolateral stripes, whereas some individuals of E. schultei and E. wiensi have dark dorsolateral stripes. Some specimens of E. bearsei and E. petrobardus have pale spots on the dorsum. Facial markings usually consist of dark labial bars, but these are absent in some species (Table 2). Eleutherodactylus cuneirostris and some specimens of E. peruvianus have a pale labial stripe. A dark canthal stripe is evident in most species and is especially notable in E. acuminatus and E. galdi. The canthal stripe is incorporated into a dark face mask encompassing the loreal region in E. peruvianus. The coloration on the venter in most species is cream (with or without dark flecks) to brown. However, distinc- tive coloration is present in a few species—bright yellow (in life) in Eleutherodactylus citriogaster, uniform black in E. melanogaster, black on the ventral surfaces of the hind limbs in E. atrabracus, and brown or black marbling or reticula- tion (e.g., E. ardalonychus, infraguttatus, muscosus, and ver- sicolor). Whereas the groin is colored like the flanks in most Eleutherodactylus, distinctive markings are present in some species. Commonly these are pale spots, which are white, yellow, or red in life; such spots are characteristic of most species in the Eleutherodactylus orestes Group, but also ex- ist in E. cajamarcensis, condor, lirellus, muscosus, and rufioculis. The groin has dark reticulation in E. phoxocephalus and is black in E. cryptomelas. In preserved specimens of Eleutherodactylus, the pos- terior surfaces of the thighs usually are varying shades of tan to brown, with or without pale flecks or spots that are cream to red in life (Table 2). Notable exceptions are dark reticulations (E. phoxocephalus, quaquaversus, and E. rhodoplichus), pale mottling (E. ardalonychus, citriogaster, and infraguttatus), or black with white spots (E. lymani). Sizes and proportions.—The sizes of adults of Eleutherodactylus are highly variable. With the exception of some species in the Eleutherodactylus conspicillatus Group, most species in the Andes of northern Peru have SVLs of less than 35 mm (Table 3). Considerable sexual dimorphism in size is evident in most species of Eleutherodactylus, in which females usually are at least one third larger than males. In the species in the Andes of northern Peru, SVLs SctentiFic PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS 14 yeory] UO sy~paTj sjods 931M sjods 10 oUON Avi3 YIM ayy YIM org poyrewup, sjods y1eq = suorAayp yAeG SOK 4 Sak mouth] *J Of! UL MOIJOA Sy99|} YOR|G seq jeuocBbeip H 1epndeos jods utor5y YIM ureaID uUMOIq WIOFTUQ jods ajeg jure peoig SaX SOK ou ‘4juIey Sn]]aat] “J yeoryy Avis sypopy Avis sjods ayed qjeus uo adiys ayy AA TM 9M ATTAg YIM uMoIg payzeuup, seq [euOseIq sSuoIAaYyD y1eGq Sa ou ‘Sax SOK sapUvyjUv] “FZ syxpayj UMOIqG sjods ureas9 SUOIAIYD OM} auON UTM uleaID WIM uUMoIg payreuuy, Jeg [euoseiq ‘M ze[ndeasg Jeuoseiq sax SaK SULADYIADY "J OI] Ul por Surpqieul UMOIq SurpjJOU UIeaID Ajiortaysod sjods Ysry IoWeyUy YIM weaID YIM uUMOIg payzeuupy, = umMoig yyIMuey syods sepndeag Sox ON Sax snyojynsvsful “J JOU 10 Yeas UOIAaYpD [eIDesS auoN uMOIq ULIOFTU UMOIg UOJ, ~—- payAeLU), Jeuo8erp aug = “jp eTNdeag Sax SOK Sax snqdwoout “J Ost] Ur saysep yep uaei3 umnsi10q urvald WHOFTUA Weal WIOsUA = payreutuA) = Wea WOJIU, = JIM RAID MOLILN ON Sax 1pjvs -J adiys Spay UMOIq sypayj weaID SUOIAYp [eIIeS [eyyued asnyyiq YIM uel, YIM uMOIg poxyreurup, sieq [euOSeIq ‘YH 10 (A Iepndvas Sax Say Say SN4jStoXa “J uOTENIVeL UMOIq sjods adiys [erqey apeg = yurey YIM wear UMOIg ULIOFIUA, = payreuU) ue} WOFTUE) yaep Auepy SOK ON ON SLAJSOAIIUNI “FZ uoQepNdyer UMO1q UOIAVYp [eIDeS PTPIXe ur yoeyg YIM wWeaiD sprig sprig wreaip ‘M te[ndeag jeuoseiq sax Sax sujauiojdhud “J Aqjaq uo SurpyoUL auoN uModg ‘JeoIy] UMOIg sods weaid sjods afeg sieq][euoseiq suorAayd yeq Sax Sox Sax AOPUuod “J alos ut adiys [e49}e]OSIOp ae umoig UMOIg UOJ, = payreuTu, «= Ava 1ouUMoIg = Av3 10 UMOIg ON ON ou “ale g snjAjoupojo2 “Jy at] Ut MOTTA WYZIIq JayUsA —«4JLO.TY] UO SyYpaT] Ye SulyOW weet payrzeuup sieq [euOseIq suOoIAeYyD UMOIg Sax Sax Sax AdaJSVSOLAJ1I “J azt] UT MOTTA spay s}ods aed saysep Yep YIM sjods a]eg UMOIG YIM FeJ YIM uey, sjods aye UP} WIOFTUA, 10 Ue} WOFTUA ou /sax Sax Sax snpidsoyjnao “J yl] UI par sjods sjods 914M sjods ay1yM sjods y1ep saypjo[q sysry] pue uloIg = Yep YJIM UeaID YIM KI YIM »peig WIM uey, yrep [peus qurey que sax sisuaodvulnlDo “J OFT] UL 66 ut sypay sjods uteai9 UuOJAVYp IO use13 UNsIOq = UMOI Y}IM) wWRaID YIM uMoIg payreuun, ue} wuoyTuQ, = sjods yep May sjodg Sax SaX snaovljauo4g “J autos ut ATTesiop sypayy wuva1d sypayj weed = say} 0Tq yep sjods ayed a8ie7] YIM uMOIg UMOIg WIOFUA = payreuu) YIM UMOIg ‘M re[ndeasg sax sax Sax lasavaq “Yq uoeMoyer uorAayp ajed auoN uMOIq JUTey sjods gjed Jews payrzewupy ue} uO, Jods yep [erpayy yurey ures qurey wnaodngiav “J squirt] pury jo uoleEpnoyer JoJUdA UO YoLTg uMoig umoig wuostUA, jods aed a81eq] UMOIg WIOJTUQ UMOI WHIOJTUQ ON ON ON SNIVAQUAJD “FJ SOsIp [e}I81p uo UOIAVY [eIIeS sa8poa y1eq suoyeNyer uMoIg SUIJOW WeaID = payreWUp siveqeuoseiq ‘pq aejndeog jeuoseiq sax sax SnYyoAuo[Mpav “FZ adrays yreur uerpeut ayed [esioppIyy ue} WOsIUA ue} WIOFTUA payreuupy, UMOIg YIRq Yep yIM geg qurey ON say snjijoupoiwav “J ost] UT por -a3ueIO WNSIOG SYDa]J YLT YIM aTeq ajed waojtuq peyreuuy, ayed wuojytuq aged wuo0jruq ON ON ON SndaulauD “FZ odiys [euyjueo yperg aed wos aqed wuojstuQ paxyreuuQ ajed wuojtuq =—ayed wuoyruq ON ON qurey SNJDUIMNIV “J sSuryreur Ajjaq pue syst} UuIOI5 syurpy Apogq jo sieq sieq Ieq satads Iay1O OIL IOI19}S0,] wms10q quury rerqey [ey qro18}U] ‘9SIMAaYJO pajou ssayun ‘suswdads paasasaid Jo yey} st UOLIO[OD “Nag U1ay.IOU Jo sapuy ayy WO s/jijovposayjnaly JO UOYeIOJOD “Z 9{qeL RODACTYLUS IN THE ANDES OF NORTHERN PERU ELEUTHE auos ut adiys sjods umoiq sjods urea Surp}jOUr yep [ero}L[OsIOp ye Iepnsaiy YIM uMOIg poayreuuy YIM wear sjods y1eq Sax SaX SOX 1suaiand “J aulos UT syuey uonPE[NoVer spay ureard s1eq [PITIOA uo sjods ayy, \pr]q 10 umoIg UQIM uUMOIg payrewup, Ioyeuoseiq — saypjoyq Yue Sax saX Sax 40]0I1SAaQ “J auwios ul adiys spay uUMOIq syeays SuUOIAaYO [ero}LOsSIOp eg YIM wIeatD UMOIg WIOFTUA, paxyeuTUA, euoSeiq ‘jj to’H ‘xX Iendeog [euoseiq sax a[ Suey, xophyjousass *Y sypayy UMOIq UOIAaYP [RIDeS auoN YIM utealD UMOIg UOJ payreUUug uMoIg geq /syareut sepndeag SaX SOK SOK snqidipuasas *7 autos ut adiys syeoljs yep [era}e[OsIOp xe TUM ajed wos payreutuy, WIM [eg ayed wuostuq ou juIeJ ON ou ‘sax layjnyos *y UMOIG YIM sypapy urvaro sjods y1ep aft] UL pat Sty paxpepy ApIAvayy WIM uMoIg sjods wea = sjods weary ~~ aejndeds |Jeug ques ON SaX synaouna J aft] UL par Aype19qe] s}ods syearys peioes syieul 1eaUurT] sypoyy YIM ue, ue} WIOFUA payreuup, yrep [pews pue azejndeas Sax ques Sax SNYISOpoy. “J awios ut adiys spay yep uoyEnoyar uOQLMWeI IO —- syAeUT yep [e1a}e]OsIOp geg YIM urea yaeq payreuruy, sieq]euoseiq repnSar ypeuig ou ‘Sax ON aysueny snyaydopory. “J autos UI suoyepnonal SUOIAVY 10 sjods [equa,, TUM umoig payzeurup, ue} WIOFUQ, sjods y1eq jeuoseiIq ON sax snsdaavnbunb -7 OUOS UT uMolq IO yoo sadiyjs [e1a}ZjOsI0q + UMOIG WAOFTUQ wieaId ULIOJIUQ, = payeWUA, ue} WIOFTUA a8] 10 Xx Sax ON Sax suadaasold “J uoyeNoyer syieul UMOIG jods spay dUuON IO SyDoTF Ye YIM wreaID ajed a81e7] JUON yrep [pews Jeuoseiq oN jurey simsuid *7 QUIOS UT WNsIOp uonepnoyar uonepnoyar uOoHYNHeI = —- sYAeU Yep Ay uo sjods aeg wieaI ULIOJIUA yaeq ued sep 10 suoN -N8arI1 10 aUON ou 4urej = ON ou ‘sax snjvoydasoxoyd *y QWs UT WIMsI0p spay YOrq sjods ureaso syealys uo sjods ayn, UIM 9yTY WIM uMoIg payreurup, sieq [euoseIq yAep aepnSa.y jeuoseiq sax SOK snpavgoijad *y Aqjesiop syxpoyj UMOIq sypayy ajed pause}jedun auog UQIM wRalD YIM UMOIg paxreuuy, sivg [euOSeIq suoIAaYD yeq jeuosriq sax ou ‘sax snupianiad "y spay PLIG uMmolq umMoIq syeaijs 10 sjods auoN [fetus YIM uel ayed wu0j1uqQ payer, ayed wu0jymQ = ype, tepndeog Jeuoseiq sax Sax snajdouasad *J UMOIg JIM syeays auON paxpayy ATIAvap] UMOIG WIOJTUA, = payeMUA, §= UMOIG WIOJTUQ, 10 sjods yaeq Sax sax ou ‘sox 1yoad “J at] ut adtys [eujues urea ue} UOJ UMOIg WIOJUA, = payzeuUA, «= UMoq WAOJTUQ, UMOIq WHIOFIUA, ON ON ON soyrwjud “J OUIOS UT sjods umoiq suOIAayp yurey 10 payjodsun 13}u0,, UIIM FT UMOIG UOJ, «= payxreuTU, ue} ULIOJTU, §~—- IO [A AeTNd vag jeuoseiq sax MOLILN 1uapuayao “J uMOJg YIM sieq wieald uonP[NoVaL 10 syea]s 10 AFI] UT par SLIT paxpayy ApiAvapy WIM uMorg payreuup sieq[euoseiq — sayajoyq ye jeuoseiq sax ou ‘sax snjiyjdoydau “gy squiy pury uonPepNoyel UMOIq sjods uvaid sjods ay1YM ~— SUOTRNOTULIAA japun sjods ajeg UTM urea WIM uMoIg sjods avg WIM UMOIg —-9}TYM YIM YEG SaK Sa yurey SNSOOSNML “J Jeo] UO SyDaTJ sypayy urvara SUOIADY 10 /pue auoN umoag /ueaID UTM uMoIg payreurup sieq[euoseiq yeu padeys-x qurey yurey yurey Sayuqujaul “J apt] UL MOTTA uro13 ut sjods SPR]q WO; yoryq wos, sjods avg SPL]G UOJ = yoeyq wWuAO;Up, ON ON ON 4ajSvsouvjaul “J ssurieu Ajjaq pue sysiyy UlOID syuepy Apog jo sieq sieq Ieq satoadg IIYIO 1O110}SOg uinsi10q jeiqe] [eyq4to19; Uy jeory |, quar] ponunuos Z afqey, 16 ScrentTiFic Papers, NATURAL History Museum, THE UNIVERSITY OF KANSAS Table 3. Body sizes (SVL in mm) of Eleutherodactylus in the Andes of northern Peru. Values include range (mean, sample size), SD = sexual dimorphism. Unless noted otherwise, measurements are for Peruvian specimens only. Species Males Females SD E. acuminatus 21.3-21.5 (21.4, 2) 24.3 1.14 E. anemerus 20.4 — — E. araiodactylus == 24.5 — E. ardalonychus 19.7-21.9 (20.8, 2) 27.4 ie E. atrabracus 18.9 PY 1.20 E. avicuporum 25.2 31.0-34.3 (32.4,6) 1.29 E. bearsei 22.7-25.5 (24.0, 3) 38.0-38.8 (38.4),2) 1.60 E. bromeliaceus' 16.7—23.2 (20.8, 14) 22.9-28.1 (26.5,5) 1.27 E. cajamarcensis 22.6-25.9 (24.7, 5) 27.0-32.0 (29.2,14) 1.18 E. ceuthospilus 19.9-25.8 (21.3, 35) 23.5-26.7 (25.2,8) 1.18 E. citriogaster 31.6-41.3 (37.0, 9) 42.0-51.0 (44.4,4) 1.20 E. colodactylus 16.8-19.6 (18.3, 19) 19.1-23.2 (21.5,9) 1.17 E. condor* 32.1-39.5 (36.5, 9) 52.6-63.2 (58.7,3) 1.61 E. cryptomelas* 28.2—-30.2 (29.2, 4) 38.6 1S: E. cuneirostris — 29.1 _ E. exoristus 15.0-16.9 (16.2, 4) 21.3-23.5 (22.7,6) 1.40 E. galdi* 17.1-24.8 (21.0, 8) 28.1-34.0 (31.7,11) 1.51 E. incomptus? 15.6-18.8 (17.5, 10) 23.7-25.9 (24.5,7) 1.40 E. infraguttatus 15.8 22.9-24.3 (23.6,3) 1.49 E. karcharias _ 30.4 —_— E. lanthanites° 21.7-26.0 (23.5, 20) 27.5-44.8 (36.2,32) 1.54 E. lirellus 14.1-17.0 (15.3, 13) 19.4-24.0 (21.5,12) 1.39 E. lymani 39.7-45.3 (41.9, 4) 48.4-66.7 (58.7,4) 1.40 E. melanogaster 20.6—22.8 (21.9, 3) 24.2-24.7 (24.5,2) 1.12 E. metabates 29.9-32.2 (31.1, 2) — — E. muscosus — .- 29.6-46.1 (37.8, 4) — E. nephophilus = 24.6-34.0 (29.8, 5) = E. ockendeni’ 16.9-21.2 (19.1, 3) 24.6-31.5 (28.0,12) 1.47 E. pataikos = 21.6 = E. pecki® 15.3-18.7 (17.3, 3) 25.2 1.46 E. percnopterus 2D=25:2)(22:0;10) 25.9 1.15 E. peruvianus® 26.1-30.0, 28.1, 8) 36.0-42.5 (39.1,9) 1.39 E. petrobardus 27.0-30.7 (28.5, 7) = = E. phoxocephalus 22.7-27.8 (26.2, 5) 38.9 1.48 E. pinguis — 28.4—29.8 (28.9, 3) — E. proserpens” 15.2—21.0 (18.6, 11) 20.2—23.5 (22.0,4) 1.18 E. quaquaversus'” 19.6—22.5 (20.7, 29) 24.6-31.3 (27.3,20) 1.32 E. rhodoplichus 21.8-28.9 (25.8, 17) 30.1-34.2 (32.5,4) 1.26 E. rhodostichus 19.3 — — E. rufioculis 18.1 20.6 1.14 E. schultei 23.5-26.5 (25.0, 10) 28.4-34.0 (31.8,5) 1.27 E. serendipitus 20.4-21.2 (20.8, 2) = = E. sternothylax 18.3-29.1 (24.2, 39) 28.3-36.7 (32.1,6) 1.33 E. versicolor 21.8-24.7 (23.3, 2) 26.0-27.3 (26.5,3) 1.14 E. wiensi 27.8-33.0 (30.7, 6) 37.0 1.21 ‘Provincia Morona-Santiago, Ecuador (Lynch, 1979) *Including specimens from Provincia Morona-Santiago, Ecuador (Lynch and Duellman, 1980) *Provincias Loja and Santiago-Zamora, Ecuador (Lynch, 1979) ‘Cordillera Oriental, Ecuador, and Cordillera del Condor (Lynch and Duellman, 1980) ‘Provincia Napo, Ecuador (Lynch and Duellman, 1980) "Upper Amazon Basin (Lynch, 1980) 7Amazonian Peru (Lynch, 1980) ‘Including specimens from the Cordillera de Cutucu, Ecuador (Duellman and Lynch, 1988) *Cordillera Oriental, Ecuador (Lynch, 1979) Cordillera Oriental, Ecuador (Lynch and Duellman, 1980) of females are 1.12-1.61 (x = 1.25) greater than those of males. The three groups of Eleutherodactylus in the Andes of northern Peru have distinctly different proportional hind- limb lengths (Fig. 4). In members of the Eleutherodactylus orestes Group, the mean tibia/SVL ratio is 37.0-39.6 (x = 38.4, n = 4), whereas members of the Eleutherodactylus conspicillatus Group have much longer hind limbs; the mean tibia/SVL ratio is 60.4-65.6 (X = 62.2, n = 9). Ratios among species in the Eleutherodactylus unistrigatus Group are intermediate—43.3-56.0 (Xx = 49.4, n = 31). The lowest ratios (43.3 and 44.1) are for two bromeliad-dwelling spe- cies, E. colodactylus and E. proserpens; elimination of those two species gives a range of 47.5 (E. phoxocephalus, also a bromeliad-dweller), to 56.0 (E. rufioculis and E. wiensi) with a mean of 49.9. ise) [o) ine) ol Tibia Length (mm) a 8 @ E£. conspicillatus group — — WV E- nigrovittatus group =a fo) A E. orestes group @ E. wnistrigatus group Head Width (mm) 15 20 25 30 35 40 45 50 55 Snout-vent Length (mm) Fig. 4. Correlations of tibia length (A) and head width (B) with snout-vent length in Eleutherodactylus in the Andes of northern Peru. Symbols for the species groups are the same in both graphs. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU IDENTIFICATION OF SPECIES Key TO THE SPECIES This key emphasizes those external features usually independent of maturity and sex; however, juveniles are always difficult to identify, because the distinctness of the tympanic membrane and annulus increases with age, and many features of color patterns on the venter and concealed surfaces of the limbs develop with age. This key is only a guide; it is necessary to confirm the identifications by re- ferring to the diagnoses and descriptions. 1G 10. le ip. OER aon cera thane ep lll oecceeceeeecencestacenceeceseceeeerereree == 2 Toes Il and V equal in length.............. E. araiodactylus . Toe V only slightly longer than Toe II] ............0 3 Toe V much longer than Toe II] ...............cccsseseeees 15 . Finger I longer than, or equal in length, to Finger II; outer fingers with greatly expanded discs Finger I shorter than Finger II; outer fingers with nar- TOV ALO UNG CURCHISCS Weerrcecrtctere eee ae ee See 2 BS KamMOnM Oe lvaaceO laters ecarasssseessaccstessersasrscesesecsaeascscsr = 5 SKeMMOMM DE hy sSTOO Elileeresteeceeeee teeter sceesecerscersrsestensnseses 6 . Fin-shaped middorsal tubercle present; interocular 180! JOURESEIAL snc cosciconrecncnasnpcqococecasecanecosecAoseNG060 E. karcharias Fin-shaped middorsal tubercle absent; interocular fold istallliyajores emt, wiealkctr...s.rccsscsscceseresos=2s E. avicuporum MloesnwiebbedsbaSalllyjtssrecsesesce E. cunetrostris Rostro no en forma de cuna; region labial usualmente con barras; patron dorsal no compuesto de muchas MANChaS OSCULAS PEQUENIAS ...........sseccecseecereereneesenseeeens 9 . Superficies posteriores de los muslos oscuras con PUAMAtLOS PAlidOs: <..<.cce-c-sccencesecencesececeecscecnecesveecesreeesseses 10 Superficies posteriores de los muslos con monteado (DIREUTRS) V7 [BAVC aco -cecncencnasnnnpco coe roc53053030¢ 0c c0gE OP ACCOCOOOREE 11 . Garganta oscura; vientre del cuerpo y los miembros posteriores con monteadO OSCUTO «0... E. condor Garganta palida; puntillos 0 reticulaciones oscuras en Lab@ataltal yaCl PCCM Oleceseneereeceeetesscarcesaes E. peruvianus Superficies posteriores de los muslos negras con monteado crema; marcas oscuras en el margen de la FBR Hl OYE) sccnccbccccececccene rose CerOTp Lon SCARDCERTEDOSSCOSS E. lymani Superficies posteriores de los muslos café oscuro con monteado café palido; vientre palido (amarillo en vida) con manchas difusas de café oscuro en la garganta ... ~epeeececstaneegacesncasincuone conan sececonobacresoneqsesacoc00s500" E. citriogaster Dorso y vientre negro Dorso café; vientre variable E. melanogaster Superficie ventral de los miembros posteriores negra E. atrabracus Superficie ventral de los miembros posteriores crema ©) CUS CIRO) escort pesca ere co eB asce one oes on See R aC Eo eaS oo SEE: 14 Mancha palida grande en la ingle; superficies posteriores de los muslos crema con marcas café....... E. pinguis Sin mancha palida en la ingle; superficies posteriores de los muslos uniformemente café E. pataikos Membrana timpanica no diferenciada ...............0+- 16 Membrana timpanica distintiva con anillo timpanico bien defindo ....... Tee CDCR Co SCPEECI EEs R S 22 ANTI (CaM RUA EO GIOSEIMUD ro sococosaacocoasaccaconc aso UCR 7 Anillo timpanico visible debajo de la piel o evidente solamente ventralmente Dedos d la mano cortos y tiesos, con discos redondos; barras de los labios y miembros ausentes; sin mancha palicdatenwl avingl eee nerecrseetescersesrtcrssces E. colodactylus Dedos de la mano largos, delgados, con discos elipticos; barras de los labios y miembros presentes; mancha palida en la ingle... E. lirellus Mubéreuloypresente em El CalOM <.2.0.0554000m 50 e. avicuporum ). citriogaster E. karcharias E. metabates E. peruvianus Fig. 8. Localities of known occurrence of five species in the Eleutherodactylus conspicillatus group in the Andes of southern Ecuador and northern Peru. lateral fringes; toes webbed basally; discs on toes about equal in size to those on outer fingers; tip of Toe V extend- ing to distal edge of pentultimate subarticular tubercle on Toe IV; tip of Toe III extending to middle of pentultimate subarticular tubercle on Toe IV (Fig. 6A); when hind limbs flexed perpendicular to axis of body, heels broadly over- lapping; shank 58.0-63.9 (x = 60.7)% SVL. Coloration in preservative: Dorsum brown with darker brown markings consisting of small round spot middorsally about midway between scapular region and sacrum, canthal and supratympanic stripes, minute spots posterolaterally from scapular region (2 specimens), row of minute spots just ventral to dorsolateral fold (1 speci- men); diffuse brown marking consisting of interorbital bar, labial bars that extend onto margin of lower lip, and nar- row transverse bars on forearms, thighs, shanks, and tarsi; one individual with diffuse brown chevron posterior to sacrum, and one with diffuse brown spot posterior to sacrum followed by diffuse transverse mark posteriorly; flanks tan. Posterior surfaces of thighs dark brown with small tan spots, especially distally. Throat and belly tan with diffuse brown reticulation; ventral surfaces of thighs and shanks creamy tan with diffuse brown reticulations and flecks; ventral surfaces of tarsi black, bordered by nar- row creamy tan stripes. Coloration in life: Unknown. Measurements of holotype: SVL 33.2, tibia length 20.1, foot length 17.7, head width 13.1, head length 13.9, IOD 4.1, upper eyelid width 2.4, E-N 4.0, eye, 3.5, tympanum 2.0. Distribution and habitat.—Eleutherodactylus avicuporum is known only from elevations of 1700-2030 m on the western slopes of the Cordillera Colan in northern Peru (Fig. 8). All individuals were on the ground in humid montane forest. Etymology.—The specific name is derived from the Latin avicupis meaning bird catcher, and the genitive plu- ral suffix -orwm. The name is used in recognition of the efforts expended by ornithologists from the Museum of Zoology, Louisiana State University, collecting amphibians in the Cordillera Colan. Remarks.—The five juveniles have SVLs of 14.0-19.3 (x = 17.4) mm. The interocular dermal ridge is weak in three individuals and absent in two, both of which are males. The coloration is like that of the adults, except that in three of the juveniles a diffuse brown chevron in present immediately posterior to the middorsal dark brown spot. Eleutherodactylus citriogaster Duellman Eleutherodactylus citriogaster Duellman, 1992b:24. Holotype: KU 212277, adult female, from Cataratas Ahuashiyacu, 730 m, 14 km [by road] NE Tarapoto, Departamento San Martin, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) conspicillatus Group having (1) skin on dorsum shagreen, lacking tubercles, that on venter smooth; discoidal fold evident; dorsolateral folds indistinct; (2) tym- panic membrane and tympanic annulus prominent, round, its length about 30% length of eye; (3) snout subacuminate in dorsal view, rounded in profile; canthus rostralis angu- lar; (4) upper eyelid lacking tubercles, slightly narrower than or equal to IOD; cranial crests absent; (5) vomerine odontophores triangular, prominent; (6) males having vo- cal slits and nuptial pads; (7) Finger I longer than II; discs broad, round; (8) fingers lacking lateral fringes; (9) ulnar tubercles present distally; (10) heel and tarsus lacking tu- bercles; (11) inner metatarsal tubercle oval, 3x oval outer metatarsal tubercle; plantar supernumerary tubercles nu- merous; (12) toes lacking lateral fringes; webbing basal; Toe V longer than III but not extending to distal subarticular tubercle of Toe IV; discs as large as those on fingers; (13) dorsum grayish brown with or without nar- row brown chevrons and interorbital bar; venter cream with brown flecks on throat and thighs; (14) SVL in males 31.6-41.3 mm, in females 42.0-51.0 mm. In general appearance, Eleutherodactylus citriogaster (Fig. 9) closely resembles two other species in the E. conspicillatus ‘TENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Eleutherodactylus citriogaster, KU 212277, female, 42.0 mm SVL. Eleutherodactylus melanogaster, KU 212321, female, 24.7 mm SVL. Eleutherodactylus pataikos, KU 212320, female, 21.6 mm SVL. : cs Eleutherodactylus anemerus, KU 219798, male, 20.4 mm SVL (ERW). Eleutherodactylus ardalonychus, KU 212301, female, 27.4 mm SVL. Eleutherodactylus ceuthospilus, KU 219776, male, 22.2 mmS (ERW). Eleutherodactylus infraguttatus, KU 212297, Fig. 9. Eight species of Eleutherodactylus from the Andes of northern Peru. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 5) re POU ES id = ts ty Sgt © A ; a as ° Eleutherodactylus nephophilus, KU 212307, female, 28.3 mm SVL. Eleutherodactylus petrobardus, KU 212292, male, 28.7 mm SVL. 2 - ae Eleutherodactylus serendipitus, KU 181279, male, 20.4 mm SVL. Eleutherodactylus wiensi, KU 219795, male, 32.1 mm SVL (ERW). Fig. 10. Eight species of Eleutherodactylus from the Andes of northern Peru. 26 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Fig. 11. Ventral color patterns of three large species in the Eleutherodactylus conspicillatus group. A. E. citriogaster, KU 212287; B. E. condor, KU 147020; C. E. lymani, KU 181265. From Duellman (1992b). Group in northern Peru and southern Ecuador—E. condor and E. lymani. The three species are distinguishable on the basis of color pattern on the posterior surfaces of the thighs—bold black and white spots or reticulations in E. lymani, brown with pale flecks in E. condor, and dark brown and tan mottling in E. citriogaster (Figs. 11A, 12A). Fur- thermore, E. citriogaster has a round, instead of bifid, pal- mar tubercle. Other members of the E. conspicillatus Group are smaller; of those in the upper Amazon Basin and lower slopes of the Andes, E. conspicillatus, malkint, metabates, and peruvianus have the posterior surfaces of the thighs dark brown with small cream to red spots, and two of these (E. malkini and E. metabates) have basal webbing between the toes, whereas E. fenestratus, lanthanites, and vilarsi have uniformly dark brown posterior surfaces of the thighs. Furthermore, E. lanthanites has median pale stripe on a dark throat and a prominent tubercle on the heel, and E. vilarsi has a bifid palmar tubercle. Other small members of the E. conspicillatus Group in the region differ in structure and coloration. Eleutherodactylus buccinator has pink spots on the groin on the hidden surfaces of the thigh (Rodriguez, 1994); E. karcharias has a prominent fin-shaped middorsal tubercle, and E. avicuporum and E. skydmainos have an interocular fold. In E. avicuporum and E. karcharias, the skin on the belly is areolate. Description.—The description by Duellman (1992b) is adequate and contains all information known about this species. Distribution and habitat.—This species is known only from elevations of 600-800 m in lower humid montane forest on the slopes of a ridge northeast of Tarapoto, Departamento San Martin, Peru (Fig. 8). All adults were observed at night on rocks in and along cascading streams. Remarks.—Duellman (1992b) overlooked the fact that the presence of a round, instead of bifid, palmar tubercle Fig. 12. Color patterns on the posterior surfaces of the thighs in three large species in the Eleuthero- dactylus conspicillatus group. A. E. citriogaster, KU 212287. B. E. condor, KU 147020. C. E. lymani, KU 181265. From Duellman (1992b). is shared in Eleutherodactylus only with E. zeuctotylus, a member of the Eleutherodactylus conspicillatus Group in northeastern South America (Lynch and Hoogmoed, 1977). Eleutherodactylus condor Lynch and Duellman Eleutherodactylus condor Lynch and Duellman, 1980:18. Holotype, KU 146992, a juvenile female, from the Rio Piuntza, Cordillera del Condor, Provincia Morona-Santiago, Ecuador. Diagnosis.—A member of the Eleutherodactylus conspicillatus Group having (1) skin on dorsum shagreen, that on venter smooth; discoidal fold evident; dorsolat- eral folds present; (2) tympanic membrane smooth, and tympanic annulus prominent, nearly round, its length about % length of eye; (3) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid lacking tubercles, slightly narrower than or equal to IOD; cranial crests pal- pable in large females; (5) vomerine odontophores promi- nent, triangular; (6) males possessing vocal slits and nonspinous nuptial pads; (7) Finger I longer than II; discs expanded, about twice width of digit proximal to pad; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles; (11) inner metatar- sal tubercle elongate, 6-8x round outer metatarsal tubercle; supernumerary plantar tubercles few or absent; (12) toes bearing narrow lateral fringes; webbing absent; Toe V ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 27 slightly longer than III; discs slightly smaller than those on outer fingers; (13) dorsum brown with darker brown markings—labial bars, interorbital bar, chevrons; venter cream with brown blotches; throat heavily pigmented with brown; posterior surfaces of thighs brown with cream or white spots; (14) SVL in males 32.1-39.5 mm, in females 52.6-62.2 mm. In general appearance, Eleutherodactylus condor is most similar to E. citriogaster and E. lymani. The venter in E. con- dor is creamy white with dense gray pigmentation on the throat, belly, and ventral surfaces of the thighs (Figs. 11B, 12B); the posterior surfaces of the thighs are brown with pale spots. In contrast, in E. lymani, the venter is essen- tially unpigmented, except for dark bars on the margin of the lower lips and posterolaterally on the throat; the pos- terior surfaces of the thighs are black with bold cream mottling. In E. citriogaster, the venter is yellow with brown or gray mottling on the throat, laterally on the belly, and on the ventral surfaces of the thighs; the posterior surfaces of the thighs are mottled dark brown and creamy tan. Fur- thermore, the palmar tubercle is round, instead of bifid, in E. citriogaster. Other members of the E. conspicillatus Group are smaller; of those in the upper Amazon Basin and lower slopes of the Andes, E. conspicillatus, malkini, metabates and peruvianus have the posterior surfaces of the thighs dark brown with small cream to red spots, and two of these (E. malkini and E. metabates) have basal webbing between the toes, whereas E. fenestratus, lanthanites, and vilarsi have uniformly dark brown posterior surfaces of the thighs. Furthermore, E. lanthanites has median pale stripe on a dark throat and a prominent tubercle on the heel, and E. vilarsi has proportionately shorter hind limbs and larger tym- pana. Other small members of the E. conspicillatus Group in the region differ in structure and coloration. Eleutherodactylus cuneirostris has a wedge-shaped snout and many small dark spots on the dorsum, and E. buccinator has pink spots on the groin on the hidden surfaces of the thigh (Rodriguez, 1994); E. karcharias has a prominent fin- shaped middorsal tubercle, and E. avicuporum and E. skydmainos have an interocular fold. Eleutherodactylus avicuporum and E. karcharias differ from other members of the group in the region by having areolate skin on the belly. Distribution and habitat.—The few records for Eleutherodactylus condor suggest a rather limited elevational distribution on mountain ranges to the east of the Cordil- lera Oriental of the Andes (Fig. 13). The species is known from 1975 m in the Cordillera de Cutucti (Duellman and Lynch, 1980), the type locality at 1550 m* and Coangos at 1500-1600 m (Almendariz, 1997) on the western slope of the Cordillera del Céndor, and from 1750 m on the eastern *Lynch and Duellman (1980) erroneously gave the elevation of the type locality as 1830 m. < 1000 m 1000-2000 m 2000-3000 m 30004000 m B >4000m Fig. 13. Localities of known occurrence of four species in the Eleutherodactylus conspicillatus group and one species in the Eleuthero- dactylus nigrovittatus group (E. araiodactylus) in the Andes of southern Ecuador and northern Peru. slope of the Cordillera del Condor. All individuals have been taken in humid montane forest. Remarks.—This first Peruvian record is based on USNM 525437, an adult female having a SVL of 60.2 mm. The specimen was obtained at a site on the upper Rio Comainas, at the base of Cerro Machinaza in the Cordil- lera del Condor, by Thomas S. Schulenberg on 6 August 1994. Eleutherodactylus cuneirostris new species Holotype—LSUMZ 39369, an adult female, from 12 km [by trail] E La Peca (05°36' S, 78°19' W, 1700 m), Provincia Bagua, Departamento Amazonas, Peru, obtained on 11 June 1978 by L. J. Barkley. Diagnosis.—A member of the Eleutherodactylus conspicillatus Group having (1) skin on dorsum smooth; that on venter smooth; discoidal fold evident; dorsolat- eral folds absent; (2) tympanic membrane smooth, and tympanic annulus prominent, slightly higher than long, its length slightly more than % length of eye; (3) snout long, rounded in dorsal view, protruding well beyond margin of lower lip and acutely rounded in profile; (4) upper eye- lid lacking tubercles, narrower than IOD; cranial crests 28 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS absent; (5) vomerine odontophores prominent, oblique; (6) condition of vocal slits and nuptial pads unknown; (7) Finger I equal in length to II; discs on outer fingers ex- panded, truncate, about twice width of digit proximal to pad; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tarsus lacking tubercles; inner edge of tarsus bearing low, elliptical tubercle dis- tally; (11) inner metatarsal tubercle elliptical, about 3x subconical outer metatarsal tubercle; supernumerary plan- tar tubercles absent; (12) toes lacking lateral fringes; web- bing absent; Toe V slightly longer than III; discs slightly smaller than those on outer fingers; (13) dorsum tan with small dark brown spots; venter creamy tan with diffuse brown reticulations on belly and ventral surfaces of thighs; posterior surfaces of thighs brown; (14) SVL in one female 29.1 mm. Eleutherodactylus cunetrostris differs from all other mem- bers of the Elewtherodactylus conspicillatus Group in the re- gion by having a long, wedge-shaped snout and a con- spicuous pale labial stripe. Eleutherodactylus peruvianus also has a pale labial stripe, but the dorsum usually has dark chevrons (otherwise uniform tan, red, or green); the throat has dense dark flecks, and there are no reticulations on the belly. Of other small members of the E. conspicillatus Group, E. fenestratus, lanthanites, and vilarsi have uniformly dark brown posterior surfaces of the thighs. Furthermore, E. lanthanites has median pale stripe on a dark throat and a prominent tubercle on the heel, and E. vilarsi has propor- tionately shorter hind limbs and larger tympana. Other small members of the E. conspicillatus Group in the region differ in structure and coloration. Eleutherodactylus bucci- nator has pink spots in the groin and on the hidden sur- faces of the thighs (Rodriguez, 1994); E. karcharias has a prominent fin-shaped middorsal tubercle and areolate skin on the venter, and E. avicuporum and E. skydmainos have an interocular fold. Eleutherodactylus metabates lacks a pale labial stripe and has X- or chevron-shaped marks on the back. In the larger species in the E. conspicillatus group in the region, the venter in E. condor is creamy white with dense gray pigmentation on the throat, belly, and ventral surfaces of the thighs; the posterior surfaces of the thighs are brown with pale spots. In E. lymani, the venter is es- sentially unpigmented, except for dark bars on the mar- gin of the lower lips and posterolaterally on the throat; the posterior surfaces of the thighs are black with bold cream mottling. In E. citriogaster, the palmar tubercle is round (in- stead of bifid), and the venter is yellow with brown or gray mottling on the throat, laterally on the belly, and on the ventral surfaces of the thighs; the posterior surfaces of the thighs are mottled dark brown and creamy tan. Description.— (n = 1 female). Head as wide as body. HW 40.5% SVL; HL 44.6% SVL; Snout long, rounded.in dorsal view, protruding well beyond margin of lower lip and acutely rounded in profile, giving wedge-shaped ap- pearance (Fig. 5B); E-N 97.1% length of eye; nostrils nota- bly protuberant, directed laterally; canthus rostralis angu- lar, straight; loreal region noticeably concave; lips rounded; upper eyelid lacking tubercles; upper eyelid width 70.6% IOD; supratympanic fold weak; abruptly curving down- ward behind tympanum, not obscuring edge of tympa- num; side of head inclined ventrolaterally; tympanic mem- brane prominent; tympanic annulus slightly higher than long; length of tympanic annulus 55.9% length of eye; two low, rounded postrictral tubercles posteroventral to tym- panum. Choanae large, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores prominent, oblique, narrowly separated medially, located well behind posterior edges of choanae, each odontophore bearing five teeth. Tongue about twice as long as wide, barely notched posteriorly; posterior half not adherent to floor of mouth. Skin on dorsum smooth; dorsolateral fold absent; skin on flanks and venter smooth, except coarsely areolate on proximal posteroventral surfaces of thighs; discoidal fold prominent; cloacal sheath short; large tubercles in cloacal region absent. Ulnar tubercles absent. Thenar tubercle el- evated, elliptical, slightly smaller than bifurcate palmar tubercle; palmar supernumerary tubercles absent; subarticular tubercles prominent, subconical; fingers lack- ing lateral fringes; Finger I equal in length to II. Discs on Fingers I and II small, round. Discs on outer fingers trun- cate, about twice width of digit proximal to pad; all fin- gers having ventral pads well defined by circumferential grooves. Upper surfaces of hind limbs smooth; heel and outer edge of tarsus lacking tubercles; inner edge of tarsus bearing small, elliptical tubercle distally; inner metatarsal tubercle elevated, elliptical, about 2x subconical outer metatarsal tubercle; plantar supernumerary tubercles ab- sent; subarticular tubercles small, subconical; toes lacking lateral fringes; toes webbed basally; discs on toes about equal in size to those on outer fingers; tip of Toe V extend- ing to distal edge of penultimate subarticular tubercle on Toe IV; tip of Toe III] extending to base of penultimate subarticular tubercle on Toe IV (Fig. 6B); when hind limbs flexed perpendicular to axis of body, heels overlap by about one-third length of shank; shank 65.6% SVL. Coloration in preservative: Dorsum tan with dark brown markings consisting of many small round or ovoid spots on dorsum of body and flanks (Fig. 7B). Canthal and supratympanic stripes; transverse mark above vent and small spots distally on anterior surfaces of thighs; narrow transverse bars on limbs barely evident; broad creamy tan labial stripe. Posterior surfaces of thighs dull brown; throat creamy tan with minute brown flecks; belly and ventral surfaces of thighs creamy tan with diffuse, faint brown reticulations. Ventral surfaces of tarsi black. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 29 Coloration in life: Unknown. Measurements of holotype: SVL 29.1, tibia length 19.1, foot length 15.6, head width 11.8, head length 13.0, IOD 3.4, upper eyelid width 2.4, E-N 3.3, eye, 3.4, tympanum 1.9. Distribution and habitat.—Eleutherodactylus cuneirostris is known only from 1700 m in humid montane forest on the western slopes of the Cordillera Colan in northern Peru (Fig. 13). Etymology.—The specific name is an adjective from the Latin cuneus meaning wedge and the Latin rostrum mean- ing snout. The name refers to the wedge-shaped snout. Remarks.—Among species of Eleutherodactylus in Ec- uador and Peru, the long, wedge-shaped snout of this spe- cies is approached only by the long, subacuminate snout in E. longirostris, a member of the Eleutherodactylus (Craugastor) fitzingeri group in the Chocoan region. Eleutherodactylus karcharias Flores and Rodriguez Eleutherodactylus karcharias Flores and Rodriguez, 1997:392. Holotype, MCZ 89075, immature female, from Alva, 1000m, between Chachapoyas and Bagua Grande, Departamento Amazonas, Peru. Diagnosis.—A member of the Eleutherodactylus conspicillatus Group having (1) skin on dorsum shagreen with scattered tubercles, especially on flanks, prominent fin-shaped middorsal tubercle; skin on venter areolate; discoidal fold prominent; dorsolateral and flank folds present; (2) tympanic membrane smooth, and tympanic annulus prominent, slightly higher than long, its length about /% to % length of eye; (3) snout rounded in dorsal view and in profile; (4) upper eyelid lacking tubercles, broader than IOD; cranial crests present; (5) vomerine odontophores prominent, triangular; (6) condition of vo- cal slits and nuptial pads unknown, (7) Finger I equal in length to II; discs expanded, about twice width of digit proximal to pad; (8) fingers bearing broad lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tu- bercles; (11) inner metatarsal tubercle oval to elongate, 2— 6x round outer metatarsal tubercle; (12) toes bearing broad lateral fringes; webbing basal; Toe V slightly longer than III; discs nearly equal in size to those on outer fingers; (13) dorsum pale grayish tan to brown with darker brown markings—interorbital and labial bars, canthal and supratympanic stripes, W-shaped mark in scapular region, two dorsal chevrons (anterior one enclosing black spot with middorsal tubercle), triangular mark enclosing vent, and diagonal bars on limbs; venter cream with brown flecks, most dense on chest and throat; posterior surfaces of thighs brown with cream flecks or spots; (14) SVL in subadult female 30.4. Except for three species, all members of the Eleutherodactylus conspicillatus Group have smooth venters. By having an areolate venter, Eleutherodactylus karcharias resembles E. caprifer of the Chocoan Region of southern Colombia and Ecuador and E. avicuporwm of the Cordil- lera Colan. The latter differs from E. karcharias by lacking a fin-shaped middorsal tubercle and distinct dorsal mark- ings, and the former differs by having a broad brown dor- solateral stripe and many dark chevrons on the dorsum of the body and brown stripes on the throat. Description.—Nothing can be added to the description by Flores and Rodriguez (1997). Distribution and habitat.—Eleutherodactylus karcharias is known froma single locality at an elevation of 1000 m in the valley of the Rio Utcubamba (tributary of the Rio Maranon) in the northern part of the Cordillera Central in northern Peru (Fig. 8). The specimens were “taken from moist leaf litter in cloud forest” (Flores and Rodriguez, 1997). Remarks.—This species is known from only seven immature females. Eleutherodactylus lanthanites Lynch Eleutherodactylus lanthanites Lynch, 1975b:10. Holotype, KU 146144, an adult female, from Santa Cecilia, 340 m, Provincia Napo, Ecuador. Diagnosis.—A member of the Eleutherodactylus conspicillatus Group having (1) skin on dorsum finely tu- berculate with scattered larger tubercles, that on venter smooth; discoidal fold prominent; dorsolateral folds ab- sent; (2) tympanic membrane smooth, and tympanic an- nulus prominent, nearly round, its length about % length of eye; (3) snout long, subacuminate in dorsal view, rounded in profile; (4) upper eyelid lacking tubercles, nar- rower than IOD; cranial crests absent; (5) vomerine odontophores prominent, oblique; (6) males possessing vocal slits and nonspinous nuptial pads; (7) Finger I longer than II; discs expanded, about twice width of digit proxi- mal to pad; (8) fingers lacking lateral fringes; (9) ulnar tu- bercles absent; (10) heel bearing conical tubercle; outer edge of tarsus bearing row of indistinct tubercles; inner tarsal tubercles and fold absent; (11) inner metatarsal tubercle elongate, about 5x conical outer metatarsal tubercle; su- pernumerary plantar tubercles few in number; (12) toes lacking lateral fringes; webbing absent; Toe V slightly longer than III; discs equal in size to those on outer fin- gers; (13) dorsum brown with darker brown markings (usually chevrons); venter cream with dark flecks; throat heavily pigmented with brown or black, defining median white stripe; posterior surfaces of thighs brown with cream flecks; (14) SVL in males 21.7—27.9 mm, in females 27.5— 45.4 mm (Lynch and Duellman, 1980). Eleutherodactylus lanthanites differs from all other mem- bers of the Eleutherodactylus conspicillatus Group, except E. gutturalis, by having a dark throat with a median white stripe. The latter species, which inhabits the Guianan Re- gion, differs from E. lanthanites by lacking a tubercle on the heel and by having proportionately longer hind limbs (Hoogmoed et al., 1977). 30 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Distribution and habitat.—Eleutherodactylus lanthanites is widespread in the upper Amazon Basin in southern Colombia, Ecuador, extreme western Brazil, and northern Peru (Lynch, 1980; Duellman and Mendelson, 1995). The species ascends the eastern slopes of the Andes to elevations of 1440 m in southern Colombia and 1490 m in Ecuador (Lynch and Duellman, 1980). In northern Peru, the species has been taken only at Venceremos at an eleva- tion of 1630 m, Departamento San Martin (Fig. 13). The single individual was on the ground by day in humid montane forest. Remarks.—A juvenile male has a SVL of 19.0 mm. In life, the dorsum was tan with dull red markings. The ante- rior, ventral, and posterior surfaces of the thighs were or- ange, and the face mask was dark brown. The belly was yellow with black flecks, and the throat was black with cream spots and a median white stripe. The iris was bronze with a median, horizontal, red streak. This juvenile is as- signed to Eleutherodactylus lanthanites on the basis of the throat pattern, which is unique to this species in the Eleutherodactylus conspicillatus Group. Eleutherodactylus lymani Barbour and Noble Eleutherodactylus lymani Barbour and Noble, 1920:403. Holotype: MCZ 5422, young female, from Perico, Departamento Cajamarca, Peru. Eleutherodactylus carrioni Parker, 1932:23. Holotype: BM 1947.2.15.99, adult female, from Loja, Provincia Loja, Ecuador. Synonymy fide Lynch, 1969:263. Eleutherodactylus lymani—Lynch, 1969:263. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) conspicillatus Group having (1) skin on dorsum shagreen with scattered tubercles, that on venter smooth; discoidal fold prominent; dorsolateral folds present; (2) tympanic membrane and tympanic annulus prominent, round, its length about % length of eye; (3) snout subacuminate in dorsal view, rounded in profile; canthus rostralis angular; (4) upper eyelid lacking tu- bercles, usually narrower than IOD; cranial crests absent; (5) vomerine odontophores subtriangular in outline, prominent; (6) males lacking vocal slits; nuptial pads present; (7) Finger I longer than II; discs small; (8) fingers bearing lateral keels; (9) ulnar tubercles absent; (10) heel and outer edge of tarsus lacking tubercles; inner surface of tarsus bearing fold; (11) inner metatarsal tubercle elon- gate, 4x round outer metatarsal tubercle; plantar supernu- merary tubercles absent; (12) toes bearing lateral fringes; webbing absent; Toe V slightly longer than III; discs as large as those on fingers; (13) dorsum brown with dark brown to black markings; venter cream to white; groin and pos- terior surfaces of thighs black with white spots or reticula- tions; (14) SVL in males 25.7-45.3 mm, in females 48.4— 69.3 mm. In size and general appearance, Eleutherodactylus lymant closely resembles two other species in the E. conspicillatus Group in northern Peru and southern Ecuador—E. condor and E. citriogaster. The three species are distinguishable on the basis of color pattern on the posterior surfaces of the thighs—bold black and white spots or reticulations in E. lymant, brown with pale flecks in E. condor, and dark brown and tan mottling in E. citriogaster (Figs. 11C, 12C); further- more, E. citriogaster has a round, instead of bifid, palmar tubercle. Other members of the E. conspicillatus Group are smaller; of those in the upper Amazon Basin and lower slopes of the Andes, E. conspicillatus, malkini, metabates, and peruvianus have the posterior surfaces of the thighs dark brown with small cream to red spots, and two of these (E. malkini and E. metabates) have basal webbing between the toes, whereas E. fenestratus, lanthanites, and vilarsi have uniformly dark brown posterior surfaces of the thighs. Furthermore, E. lanthanites has median pale stripe on a dark throat and a prominent tubercle on the heel, and E. vilarsi has proportionately shorter hind limbs and larger tym- pana. Other small members of the E. conspicillatus Group in the region differ in structure and coloration. Eleutherodactylus cuneirostris has a wedge-shaped snout and many small dark spots on the dorsum, and E. buccinator has pink spots in the groin and on the hidden surfaces of the thighs (Rodriguez, 1994); E. karcharias has a prominent fin-shaped, middorsal tubercle, and E. avicuporum and E. skydmainos have an interocular fold. Two members of the E. conspicillatus Group in the region (E. avicuporum and E. karcharias) have areolate skin on the belly. Description.—Lynch (1969) provided a thorough rede- scription of the species, which was augmented by Lynch and Duellman (1997). Distribution and habitat—Eleutherodactylus lymant has a reasonably broad distribution on the drier Pacific slopes and lowlands of southern Ecuador and northwestern Peru (Fig. 13). Most records are at elevations of less than 1600 m, but it has been found as high as 3000 m in subparamo in Provincia Loja, Ecuador. Three specimens (KU 196464, 196509; LSUMZ 19553) are from 28 km N of Santa Cruz, 725 m, Provincia Jaén, Departamento Cajamarca, Peru; this village is on the road between Jaén and Bellavista in the valley of the Rio Maranon. The frogs were collected by Richard Thomas on 8 July 1968; they were under rocks at the edge of a stream in thorn forest with trees 12-18 m high. LSUMZ 32460 is from 20 km SW of Chiriaco, Provinica Bagua, Departamento Amazonas, Peru; this lo- cality also is in the valley of the Rio Maranon. Lynch (1969:266) listed MCZ 5436 as being from Palambla in Departamento Cajamarca. We have been un- able to find a Palambla in Departamento Cajamarca, but a village by that name exists at 5°23' S, 79°37' W in Departamento Piura. Although no place by that name was cited in Noble’s (1921) account of his travels in northern ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 31 Peru, the village is in the immediate vicinity of the route described by Noble; furthermore, Barbour and Noble (1920:395) stated: “The towns of Huancabamba and Palambla are on the western range of the Andes, on the border of Piura.” Duellman and Wild (1993) recorded Eleutherodactylus lymani from Canchaque, 1120 m; Palambla is about 2 km south of Canchaque and at the same eleva- tion. Remarks.—Of the specimens reported herein, one (KU 196464) is a subadult male having a SVL of 34.6 mm and the other (LSU 19553) is a female having a SVL of 47.0 mm. Eleutherodactylus metabates new species Holotype.—KU 186504, an adult male, from 20 km [by road] SW of Chiriaco (05°13' S, 78°17' W, 525 m), Provincia Bagua, Departamento Amazonas, obtained by Richard Thomas on 16 December 1974. Paratypes.—LSUMZ 32460, an adult male from the type locality. Diagnosis.—A member of the Eleutherodactylus conspicillatus Group having (1) skin on dorsum smooth weakly tuberculate, that on venter smooth; discoidal fold prominent; dorsolateral folds absent; (2) tympanic mem- brane smooth, and tympanic annulus prominent, slightly higher than long, its length slightly less than 2 length of eye; (3) snout moderately long, rounded in dorsal view and in profile; (4) upper eyelid with numerous, small, low tubercles, as wide as IOD; cranial crests absent; (5) vomer- ine odontophores prominent, oval; (6) males possessing vocal slits, lacking nuptial pads; (7) Finger I slightly longer than II; discs on outer fingers expanded, nearly truncate, more than twice width of digit proximal to pad; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tarsus lacking tubercles; inner edge of tarsus bearing distinct fold distally; (11) inner meta- tarsal tubercle elevated, elliptical, about 10x subconical outer metatarsal tubercle; supernumerary plantar tubercles absent; (12) toes bearing broad lateral fringes; webbing basal; Toe V slightly longer than III; discs slightly smaller than those on outer fingers; (13) dorsum brown with faintly darker X- or chevron-shaped marks; venter cream with diffuse brown flecks on throat; posterior surfaces of thighs brown with small cream flecks; (14) SVL in two males 29.9— 32.2 (x = 31.1) mm; females unknown. Eleutherodactylus metabates is most similar to E. conspicillatus and E. peruvianus, both of which have small, pale flecks on the posterior surfaces of the thighs, but both species lack webbing and usually have dark face masks; moreover, E. peruvianus has dark mottling on the throat and chest. The structurally similar E. malkini has more webbing between the toes and black and cream mottling on the posterior surfaces of the thighs. Of other small mem- bers of the E. conspicillatus Group, E. fenestratus, lanthanites, and vilarsi have uniformly dark brown posterior surfaces of the thighs. Furthermore, E. lanthanites has median pale stripe on a dark throat and a prominent tubercle on the heel, and E. vilarsi has proportionately shorter hind limbs and larger tympana. Other small members of the E. conspicillatus Group in the region differ in structure and coloration. Eleutherodactylus buccinator has pink spots in the groin and on the hidden surfaces of the thighs (Rodriguez, 1994); E. karcharias has a prominent middor- sal tubercle, and E. avicuporum and E. skydmainos have an interocular fold. Eleutherodactylus avicuporum and E. karcharias have areolate skin on the belly, and E. cuneirostris has a wedge-shaped snout and many round, dark brown spots on the dorsum. In the larger species in the E. conspicillatus group in the region, the venter in E. condor is creamy white with dense gray pigmentation on the throat, belly, and ventral surfaces of the thighs; the posterior sur- faces of the thighs are brown with pale spots. In E. lymant, the venter is essentially unpigmented, except for dark bars on the margin of the lower lips and posterolaterally on the throat; the posterior surfaces of the thighs are black with bold cream mottling. In E. citriogaster, the palmar tubercle is round (instead of bifid), and the venter is yellow with brown or gray mottling on the throat, laterally on the belly, and on the ventral surfaces of the thighs; the posterior sur- faces of the thighs are mottled dark brown and creamy tan. Description.—(n = 2 males). Head wider than body; HW 37.1-37.3 (X = 37.1)% SVL; HL 41.8-42.2 (x = 42.0)% SVL; snout moderately long, rounded in dorsal view and in profile (Fig. 5C); E-N 86.0-92.7 (x = 89.4)% length of eye; nostrils barely protuberant, directed dorsolaterally; canthus rostralis angular, straight; loreal region nearly flat; lips rounded; upper eyelid bearing small, rounded tu- bercles; upper eyelid width 100.0-103.4 (x = 101.7)% IOD; supratympanic fold moderately heavy; abruptly curving downward behind tympanum, obscuring dorsal and posterodorsal edge of tympanum; side of head nearly ver- tical; tympanic membrane prominent; tympanic annulus slightly higher than long; length of tympanic annulus 41.3— 41.5 (x = 41.4)% length of eye; two prominent postrictral tubercles posteroventral to tympanum. Choanae small, round, partly concealed by palatal shelf of maxillary arch; vomerine odontophores prominent, oval, moderately sepa- rated medially, located well behind posterior edges of choa- nae, each odontophore bearing 5-6 (x = 5.5) teeth; tongue nearly twice as long as wide, barely notched posteriorly; posterior one third not adherent to floor of mouth. Skin on dorsum weakly tuberculate; dorsolateral fold absent; skin on flanks tuberculate; skin on venter smooth; discoidal fold prominent; cloacal sheath short; large tu- bercles in cloacal region absent. Ulnar tubercles absent; 32 ScreNTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS thenar tubercle elevated, elliptical, slightly smaller than bifid palmar tubercle; palmar supernumerary tubercles absent; subarticular tubercles prominent, subconical; fin- gers bearing lateral fringes; first finger slightly longer than second; discs on Fingers I and II small, round; discs on outer fingers nearly truncate, more than twice width of digit proximal to pad; all fingers having ventral pads well defined by circumferential grooves. Upper surfaces of hind limbs smooth; heel and outer edge of tarsus lacking tu- bercles; inner edge of tarsus bearing distinct fold distally; inner metatarsal tubercle elevated, elliptical, about 10x subconical outer metatarsal tubercle; plantar supernumer- ary tubercles absent; subarticular tubercles small, rounded; toes bearing broad lateral fringes; toes webbed to basal subarticular tubercles; discs on toes slightly smaller than those on outer fingers; tip of Toe V extending to middle of penultimate subarticular tubercle on Toe IV; tip of Toe III extending to base of penultimate subarticular tubercle on Toe IV (Fig. 6C); when hind limbs flexed perpendicular to axis of body, heels overlapping by about % length of shank; shank 62.2-62.4 (X= 62.3)% SVL. Coloration in preservative: Dorsum brown with faint darker brown markings consisting of interorbital bar, la- bial bars, supratympanic stripe (but no canthal stripe), X- or chevron-shaped marks on the back, and barely discern- ible transverse bars on limbs (Fig. 7C); flanks tan with dif- fuse brown diagonal bars; axilla cream with small brown spots; posterior surfaces of thighs brown with cream flecks; venter cream with diffuse brown flecks on throat. Coloration in life: Unknown. Measurements of holotype: SVL 32.2, tibia length 20.1, foot length 17.0, head width 12.0, head length 13.6, IOD 3.2, upper eyelid width 3.2, E—-N 3.5, eye, 4.6, tympanum 1.9. Distribution and habitat—Eleutherodactylus metabates is known only from the type locality at an elevation of 525 m in the Rio Maranon Valley in Departamento Amazonas, Peru (Fig. 8). This region supports thorn forest. Etymology.—The specific name is a Greek noun mean- ing leaper. The name is applied to this long-legged species that presumably is capable of lengthy jumps. Remarks.—The paratype (adult male having a SVL of 29.9 mm) differs from the holotype only by having a more pallid coloration. Eleutherodactylus peruvianus (Melin) Hylodes peruvianus Melin, 1941:43. Holotype, NHMG 490, an adult fe- male, from Roque, Departamento San Martin, Peru Eleutherodactylus peruvianus—Gorham, 1966:91. Diagnosis.—A member of the Eleutherodactylus conspicillatus Group having (1) skin on dorsum shagreen with scattered tubercles, that on venter smooth; discoidal fold prominent; dorsolateral folds present; (2) tympanic membrane smooth, and tympanic annulus prominent, nearly round, its length about % length of eye; (3) snout rounded to subacuminate in dorsal view, rounded in pro- file; (4) upper eyelid lacking tubercles, broader than IOD; cranial crests absent; (5) vomerine odontophores promi- nent, triangular; (6) males possessing vocal slits and white, nonspinous nuptial pads; (7) Finger I longer than II; discs expanded, about twice width of digit proximal to pad; (8) fingers bearing lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles, except for small tu- bercle on inner edge of tarsus; (11) inner metatarsal tu- bercle oval, about 6x subconical outer metatarsal tubercle; supernumerary plantar tubercles few in number; (12) toes bearing lateral fringes; webbing absent; Toe V slightly longer than III; discs equal in size to those on outer fin- gers; (13) dorsum usually brown with darker brown chev- rons; venter cream with brown flecks, most dense on throat; posterior surfaces of thighs brown with cream spots; (14) SVL in males 29.2-35.8 mm, in females 38.6-46.4 mm (Lynch and Duellman, 1980). Eleutherodactylus peruvianus is most similar to E. conspicillatus, malkini, and metabates, all of which have pale spots on the posterior surfaces of the thighs. The structur- ally similar E. conspicillatus lacks dark pigmentation on the throat and chest and lacks labial bars, whereas E. malkini and E. metabates have distinct webbing between the toes (absent in E. peruvianus), and E. malkini has black and cream mottling on the posterior surfaces of the thighs. Of other small members of the E. conspicillatus Group, E. fenestratus, lanthanites, and vilarsi have uniformly dark brown poste- rior surfaces of the thighs. Furthermore, E. lanthanites has median pale stripe on a dark throat and a prominent tu- bercle on the heel, and E. vilarsi has proportionately shorter hind limbs and larger tympana. Other small members of the E. conspicillatus Group in the region differ in structure and coloration. Eleutherodactylus buccinator has pink spots in the groin and on the hidden surfaces of the thighs (Rodriguez, 1994), and E. cuneirostris has many small, dark spots on the dorsum; E. karcharias has a prominent mid- dorsal tubercle, and E. avicuporum and E. skydmainos have an interocular fold (Flores and Rodriguez, 1997). In the larger species in the E. conspicillatus Group in the region, the venter in E. condor is creamy white with dense gray pigmentation on the throat, belly, and ventral surfaces of the thighs; the posterior surfaces of the thighs are brown with pale spots. In E. lymani, the venter is unpigmented, except for dark bars on the margin of the lower lips and posterolaterally on the throat; the posterior surfaces of the thighs are black with bold cream mottling. In E. citriogaster, the palmar tubercle is round (instead of bifid), and the venter is yellow with brown or gray mottling on the throat, laterally on the belly, and on the ventral surfaces of the thighs; the posterior surfaces of the thighs are mottled dark brown and creamy tan. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 33) Distribution and _ habitat.—Eleutherodactylus peruvianus is widespread in the upper Amazon Basin in western Brazil, Ecuador, and Peru (Lynch, 1980; Lynch and Duellman, 1980). Herein, we record it at elevations of 950 and 1080 m on the eastern slopes of the Cordillera Central in northern Peru—respectively, KU 217313 from 15.4 km SW of Zapatero and KU 212291 from Abra Tangarana, 7 km [by road] NE San Juan de Pacaysapa, both in Provincia Lamas, Departamento San Martin (Fig. 8). The latter was on the ground by a stream during the day. The species as- cends the eastern slopes of the Andes to elevations of nearly 2000 m in Ecuador and Peru (Lynch and Duellman, 1980). It has been reported from elevations of 1700-1975 m in the Cordillera de Cutucti (Duellman and Lynch, 1988), from 1500-1830 m on the western slopes of the Cordillera del Condor (Almendariz, 1997; Lynch and Duellman, 1980), and from 665-1750 m on the eastern slopes of the Cordil- lera del Condor in Peru (Reynolds and Icochea, 1997). Two of the latter were on vegetation at night, and one was ina bromeliad; all were in humid montane forest. Remarks.—Two specimens (USNM 525438-39) from the upper Rio Comainas, 1750 m, base of Cerro Machinaza, Departamento Amazonas, are adult females having SVLs of 37.8 and 37.2 mm, respectively. A third individual (USNM 525440) from the Puesto Vigilancia, Rio Comainas, 665 m, Departamento Amazonas, is a male having a SVL of 28.7 mm. In USNM 525438, the dorsum is uniform olive tan, and the flanks are brown, whereas in the other speci- mens, the dorsum is tan with brown chevrons. In the fe- males, the throat and chest have dark gray mottling, whereas only small gray flecks are present on the throat and chest of the male. The coloration of living individuals was: USNM 525438—Dorsum reddish brown; flanks and legs brown; dark canthal mask; chin mottled brown-yel- low; chest and belly yellow with brown spotting; under- side of legs yellow with brown spotting; iris bronze (R. P. Reynolds field notes, 17 July 1994). USNM 5254440—Dor- sum tan with light brown chevrons; dark canthal stripe; legs barred; rear of thighs brown with yellow spots; chin green; belly cream (Robert P. Reynolds field notes, 31 July 1994). The specimen from Abra Tangarana is a juvenile with a SVL of 23.5 mm. In life, the dorsum was dull olive-tan with brown chevrons, and the posterior surfaces of the thighs were brown with pale red flecks; the ventral sur- faces of the limbs were pale yellow (WED field notes, 5 February 1989). ELEUTHERODACTYLUS NIGROVITTATUS GROUP This group was defined by Lynch (1989) as having the skin on the venter smooth, dorsolateral folds present or absent, Finger I longer than Finger II, discs on digits not expanded, and vomerine odontophores present. Lynch and Duellman (1997) added relative length of the toes—Toe III longer than Toe IV. The inclusion of the new species de- scribed below necessitates a slight modification of the defi- nition—Finger I longer than, or equal in length to, Finger II, and Toe IV shorter than, or equal in length to Toe III. In addition to the species described herein, the group includes four species—E. latens and E. manipus in the Andes of Colombia, E. elassodiscus in the Cordillera Oriental in Ecuador, and E. nigrovittatus in the upper Amazon Basin and lower slopes of the Andes in Colombia, Ecuador, and Peru. (See map in Lynch et al., 1997.) Eleutherodactylus araiodactylus new species Holotype.—UF 40764, subadult female, from 24 km [by road] SW Leimebamba (06°40' S, 77°50' W, 3370 m), Provincia Chachapoyas, Departamento Amazonas, Peru, obtained on 25 April 1972 by Fred G. Thompson. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) nigrovittatus Group having (1) skin on dorsum smooth; that on venter smooth; discoidal fold evi- dent; dorsolateral folds present; (2) tympanic membrane smooth, and tympanic annulus prominent, nearly round, its length about % length of eye; (3) snout moderately long, rounded in dorsal view and in profile; (4) upper eyelid lacking tubercles, narrower than IOD; cranial crests absent; (5) vomerine odontophores prominent, elongately oval; (6) condition of vocal slits and nuptial pads unknown, (7) Finger I equal in length to II; discs on outer fingers nar- row, acutely rounded; (8) fingers bearing lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tu- bercles; (11) inner metatarsal tubercle elliptical, about 2x subconical outer metatarsal tubercle; supernumerary plan- tar tubercles few; (12) toes bearing lateral fringes; webbing absent; Toes V and III equal in length; discs equal to those on outer fingers; (13) dorsum tan with middorsal brown marking; venter and posterior surfaces of thighs tan; (14) SVL in one female 24.5 mm. Eleutherodactylus araiodactylus is readily distinguished from all other members of the genus in the Andes of north- ern Peru by having a combination of smooth skin on the venter and undilated discs on the digits. The only other species in the region having smooth skin on the venter are members of the Elewtherodactylus conspicillatus Group, all of which have greatly expanded digital discs. From other members of the Eleutherodactylus nigrovittatus Group, E. araiodactylus differs by having Fingers I and II of equal length (instead of Finger I > II), Toes II] and IV of equal length (instead of Toe III > V), digits acutely rounded in- stead of nearly pointed, and a diagonal dermal ridge posteroventral to the tympanum. Moreover, E. araiodactylus differs from E. elassodiscus by having dorsolateral folds and lateral fringes on the digits; the latter character also differ- entiates E. araiodactylus from E. nigrovittatus, a species with proportionately shorter fingers and more robust body. The 34 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Fig. 14. Dorsal and lateral views of the head of Eleutherodactylus araiodactylus, UF 40764. Scale bar = 5 mm. presence of lateral fringes on the digits also distinguishes E. araiodactylus from E. manipus. Description.— (n = 1 female). Head as wide as body. HW 40.0% SVL; HL 44.0% SVL; Snout moderately long, rounded in dorsal view and in profile (Fig. 14); E-N 89.7% length of eye; nostrils slightly protuberant, directed dorsolaterally; canthus rostralis rounded, slightly curved; loreal region noticeably concave; lips rounded; upper eye- lid lacking tubercles; upper eyelid width 71.0% IOD; supratympanic fold moderate; abruptly curving down- ward behind tympanum, barely obscuring upper edge of tympanum; side of head nearly vertical; tympanic mem- brane evident; tympanic annulus distinct; length of tym- panic annulus 48.3% length of eye; prominent diagonal dermal ridge (possibly coalesced postrictal tubercles) posteroventral to tympanum. Choanae small, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores prominent, oval, widely separated medially, located behind posterior edges of choanae, each odontophore bearing three teeth. Tongue about 1.5x as long as wide, not notched posteriorly; posterior half not adherent to floor of mouth. Skin on dorsum smooth; dorsolateral fold low, extend- ing from level of axilla to groin; skin on flanks pustular, venter smooth, except coarsely areolate on proximal posteroventral surfaces of thighs; discoidal fold prominent; cloacal sheath short; large tubercles in cloacal region ab- sent. Ulnar tubercles absent. Thenar tubercle elevated, el- liptical, about twice size of inner palmar tubercle; palmar tubercle elevated, completely divided, outer tubercle about 2 size of inner; palmar supernumerary tubercles absent; subarticular tubercles large, round; fingers bearing lateral fringes; first finger equal in length to second. Discs on Fin- gers I and II small, round; discs on all fingers narrow, round, barely wider than digit proximal to pad (Fig. 15); Fingers I-IV having ventral pads weakly defined by cir- cumferential grooves. Upper surfaces of hind limbs smooth; heel tarsus lacking tubercles; inner metatarsal tu- Fig. 15. Hand and foot of Eleutherodactylus araiodactylus, UF 40764. Scale = 5 mm. bercle elevated, elliptical, about 2x subconical outer meta- tarsal tubercle; few plantar supernumerary tubercles on proximal segments of Toes IV and V; subarticular tubercles small, subconical; toes bearing lateral fringes; toes not webbed; discs on toes about equal in size to those on fin- gers; tips of Toes III and V extending to distal edge of penultimate subarticular tubercle on Toe IV (Fig. 15); when hind limbs flexed perpendicular to axis of body, heels over- lap by about 4 length of shank; shank 54.3% SVL. Coloration in preservative: Dorsum between dorsolat- eral folds tan with narrow, median cream stripe from snout to vent; on body, stripe bordered by irregular brown mark- ings 2-3x width of stripe (Fig. 16). Brown interorbital bar, and canthal and supratympanic stripes present; short, dark brown stripe from upper eyelid extending posteromedially onto occiput; labial bars absent. Flanks brown; groin un- marked; forelimbs tan with two brown spots on inner sur- face of forearm; hind limbs brown with faint, narrow, darker diagonal marks; venter and posterior surfaces of thighs uniform tan. Coloration in life: Unknown. Measurements of holotype: SVL 24.5, tibia length 13.3, foot length 13.5, head width 9.8, head length 9.8, IOD 3.1, upper eyelid width 2.2, E—-N 2.6, eye, 2.9, tympanum 1.4. Distribution and _ habitat.—Eleutherodactylus araiodactylus is known only from 3370 m in very humid ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 35) montane forest in the northern part of the Cordillera Cen- tral in Peru (Fig. 13). Etymology.—The specific name derived from the Greek araios, meaning thin, and the Greek daktylos, meaning fin- ger or toe. The name applies to the extremely slender dig- its of this species. Remarks.—This is one of the most distinctive species of Eleutherodactylus in northern Peru; consequently, we have no qualms about naming it on the basis of a single specimen. The presence of a middorsal pale stripe is a color polymorphism in many species of Eleutherodactylus; its presence in the holotype probably is not indicative of the species. ELEUTHERODACTYLUS ORESTES GROUP Inclusion in the Eleutherodactylus orestes Group of the four species described herein necessitates a slight modifi- cation of the definition of the group as given by Lynch and Duellman (1997). These are that vocal slits and vomer- ine odontophores are present (E. orestes, simonbolivari, and vidua), absent (E. melanogaster and E. pataikos), vomerine odontophores absent and vocal slits present (E. atrabrachus), or vomerine odontophores present and vocal slits un- known (E. pinguis). Furthermore, the distribution of the group needs to be changed. Members of the group inhabit high-altitude cloud forests in the Cordillera Occidental in western Ecuador (E. simonbolivari), paramo in the south- ern part of the Cordillera Oriental in Ecuador (E. orestes and E. vidua), and supra-treeline habitats in the Cordillera Colan (E. atrabracus) and the northern parts of the Cordil- lera Central (E. melanogaster and E. pataikos) and Cordil- lera Occidental (E. pinguis) in Peru. Eleutherodactylus atrabracus new species Holotype.—LSUMZ 49144, adult female from the Cor- dillera Colan, east of La Peca (05°36' S, 78°22' W, 2963 m), Provincia Bagua, Departamento Amazonas, Peru, obtained on 4 September 1978 by Morris D. Williams. Paratype-—LSUMZ 45090, adult male from the Cor- dillera Colan, east of La Peca, 3330 m, obtained on 31 Au- gust 1978 by Thomas S. Schulenberg. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) orestes Group having (1) skin on dorsal surfaces finely shagreen, that on throat, belly, and ventral surfaces of thighs coarsely areolate; discoidal fold weak posteriorly; dorsolateral folds absent; (2) tympanic mem- brane differentiated, tympanic annulus round; its diam- eter slightly about /% length of eye; (3) snout moderately long, bluntly rounded in dorsal view and in profile, in- clined anteroventrally from level of nostrils; (4) upper eye- lid narrower than IOD, lacking tubercles; cranial crests absent; (5) vomerine odontophores absent; (6) vocal slits present; nuptial excrescences absent; (7)Finger I shorter than II; discs on outer fingers rounded, barely wider than digit proximal to pad; (8) fingers bearing lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tar- sus lacking tubercles; low, thick fold on distal part on in- ner surface of tarsus; (11) inner metatarsal tubercle el- evated, ovoid, about 5x round outer metatarsal tubercle; plantar surfaces coarsely areolate; (12) toes bearing lateral fringes; webbing absent; Toe V slightly longer than III; discs as large as those on outer fingers; (13) dorsum nearly uni- form brown; throat and belly creamy tan with fine brown reticulations; ventral surfaces of hind limbs black; groin and anterior surfaces of thighs dark brown with large tan spots; (14) SVL in one adult male 18.9, in one adult female 22.7 mm. Eleutherodactylus atrabrachus differs from all other members of the E. orestes Group by having a pale belly and the ventral surfaces of the hind limbs black; further- more, it differs from E. pataikos by having large pale spots in the groin and on the anterior surfaces of the thighs (Fig. 17). Eleutherodactylus atrabracus differs from E. melanogaster and E. simonbolivari by having a pale, instead of black, belly, and from E. pinguis by having the ventral surfaces of the hind limbs black and by lacking vomerine teeth. Two other species in the group, E. orestes and E. vidua, both of which have vomerine odontophores, also have pale spots in the groin; in the latter, the spots are large and lie between di- agonal brown bars on the posterior half of the flanks, whereas in E. vidua, the groin is black with small white spots. Description.—(n = 1 male, 1 female). Head narrower than body; HW 34.9-36.1 (x = 35.5)% SVL; HL 34.9-40.5 (x = 37.7)% SVL; snout moderately long, bluntly rounded in dorsal view; in profile, inclined anteroventrally from level of nostrils to margin of lip; E-N 90.0-100.0 (x = 95.0)% length of eye; nostrils slightly protuberant, directed later- ally; canthus rostralis weakly angular, slightly curved; lo- real region noticeably concave; lips rounded; upper eye- lid lacking tubercles; upper eyelid width 71.4-84.0 (x = 77.7)% IOD; cranial crests absent; supratympanic fold weak, barely obscuring upper edge of tympanum; side of head nearly vertical; tympanic membrane differentiated; tympanic annulus round; length of tympanic annulus 39.1— 50.0 (x = 44.6)% length of eye; postrictal tubercles low, dif- fuse. Choanae small, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores absent; tongue much longer than wide, shallowly notched posteriorly, pos- terior half not adherent to floor of mouth. Skin on dorsum finely shagreen, that on venter ar- eolate; discoidal fold barely evident posteriorly; dorsolat- eral folds absent; cloacal sheath short; enlarged tubercles in cloacal region absent. Ulnar tubercles absent; thenar tubercle ovoid, elevated, about half the size of bifid pal- mar tubercle; supernumerary palmar tubercles round; ScrenTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Fig. 17. Color P anterior surfaces of the thighs in species in the Eleutherodactylus orestes ee A. E. atrabracus, LSUMZ 49144. B. E. melanogaster, KU 218513. . E. pataikos, KU 212320. D. E. pinguis, KU 181283. E. E. simonbolivari, KU 218253. FE. vidua, KU 120083. G. E. orestes, KU 141999. Not drawn to scale. attern and skin texture in the groin and on the subarticular tubercles prominent, round; fingers bearing thick lateral fringes; Finger I shorter than II; discs on all fingers small, only slightly wider than width of digits; all fingers having ventral pads weakly defined by circumfer- ential grooves. Upper surfaces of hind limbs coarsely shagreen; heel and outer edge of tarsus lacking tubercles; inner edge of tarsus bearing low, thick fold distally; inner metatarsal tubercle elevated, ovoid, about 5x round outer metatarsal tubercle; plantar surfaces areolate; subarticular tubercles, round; toes bearing thick lateral fringes; web- bing absent; discs on toes about equal in size to those on fingers; tip of Toe V extending to proximal edge of penultimate subarticular tubercle on Toe IV; tip of Toe III not extending to that tubercle; when hind limbs flexed perpendicular to axis of body, heels barely overlapping; shank 38.6-40.5 (X = 39.6)% SVL. Coloration in preservative: Dorsum of head, body, and limbs nearly uniform brown; faint dark brown canthal and Fig. 18. Ventral color pattern Eleutherodactylus atrabracus, LSUMZ 49144. SVL = 22.7 mm. supratympanic stripes present in both specimens; in fe- male holotype, diffuse brown dorsolateral line and dis- tinct middorsal tan stripe from tip of snout to vent; inter- orbital bar, labial bars and transverse bars on limbs ab- sent. Groin and anterior surfaces of thighs dark brown with large tan spots (Fig. 17A); throat and belly creamy tan with fine brown reticulations and narrow transverse line in tho- racic region; ventral surfaces of hind limbs black (Fig. 18). Coloration in life: Unknown. Measurements of holotype: SVL 22.7, tibia length 9.2, foot length 9.5, head width 8.2, head length 9.2, IOD 22.5, upper eyelid width 2.1, E-N 2.3, eye 2.3, tympanum 0.9. Distribution and habitat.—Eleutherodactylus atrabracus is known only from two localities at elevations of 2963 and 3330 m on the western slopes of the Cordillera Colan in northern Peru (Fig. 19). The paratype was in a grassy bog above treeline. Etymology.—The specific name is derived from the Latin adjective, atra, meaning black and the Latin noun, braca, meaning trousers, and refers to the black ventral surfaces of the hind limbs in this species. Eleutherodactylus melanogaster new species Holotype.—KU 212321, adult female, from the north slope of Abra Barro Negro (06°41' S, 77°51' W, 3470 m), 28 km [by road] SSW Leimebamba, Provincia Chachapoyas, Departamento Amazonas, Peru, one of a series collected on 23 January 1989 by William E. Duellman and John J. Wiens. Paratypes.—KU 212322-23 and 218513, adult males, collected with the holotype; KU 181281, an adult female, ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 3 < 1000 m 1000-2000 m 2000-3000 m SS 3000-4000 m > 4000 m @ £. atrabracus || M@ E. melanogaster .| OE. orestes \| AE. pataikos “| O E. pinguis AE. vidua Fig. 19. Localities of known occurrence of six species in the Eleutherodactylus orestes group in the Andes of southern Ecuador and northern Peru. Another member of the group, E. simonbolivari, occurs farther north in Cordillera Occidental in Ecuador. from the north slope of Abra Barro Negro, 3300 m, 25.5 km [by road] SSW Leimebamba, Provincia Chachapoyas, Departamento Amazonas, Peru, collected on 6 March 1979 by William E. Duellman. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) orestes Group having (1) skin on most surfaces coarsely areolate, nearly pustular dorsolaterally; snout, anterior surfaces of thighs and inner surfaces of shanks smooth; discoidal fold weak posteriorly, prominent as transverse thoracic fold anteriorly; dorsolateral folds absent; (2) tympanum indistinct, tympanic annulus round; its diameter about 30% length of eye in males, 60% in fe- males; (3) snout short, rounded in dorsal view and in pro- file; (4) upper eyelid narrower than IOD, lacking tubercles; cranial crests absent; (5) vomerine odontophores absent; (6) vocal slits and nuptial excrescences absent; (7) Finger I shorter than II; discs on outer fingers rounded, barely wider than digit proximal to pad; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tarsus lacking tubercles; inner edge of tarsus bearing low, indistinct fold distally; (11) inner metatarsal tubercle elevated, ovoid, about 3x round outer metatarsal tubercle; plantar surfaces areolate; (12) toes lacking lateral fringes; webbing absent; Toe V slightly longer than III; discs as large as those on outer fingers; (13) dorsum and venter black with or without pale spots in groin and on anterior and posterior surfaces of thighs and inner surfaces of shanks; (14) SVL in three adult males 20.6—-22.8 (x = 21.9 mm), in two adult females, 24.2—24.7(x = 24.5 mm). Eleutherodactylus melanogaster differs from all other members of the E. orestes Group by having the skin on the dorsum coarsely areolate and by being black dorsally and ventrally. With the exception of the nature of the skin on the dorsum, E. melanogaster most resembles the smaller E. simonbolivari, which also has pale spots in the groin and on the hidden surfaces of the hind limbs, but E. simonbolivari has vomerine odontophores and a small, nonconical tubercle on the heel. Eleutherodactylus orestes and E. pinguis also have pale spots in the groin and on the pos- terior surfaces of the thighs, but the skin on the dorsum is finely tuberculate, the dorsum is brown with darker brown markings, vomerine odontophores are present, and there is asmall, nonconical tubercle on the heel in E. simonbolivart. The sympatric E. pataikos is like E. melanogaster in lacking vomerine odontophores and a tubercle on the heel, but it differs by having a smooth dorsum that is pale brown and by lacking markings in the groin and on the posterior sur- faces of the thighs. Eleutherodactylus atrabracus also lacks vomerine teeth, but it has a finely shagreen dorsum, a pale belly, and black ventral surfaces of the hind limbs. Description.—(n = 3 males, 2 females; proportions are for males with followed those of by females). Head nar- rower than body; HW 36.0-38.3 (x = 36.9), 39.2-39.4 (x = 39.3)% SVL; HL 33.8-36.4 (x = 35.1), 36.4-37.2 (X = 36.8)% SVL; snout short, rounded in dorsal view, and in profile; E-N 64.0-71.4 ( = 67.8), 80.0-83.0 (x = 81.5) % length of eye; nostrils slightly protuberant, directed dorsolaterally; canthus rostralis rounded, curved; loreal region weakly concave; lips rounded; upper eyelid lacking tubercles; upper eyelid width 68.0-71.4 (x = 70.3), 75.9-76.7 (x = 76.3)% IOD; cranial crests absent; supratympanic fold a series of pustular granules, obscuring upper edge of tym- panum; side of head nearly vertical; tympanum indistinct; tympanic annulus thin; tympanic membrane thickened; tympanic annulus round; length of tympanic annulus 33.3— 44.0% (37.8), 60.0-66.7 (63.3) % length of eye; postrictal tubercles not discernible from other pustules; skin on top of head smooth. Choanae small, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores absent; tongue much longer than wide, not notched pos- teriorly, posterior half not adherent to floor of mouth. Males lacking vocal slits and nuptial excrescences. Skin on dorsum coarsely areolate, especially dorsolaterally; that on venter coarsely areolate; discoidal 38 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Fig. 20. Hand and foot of Eleutherodactylus melanogaster, KU 212321. Scale = 5 mm. fold barely evident posteriorly, present as distinct trans- verse thoracic fold anteriorly; dorsolateral folds absent; flanks coarsely areolate; cloacal sheath short; enlarged tu- bercles in cloacal region absent. Ulnar tubercles not dis- tinguishable; thenar tubercle ovoid, slightly elevated, about as large as ovoid palmar tubercle; supernumerary palmar tubercles not evident; subarticular tubercles prominent, round; fingers lacking lateral fringes; first finger shorter than second; discs on all fingers small, only slightly wider than width of digits (Fig. 20); all fingers having ventral pads weakly defined by circumferential grooves. Upper surfaces of hind limbs coarsely areolate; heel and outer edge of tarsus lacking tubercles; inner edge of tarsus with low fold distally; inner metatarsal tubercle elevated, ovoid, about 3x round outer metatarsal tubercle; plantar surfaces areolate; subarticular tubercles large, round; toes lacking lateral fringes; webbing absent; discs on toes about equal in size to those on fingers; tip of Toe V extending to point midway between penultimate and distal subarticular tu- bercles on Toe IV; tip of Toe III extending to distal edge of penultimate tubercle on Toe IV (Fig. 20); when hind limbs flexed perpendicular to axis of body, heels not overlap- ping; shank 36.4-39.3 X = 37.5), 37.1-37.7 (X = 37.4) % SVL. Coloration in preservative: Female holotype and one male (KU 212323) black above and below with small,.dif- fuse white spots in groin, on anterior and posterior sur- faces of thighs (Fig. 17B), and on inner surfaces of shanks; spots absent in other female (KU 181281) and in two males (KU 212322, 218513). Coloration in life: KU 212321: Dorsum dull brown with creamy tan canthal stripe bordered below by black line (Fig. 9); black postocular stripe; belly dark gray; axilla, groin, and ventral surfaces of limbs black with bright yel- low spots; iris pale gold with black flecks (WED field notes, 23 January 1989). Measurements of holotype: SVL 24.7, tibia length 9.3, foot length 10.5, head width 9.7, head length 9.0, IOD 3.0, upper eyelid width 2.3, E-N 2.0, eye 2.4, tympanum 1.6. Distribution and habitat.—Eleutherodactylus melanogaster is known only from two localities at eleva- tions of 3300 and 3470 m, respectively, on the road from Balsas to Leimebamba, near the crest of the northern part of the Cordillera Central in northern Peru (Fig. 19). All in- dividuals were under stones by day; those from 3470 m were in wet paramo, whereas the individual from 3300 m was in elfin forest; both habitats fall within the very hu- mid montane forest Etymology.—The specific name is derived from the Greek adjective melanos meaning black and the Green noun gaster meaning belly; the name is used in reference to the uniformly black ventral coloration. Eleutherodactylus pataikos new species Holotype.—KU 212320, adult female, from the north slope of Abra Barro Negro, (06°41'S, 77°51' W, 3470 m), 28 km [by road] SSW Leimebamba, Provincia Chachapoyas, Departamento Amazonas, Peru, obtained on 23 January 1989 by William E. Duellman. Diagnosis.—A member of the Elewtherodactylus (Eleutherodactylus) orestes Group having (1) skin on dor- sum smooth with few low tubercles dorsolaterally and on hind limbs, that on venter coarsely areolate; discoidal fold weak; dorsolateral folds absent; (2) tympanum indistinct, tympanic annulus distinguishable anteroventrally, round; its diameter less than % length of eye; (3) snout short, rounded in dorsal view and in profile; (4) upper eyelid narrower than IOD, lacking tubercles; cranial crests absent; (5) vomerine odontophores absent; (6) condition of vocal slits and nuptial excrescences unknown, (7) Finger I shorter than IJ; discs on outer fingers rounded, barely wider than digit proximal to pad; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tar- sus lacking tubercles; inner edge of tarsus bearing single, low tubercle; (11) inner metatarsal tubercle flat, ovoid about 4x round outer metatarsal tubercle; supernumerary tu- bercles low, diffuse; (12) toes lacking lateral fringes; web- bing absent; Toe V slightly longer than III; discs as large as ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 39 Fig. 21. Dorsal and lateral views of the head of Eleutherodactylus pataikos, KU 212320. Scale bar = 5 mm. those on outer fingers; (13) dorsum uniform pale brown; venter tan; posterior surfaces of thighs brown; (14) SVL in one adult female 21.6 mm. Eleutherodactylus pataikos differs from other members of the E. orestes Group by lacking distinctive markings. Eleutherodactylus atrabracus, orestes, pinguis, simonbolivari, and melanogaster have large pale spots in the groin, and the latter, which occurs sympatrically with E. pataikos, has coarsely areolate skin on the dorsum, which, like the ven- ter, is black. Eleutherodactylus pinguis and E. atrabracus also differ by having a prominent tympanum and tympanic annulus; the former also differs by having the ulnar tu- bercles coalesced into a fold, and the latter also differs by having a pale belly and black ventral surfaces of the hind limbs. Eleutherodactylus orestes and E. vidua have cream spots at least distally on the posterior surfaces of the thighs, which are uniform pale brown in E. pataikos; the latter (E. pataikos) also differs by lacking vomerine odontophores, a feature shared only with E. atrabracus and E. melanogaster in the E. orestes Group. Description.—(n = 1 female). Head narrower than body; HW 39.4% SVL HL 36.1 % SVL; snout short, rounded in dorsal view, and in profile (Fig. 21); E-N 71.4% length of eye; nostrils slightly protuberant, directed dorsolaterally; canthus rostralis rounded, nearly straight; loreal region weakly concave; lips rounded; upper eyelid lacking tu- bercles; upper eyelid width 70.8% IOD; cranial crests ab- sent; supratympanic fold diffuse, obscuring upper edge of tympanum; side of head nearly vertical; tympanic an- nulus thin, evident only anteroventrally; tympanic mem- brane not pustular or thickened; tympanic annulus round; length of tympanic annulus 33.3% length of eye; two large, round postrictal tubercles, posteroventral to tympanic an- nulus; skin on head smooth. Choanae small, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores absent; tongue much longer than wide, not notched posteriorly, posterior half not adherent to floor of mouth. Males unknown. Skin on dorsum smooth with few low tubercles dorsolaterally and on hind limbs, that on venter coarsely areolate; discoidal fold weak; dorsolateral folds absent; flanks tuberculate; cloacal sheath short and enlarged tu- bercles in cloacal region absent; skin on throat smooth, on belly coarsely granular; discoidal fold weak. Ulnar tu- bercles absent; thenar tubercle ovoid, flat, about as large as ovoid palmar tubercle; supernumerary palmar tubercles few, large, round; subarticular tubercles prominent, round; fingers lacking lateral fringes; Finger I shorter than LI; discs on all fingers small, only slightly wider than width of dig- its; all fingers having ventral pads weakly defined by cir- cumferential grooves. Upper surfaces of hind limbs smooth with scattered small tubercles; heel and outer edge of tar- sus lacking tubercles; inner edge of tarsus bearing small, indistinct tubercle; inner metatarsal tubercle flat, ovoid about 4x round outer metatarsal tubercle; supernumerary tubercles low, diffuse; subarticular tubercles large, round; toes lacking lateral fringes; webbing absent; discs on toes about equal in size to those on fingers; tip of Toe V extend- ing to point midway between penultimate and distal subarticular tubercles on Toe IV; tip of Toe III extending to distal edge of penultimate tubercle on Toe IV; when hind limbs flexed perpendicular to axis of body, heels not over- lapping; shank 37.0% SVL. Coloration in preservative: Dorsum, flanks and hid- den surfaces of limbs dull, pale brown; venter, dull tan. No markings evident (Fig. 17C). Coloration in life: Dorsum pale brown with narrow, diffuse cream stripe on canthus and outer edge of eyelid, and narrow, dark brown supratympanic stripe (Fig. 9); venter creamy gray; groin dark gray; iris dull bronze with black flecks. Measurements of holotype: SVL 21.6, tibia length 8.0, foot length 8.8, head width 8.5, head length 7.8, IOD 2.4, upper eyelid width 1.7, E-N 1.5, eye 2.1, tympanum 0.7. Distribution and habitat.—The species is known only from one locality at 3470 m on the road from Balsas to Leimebamba, near the crest of the northern part of the Cordillera Central in northern Peru (Fig. 19). The only known specimen, an adult female, was under a rock in very humid montane forest by day. Etymology.—The specific name pataikos is a Greek noun in apposition for an odd-shaped dwarflike Phoenician deity. The name refers to the small but Rubenesque stature of this frog. Remarks.—Loath though we are to describe a new species of Eleutherodactylus from a single specimen, this individual certainly is distinctive in the Eleutherodactylus orestes Group. 40, ScIeNTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Eleutherodactylus pinguis new species Holotype.—KU 181283, an adult female from 23 km [by road] SW Celendin (06°58' S, 78°06' W, 3050 m), Provincia Celendin, Departamento Cajamarca, Peru, one of a series collected by David C. Cannatella, William E. Duellman, and Thomas J. Berger on 8 March 1979. Paratypes.—KU 181282 collected with the holotype, and KU 181284 from 33 km [by road] SW Celendin, 3200 m, Provincia Celendin, Departamento Cajamarca, Peru. Referred specimen.—UF 40766 from 57 km [by road] N Cajamarca, 3760 m, Provincia Cajamarca, Departamento Cajamarca, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) orestes Group having (1) skin on dorsal surfaces finely areolate, that on throat, belly, and ventral surfaces of thighs coarsely areolate; discoidal fold weak posteriorly, prominent as transverse thoracic fold anteri- orly; dorsolateral folds absent; (2) tympanum membrane differentiated, tympanic annulus round; its diameter slightly more than % length of eye; (3) snout moderately long, acutely rounded in dorsal view and bluntly rounded in profile; (4) upper eyelid narrower than IOD, lacking tubercles; cranial crests absent; (5) vomerine odontophores prominent, oblique; (6) condition of vocal slits and nup- tial excrescences unknown; (7) Finger I shorter than II; discs on outer fingers rounded, barely wider than digit proxi- mal to pad; (8) fingers lacking lateral fringes; (9) ulnar tu- bercles coalesced into short fold; (10) heel and outer edge of tarsus lacking tubercles; low, thick fold on distal part on inner surface of tarsus; (11) inner metatarsal tubercle flat, ovoid, about 3x subconical outer metatarsal tubercle; plan- tar surfaces coarsely areolate; (12) toes bearing lateral fringes; webbing absent; Toe V slightly longer than III; discs as large as those on outer fingers; (13) dorsum brown; ven- ter tan with brown flecks or reticulations; groin and proxi- mal anterior surfaces of thighs dark brown with large cream spots; (14) SVL in three adult females 28.4—29.8 (x= 28.9 mm). Eleutherodactylus pinguis differs from other members of the E. orestes Group by having the ulnar tubercles coa- lesced into a low fold; it differs from E. melanogaster and E. simonbolivari by having a pale, as opposed to a dark, ven- ter, and from E. atrabracus, melanogaster, and pataikos by having vomerine teeth. Eleutherodactylus pinguis also dif- fers from E. pataikos by having a prominent tympanum and tympanic annulus and distinctive markings in the groin and pale spots on the posterior surfaces of thighs, which are uniform pale brown in E. pataikos. Two other species in the group, E. orestes and E. vidua, also have pale spots in the groin; in the latter, the spots are large and lie between diagonal brown bars on the posterior half of the flanks, whereas in E. vidua, the groin is black with small white spots. Description.—(1 = 3 females). Head narrower than body; HW 36.5-38.0 (x =37.1)% SVL; HL 34.2-36.1 (x = 35.1)% SVL; snout moderately long, acutely rounded in dorsal view, and bluntly rounded in profile; E-N 75.9-92.8 (x= 83.0)% length of eye; nostrils slightly protuberant, di- rected dorsolaterally; canthus rostralis weakly angular, slightly curved; loreal region noticeably concave; lips rounded; upper eyelid lacking tubercles; upper eyelid width 56.4—-65.7 (x = 59.7)% IOD; cranial crests absent; supratympanic fold weak, granular, obscuring upper edge of tympanum; side of head nearly inclined ventrolaterally; tympanic membrane weakly differentiated; tympanic an- nulus round; length of tympanic annulus 53.6-55.2 (Xx = 54.6)% length of eye; postrictal tubercles not discernible from other tubercles. Choanae small, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores prominent, oval in outline, oblique, widely separated me- dially, located behind posterior edges of choanae, each odontophore bearing 3 or 4 (x = 3.8 teeth); tongue much longer than wide, shallowly notched posteriorly, posterior half not adherent to floor of mouth. Males unknown. Skin on dorsum finely areolate, more coarsely areolate laterally; that on venter coarsely areolate; discoidal fold barely evident posteriorly, present as distinct transverse thoracic fold anteriorly; dorsolateral folds absent; cloacal sheath short; enlarged tubercles in cloacal region absent. Ulnar tubercles coalesced so as to form low ridge distally on forearm; thenar tubercle ovoid, slightly elevated, some- what smaller than bifurcate palmar tubercle; supernumer- ary palmar tubercles not evident; subarticular tubercles prominent, round; fingers bearing thick lateral fringes; Fin- ger I shorter than II; discs on all fingers small, only slightly wider than width of digits; all fingers having ventral pads weakly defined by circumferential grooves. Upper surfaces of hind limbs coarsely areolate; heel and outer edge of tar- sus lacking tubercles; inner edge of tarsus bearing low, thick fold distally; inner metatarsal tubercle flat, ovoid, about 3x subconical outer metatarsal tubercle; plantar surfaces areolate; subarticular tubercles large, round; toes bearing thick lateral fringes; webbing absent; discs on toes about equal in size to those on fingers; tip of Toe V extending to distal edge of penultimate subarticular tubercle on Toe IV; tip of Toe II not extending to that tubercle; when hind limbs flexed perpendicular to axis of body, heels not over- lapping; shank 35.4—40.4 (X= 38.1)% SVL. Coloration in preservative: Dorsum of head, body, and limbs brown with faintly darker brown markings (absent in KU 181284); markings consisting of faint interorbital mark, numerous small irregular flecks on body and faint ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 4] diagonal crossbar on shank. Dark brown markings in groin and on proximal anterior surfaces of thighs, nearly enclos- ing large cream spots (Fig. 17D); distal portion of poste- rior surfaces of thighs and inner surfaces of shanks cream with large, dark brown markings. Venter creamy tan with minute brown flecks on throat and belly in KU 181282, larger clusters of flecks, especially on belly, in KU 181283, and fine brown reticulations in KU 181284. Coloration in life: KU 181284: Dorsum dull red becom- ing dull green on upper flanks; venter dull yellow reticu- lated with grayish brown; iris dull reddish bronze (Fig. 9; WED field notes, 08 March 1989). Measurements of holotype: SVL 28.4, tibia length 10.7, foot length 11.4, head width 10.8, head length 9.7, IOD 3.5, upper eyelid width 2.2, E-N 2.2, eye 2.9, tympanum 1.6. Distribution and habitat.—Eleutherodactylus pinguis is known from three localities in the northern part of the Cordillera Occidental in Peru (Fig. 19). Two localities are at elevations of 3050 and 3200 m on the road between Cajamarca and Celendin, on the eastern slopes of Abra Comulica. All individuals were under stones by day. Those from 3050 m were in a wet grassy area, whereas the indi- vidual from 3200 m was in a bunch grass—Baccharis asso- ciation. Another specimen is from an elevation of 3760 m north of Cajamarca; it was under a rock ina grassy region by day. Etymology—the specific name is a Latin adjective meaning fat; the name refers to the corpulent habitus of this species. Remarks.—A subadult female (UF 40766) having a snout-vent length of 24.0 mm is like the type series in struc- tural features. The color pattern is faded, so there are no markings evident in the groin or on the thighs; no dark flecks are present on the venter. ELEUTHERODACTYLUS UNISTRIGATUS GROUP As noted by Lynch and Duellman (1997), members of this group are characterized by areolate skin on the venter and Toe V being much longer than Toe III. This is the larg- est group of Eleutherodactylus and now contains more than 175 species, of which 31 are known from the Andes in northern Peru. Eleutherodactylus acuminatus Shreve Eleutherodactylus acuminatus Shreve, 1935:217. Holotype, MCZ 19951 from Canelos, Provincia Pastaza, Ecuador. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum smooth, that on belly and proximal posteroventral surfaces of thighs coarsely areolate; skin on other ventral surfaces smooth; discoidal fold barely evident posteriorly; dorsolateral folds absent; (2) tympanic membrane not dif- ferentiated; tympanic annulus distinguishable under skin, 1000 m 1000-2000 m 2000-3000 m 3000-4000 m = >4000m E. acuminatus E. anemerus E. ardalonychus E. bearsei E. bromeliaceus E. cajamarcensis Fig. 22. Localities of known occurrence of six species in the Eleutherodactylus unistrigatus group in the Andes of southern Ecuador and northern Peru. round, its length less than % length of eye; (3) snout long, acuminate in dorsal view, truncate and posteriorly inclined in profile; (4) upper eyelid lacking tubercles, much nar- rower than IOD; cranial crests absent; (5) vomerine odontophores low, ovoid; (6) males lacking vocal slits and nuptial pads; (7) Finger I shorter than II; discs on outer fingers elliptical; (8) fingers lacking lateral fringes; (9) ul- nar tubercles absent; (10) heel lacking tubercles; outer edge of tarsus bearing two or three low subconical tubercles; inner edge of tarsus lacking tubercles or fold; (11) inner metatarsal tubercle elevated, elliptical, about 4x conical outer metatarsal tubercle; plantar supernumerary tubercles absent; (12) toes lacking lateral fringes; webbing absent; Toe V much longer than III; discs nearly as large as those on outer fingers; (13) dorsum creamy tan (green in life) with black canthal and supratympanic stripes and narrow interorbital bar; limb bars absent; venter and hidden sur- faces of thighs cream; (14) SVL in males 17.1—22.6 mm, in females 25.6-31.3 (Lynch, 1980). This nearly uniformly green species with an acuminate snout, broad head, and concealed tympana is readily dis- tinguished from all other members of the genus in the re- gion. The only other predominately green species is E. galdi, 42 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS which has flared lips, cranial crests, distinct tympanum, large truncate discs, conical tubercles along the tarsus, and bars on the limbs. Description.—The description by Shreve (1935) was augmented by Duellman (1978c). Distribution and habitat.—This primarily Amazonian species invades the lower reaches of the Andes in south- ern Ecuador and northern Peru. It is known from San José, 830 m, Provincia Morona-Santiago, Ecuador (KU 147976) on the western slopes of the Cordillera del Condor and from 15.4 km [by road] SW Zapateros, 950 m, Provincia Lamas, Departamento San Martin, Peru (KU 217308-10) on the eastern slopes of the Cordillera Central (Fig. 22). The latter were in arboreal bromeliads by day. Remarks.—Two of the specimens from the Cordillera Central are males with SVLs of 21.3 and 21.5 mm; the other is a female having a SVL of 24.3 mm. Although this spe- cies is abundant in the lowland rainforest in Amazonian Ecuador, it has been reported only twice from Amazonian Peru—Rio Aguayita, Departamento Loreto (Lynch, 1980), and San Jacinto, Departamento Loreto (Duellman and Mendelson, 1995). Additional records are from Departamento San Martin—Rio Cainarache, 330 m, 33 km [by road] NE Tarapoto (KU 209466), and Rio Shilcayo, 500 m, near Tarapoto (KU 209467). Eleutherodactylus anemerus new species Holotype.—KU 219798, adult male, from El Tambo (05°21' S, 79°33' W, 2770 m, 31 km [by road] ENE of Canchaque, Provincia Huancabamba, Departamento Piura, Peru, obtained on 9 January 1991 by Erik R. Wild. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum finely tuberculate, that on venter areolate; discoi- dal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus prominent, round, its length slightly more than % length of eye; (3) snout short, acuminate in dorsal view, truncate and posteriorly inclined in profile; tubercle on tip of snout; canthus rostralis rounded; (4) upper eyelid lacking tubercles, narrower than IOD; cranial crests absent; (5) vomerine odontophores absent; (6) males having vocal slits but lacking nuptial pads; (7)Finger I shorter than II; discs moderately small, nearly truncate; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles low, diffuse; (10) heel lacking tubercles; outer edge of tarsus bearing low tubercles; inner edge of tarsus lacking tubercles or fold; (11) inner metatarsal tu- bercle broadly ovoid, about 5x subconical outer metatar- sal tubercle; plantar supernumerary tubercles few, basal only; (12) toes bearing narrow lateral fringes; webbing absent; Toe V much longer than III; discs about equal in size to those on outer fingers; (13) dorsum uniform yel- lowish tan; venter creamy white with minute black flecks; (14) SVL in male 20.4 mm. No other species of Eleutherodactylus in the region has an unmarked orange-red dorsum and yellow flanks. The dorsal coloration is reminiscent of some individuals of E. chalceus, a member of the Eleutherodactylus diastema Group in Chocoan Colombia and Ecuador. (See Lynch and Duellman, 1997:plate 8.) However, E. chalceus differs from E. anemerus in a number of features—areolate skin on the dorsum, absence of a tympanum, papillate discs on the fingers, and Toe V shorter than Toe III. Five other species of Eleutherodactylus in the Andes of northern Peru lack vomerine odontophores; three of these (E. atrabracus, melanogaster, and pataikos) robust-bodied members of the E. orestes Group that have small terminal discs on the fin- gers, short hind limbs, and Toe V only slightly longer than Toe III. The other two (E. incomptus and E. percnopterus) are members of the E. unistrigatus Group and have dis- tinctive dark markings on the dorsum. Description.—(n = 1 male). Head as wide as body; HW 36.3% SVL; HL 35.3% SVL. Snout moderately short, acumi- nate in dorsal view, truncate and slightly incline posteroventrally in profile, with subconical tubercle on tip; E-N = length of eye; nostrils noticeably protuberant, di- rected dorsolaterally; canthus rostralis rounded, slightly curved; loreal region weakly concave; lips rounded; up- per eyelid width 77.0% IOD, lacking tubercle; cranial crests absent; supratympanic fold weak, obscuring upper edge of tympanic annulus; side of head inclined ventrolater- ally; tympanic membrane present; tympanic annulus round; length of tympanic annulus 55.0% length of eye; three small, subconical postrictal tubercles below and posteroventral to tympanum. Choanae small, round. not concealed by palatal shelf of maxillary arch; vomerine odontophores absent; tongue nearly twice as long as wide, distinctly notched posteriorly posterior half not adherent to floor of mouth; vocal slits small, extending posterolaterally from lateral base of tongue; vocal sac single, median, subgular. Skin on dorsum finely tuberculate; dorsolateral fold absent; skin on throat, belly, and ventral surfaces of thighs areolate; skin on other ventral surfaces smooth; discoidal fold prominent; cloacal sheath short; large tubercles in anal region absent. Ulnar tubercles low, diffuse; thenar tubercle elevated, elliptical, slightly larger than bifurcate palmar tubercle; palmar supernumerary tubercles absent; subarticular tubercles moderately small, round; fingers bearing narrow lateral fringes; first finger shorter than second; discs on fingers nearly truncate, nearly 14x width of digit proximal to pad; all fingers having ventral pads well defined by circumferential grooves. Upper surfaces of hind limbs smooth with small tubercles; heel lacking ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 43 tubercle; outer edge of tarsus bearing low tubercles; inner edge of tarsus lacking tubercles or fold; inner metatarsal tubercle elevated broadly ovoid, about 5x subconical outer metatarsal tubercle; plantar supernumerary tubercles few, basal; discs on toes about equal in size to those on fingers; toes bearing narrow lateral fringes; toes unwebbed; tip of Toe V extending to distal edge of distal subarticular tu- bercle on Toe IV; tip of Toe III extending to middle of penultimate subarticular tubercle on Toe IV; when hind limbs flexed perpendicular to axis of body, heels broadly overlapping; shank 52.5% SVL. Coloration in preservative: Dorsum uniform tan; can- thal and supratympanic stripes, labial and interorbital bars, and transverse marks on limbs absent; venter creamy tan with minute black flecks on throat and belly. Coloration in life: Dorsum of head, body, and limbs orange-red; flanks, ventral surfaces of limbs, upper lips, loreal region, tympanum, and vocal sac yellow; rest of venter cream; tubercles on dorsum, flanks, and venter white; iris bronze with black reticulations and median, horizontal red streak (Fig. 9; Erik R. Wild, field notes, 8 January 1991). Measurements of holotype: SVL 20.4, tibia length 10.7, foot length 9.9, head width 7.5, head length 7.2, IOD 2.6, eyelid width 2.0, E—-N 2.0, eye 2.0, tympanum 1.1. Distribution and habitat.—This species is known only from 2770 m in humid montane forest on the western slopes of the Cordillera de Huancabamba (Fig. 22). The male was calling from a red leaf 0.5 m above the ground at night. Etymology.—The specific epithet is a Greek adjective, anemeros, meaning wild. The name is used in reference to Erik R. Wild, collector of the only known specimen. Remarks.—Duellman and Wild (1993) recognized this species as being distinct from others in the Cordillera de Huancabamba and were reluctant to name the species on the basis of a single specimen. After our review of the Eleutherodactylus of the Andes of northern Peru, we are convinced that the specimen cannot be associated with any species described previously. Eleutherodactylus ardalonychus new species Holotype.—KU 212301, an adult female, from the Rio Cerranayacu (05°46' S, 77°27' W, 1200 m), 76 km [by road] NW Rioja, Provincia Rioja, Departamento San Martin, Peru, obtained on 2 February 1989 by William E. Duellman. Paratypes.—KU 212299, adult male, from the west slope of Abra Tangarana, 1080 m, 7 km NE [by road] San Juan de Pacaysapa, Provincia Lamas, Departamento San Martin, Peru; KU 212300, adult male, from 8 km [by road] NE Tarapoto, 680 m, Provincia San Martin, Departamento San Martin, Peru. Referred specimen.—KU 212310, juvenile, from the type locality. Diagnosis.—A member of the Elewtherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum smooth with scattered low tubercles on posterior part of body, that on venter weakly areolate; discoidal fold barely evident posteriorly; dorsolateral folds absent; (2) tympanic membrane not differentiated; tympanic annu- lus distinguishable under skin, round, its diameter about ¥% length of eye; (3) snout long, rounded in dorsal view and in profile; (4) upper eyelid narrower than IOD, lack- ing tubercles; cranial crests absent; (5) vomerine odontophores low, ovoid; (6) males having vocal slits and unpigmented nuptial pads; (7) Finger I shorter than II; discs on outer fingers rounded, about twice width of digit proximal to pad; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tar- sus lacking tubercles; inner edge of tarsus bearing low, elongate tubercle; (11) inner metatarsal tubercle elevated, elliptical, about 10x low, round outer metatarsal tubercle; supernumerary plantar tubercles numerous; (12) toes bear- ing narrow lateral fringes; webbing absent; Toe V much longer than III; discs nearly as large as those on outer fin- gers; (13) dorsum brown with darker brown markings; venter tan with dark brown flecks or reticulations; poste- rior surfaces of thighs brown, with or without cream flecks or mottling; (14) SVL in two adult males 19.7 and 21.9 mm and an adult female 27.4 mm. The only other species in the region that can be con- fused with Eleutherodactylus ardalonychus is E. versicolor, which also has diagonal bars on the flanks and brown re- ticulations on the venter, but the reticulations are much coarser than those in E. ardalonychus. Furthermore, E. ver- sicolor differs by having a shagreen dorsum with scattered fine tubercles, a prominent tympanum, and elliptical, in- stead of rounded, discs on the outer fingers. Description.—(n = 2 males, 1 female; proportions are for the males with mean in parentheses, followed by those of the female). Head noticeably wider than body; HW 37.4— 38.1 (x = 37.8), 38.0% SVL; HL 38.4-39.1 (x = 38.8), 39.1% SVL; snout long, rounded in dorsal view and in profile; E-N 92.3-100.0 (x = 96.2), 96.8% length of eye; nostrils slightly protuberant, directed laterally; canthus rostralis rounded, nearly straight; loreal region concave; lips not flared; upper eyelid with lacking tubercles; upper eyelid width 73.9-76.0 (x = 75.0), 70.8% IOD; cranial crests ab- sent; supratympanic fold weak, not obscuring upper edge of tympanic annulus; side of head nearly vertical; tym- panic annulus thin; tympanic membrane not pustular or thickened; tympanic annulus round; diameter of tympanic annulus 40.0, 41.7% length of eye; single, enlarged postrictal tubercle, posteroventral to tympanic annulus; skin on head smooth except for single, rounded tubercle 44 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS between orbits in KU 212301; choanae large, nearly round, not concealed by palatal shelf of maxillary arch; vomerine odontophores, low, oblique, posteromedian to choanae, oval in outline, each slightly smaller than choanae, sepa- rated medially by distance greater than width of odontophore, bearing 2-2 or 2-3 teeth in males, 3-4 in fe- male; tongue longer than wide, its posterior border shal- lowly notched, posterior half not adherent to floor of mouth. Male with vocal slits extending posterolaterally from midlateral base of tongue; vocal sac single, median, subgular; unpigmented nuptial pads present. Dorsum of head, body, and limbs smooth with scat- tered low tubercles on posterior part of body, small, subconical tubercles; dorsolateral folds absent; flanks finely tuberculate; cloacal sheath short; skin on throat weakly areolate, on belly coarsely areolate; discoidal fold barely evident posteriorly. Ulnar tubercles absent; thenar tubercle ovoid, about 2 size of bifid palmar tubercle; su- pernumerary palmar tubercles few, minute; subarticular tubercles prominent, round; fingers bearing distinct lat- eral fringes; Finger I shorter than II; disc on thumb barely expanded; disk on Finger II slightly larger; discs on Fin- gers II-IV broadly rounded, more than twice width of digits; all fingers having ventral pads defined by circum- ferential grooves. Upper surfaces of hind limbs smooth; heel and outer edge of tarsus lacking tubercles; inner edge of tarsus bearing low, elongate tubercle; inner metatarsal tubercle elevated, elliptical, about 10x low, round, outer metatarsal tubercle; supernumerary plantar tubercles nu- merous; subarticular tubercles round, subconical; toes bearing narrow lateral fringes; webbing absent; discs on toes equal in size to those on fingers; tip of Toe V extend- ing to distal edge of distal subarticular tubercle on Toe IV; tip of Toe III extending to distal edge of penultimate subarticular tubercle on Toe IV; when hind limbs flexed perpendicular to axis of body, heels barely overlapping; shank 51.8-52.5 (x= 51.2), 50.0% SVL. Coloration in preservative: Dorsum tan with dark brown markings—KU 212264; elongate mark on snout, interorbital bar, canthal stripe. Supratympanic stripe, la- bial bars, W-shaped mark in scapular region (absent in KU 212229); chevron in sacral region (absent in KU 212229 and 212310). Diagonal bars on flanks and limbs, digits cream with dark brown transverse bars, discs cream with dark brown on distal edges. Posterior surfaces of thighs brown with cream mottling in KU 212229-300 and cream flecks in KU 212310. Venter creamy tan with fine dark brown reticulations (brown flecks in KU 212301). Coloration in life: KU 212301, female: Dorsum reddish brown with dark brown marking; venter and anterior and posterior surfaces of thighs gray; iris dull bronze with median, horizontal, red streak (Fig. 9). KU 212299, male: Dorsum tan, black dorsolaterally with greenish-tan bars on flanks; dorsal surfaces of limbs olive-brown with yel- lowish-tan bars; axilla, groin, and ventral surfaces of hind limbs salmon; throat and belly dull yellow with black flecks; salmon suffusion on belly; iris reddish copper. KU 212300, male: Dorsum mottled reddish brown and dark brown; flanks cream with diagonal cream bars; venter cream with gray mottling; iris bronze with median, hori- zontal, red streak. Measurements of holotype: SVL 27.4, tibia length 13.7, foot length 11.6, head width 10.4, head length 10.7, IOD 2.4, upper eyelid width 1.7, E-N 3.0, eye 3.0, tympanum 1.2. Distribution and habitat——The species is known from three localities at 680-1200 m on the east slope of the north- ern part of the Cordillera Central in northern Peru, where they were found in lower humid montane forest (Fig. 22). The juvenile was on the ground by day, and one male was in the axil of an elephant ear plant (Xanthosoma) by day. The female was on the leaf of an elephant ear plant at night, and a male was on the leaf of an herb at night. Etymology.—The specific name is derived from the Greek ardalos meaning dirty and the Greek onychos mean- ing fingernail; the name is applied in reference to the ir- regular, dark periphery of the discs on the fingers. Remarks.—One juvenile male (KU 212310 ) hasaSVL of 14.4 mm. In life, the dorsum was black with a tan mid- dorsal stripe broken by a black interorbital bar and at the level of the forelimbs. The flanks and dorsal surfaces of the thighs were marked with yellow bars; the throat and chest were black with pale yellow flecks, and the belly and groin were cream with yellow flecks. The iris was bronze-brown. Eleutherodactylus bearsei Duellman Eleutherodactylus bearsei Duellman, 1992a:2. Holotype: KU 212268, adult female, from Cataratas Ahuashiyacu, 730 m, 14 km [by road] NE Tarapoto, Provincia San Martin, Departamento San Martin, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum shagreen, bearing scattered low tubercles in males, that on venter areolate; discoidal fold evident only poste- riorly; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus prominent, vertically ovoid, its length about 30% length of eye; (3) snout acutely rounded in dorsal view, bluntly rounded in profile; canthus rostralis angular; (4) upper eyelid lacking small tubercles, broader than IOD; cranial crests absent; (5) vomerine odontophores transverse, prominent; (6) males having vocal slits but lack- ing nuptial pads; (7) Finger I shorter than II; discs broad; (8) fingers bearing lateral fringes; (9) ulnar tubercles dif- fuse; (10) heel lacking tubercles; outer edge of tarsus bear- ing tubercles; inner edge of lacking tubercles or fold; (11) inner metatarsal tubercle oval, 8-10 subconical outer ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 45 metatarsal tubercle; supernumerary tubercles diffuse, present only proximally; (12) toes bearing lateral fringes; webbing absent; Toe V much longer than III; discs nearly as large as those on fingers; (13) dorsum brown with darker brown marks on back and transverse bars on limbs; ven- ter brown with cream flecks; (14) SVL in males 22.7—25.5 mm, in females 38.0-38.8 mm. Only three species of Eleutherodactylus in the Andes of northern Peru have pale spots or flecks on brown flanks. In Eleutherodactylus bearsei, cream flecks are present on the flanks; the posterior surfaces of the thighs are uniform brown. The flanks have cream (E. rufioculis) or white (E. muscosus) spots, and the posterior surfaces of the thighs have cream spots in E. muscosus. The latter and E. rufioculis have smooth skin on the dorsum (shagreen and tubercu- late in E. bearsei); the former also differs from E. bearsei by having a conical tubercle on the heel, and the latter differs by lacking a tympanic membrane. Description.—The description by Duellman (1992a) is adequate. Distribution and habitat.—This species is known from the type locality and 30 km [by road] SW Zapatero (ca. 10 km NE San José de Sisa), 500 m, Provincia Lamas, Departamento San Martin, Peru (Fig. 22). Adults were found on mossy boulders and juveniles on herbaceous vegetation at night in deep ravine at elevations of 500 and 730 m in lower humid montane forest. Remarks.—As noted by Duellman (1992a), Eleuth- erodactylus bearsei can be grouped phenotypically with three other species (E. diadematus, eurydactylus, and platydactylus) in the upper Amazon Basin and on the lower Amazonian slopes of the Andes. Eleutherodactylus bromeliaceus Lynch Eleutherodactylus bromeliaceus Lynch, 1979:12. Holotype: USNM 199731, adult female, from between Sapote and Suro Rancho, Provincia Morona-Santiago, Ecuador. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum smooth, that on venter coarsely areolate; discoi- dal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus evident, round, its length about 30% length of eye; (3) snout subacuminate in dorsal view, acutely rounded in profile; canthus rostralis rounded; (4) upper eyelid bearing small tubercles, slightly narrower than IOD; cranial crests absent; (5) vomerine odontophores oblique, prominent; (6) males having vocal slits but lacking nuptial pads; (7) Finger I shorter than II; discs elliptical; (8) fingers bearing lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tarsus bear- ing low tubercles; inner edge of tarsus bearing one tubercle; (11) inner metatarsal tubercle oval, 3x elongate outer meta- tarsal tubercle; supernumerary tubercles numerous; (12) toes bearing lateral fringes; webbing absent; Toe V much longer than III; discs slightly smaller than those on fin- gers; (13) dorsum tan with brown flecks or blotches on dorsum; venter cream, with brown flecks in adult females; (14) SVL in males 16.7—23.2 mm, in females 22.9-28.1 mm. This species with a broad head and pointed snout has small tubercles on the upper eyelid and on the heel. Other species with this combination of characters are Eleutherodactylus ceuthospilus, rhodoplichus, and schultet. Eleutherodactylus ceuthospilus differs by having the skin on the dorsum shagreen and having a brown (as opposed to green) dorsum in life and pale spots in the groin; E. rhodoplichus differs by lacking vomerine odontophores and labial bars and by having dark diagonal bars or reticula- tions on the flanks and dark reticulations on the posterior surfaces of the thighs, whereas E. schultei differs by hav- ing a uniformly pale dorsum (as opposed to dark spots) and dark streaks on the flanks, and by lacking labial bars, pale spots on the posterior surfaces of the thighs, and dark flecks on the venter. Description.—The description by Lynch (1979) is ad- equate. Distribution and habitat.—The distribution of this spe- cies is discontinuous. It is known from the Amazonian slopes of the Cordillera Oriental (1707-2622 m), northern (1500-1600 m) and western (1830 m) slopes of the Cordil- lera del Condor, and the Cordillera de Cutucu (1700 m) in southern Ecuador, and from Abra Pardo de Miguel (2180 m) in the northern part of the Cordillera Central, Departamento San Martin, Peru (Fig. 22). All localities are in humid montane forests, where individuals were found on low vegetation at night and in bromeliads by day. At Abra Pardo de Miguel, a female was on the leaf of a herb and a male was calling from the leaf of a bush on the night of 31 January 1989; the call is a soft “peep.” Remarks.—Previously, this species has been recorded only from Ecuador (Almendariz, 1997; Duellman and Lynch, 1988; Lynch, 1980). The two Peruvian specimens (KU 212213-14) are a subadult female having a SVL of 22.7 mm and an adult male having a SVL of 23.0 mm, respec- tively. In life, the dorsum of the female was mottled green and brown, and the venter was gray; the dorsum of the male was tan with brown mottling, and the flanks, groin and vocal sac were yellow and the belly cream. In both individuals, the iris was red to reddish brown. Eleutherodactylus cajamarcensis Barbour and Noble Eleutherodactylus cajamarcensis Barbour and Noble, 1920:404. Holotype: MCZ 5407, adult male, from ruins near Huambos, Departamento Cajamarca, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus group having (1) skin on dorsum shagreen, bearing ill-defined rows of pustules, that 46 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS on venter areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic an- nulus prominent, round, its length no more than % length of eye; (3) snout rounded in dorsal view and in profile; canthus rostralis angular; (4) upper eyelid bearing small tubercles, slightly narrower than IOD; cranial crests ab- sent; (5) vomerine odontophores oblique, not prominent; (6) males having vocal slits and nuptial pads; (7) Finger I shorter than II; discs small, round; (8) fingers lacking lat- eral fringes; (9) ulnar tubercles indistinct or absent; (10) heel and outer edge of tarsus lacking tubercles; inner edge of tarsus usually bearing one tubercle; (11) inner metatar- sal tubercle oval, 46x oval outer metatarsal tubercle; plan- tar supernumerary tubercles numerous; (12) toes bearing lateral fringes; webbing absent; Toe V much longer than Toe III; discs as large as those on fingers; (13) dorsum gray to pale brown with darker brown markings; venter cream with brown or gray spots; (14) SVLin males 19.2—24.1 mm, in females 27.1—-33.8 mm. The only other members of the Eleutherodactylus unistrigatus Group in the Andes of northern Peru that have pale spots in the groin are E. ceuthospilus, lirellus, muscosus, and rufioculis. In all of these, the pale spots are not set ina black field, as in E. cajamarcensis. Furthermore, the larger E. muscosus has a conical tubercle on the heel and white (instead of brown) spots on the flanks, whereas the smaller E. rufioculis lacks a tympanic membrane and tubercles on the upper eyelid, and has cream (instead of brown) spots on the flanks. The much smaller E. lirellus lacks a tympanic membrane and annulus. Eleutherodactylus ceuthospilus dif- fers by lacking brown spots on the flanks and having dark flecks (instead of spots) on the venter; the spots in the groin are bright yellow in life. In E. cajamarcensis, the large spots in the groin are red in life. Description.—Lynch (1969) provided a thorough rede- scription of the species. Coloration of living individuals from Peru was described by Duellman and Wild (1993) and of individuals from southern Ecuador by Lynch and Duellman (1997). Distribution and habitat.—This species is widely dis- tributed in the mountains of the Huancabamba Depres- sion and around the Cuenca de Loja, where it occurs at elevations of 1800-3100 in tropical dry forest, humid mon- tane forest, and subparamo (Fig. 22). A few records exist for the Pacific slopes of the Cordillera Occidental in Peru (Huambos, Departamento Cajamarca) and in Ecuador (Luz Maria, Provincia Azuay). Most individuals have been found under rocks and logs or in arboreal or terrestrial bromeliads by day; few have been observed perched on low vegetation at night. Remarks.—Eleutherodactylus cajamarcensis is the most widespread species of the genus in western Peru and ad- jacent Ecuador. Eleutherodactylus ceuthospilus Duellman and Wild Eleutherodactylus ceuthospilus Duellman and Wild, 1993:8. Holotype: KU 219775, adult male, from 15.8 km [by road] ENE Canchaque, 1800 m, Departamento Piura, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum shagreen with minute, low, round tubercles and lacking folds, that on venter areolate; discoidal fold evi- dent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus prominent, round, its length about % length of eye; (3) snout acutely rounded in dorsal view and in profile; canthus rostralis rounded; (4) upper eyelid bearing small tubercles, narrower than IOD; cranial crests absent; (5) vomerine odontophores low, round, concealed in buccal mucosa; (6) males having vocal slits but lacking nuptial pads; (7) Finger I shorter than II; discs large, ellip- tical; (8) fingers bearing lateral fringes; (9) ulnar tubercles low, diffuse; (10) heel lacking tubercles; outer edge of tar- sus with few, low, round tubercles; inner edge of tarsus bearing low tubercles or fold; (11) inner metatarsal tubercle oval, 6x subconical outer metatarsal tubercle; plantar su- pernumerary tubercles numerous; (12) toes bearing lateral fringes; webbing absent; Toe V much longer than III; discs nearly as large as those on fingers; (13) dorsum gray to pale brown, usually with small brown flecks or streaks; venter cream with minute dark flecks; (14) SVL in males 19.0-25.8 mm, in females 23.5—26.7 mm. Eleutherodactylus ceuthospilus most closely resembles E. cajamarcensis, which differs by having dark (instead of pale) spots on the flanks and dark spots (instead of flecks) on the venter; moreover, the spots in the groin are dull red in life in E. cajamarcensis and bright yellow in E. ceuthospilus (Fig. 9). Other members of the Elewtherodactylus unistrigatus group in the region that have pale spots in the groin are the larger E. muscosus and the smaller E. lirellus and E. rufioculis; both of the small species lack a tympanic mem- brane (annulus also absent in E. lirellus), whereas E. muscosus differs in color pattern (white reticulations on the dorsum) and by having a conical tubercle on the heel. Description.—The description given by Duellman and Wild (1993) is adequate. Distribution and habitat.—This species is known from humid montane forest at elevations of 1735-2870 m on the western slopes of the Cordillera de Huancabamba, and from a site at 12 km [by road]W of Lamas (1500 m) on the Pacific slope of the Cordillera Occidental in northern Peru (Fig. 23). Males call from low vegetation in cloud forest at night; adults of both sexes have been found in bromeliads by day. Remarks.—Richard Thomas (field notes, 8 December 1974) described the call as ratchetlike, rapid sequence of three (rarely 2 or 4) clicks repeated at intervals of 15-20 sec. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 47 < 1000 m 1000-2000 m 2000-3000 m SH 3000-4000 m Sa >4000m ©. ceuthospilus . colodactylus . cryptomelas . ExOristus . galdi 2. Incomptus Fig. 23. Localities of known occurrence of six species in the Eleutherodactylus unistrigatus group in the Andes of southern Ecuador and northern Peru. Eleutherodactylus colodactylus Lynch Eleutherodactylus colodactylus Lynch, 1979:15. Holotype: KU 142151, adult female, from Abra de Zamora, 2800 m, 13.5 km [by road] E Loja, Provincia Loja, Ecuador. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum and venter areolate; discoidal fold and dorsolat- eral folds absent; (2) tympanic membrane and tympanic annulus absent; (3) snout rounded in dorsal view, bluntly rounded in profile; canthus rostralis rounded; (4) upper eyelid bearing one or two small but prominent tubercles, narrower than IOD; cranial crests absent; (5) vomerine odontophores oval, concealed; (6) males lacking vocal slits and nuptial pads; (7) Finger I shorter than II; discs small, round; (8) fingers having lateral fringes; (9) ulnar tubercles absent; (10) heel bearing single, conical tubercle; tarsus lacking tubercles and fold; (11) inner metatarsal tubercle oval, 4-5x round outer metatarsal tubercle; plantar super- numerary tubercles numerous, continuing onto toes; (12) toes bearing lateral fringes; webbing basal; Toe V longer than III; discs about as large as those on fingers; (13) dor- sum brown with pale dorsolateral stripes or interorbital bar; venter cream with brown flecks; (14) SVL in males 14.0-20.7 mm, in females 16.5—25.8 mm. In the region under consideration, only Elewtherodactylus proserpens is like E. colodactylus in having short, stocky fin- gers, but E. colodactylus differs from E. proserpens by lack- ing a cloacal sheath, tympanic membrane and annulus, a papilla on the tip of the snout, and prominent vomerine odontophores. In other Eleutherodactylus in the region hav- ing digital pads that are only slightly broader than the digit proximal to the pad (E. atrabracus, melanogaster, pataikos, and pinguis), the fingers are proportionately longer and more slender than those in E. colodactylus and E. proserpens; moreover, in those robust-bodied species, Toe V is only slightly longer than Toe III. The absence of a tympanum and tympanic annulus distinguishes E. colodactylus from all other species in the region, except E. lirellus; the latter differs by having vocal slits, labial and limbs bars, a yel- low spot in the groin, black flecks on the venter, and long fingers bearing elliptical discs. Description.—The original description by Lynch (1979) was augmented with data from Peruvian specimens by Duellman and Wild (1993). Distribution and habitat.—This species is known from two disjunct regions—Amazonian slopes (2195-3140 m) of the Cordillera Oriental in Provincia Morona-Santiago, Ecuador; the crest and Amazonian slopes (2710-2800 m) of the Abra de Zamora in the southern Cordillera Orien- tal; and the crest and upper eastern slopes of the Cordil- lera de Huancabamba in northern Peru (Fig. 23). All indi- viduals for which habitat data are available were found in terrestrial and arboreal bromeliads by day in humid mon- tane forest or subparamo. Eleutherodactylus cryptomelas Lynch Eleutherodactylus cryptomelas Lynch, 1979:21. Holotype: KU141992, imma- ture female, from 15 km [by road] E of Loja, 2710 m, Provincia Morona-Santiago, Ecuador. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum shagreen, bearing conical warts and ridges, that on venter areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic an- nulus prominent, its length 30-40% length of eye; (3) snout subacuminate in dorsal view, rounded in profile; canthus rostralis rounded; (4) upper eyelid bearing tubercles, slightly broader than IOD; cranial crests absent; (5) vomer- ine odontophores round to oval, prominent; (6) males lack- ing vocal slits and nuptial pads; (7) Finger I shorter than II; discs broad, elliptical; (8) fingers having weak lateral fringes; (9) ulnar tubercles prominent; (10) heel and outer edge of tarsus bearing conical tubercles; inner edge of tar- sus bearing indistinct tubercle; (11) inner metatarsal tu- bercle oval, 4-6x conical outer metatarsal tubercle; plan- tar supernumerary tubercles numerous; (12) toes bearing lateral fringes; webbing absent; Toe V longer than III; discs 48 ScrenTiFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS smaller than those on fingers; (13) dorsum gray to brown sparse darker markings; venter white to cream with brown reticulations; (14) SVL in males 28.2—30.2 mm, in female 38.6 mm. Eleutherodactylus cryptomelas is readily distinguished from all other members of the Eleutherodactylus unistrigatus Group in the region by having the groin and hidden sur- faces of the thighs uniform black. The only other species having extensive black coloration are short-legged mem- bers of the Eleutherodactylus orestes group—E. atrabracus with black on the undersides of the hind limbs and E. melanogaster, which is nearly uniform black dorsally and ventrally; both species have pale yellow spots in the groin. Description.—The original description by Lynch (1979) is adequate, but coloration in life was expanded by Duellman and Wild (1993). Distribution and habitat.—This species is known from elevations of 2470-2710 m on the Amazonian slopes of the Cordillera Oriental and the ridges (3000-3100 m) north of the Cuenca de Loja in Ecuador, and at elevations of 2770- 2820 m on the western slope of Cordillera de Huancabamba in Peru (Fig. 23). In Ecuador, the frogs were found in ter- restrial bromeliads or under rocks in paramo and subparamo by day. In Peru, the frogs were in humid mon- tane forest; one was under a rock by day, and another was in a tree at night Eleutherodactylus exoristus new species Holotype.—KU 147051, an adult female, from the Rio Piuntza (03°52' S, 78°15' W, 1550 m), western slope of the Cordillera del Condor, Provincia Morona-Santiago, Ecua- dor, one of a series collected on 3 January 1972 by John E. Simmons, Robert Fiske, and Bruce MacBryde. Paratypes.—KU 147048-49, 147053, 147056, adult males, and 147047, 147052, 147054—55, 147058, adult fe- males, from the type locality, 3-6 January 1972. Referred specimens.—KU 147050, 147057, juveniles, from the type locality; USNM 525442, an adult female, from Puesto Vigilancia Comainas, 665 m, Rio Comainas, Departamento Amazonas, Peru; USNM 525448, 525462, 525470 from Alfonso Ugarte, 1138 m, upper Rio Comainas, Departamento Amazonas, Peru. Diagnosis.—A member of the Eleutherodactylus (unistrigatus) unistrigatus Group having (1) skin on dor- sum tuberculate with small conical tubercles in dorsolat- eral rows, that on venter areolate; discoidal fold evident; dorsolateral folds absent; (2) tympanic membrane differ- entiated; tympanic annulus prominent, round, its diam- eter slightly less than 2 length of eye; (3) snout moder- ately long, acutely rounded in dorsal view, bluntly rounded in profile; (4) upper eyelid slightly narrower than IOD, bearing many conical tubercles; cranial crests absent; (5) Fig. 24. Dorsal and lateral views of the head of Eleutherodactylus exoristus, KU 147051. Scale bar = 5 mm. vomerine odontophores low, ovoid, oblique; (6) males hav- ing vocal slits but lacking nuptial pads; (7) Finger I shorter than II; discs on outer fingers elliptical, about twice width of digit proximal to pad; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tar- sus lacking tubercles; inner edge of tarsus bearing low, elongate tubercle distally; (11) inner metatarsal tubercle elevated, elliptical, about 8x subconical outer metatarsal tubercle; supernumerary plantar tubercles few, low; (12) toes lacking lateral fringes; webbing absent; Toe V much longer than III; discs nearly as large as those on outer fin- gers; (13) dorsum brown with darker brown markings; flanks pale tan, usually with diffuse diagonal bark bars; venter tan with dark brown flecks; posterior surfaces of thighs brown with small cream flecks ventrally; (14) SVL in four adult males 15.0-16.9 (x = 16.2) mm and six adult females 21.3-23.5 (x= 22.7) mm. Eleutherodactylus exoristus differs from all other species in the region by having flared lips and tubercles arranged in sinusoidal dorsolateral rows on the body. This species is most similar to E. percnopterus and E. serendipitus, both of which lack lateral fringes on the digits and tubercles on the heels and have tubercles on the dorsum, but the tu- bercles are lower and not arranged in dorsolateral rows; furthermore, these species lack conical tubercles on the upper eyelids, are slightly larger, and vomerine odontophores are absent in E. percnopterus. Other species in the region having tubercles on the dorsum are E. anemerus, which lacks dorsal markings and has a tubercle on the tip of the snout, the much larger E. bearsei, which has a rounded snout, lateral fringes on the digits, and no tubercles on the upper eyelids, and E. lanthanites, which has a smooth venter, first finger longer than the second, and fifth toe only slightly longer than the third. Description.—(1 = 4 males, 6 females). Head slightly wider than body; HW 36.2-42.0 (x= 38.7% SVL; HL 40.0— 46.0 (X = 42.8)% SVL; snout moderately long, acutely rounded in dorsal view, bluntly rounded in profile (Fig. 24); E-N 74.0-93.3 (X= 83.9)% length of eye; nostrils slightly ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 49 protuberant, directed laterally; canthus rostralis distinctly angular, slightly curved; loreal region concave; lips flared; upper eyelid bearing many conical tubercles; upper eye- lid width 81.5-91.3 (X= 85.8)% IOD; cranial crests absent; supratympanic fold weak, barely obscuring upper edge of prominent tympanic annulus; side of head nearly verti- cal; tympanic membrane not pustular or thickened; tym- panic annulus round, its diameter 37.9-52.3 (X = 43.7)% length of eye; two or three conical postrictal tubercles posteroventral to tympanic annulus. Choanae small, oval, not concealed by palatal shelf of maxillary arch; vomerine odontophores, low, oblique, medial to posterior edges of choanae, oval in outline, narrowly separated medially, each bearing three teeth in males (odontophore absent on one side in each of two males), 3-6 (x= 5.8) in females; tongue slighily more than twice as long as wide, its posterior bor- der shallowly notched, posterior half not adherent to floor of mouth. Males with vocal slits extending posterolaterally from midlateral base of tongue; vocal sac single, median, subgular; nuptial pads absent. Dorsum of head, body, and limbs bearing conical tu- bercles; in most individuals tubercles forming shallow, si- nusoidal rows extending from posterior border of orbit to point posterior to sacrum; numerous low tubercles on dor- sum of snout; dorsolateral folds absent; flanks finely tu- berculate; skin on throat, belly, and ventral surfaces of thighs areolate; discoidal fold evident; cloacal sheath short; enlarged tubercles lateral to vent absent. Ulnar tubercles absent; thenar tubercle low, ovoid, nearly as large as low, bifid palmar tubercle; supernumerary palmar tubercles few, subconical; subarticular tubercles prominent, subconical; fingers lacking lateral fringes; Finger I shorter than II; disc on thumb moderately expanded; discs on other fingers elliptical, about twice width of digits proximal to pad; all fingers having ventral pads defined by circumfer- ential grooves. Heel and outer edge of tarsus lacking tu- bercles; inner edge of tarsus bearing low, elongate tubercle distally; inner metatarsal tubercle elevated, elliptical, about 8x subconical outer metatarsal tubercle; supernumerary plantar tubercles few, low; subarticular tubercles low, round; toes lacking lateral fringes; webbing absent; discs on toes nearly as large as those on fingers; tip of Toe III extending to middle of distal subarticular tubercle on Toe IV; Toe V extending to point midway between penultimate and distal subarticular tubercles on Toe IV; when hind limbs flexed perpendicular to axis of body, heels overlap- ping by about 20% length of shank; shank 52.1-59.3 (x = 54.3)% SVL. Coloration in preservative: Dorsum of head, body, and limbs reddish brown with darker brown to black mark- ings (Fig. 25) consisting of distinct labial bars and trans- verse bars on limbs and diffuse canthal stripe in all indi- viduals, interorbital bar (8 individuals), narrow Fig. 25. Dorsal color pattern of Eleutherodactylus exoristus, KU 147051. SVL = 23.4 mm. supratympanic stripe (5), W- or H-shaped mark in scapu- lar region (4), paravertebral streaks on anterior part of body (6), diffuse sacral chevrons (3); one individual (KU 147050) with pale middorsal stripe, and one (KU 147058) with large cream blotch in sacral region. Flanks cream or tan with (8) or without (2) vertical to slightly diagonal bars; posterior surfaces of thighs brown with intrusion of tan dorsally and minute pale flecks ventrally; venter tan with brown flecks, most dense on throat and chest; two individuals with pale tan heels. Coloration in life: Dorsum brown to reddish brown with darker brown markings; one with white middorsal stripe and one with yellow blotch in sacral region; flanks yellow with brown bars; venter gray with or without white flecks (John E. Simmons field notes, 3-6 January 1972). Measurements of holotype: SVL 23.4, tibia length 12.5, foot length 10.5, head width 9.0, head length 9.6, IOD 2.3, upper eyelid width. 2.1, E-N 2.8, eye 3.0, tympanum 1.3. Distribution and habitat.—This species is known from one locality at 1550 m on the western slopes of the northern part of the Cordillera del Condor and two localities along the Rio Comainas (665 and 1138 m) on the eastern slopes of the southern part of the Cordillera del Condor (Fig. 23). All individuals were on low vegetation in humid montane forest at night. Etymology.—The specific name is a Greek adjective, exhoristos, meaning exiled. The name is applied in the sense of the species being restricted to the isolated Cordillera del Condor. 50 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Remarks.—The two juveniles from the type locality have SVLs of 13.3 and 14.0 mm. They are colored like the adults; one has bars on the flanks, and in the other the flanks are uniform tan. Three subadult females from 1138 m on the eastern slopes of the Cordillera del Condor have SVLs of 17.4-19.4 (x= 18.7) mm. Structurally, they are like the topotypes, except that USNM 525462 is less tubercu- late on the dorsum. That individual has short paraverte- bral dark streaks and plain flanks; USNM 525470 has a diffuse middorsal blotch and plain flanks, whereas USNM 525448 has a dark chevron in the scapular region, a trans- verse bar in the sacral region, and barred flanks. In life, the last two individuals had “clear green” or “clear blue green” venters (Robert P. Reynolds field notes, 21 and 23 July 1994). The largest and most tuberculate specimen is an adult female (USNM 525442) having a SVL of 25.6 mm. It has a dark W-shaped mark in the scalar region, dark bars on the flanks, and faint brown reticulations on the poste- rior part of the belly. In life, the dorsum was brown with darker mottling, and the flanks were tan with brown bars; the venter was cream with brown spots, and the under- sides of the thighs were reddish (Robert P. Reynolds field notes, 3 August 1994). Eleutherodactylus galdi Jiménez de la Espada) Pristimantis galdi Jimenez de la Espada, 1870:62. Types, MNCN (now lost) from San José de Moti, Provincia Napo, Ecuador. Hylodes festae Peracca, 1904:28. Holotype: MSNT AN413, ex. 3776, adult male, from San José de Cuchipamba, Provincia Morona-Santiago, Ecuador. Synonymy fide Lynch, 1974b:16. Hylodes margaritifer Boulenger, 1912:189. Syntypes, BM 1912.11.1.54-55 (= 1947.2.16.78-79), from El Topo, Rio Pastaza, Provincia Tungurahua, Ecuador. Synonymy fide Lynch, 1969:270. Eleutherodactylus festae—Peters, 1955:348. Eleutherodactylus margaritifer—Peters, 1955:349. Eleutherodactylus galdi—Peters, 1955:350; Lynch, 1969:270; Lynch, 1974b:16. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum smooth to finely shagreen with scattered small tubercles, that on venter coarsely areolate, usually with scattered larger warts; discoidal fold evident; dorsolateral folds absent; (2) tympanic membrane differentiated, and tympanic annulus prominent, nearly round, its length 4— Y length of eye; (3) snout long, acuminate in dorsal view, truncate in profile, with slightly swollen tip; (4) upper eye- lid bearing conical tubercle, narrower than IOD; cranial crests present, serrate in large females; (5) vomerine odontophores prominent, oblique; (6) males possessing vocal slits; nuptial pads absent; (7) Finger I shorter than II; discs on outer fingers expanded, truncate, more than twice width of digit proximal to pad; (8) fingers bearing weak lateral fringes; (9) ulnar tubercles conical; (10) heel, outer edge of tarsus, and outer edge of foot bearing conical tu- bercles; inner edge of tarsus lacking tubercles; (11) inner metatarsal tubercle elliptical, 56x ovoid outer metatarsal tubercle; supernumerary plantar tubercles absent; (12) toes bearing weak lateral fringes; webbing absent; Toe V much longer than III; discs smaller than those on outer fingers; (13) dorsum cream (green in life) with dark brown canthal and supratympanic stripes, interorbital and limb bars, and sparse spotting on dorsum; venter cream with minute dark flecks and usually dark streaks on throat; posterior sur- faces of thighs cream; (14) SVL in males 17.1—24.8 mm, in females 28.1—-34.0 mm (Lynch and Duellman, 1980). The green (in life) or cream (in preservative) coloration in combination with an acuminate snout, flared lips, greatly expanded, truncate discs on the outer fingers, row of coni- cal tubercles extending from the heel to the outer edge of the foot, and serrate cranial crests (in large females) im- mediately distinguish Eleutherodactylus galdi from all other species in the region. The only other predominately green species is E. acuminatus, which has rounded lips and ellip- tical discs and lacks tubercles along the tarsus, cranial crests, and bars on the limbs; moreover, E. acuminatus has an undifferentiated tympanic membrane. Elewtherodactylus quaquaversus also has a conical tubercle on the upper eye- lid and a conical tubercle on the heel, but it lacks a tympa- num, cranial crests, tubercles along the outer edge of the tarsus and foot, and large, truncate discs on the fingers. Description.—Adequate descriptions and illustrations were provided by Lynch (1969) and Lynch and Duellman (1980). Distribution and habitat.—In Ecuador, this gaudy arboreal species is known in humid montane forests at el- evations of 1000-1740 m on the eastern face of the Cordil- lera Oriental (Lynch and Duellman, 1980), 1700-1975 m in the Cordillera de Cutuct (Duellman and Lynch (1988), and 1500-1550 in the Cordillera del Condor (Almendariz, 1997; Lynch and Duellman, 1980). The only Peruvian record is from 12 km [by trail] east of La Peca, 1700 m on the west- ern slope of the Cordillera Colan, Provincia Bagua, Departamento Amazonas (Fig. 23). The specimen was along a stream in humid montane forest. Remarks.—The single Peruvian specimen (LSUMZ 39362) is a juvenile having a SVL of 13.9 mm. In this small specimen, cranial crests and ventral warts are not evident. The dorsum is yellowish tan with dark brown canthal and supratympanic stripes, narrow interorbital bar, pair of di- agonal (directed posteromedially) streaks in the scapular region, chevron in sacral region, and limb bars. Several short, dark brown streaks are present on the posterior part of the throat. Eleutherodactylus incomptus Lynch and Duellman Eleutherodactylus incomptus Lynch and Duellman, 1980:35. Holotype: KU 143484, adult male, from 16.5 km [by road] NNE Santa Rosa, 1700 m, Provincia Napo, Ecuador. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 5) Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum smooth with low, flat tubercles, that on venter coarsely areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane smooth, and tym- panic annulus prominent, nearly round, its length about 30% length of eye; (3) snout moderately short, rounded in dorsal view and in profile; (4) upper eyelid with or with- out tubercles, as wide as IOD; cranial crests absent; (5) vomerine odontophores not visible except in large females, low, oblique; (6) males possessing vocal slits and nonspinous nuptial pads; (7) Finger I shorter than II; discs on outer fingers expanded, rounded, about twice width of digit proximal to pad; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tarsus lacking tubercles; inner edge of tarsus bearing one or two tubercles; (11) inner metatarsal tubercle oval, 56x subconical outer metatarsal tubercle; supernumerary plantar tubercles numerous; (12) toes bearing narrow lat- eral fringes; webbing absent; Toe V much longer than III; discs equal in size to those on outer fingers; (13) dorsum brown with darker brown markings—interorbital bar, W- shaped or diagonal shaped marks in scapular region, and labial and limb bars; venter brown; posterior surfaces of thighs dark brown; (14) SVL in males 15.6-18.8 mm, in fe- males 23.7—25.9 mm (Lynch and Duellman, 1980). Eleutherodactylus incomptus is most easily confused with E. percnopterus, which is similar in size and also lacks vomerine odontophores; the latter differs from E. incomptus by having the snout subacuminate in dorsal view, discs on fingers nearly truncate, digits lacking lateral fringes, inner edge of tarsus lacking tubercles, and males lacking nuptial pads. Furthermore, in E. percnopterus, the only dis- tinct dorsal markings consist of a pair of small black spots or streaks in the scapular region, and the venter is cream. The only other member of the Eleutherodactylus unistrigatus Group in the Andes of northern Peru that lacks vomerine odontophores is E. anemerus, a frog that lacks dorsal mark- ings and has an orange-red dorsum, yellow flanks, and a prominent tubercles on the snout. Three other species (E. atrabracus, melanogaster, and pataikos) are robust-bodied members of the E. orestes Group that have small terminal discs on the fingers, short hind limbs, and Toe V only slightly longer than Toe III. In its absence of distinctive features, Eleutherodactylus incomptus is like E. pecki, which differs from the former by having vomerine odontophores, a small tubercle on the heel and distinctly larger digital discs. It also resembles E. exoristus, which differs by hav- ing finely tuberculate skin on the dorsum, tubercles on the upper eyelid, cream flecks on the posterior surfaces of the thighs, and dark flecks on a cream venter. Distribution and habitat—Eleutherodactylus incomptus has been reported from elevations of 1370-1910 m on the Amazonian slopes of the Cordillera Oriental in Ecuador (Lynch and Duellman, 1980). The present specimens (KU 217319-20) represent the first records of the species from Peru; they were in terrestrial bromeliads (Aechmea) by day at Santa Rosa de la Yunga, 19 km north of Pongo de Rentema on the Rio Maranon, Provincia San Ignacio, Departamento Cajamarca, by W. Razzetto and W. A. Alarcon on 15 July 1989. This locality is at about 1300 m on the southern slopes of the Cordillera del Condor (Fig. 23). Remarks.—The two specimens are a subadult female having a SVL of 18.2 mm and a juvenile having a SVL of 13.9 mm. Both specimens have the dark brown W-shaped mark in the scapular region. Eleutherodactylus infraguttatus sp. nov. Holotype.—KU 212297, adult female, from the east slope of Abra Pardo de Miguel (05°46'S, 77°42' W, 2180 m), Provincia Rioja, Departamento San Martin, Peru, one of a series of five specimens collected on 31 January 1989 by William E. Duellman. Paratypes.—KU 212298, 212314-16 from the type lo- cality, and KU 217317 from 14 km [by road] west of Venceremos, 2000 m, Provincia Rioja, Departamento San Martin, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum smooth with scattered, small tubercles, that on venter areolate; discoidal fold absent; dorsolateral folds absent; (2) tympanic membrane evident, and tympanic annulus thin, not prominent, higher than long, its length Y%4—Ys length of eye; (3) snout short, rounded in dorsal view and in profile; (4) upper eyelid with small tubercles poste- riorly, about as wide as IOD; cranial crests absent; vomer- ine odontophores low, oval in outline; (6) males possess- ing vocal slits but lacking nuptial pads; (7) Finger I shorter than II; discs on outer fingers nearly truncate, more than twice width of digit proximal to pad; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles prominent; (10) heel and inner edge of tarsus lacking tubercles; (11) inner metatarsal tubercle ovoid, about 5x round outer metatar- sal tubercle; supernumerary plantar tubercles evident; (12) toes bearing narrow lateral fringes; webbing absent; Toe V slightly longer than III; discs equal in size to those on outer fingers; (13) dorsum brown with diffuse darker mark- ings; venter cream with dark brown mottling; throat brown with cream mottling; groin and posterior surfaces of thighs cream to tan with brown bars; (14) SVL in one male 15.8 mm, in three females 22.9-24.3 mm (X= 23.6). Eleutherodactylus infraguttatus has more distinct dark mottling on the belly than any other member of the Eleutherodactylus unistrigatus Group in the region. Four other species have dark reticulations on the belly, but all have dark labial bars (absent in E. infraguttatus). Further- Nn more, of these species, E. ardalonychus differs by lacking tubercles of the upper eyelid and diagonal bars on the flanks, and E. muscosus differs by having a conical tubercle on the heel and pale spots on the flanks; E. versicolor lacks vocal slits and has distinct bars on the flanks, whereas in E. cryptomelas, the groin and hidden surfaces of the thighs are black. Description.—(1 = 4; 1 male, 3 females; proportions are for male, followed by range and mean in parentheses of three females). Head as wide as body in male and in mature females, wider than body in gravid females; HW 39.2, 37.4-38.9% (X = 38.1) SVL; HL 37.9, 37.1-40.1% (x = 38.8) SVL; snout short, rounded in dorsal view, and in pro- file; E-N 68.3, 60.0-74.6% (x = 68.9)% length of eye; nos- trils slightly protuberant, directed laterally; canthus rostralis weakly angular, barely concave; loreal region weakly concave sloping abruptly to lips; lips not flared; upper eyelid with small tubercles posteriorly; upper eye- lid width 63.6, 70.3-89.0% (X = 76.9)% IOD; cranial crests absent; supratympanic fold weak, barely obscuring upper edge of tympanic annulus; side of head nearly vertical; tympanic annulus thin; tympanic membrane not pustular or thickened; tympanic annulus slightly higher than long; length of tympanic annulus 42.1, 26.9-33.3% (X = 30.7)% length of eye; postrictal tubercles small, subconical, posteroventral to tympanic annulus; skin on head smooth. Choanae small, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores posteromedian to choanae (absent in one female), oval in outline, each about twice size of choana, separated medially by distance less than width of odontophore, each bearing two teeth in male and 4-6 teeth in female in single row; tongue longer than wide, its posterior border deeply notched, posterior half not adherent to floor of mouth; vocal slits short, postero- lateral to base of tongue; vocal sac single, median, subgular. Dorsum smooth with scattered, small, subconical tu- bercles and faint dermal ridge from posteromedian bor- der of eyelid to scapular region (most evident in male); dorsolateral folds; flanks smooth; cloacal sheath and en- larged tubercles in cloacal region absent; skin on throat and belly areolate; discoidal fold absent. Upper surfaces of arms smooth; ulnar tubercles round; thenar tubercle ovoid, smaller than bifid palmar tubercle; supernumerary palmar tubercles prominent, round, smaller than round, non conical subarticular tubercles; fingers bearing narrow lateral fringes; Finger I shorter than II; disc on thumb not expanded; discs on Fingers II-IV nearly truncate, twice width of digits; all fingers having ventral pads defined by circumferential grooves; nuptial pads absent in male. Up- per surfaces of hind limbs smooth with scattered subconical tubercles; heel tubercles absent; outer edge of tarsus lacking tubercles; inner metatarsal tubercle nearly round, flat, about 5x subconical outer metatarsal tubercle; 2 ScIeENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS < 1000m 1000-2000 m. 2000-3000 m 3000-4000 m > 4000 m Kilometers . iInfraguttatus . lirellus . MUSCOSUS . nephophilus . ockendeni E. pecki Fig. 26. Localities of known occurrence of six species in the Eleutherodactylus unistrigatus group in southern Ecuador and northern Peru. supernumerary plantar tubercles small, subconical; subarticular tubercles round, not conical; toes bearing lat- eral fringes; webbing absent; discs on toes nearly as large as those on fingers; tip of Toe V extending to distal edge of distal subarticular tubercle on Toe IV; tip of Toe III not ex- tending to that tubercle; when hind limbs flexed perpen- dicular to axis of body, heels barely overlapping; shank 49.4%, 46.7-50.8 (x = 49.0)% SVL. Coloration in preservative: Dorsum brown with darker brown canthal stripe, supratympanic stripe, interorbital bar, scapular marks, and transverse bars on limbs; flanks creamy tan with brown suffusion or spots posteriorly; an- terior and posterior surfaces of thighs brown, mottled with cream; venter cream with brown marbling in females, pri- marily brown in male. Coloration in life: Dorsum mottled dull olive and tan; flanks yellow and brown (Fig. 9); anterior surfaces of thighs red; venter greenish yellow with brown mottling; iris dull bronze with median, horizontal, brown streak (WED field notes on holotype, 31 January 1988). Measurements of holotype: SVL 22.9, tibia length 10.7, foot length 9.9, head width 8.9, head length 8.5, IOD 2.8, upper eyelid width 2.0, E-N 1.8, eye 2.5, tympanum 0.8. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 53 Distribution and habitat——The species is known only from two localities at 2000 and 2180 m along the road from Abra Pardo de Miguel to Moyobamba on the east slope of the northern part of the Cordillera Central in northern Peru (Fig. 26). All individuals were on leaves on low vegetation (<1m) above the ground in humid montane forest at night. Etymology.—The specific name is derived from the Latin prefix infra— meaning underside and the Latin adjec- tive guttatus meaning dappled; the name is applied in ref- erence to the dark mottling on the venter. Remarks.—Two juvenile females (KU 212314-15) have SVLs of 17.4 and 18.2 mm. Eleutherodactylus lirellus Dwyer Eleutherodactylus lirellus Dwyer, 1995:247. Holotype: KU 212240, adult female, from western slope of Abra Tangarana, 1080 m, 7 km [by road] NE San Juan de Pacaysapa, Provincia Lamas, Departamento San Martin, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum coarsely areolate with scattered small tubercles, that on finely areolate; discoidal fold prominent; dorsolat- eral folds absent; (2) tympanic membrane tympanic annu- lus not visible; (3) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid bearing many small tubercles, narrower than IOD; cranial crests absent; (5) vomerine odontophores prominent, oblique; (6) males possessing vocal slits; nuptial pads absent; (7) Finger I shorter than II; discs expanded, about twice width of digit proximal to pad; (8) fingers bearing lateral fringes; (9) ul- nar tubercles absent; (10) heel bearing two small tubercles; outer edge of tarsus bearing row of tubercles; inner tarsal tubercles and fold absent; (11) inner metatarsal tubercle elliptical, 4-5x subconical outer metatarsal tubercle; super- numerary plantar tubercles proximally; (12) toes bearing lateral fringes; webbing absent; Toe V much longer than III; discs nearly as large as those on outer fingers; (13) dor- sum brown with darker brown markings; groin brown with one pale spot; venter cream with dark flecks; poste- rior surfaces of thighs dark brown with cream flecks; (14) SVL in males 14.1-17.0 mm, in females 19.4—24.0 mm (Dwyer, 1995). The absence of a tympanum and tympanic annulus dis- tinguishes Eleutherodactylus lirellus from all other species in the northern Andes, except E. colodactylus, which dif- fers by lacking vocal slits and having short fingers with round discs. Furthermore, E. colodactylus lacks labial and limb bars and a yellow spot in the groin. The latter feature is shared by several species in the upper Amazon Basin. Two of these (E. toftae and E. variabilis) have a distinct tym- panic annulus; E. carvalhoi and E. croceoinguinis lack vocal slits and lateral fringes on the fingers and toes, and E. imitatrix has small yellow spots on the anterior surfaces of the thighs. Andean species with pale spots in the groin are E. ceuthospilus, muscosus, and rufioculis, all of which have tympanic membranes and distinct tympanic annuli, ex- cept E. rufioculis, in which the tympanic membrane is ab- sent, and the annulus is evident only ventrally. In the north- ern Andes, other species having yellow or pale spots in the groin are members of the Eleutherodactylus orestes Group; these small frogs have robust bodies, digits with narrow terminal discs, and Toe V only slightly longer than Toe III. Description.—The description by Dwyer (1995) is adequate. Distribution and habitat.—Eleutherodactylus lirellus is known from three localities at elevations of 470-1200 m on ridges of the eastern slopes of Cordillera Central in northern Peru (Fig. 26). All individuals were found at night on low vegetation in lower montane rainforest. Remarks.—Dwyer (1995) provided a detailed morpho- metric analysis of Eleutherodactylus lirellus and five Ama- zonian species that have yellow spots in the groin. How- ever, there is no evidence that the presence of yellow spots in the groin is a synapomorphy. Eleutherodactylus muscosus new species Holotype.—KU 219482, adult female, the east slope of Abra Pardo de Miguel (05°46' S, 77°41' W, about 1800 m), Provincia Rioja, Departamento San Martin, Peru, one of a series of four specimens collected on 30 July 1981 by Rainer Schulte. Paratypes.—KU 209479-80, subadult females, and 209481, an adult female, collected with the holotype. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum smooth with scattered, small tubercles, that on venter areolate; discoidal fold not evident; dorsolateral folds absent; (2) tympanic membrane and tympanic an- nulus distinct, slightly higher than long, its length about 30% length of eye; (3) snout long, bluntly rounded in dor- sal view and rounded in profile; (4) upper eyelid narrower than IOD, with one or two round tubercles posteriorly; cranial crests absent; (5) vomerine odontophores promi- nent, slightly oblique; (6) males unknown; (7) Finger | shorter than II; discs on outer fingers broadly expanded, nearly truncate, more than twice width of digit proximal to pad; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles low, diffuse; (10) heel bearing conical tubercle; outer edge of tarsus bearing one or two tubercles proxi- mally, inner edge of tarsus lacking tubercles; (11) inner metatarsal tubercle elevated, ovoid, about 3x subconical rounded outer metatarsal tubercle; supernumerary plan- tar tubercles prominent, in single row on each digit; (12) toes bearing narrow lateral fringes; webbing basal, between 54 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Toes II and IV, and IV and V; Toe V much longer than III; discs slightly smaller than those on outer fingers; (13) dor- sum with darker brown markings and with or without narrow, irregular white venation; throat and chest tan with brown reticulations or blotches; belly, groin, posterior sur- faces of thighs, and ventral surfaces of hind limbs dark brown with cream spots; (14) SVL in four females 29.6— 46.1 mm (x = 37.8). In coloration, Eleutherodactylus muscosus is unlike any other species of the genus in the Andes of northern Peru. The pale vermiculations on the dorsum are similar to those exhibited by some specimens of E. spinosus (Lynch, 1979:fig. 19), a species that differs in being smaller (females 28.3— 34.5 mm) and having a snout that is truncate in profile, elongate tubercles on the upper eyelid, low cranial crests, and subconical ulnar tubercles. Only two other species in the region have tubercles on the upper eyelid and a coni- cal tubercle on the heel; of these, E. galdi differs by having an acuminate snout in dorsal view and a uniformly cream venter, and E. quaquaversus differs by having elliptical (as opposed to truncate) discs on the fingers and a white venter. Description.—(n = 4 females). Head as wide as body; HW 39.4-41.0 (x = 39.8)% SVL; HL 40.1-43.4 (X = 41.9)% SVL; snout moderately long, bluntly rounded in dorsal view, rounded in profile; E—N 80.7-91.0 (x= 86.1)% length of eye; nostrils barely protuberant, directed laterally; can- thus rostralis angular, barely concave; loreal region deeply weakly concave, sloping abruptly to rounded lips; upper eyelid with one or two rounded tubercles posteriorly; up- per eyelid width 67.8-83.3 (X = 75.7)% IOD; cranial crests absent; supratympanic fold moderately heavy, obscuring upper part of tympanum; side of head nearly vertical; tym- panic annulus thin; tympanic membrane not pustular or thickened; tympanic annulus slightly higher than long; length of tympanic annulus 31.3-38.6 (X = 35.9)% length of eye; postrictal tubercles small, subconical, posteroventral to tympanic annulus; skin on head smooth. Choanae small, oval, not concealed by palatal shelf of maxillary arch; vomerine odontophores slightly oblique, posteromedian to choanae, elongately oval in outline, each about 4x size of choana, separated medially by distance less than width of odontophore, each bearing 4-6 (x = 4.9) teeth; tongue oval, longer than wide, its posterior border not notched, posterior third not adherent to floor of mouth. Dorsum of head, body, and limbs smooth with scat- tered, small, subconical tubercles; dorsolateral folds ab- sent; flanks weakly tuberculate; cloacal sheath and en- larged tubercles in cloacal region absent; skin on throat and belly areolate; discoidal fold not evident. Ulnar tu- bercles few, low, diffuse; thenar tubercle elongately ellip- tical, about equal in size to bifid palmar tubercle; super- numerary palmar tubercles few, minute; subarticular tu- lis 2. KU 219482. Scale bar = 5 mm. Hand and foot of Eleutherodactylus muscosus, bercles prominent, round; fingers bearing narrow lateral fringes; first finger shorter than second; disc on thumb expanded, round; discs on Fingers II-IV broadly expanded, nearly truncate, more than twice width of digits (Fig. 27); all fingers having ventral pads defined by circumferential grooves. Upper surfaces of hind limbs smooth; heel bear- ing conical tubercle; outer edge of tarsus bearing one or two subconical tubercles proximally; inner surfaces of tar- sus smooth; inner metatarsal tubercle elevated, ovoid, about 3x rounded outer metatarsal tubercle; supernumer- ary plantar tubercles prominent, in single row on each digit; subarticular tubercles subconical; toes bearing narrow lat- eral fringes; webbing basal between Toes III and IV and IV and V; discs on toes slightly smaller than those on fingers; tip of Toe V extending beyond distal edge of distal subarticular tubercle on Toe IV; tip of Toe III not extend- ing to that tubercle (Fig. 27); when hind limbs flexed per- pendicular to axis of body, heels slightly overlapping; shank 51.0-56.3 (X = 53.3)% SVL. Coloration in preservative: Dorsum grayish brown with darker brown markings consisting of streaks on dor- sum of snout, labial bars, diffuse interorbital bar, supratympanic stripe, and transverse bars on limbs (Fig. 28). In two smallest specimens (KU 209479-80, SVLs 32.7 and 29.6 mm) dark transverse marks in scapular and sac- ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 3)5) Fig. 28. Dorsal and ventral color pattern of Eleutherodactlyus muscosus, KU 219462. SVL = 46.1 mm. ral regions. In two largest specimens (KU 209481-81, SVLs 41.9 and 46.1 mm) dorsum of body pale brown with white vermiculations. Throat and chest of two smallest specimens tan with brown blotches. Throat, chest, and anterior part of belly in two larger specimens tan with fine brown mark- ings nearly forming reticulations; flanks brown with white spots. Posterior part of belly, ventral surfaces of hind limbs, groin, and posterior surfaces of thighs dark brown with cream spots. Coloration in life: Dorsum green with many brown and black spots and irregular white lines resembling struc- tures made by worms or fly larvae on stones; groin and lower surfaces of hind limbs with orange-yellow spots (R. Schulte, photographs and field notes, 30 July 1981). Measurements of holotype: SVL 46.1, tibia length 23.1, foot length 24.5, head width 18.3, head length 18.5, IOD 5.9, upper eyelid width 4.0, E-N 4.2, eye 5.2, tympanum 2.0. Distribution and habitat.—This species is known only from the type locality on the eastern slopes of the north- ern part of the Cordillera Oriental in northern Peru (Fig. 26). The type locality is a stream with rocky banks in hu- mid, upper montane forest. Stones along the edge of the stream are covered with moss. The frogs were found by day in cavities between stones; according to R. Schulte, the frogs are “imitating exactly a mossy stone.” Etymology.—The specific name is a Latin adjective muscosus, meaning mossy; the name is used in reference to the appearance of the species, as well as its habitat. Remarks.—The four specimens of this distinctive spe- cies are poorly preserved; they died en route from the type locality to the collector’s home in Tarapoto. Consequently, neither the measurements nor the descriptions of certain morphological structures are as accurate as in our other descriptions. Although Schulte’s field notes imply white vermiculations on all four individuals, they are evident only in the two largest individuals; however, the distinc- tive ventral coloration is the same in all four specimens. Eleutherodactylus nephophilus new species Holotype.—KU 212306, an adult female, the east slope of Abra Pardo de Miguel (05°46' S, 77°42' W, 2180 m, Provincia Rioja, Departamento San Martin, Peru, one of a series of six specimens collected on 31 January 1989 by William E. Duellman, Michael E. Morrison, and John J. Wiens. Paratypes.—KU 212307-09 from the type locality; KU 217322, an adult female, from 14 km [by road] W Venceremos, 2000 m, Provincia Rioja, Departamento San Martin, Peru. Referred specimens.—KU 212305 and 212317, juve- nile females, from the type locality; KU 212311, juvenile, from the west slope of Abra Tangarana, 7 km [by road] NE San Juan de Pacaysapa, Provincia Lamas, Departamento San Martin, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum smooth with sinusoidal dermal ridges from orbit to scapular region and scattered, small tubercles, that on venter areolate; discoidal fold weak; dorsolateral folds absent; (2) tympanic membrane evident, and tympanic annulus distinct, slightly higher than long, its length only 20-30% length of eye; (3) snout moderately long, rounded in dorsal view and in profile; (4) upper eyelid narrower than IOD, with small tubercles posteriorly; cranial crests absent; (5) vomerine odontophores prominent, oblique; (6) males unknown, (7) Finger I shorter than II; discs on outer fingers nearly truncate, more than twice width of digit proximal to pad; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles present; (10) heel and outer edge of tar- sus bearing tubercles; inner edge of tarsus bearing tubercles tending to coalesce into low ridge; (11) inner metatarsal tubercle ovoid, about 5x subconical outer metatarsal tu- bercle; supernumerary plantar tubercles prominent; (12) toes bearing narrow lateral fringes; webbing absent; Toe V slightly longer than III; discs slightly smaller than those on outer fingers; (13) dorsum brown with or without darker markings or paler markings; venter tan with dense dark brown flecks; posterior surfaces of thighs brown with cream spots; (14) SVL in five adult females 24.6-34.0 mm (x = 29.8). Eleutherodactylus nephophilus is like the sympatric E. rufioculis in having a red iris, but it differs from that smaller species (female = 20.6 mm) in several features—tubercles and ridges on dorsum (smooth in E. rufioculis), prominent Nn tympanum (annulus visible under skin), tubercles on up- per eyelid (absent), oblique vomerine odontophores ob- lique (ovoid), lateral fringes on fingers (absent), ulnar, heel, and tarsal tubercles (absent), dark bars on limbs diagonal (transverse), interorbital bar absent (present), labial bars present (absent), and posterior surfaces of thighs dark with pale spots (pale with dark spots). Eleutherodactylus nephophilus is smaller than E. muscosus (females to 37.8 mm), which differs by having truncate discs, conical tu- bercle on the heel, and large pale spots on the flanks and lower surfaces of the hind limbs. Description.—(n = 5 females). Head slightly wider than body; HW 37.8-40.0 (x = 39.1)% SVL; HL 37.8-39.4 (x = 38.1)% SVL; snout moderately long, rounded in dorsal view, and in profile; E-N 81.8-100.0 (x = 92.5)% length of eye; nostrils slightly protuberant, directed laterally; can- thus rostralis weakly angular, barely concave; loreal region weakly concave sloping abruptly to lips; lips not flared; upper eyelid with small tubercles posteriorly; upper eye- lid width 75.8-88.9 (X = 80.4)% IOD; cranial crests absent; supratympanic fold weak, barely obscuring upper edge of tympanic annulus; side of head nearly vertical; tympanic annulus thin; tympanic membrane not pustular or thick- ened; tympanic annulus slightly higher than long; length of tympanic annulus 21.2-37.5 (xX = 28.4)% length of eye; postrictal tubercles large, subconical, posteroventral to tympanic annulus; skin on head smooth. Choanae small, oval, not concealed by palatal shelf of maxillary arch; vomerine odontophores oblique, posteromedian to choa- nae, oval in outline, each about 4x size of choana, sepa- rated medially by distance less than width of odontophore, each bearing 3-5 (x= 4.2) teeth; tongue longer than wide, its posterior border shallowly notched, posterior half not adherent to floor of mouth. Dorsum of head, body, and limbs smooth with scat- tered, small, subconical tubercles; tubercles forming sinu- soidal ridge from posteromedian edge of eyelid to scapu- lar region; single, conical tubercle on midline anterior to orbits in two individuals; dorsolateral folds absent; flanks tuberculate; cloacal sheath and enlarged tubercles in cloa- cal region absent; skin on throat and belly areolate; discoi- dal fold evident posteriorly. Ulnar tubercles few, round to subconical; thenar tubercle ovoid, about % size of bifid palmar tubercle; supernumerary palmar tubercles few, minute; subarticular tubercles prominent, subconical; fin- gers bearing narrow lateral fringes; Finger I shorter than II; disc on thumb barely expanded; discs on Fingers I-IV elliptical, nearly truncate, more than twice width of digits; all fingers having ventral pads defined by circumferential grooves. Upper surfaces of hind limbs smooth with scat- tered small tubercles; heel bearing subconical tubercle; outer edge of tarsus bearing three or four subconical tu- 6 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS bercles; low fold on distal third of inner surfaces of tarsus; inner metatarsal tubercle flat, ovoid, about 5x subconical outer metatarsal tubercle; supernumerary plantar tubercles prominent, round; subarticular tubercles round, not coni- cal; toes bearing weak lateral fringes; webbing absent; discs on toes nearly as large as those on fingers; tip of Toe V extending to distal edge of distal subarticular tubercle on Toe IV; tip of Toe III not extending to that tubercle; when hind limbs flexed perpendicular to axis of body, heels barely overlapping; shank 50.6-55.2 (x = 52.9)% SVL. Coloration in preservative: Dorsum brown with a va- riety of markings—KU 212306: broad, median grayish tan blotches (snout, occipital region, scapular region, and sac- ral region); KU 212307: broad cream middorsal stripe nar- rowly bordered by dark brown; KU 212308 and 217322: dark brown dermal ridges in occipital-scapular region; KU 212309: like KU 212308 but with top of head pale tan with faint brown markings. Dorsal surfaces of limbs brown with darker brown diagonal bars narrower than interspaces. Flanks brown with narrow creamy tan diagonal marks or reticulations, especially evident posteriorly; anterior and posterior surfaces of thighs brown with pale cream verti- cal bars dorsally and spots or reticulations ventrally; dis- tinctive dark cloacal patch absent. Canthal stripe and in- terorbital bar usually absent (canthal stripe present in KU 217322); three or four dark labial bars; dark postorbital stripe encompassing upper part of tympanum or not. En- tire venter tan heavily flecked with brown, reticulate in KU 217322; cream midventral line on chest and belly in KU 212307. Coloration in life: Dorsal coloration highly variable— KU 212306: mottled yellowish tan, olive-green, and dark dull red; KU 212307: olive with median yellow stripe and orange dorsolateral area (Fig. 10); KU 212308: dull brown; KU 212309: brown with grayish-tan head. Flanks brown (dull red in KU 212306) with cream markings; venter dull yellow to tan with reddish, dark brown, or black flecks or reticulations; iris red. Measurements of holotype: SVL 29.7, tibia length 15.9, foot length 13.4, head width 11.6, head length 11.4, [OD 3.3, upper eyelid width 2.5, E—N 3.0, eye 3.0, tympanum 1.0. Distribution and habitat.—The species is known only from three localities at 1080, 2000, and 2180 m along the road from Abra Pardo Miguel to Moyobamba on the east slope of the northern part of the Cordillera Central in north- ern Peru (Fig. 26). Five adults and two juveniles were on leaves on low vegetation (< 1 m) above the ground in hu- mid montane forest at night; one juvenile was on the ground by day. Etymology.—The specific name is derived from the Greek nephos—meaning cloud and the Greek adjective ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU >)// philia meaning fondness; the name is applied in reference to the species inhabiting cloud forest. Remarks.—Three juveniles (KU 212317, 212305, and 212311 ) have SVLs of 17.1, 16.9, and 9.1 mm, respectively. Eleutherodactylus ockendeni (Boulenger) Hylodes ockendeni Boulenger, 1912:187. Syntypes, BM 1907.5.7.19-21 (= 1947.2.16.88-90), from La Union, Rio Huacamayo, Carabaya, Departamento Puno, Peru. Hylodes hylaeformis Melin, 1941:48. Types, NHMG, from Roque, Departamento San Martin, Peru. Synonymy fide Lynch, 1980. Syrrhophus calcaratus Andersson, 1945:27. Holotype, NHRM 1941, from Rio Cosanga near Archidona, Provincia Napo, Ecuador. Synonymy fide Lynch, 1974:16. Eleutherodactylus melini Bokermann, 1958:95. Replacement name for Hylodes hylaeformis Melin, 1941, non Phyllobates hylaeformis Cope, 1875). Synonymy fide Lynch, 1974b:16. Eleutherodactylus anderssoni Lynch, 1968:292. Replacement name for Syrrhophus calcaratus Andersson, 1945, non Hylodes calcaratus Boulenger, 1908. Synonymy fide Lynch 1974b:16. Eleutherodactylus ockendeni—Lynch, 1974b:16; 1980:12. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum shagreen with or without W-shaped occipital- scapular ridges and dorsolateral folds, that on venter coarsely areolate; discoidal fold evident; (2) tympanic membrane and tympanic annulus usually prominent, its length about 4—% length of eye; (3) snout short, subacuminate in dorsal view, rounded in profile; (4) up- per eyelid usually bearing tubercles, about equal in width to IOD; cranial crests absent; (5) vomerine odontophores, oval, oblique; (6) males having vocal slits, lacking nuptial pads; (7) Finger I shorter than II; discs moderately large, elliptical; (8) fingers having narrow lateral fringes; (9) ul- nar tubercles minute; (10) heel bearing rounded tubercle; outer edge of tarsus bearing small tubercles; inner edge of tarsus lacking tubercles or fold; (11) inner metatarsal tu- bercle elliptical, 46x round outer metatarsal tubercle; plan- tar supernumerary tubercles few; (12) toes bearing lateral fringes; webbing absent; Toe V much longer than III; discs as large as those on fingers; (13) dorsum cream to brown with brown to black interorbital bar, subocular spots, supratympanic stripe, dorsal chevrons, and bar posterior to sacrum; posterior surfaces of thighs uniform brown; venter white with or without brown flecks; (14) SVL in males 16.9-21.2 mm, in females 24.6-31.5 mm (Lynch, 1980, for Amazonian Peru). The presence of a W-shaped scapular mark on an oth- erwise plain pale brown dorsum is shared with Eleutherodactylus percnopterus, which lacks vomerine odontophores and has a dark canthal stripe. Four other members of the Eleutherodactylus unistrigatus Group in the region may have a W-shaped scapular mark; three of these (E. exoristus, incomptus, and sternothylax) have diagonal bars or streaks on the flanks, whereas E. bearsei has cream flecks on the flanks, and E. cryptomelas has black in the groin and on the anterior surfaces of the thighs. Description.—The description by Lynch (1974b) was augmented by that of Lynch (1980). Distribution and habitat.—Eleutherodactylus ockendeni is distributed throughout the upper Amazon Basin from southern Colombia to southern Peru. A few records indi- cate that the species ascends to slopes of the Andes to moderate elevations. The species was reported from 1140— 1150 m on the eastern slopes of the Cordillera Oriental (Lynch and Duellman, 1980) and from Miaza, 900 m, on the western slope of the Cordillera del Condor (Almendariz, 1997) in Ecuador, and from 1100-1280 m in the Serrania de Sira, Departamento Huanuco, Peru (Duellman and Toft, 1979). A single specimen from 4 km SW of Chiriaco, 725 m, Provincia Bagua, Departamento Amazonas, Peru, documents the occurrence of E. ockendeni in the Rio Maranon Valley in the Andes of northern Peru (Fig. 26). Remarks.—The specimen (KU 196470) from the Rio Maranon Valley is a female having a SVL of 26.0 mm. Eleutherodactylus pecki Duellman and Lynch Eleutherodactylus pecki Duellman and Lynch, 1988:135. Holotype: KU 147040, adult male, from the Rio Piuntza, 1550 m, Cordillera del Condor, Provincia Morona-Santiago, Ecuador. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum smooth with small tubercles on flanks, posterior part of back, and upper surfaces of limbs, that on venter areolate; discoidal fold evident; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus prominent, its length about 32% length of eye; (3) snout short, subacuminate in dorsal view, rounded in profile; canthus rostralis rounded; (4) upper eyelid bearing tubercles, equal in width to IOD; cranial crests absent; (5) vomerine odontophores small, oval; (6) males having vocal slits and white, nonspinous nuptial pads; (7) Finger I shorter than II; discs moderately large, round; (8) fingers having weak lateral fringes; (9) ulnar tubercles absent; (10) heel bearing small tubercle; outer edge of tarsus lacking tubercles; in- ner edge of tarsus bearing elongate tubercle; (11) inner metatarsal tubercle oval, 3-4x round outer metatarsal tu- bercle; plantar supernumerary tubercles numerous; (12) toes bearing lateral fringes; webbing absent; Toe V longer than III; discs smaller than those on fingers; (13) dorsum brown, darkest laterally; venter cream; (14) SVL in males 15.3-18.7 mm, in female 25.2 mm. As noted by Duellman and Lynch (1988), Eleutherodactylus pecki is most similar to E. incomptus and the lowland E. martiae. It differs from E. incomptus by hav- ing a tubercle on the heel and tubercles on the upper eye- 58 ScrENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS lid, and it differs from E. martiae by being larger (males of E. pecki being as large as females of E. martiae) and in hav- ing a prominent tympanum. Of the other species in north- ern Peru having tubercles on the upper eyelids and small tubercles on the heels, E. bromeliaceus, cryptomelas, lirellus, nephophilus, and rhodoplichus differ from E. pecki in having the discs on the outer fingers being elliptical (much broader than long) instead of nearly rounded. Rounded terminal discs also are present in E. colodactylus and E. schultei, both of which have tubercles on upper eyelids and small tu- bercles on the heels. Eleitherodactylus colodactylus differs by lacking a tympanum, vocal slits, and nuptial pads. Eleutherodactylus schultei differs by being much larger (males > 25 mm; females > 28 mm), by having the snout inclined posteroventrally in profile (rounded in E. peck), and in coloration. Description.—The original description by Duellman and Lynch (1988) is expanded by the addition of a female (USNM 525476) having a SVL of 25.2 mm. Comparison of this specimen with the holotype and two paratypes (KU 147041—42) reveals that in the female the tubercles on the eyelids of smaller and lower than those in the type series; furthermore, the tubercles on the heels are barely discern- ible. The coloration of the female resembles the males in the type series. The dorsum is tan with three diffuse longi- tudinal, brown streaks and diffuse dark brown dorsolat- eral stripes. The flanks and hidden surfaces of the thighs are dark brown, and the venter is heavily flecked with brown. A dark brown canthal stripe and two brown labial bars are present, but an interorbital bar is absent. Distribution and habitat.—This species is known from elevations of 1700 m in the Cordillera de Cutucu, 1550 m on the western slopes of the Cordillera del Condor, and 1138 m on the eastern slopes of the Cordillera del Condor (Fig. 26). The last locality is the only record for the species in Peru; the specimen was ona leaf 2 m above the ground at night in lower montane rainforest. Remarks.—Duellman and Lynch (1988) suggested that Eleutherodactylus frater, incomptus, and pecki were allopat- ric variants of E. ockendeni in the lowlands of the upper Amazon Basin. The latter is known from elevations as high as 1150 m in the Cordillera del Dué in Ecuador, 1280 m in the Serrania de Sira in Peru (Lynch and Duellman, 1980), and 900 m on the western slope of the Cordillera del Condor in Ecuador (Almendariz, 1997). Likewise, Flores and Vigle (1994) considered E. librarius, which is sympat- ric with E. ockendeni in the upper Amazon Basin in Ecua- dor, to be closely related to E. ockendent. Eleutherodactylus percnopterus new species Holotype.—KU 217318, adult female, from Santa Rosa de la Yunga (06°05’ S, 78°43’ W, 1300 m), Provincia Jaén, Departamento Cajamarca, Peru, obtained on 15 July 1989 by Weyder Razzetto and Walter A. Alarcon. Paratypes.—KU 196505 and LSU 32462-63, adult males, from 33 km [by road] SE Ingenio, 1830 m, Provincia Chachapoyas, Departamento Amazonas, Peru. Referred specimens.—KU 196506, subadult male, from 20 km [by road] SW Chiriaco, Provincia Bagua, Departamento Amazonas, Peru; KU 209472, subadult fe- male, from pass at 2400 m, 5 km [by road] NW Mendoza, Provincia Rodriguez de Mendoza, Departamento Amazonas, Peru. USNM 525443, juvenile, from the upper Rio Comainas at base of Cerro Machinaza, 1750 m, Provincia Condoreanqui, Depart-amento Amazonas, Peru; USNM 525445-46, 525449-63 from Alfonso Ugarte, 1138 m, Provincia Condorcanqui, Departamento Amazonas, Peru. Diagnosis.—A member of the Eleuwtherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum with few small tubercles, that on venter weakly areolate; discoidal fold prominent; dorsolateral folds ab- sent; (2) tympanic membrane present; tympanic annulus distinct, round, its diameter about 40% length of eye; (3) snout moderately long, subacuminate in dorsal view, rounded in profile; (4) upper eyelid narrower than IOD, with or without low tubercles; cranial crests absent; (5) vomerine odontophores absent; (6) males having vocal slits, lacking nuptial pads; (7) Finger I shorter than II; discs on outer fingers nearly truncate, more than twice width of digit proximal to pad; (8) fingers lacking distinct lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lack- ing tubercles; (11) inner metatarsal tubercle, elevated, el- liptical, about 5x subconical outer metatarsal tubercle; su- pernumerary plantar tubercles low, rounded; (12) toes lack- ing lateral fringes; webbing absent; Toe V much longer than ILI; discs nearly as large as those on outer fingers; (13) dor- sum pale brown with dark brown marks in scapular re- gion; venter tan with fine dark brown flecks; posterior sur- faces of thighs tan; (14) SVL in three adult males 21.5—23.2 (X = 22.5) mm, in one adult female 25.9 mm. Eleutherodactylus percnopterus is most easily confused with E. incomptus, which is similar in size and also lacks vomerine odontophores (except in large females); the lat- ter differs from E. percnopterus by having the snout rounded in dorsal view, discs on fingers rounded, digits bearing narrow lateral fringes, inner edge of tarsus bearing one or two tubercles, and males having nuptial pads. Further- more, in E. incomptus, dorsal markings are more extensive and contrasting with the ground color, and the venter is brown. The only other member of the Eleutherodactylus unistrigatus Group in the Andes of northern Peru that lacks vomerine odontophores is E. anemerus, a frog that lacks ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 59 Fig. 29. Dorsal and lateral views of the head of Eleutherodactylus percnopterus, KU 217318. Scale bar = 5 mm. dorsal markings, has an orange-red dorsum, yellow flanks, and a prominent tubercles on the snout. The presence of scapular marks on an otherwise plain pale brown dorsum is shared with the primarily Amazonian E. ockendeni, which has vomerine odontophores and lacks a dark canthal stripe. In its absence of distinctive features, E. percnopterus is like E. pecki, which differs from the former by having vomer- ine odontophores, a small tubercle on the heel and a dark venter. Three other species in the region that lack vomer- ine odontophores (E. atrabracus, melanogaster, and pataikos) are robust-bodied members of the E. orestes group that have small terminal discs on the fingers, short hind limbs, and Toe V only slightly longer than Toe III. Description.—(n = 3 males, 1 female; proportions are for males, followed by those of the female). Head as wide as body; HW 35.8-36.3 (X= 36.1), 39.4% SVL; HL 35.8-37.6 (x = 37.0), 38.6% SVL; snout moderately long, subacuminate in dorsal view, rounded in profile (Fig. 29); E-N 70.1-100.0 (x = 89.6), 90.0% length of eye; nostrils barely protuberant, directed laterally; canthus rostralis rounded, curved; loreal region concave; lips slightly flared; upper eyelid with low tubercles in males; upper eyelid width 64.2-87.5 (x = 73.9), 69.4% IOD; cranial crests ab- sent; supratympanic fold weak, barely obscuring upper edge of tympanic annulus; side of head slightly inclined ventrolaterally; tympanic annulus thin; tympanic mem- brane not pustular or thickened; tympanic annulus round; diameter of tympanic annulus 37.0-44.0 (x = 41.0), 46.6% length of eye; postrictal tubercles low, diffuse. Choanae moderately large, ovoid, not concealed by palatal shelf of maxillary arch; vomerine odontophores absent; tongue longer than wide, its posterior border distinctly notched, posterior half not adherent to floor of mouth. Males with vocal slits extending posterolaterally from midlateral base of tongue; vocal sac single, median, subgular; nuptial pads absent. Dorsum of head and body smooth with scattered, small, subconical tubercles; dorsal surfaces of limbs with Fig. 30. Dorsal color pattern of Eleutherodactylus percnopterus, KU 217318. SVL = 25.9 mm. low tubercles; dorsolateral folds absent; flanks smooth; cloacal sheath and enlarged tubercles in cloacal region absent; skin on throat weakly areolate, on belly and ven- tral surfaces of thighs coarsely areolate; discoidal fold prominent. Ulnar tubercles absent; thenar tubercle broadly ovoid, elevated more than ¥% size of bifurcate palmar tu- bercle; supernumerary palmar tubercles absent; subarticular tubercles prominent, subconical; fingers lack- ing distinct lateral fringes; Finger I shorter than II; disc on thumb barely expanded; discs on fingers broadly ellipti- cal, nearly twice width of digit proximal to disc; all fingers having ventral pads defined by circumferential grooves. Heel and tarsus lacking tubercles; inner metatarsal tubercle elevated, elliptical, about 5x subconical outer metatarsal tubercle; supernumerary plantar tubercles low, rounded; subarticular tubercles subconical; toes lacking lateral fringes; webbing absent; discs on toes nearly as large as those on fingers; tip of Toe V extending to distal edge of distal subarticular tubercle on Toe IV; tip of Toe III extend- ing to distal edge of penultimate subarticular tubercle on Toe IV; when hind limbs flexed perpendicular to axis of body, heels barely overlapping; shank 55.3-56.8 (x = 55.9), 54.4% SVL. Coloration in preservative: Dorsum pale brown with narrow, dark brown canthal and supratympanic stripes, faint brown labial bars and interorbital bar, faint brown diagonal bars on limbs, and pair of small black spots or short streaks in scapular region (Fig. 30); flanks, groin, and hidden surfaces of thighs uniform pale brown; venter creamy tan with minute dark flecks on throat and belly. 60 < 1000 m 1000-2000 m 2000-3000 m 3000-4000 m “a >4000m Kilometers @E. percnopterus OE. petrobardus BE. phoxocephalus DE. proserpens “|AE. quaquaversus AE. rhodoplichus Fig. 31. Localities of known occurrence of six species in the Eleutherodactylus unistrigatus group in southern Ecuador and northern Peru. Coloration in life: KU 217318: Dorsum pale grayish brown with reddish tint on head and small black spots; ven- ter whitish gray (Rainer Schulte field notes, 15 July 1989). Measurements of holotype: SVL 25.9, tibia length 14.1, foot length 12.7, head width 10.2, head length 10.0, IOD 3.6, upper eyelid width 2.5, E-N 2.7, eye 3.0, tympanum 1.9. Distribution and habitat——The species is known from two localities at elevations of 1138 m and 1750 on the east- ern slopes of the Cordillera del Condor, and one locality at 1300 m on the southern edge of the Cordillera del Condor (Fig. 31). The species also is known at elevations of 1830 and 2400 m in the northern part of the Cordillera Central in Peru. The latter is a pass northwest of Mendoza at the headwaters of the Rio Chiriaco; the type locality also is in the Rio Chiriaco Drainage. The locality 20 km SW Chiriaco is at an elevation of less than 1000 m in the arid Rio Maranon Valley. Thus, the known elevational distribution is at least 1400 m and suggests that the species may have a continuous distribution between the Cordillera del Condor and the Cordillera Central. The holotype was in an arbo- real bromeliad by day in semiarid forest. Specimens from the eastern slopes of the Cordillera del Condor were on low vegetation at night in humid montane forest. SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Etymology.—The specific name is the Greek noun perknopteros meaning vulture; the name is in loose refer- ence to Vultur gryphus, the Andean condor, to which the Cordillera del Condor refers. Remarks.—The paratypes were collected by Richard Thomas on 21-22 December 1974. One (KU 196505) was calling. The call consists of 2-5 notes (“pink”) followed or not by an ascending series of 10 or more rachetlike clicks (Richard Thomas field notes, 21 December 1974). The sub- adult male from 20 km SW Chiriaco (KU 196506), has a SVL of 18.5 mm, and a subadult female from 5 km [by road] NW of Mendoza (KU 209472) has a SVL of 21.6 mm. Both have color patterns like those of the type series. Of the specimens from eastern slopes of the Cordillera del Condor, five subadult females have SVLs of 18.7—21.0 (x= 20.1) mm, and 13 juveniles having SVLs of 10.7-16.0 (x = 14.6) mm. The juveniles tend to be more boldly patterned than the adults. Eleutherodactylus petrobardus Duellman Eleutherodactylus petrobardus Duellman, 1991a:6. Holotype: KU 212292, adult male, from approximately 2 km [by road] W Huambos, 2500 m, Provincia Chota, Departamento Cajamarca, Peru. Diagnosis.—A member of the Elewtherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum, flanks, and limbs with small, irregularly arranged pustules, that on venter areolate; discoidal fold evident; dorsolateral folds absent; (2) tympanic membrane and tym- panic annulus distinct, round, its length about 45% length of eye; (3) snout rounded in dorsal view and in profile; canthus rostralis rounded; (4) upper eyelid lacking tu- bercles, slightly narrower than IOD; cranial crests absent; (5) vomerine odontophores oblique, prominent; (6) males having vocal slits but no nuptial pads; (7) Finger I shorter than II; discs broad; (8) fingers bearing lateral fringes; (9) ulnar tubercles low, diffuse; (10) heel bearing small tu- bercles; outer edge of tarsus lacking tubercles; inner edge of tarsus bearing elongate tubercle; (11) inner metatarsal tubercle ovoid, 3-4 conical outer metatarsal tubercle; plan- tar supernumerary tubercles small; (12) toes bearing lat- eral fringes; webbing absent; Toe V much longer than III; discs as large as those on fingers; (13) dorsum tan with irregular brown markings, with or without white spots; venter cream; (14) SVL in males 27.0-30.7 mm, unknown in females. Of the species in the Eleutherodactylus unistrigatus Group in the Andes of northern Peru, Eleutherodactylus petrobardus is most similar to the smaller E. versicolor, a species that differs by lacking vocal slits and a small tubercle on the heel, and by having brown reticulation on the venter, whereas the belly is white with black flecks in E. petrobardus. Also, E. petrobardus has more pustular skin on the dorsum than do other species in the region (Fig. 10). ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 61 Description.—The description by Duellman (1991) is adequate. Distribution and habitat.—This species is known only from the type locality in dry scrub forest on the Pacific slopes of the Cordillera Occidental in Departamento Cajamarca, Peru (Fig. 31). Individuals were in terrestrial bromeliads or calling from rock ledges at night. Eleutherodactylus phoxocephalus Lynch Eleutherodactylus phoxocephalus Lynch, 1979:29. Holotype: KU 142075, adult male, from Pilalé, 2340 m, Provincia Cotopaxi, Ecuador. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum shagreen, that on venter areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic mem- brane and tympanic annulus evident, round, its length no more than % length of eye; (3) snout rounded in dorsal view but with vertical fleshy keel, subacuminate in pro- file; canthus rostralis rounded; (4) upper eyelid lacking tubercles, narrower than IOD; cranial crests absent; (5) vomerine odontophores oblique, prominent; (6) males hav- ing vocal slits and nuptial pads; (7) Finger I shorter than II; discs broad; (8) fingers bearing lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles and folds; (11) inner metatarsal tubercle oval, 4-6x round outer metatarsal tubercle; plantar supernumerary tubercles at bases of Toes II-IV; (12) toes bearing lateral fringes; web- bing absent; Toe V much longer than III; discs as large as those on fingers; (13) dorsum gray to brown with few or no darker markings; venter cream; lower flank, groin, and concealed surfaces of limbs white with or without brown or black reticulations; (14) SVL in males 22.3-29.9 mm, in females 29.6-38.9 mm. Structurally, Eleutherodactylus phoxocephalus is unique in the region by having a vertical keel on the snout, which is difficult to discern in some specimens. However, dark reticulations in the groin and on the posterior surfaces of the thighs distinguish E. phoxocephalus from congeners in the region, except E. rhodoplichus, which differs by having a small tubercle on the heel and by lacking vomerine odontophores. Description.—The original description by Lynch (1979) is complete, and includes color in life of specimens from provincias Cotopaxi and Loja, Ecuador. Coloration of liv- ing individuals from provincias Cotopaxi and Pichincha, Ecuador, were given by Lynch and Duellman (1997) and for two individuals from Departamento Piura, Peru, by Duellman and Wild (1993). Distribution and ecology.—This species occurs in up- per humid montane forest and subparamo at elevations of 1800-3100 m on the Pacific slopes of the Cordillera Oc- cidental in Ecuador and the Andes of southern Ecuador. In Peru, it is known from only two localities—15 km [by road] ENE Canchaque, 1850 m (KU 181271) and El Tambo, 2770 m (MHNSM 15399)—on the western slopes of the Cordillera de Huancabamba, Departamento Piura, and from San Andres de Cutervo, about 1800 m, in the north- ern part of the Cordillera Occidental in Departamento Cajamarca (Fig. 31). In the Cordillera de Huancabamba, one individual was on a mossy cliff at night and another was calling at night from low vegetation. In Ecuador, many individuals have been found in the axils of elephant ear plants and arboreal bromeliads by day. Remarks.—Four specimens (KU 221710-13) were col- lected at San Andres de Cutervo, Provincia Cutervo, Departamento Cajamarca, Peru, by Alfonso Miranda on 25 June 1992 and 1 April 1993. Of these, one female has a SVL of 38.9 mm, and three males have SVLs of 26.6-27.8 (x = 27.2) mm. All specimens have a fleshy vertical keel on the snout; the keel is less pronounced in the female than in the males. The dorsum is brown (female) or tan (males) with dark brown canthal and supratympanic stripes and irregular interorbital bar. The female and one male have numerous small dark brown spots on the dorsum of the head and body anterior to the sacrum; one male has a pair of dark spots in the scapular region, and the other male has many small dark spots in the scapular region and a pair of spots in the sacral region. The venter is cream with minute dark flecks, and the groin and hidden surfaces of the thighs are uniform tan. Structurally, these specimens compare favorably with two individuals (KU 181271 and MHNSM 15399) from the Cordillera de Huancabamba and with several series from the southern part of the range in Ecuador—Provincia Azuay (KU 131281-82), Provincia Canar (KU 142118-31), Provincia Loja (KU 135460-62, 142113-17), and Provincia Zamora-Chinchipe (KU 14210412). However, there are some minor differences in coloration between the Peru- vian and Ecuadorian specimens. One of the specimens from the Cordillera de Huancabamba (MHNSM 15399) and all four from San Andres de Cutervo have minute dark flecks over the entire venter, as opposed to small brown spots or more dense flecking in the Ecuadorian specimens, most all of which have dark mottling in the groin and hid- den surfaces of the thighs. Eleutherodactylus proserpens Lynch Eleutherodactylus proserpens Lynch, 1979:32. Holotype: USNM 198484, adult female, from between Sapote and Suro Rancho, 2622 m, Provincia Morona-Santiago, Ecuador. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum and venter areolate; discoidal fold and dorsolat- eral folds absent; cloacal sheath extending onto posterior surfaces of thighs (2) tympanic membrane and tympanic 62 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS annulus prominent, round, its length no more than length of eye; (3) snout long, subacuminate in dorsal view, rounded in profile; small papilla at tip of snout; (4) upper eyelid bearing low, round tubercles, narrower than IOD; cranial crests absent; (5) vomerine odontophores low, oval; (6) males lacking vocal slits and nuptial pads; (7) Finger I shorter than II; fingers short, broad, with round discs only slightly wider than digit proximal to pad; (8) fingers bear- ing ridgelike lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles; (11) inner metatarsal tu- bercle elliptical, 2x round outer metatarsal tubercle; plan- tar supernumerary tubercles numerous, some nearly as large as subarticular tubercles; (12) toes bearing ridgelike lateral fringes; webbing basal, coalesced with lateral fringes; Toe V much longer than III; discs slightly smaller than those on fingers; (13) dorsum tan to brown with darker brown markings—interorbital bar, supratympanic stripe, limb bars, and X-shaped mark or large, elongate blotch on back—or pale dorsolateral stripe; posterior surfaces of thighs cream to brown; venter pale brown; (14) SVL in males 15.2—21.0 mm, in females 20.2—23.5 mm (Lynch, 1979). In the region under consideration, only Eleuth- erodactylus colodactylus is like E. proserpens in having short, stocky fingers, but E. proserpens differs from E. colodactylus by having a cloacal sheath, tympanic membrane and an- nulus, a tubercle on the tip of the snout, and prominent vomerine odontophores. The only other species in the re- gion with a tubercle on the tip of the snout is E. anemerus, which has a unicolor dorsum and lacks a tubercle on the heel and vomerine odontophores. In other Eleutherodactylus in the region having digital pads that are only slightly broader than the digit proximal to the pad (E. atrabracus, melanogaster, pataikos, and pinguis), the fingers are propor- tionately longer and more slender than those in E. colodactylus and E. proserpens; moreover, in those robust- bodied species, Toe V is only slightly longer than Toe III. Description.—The description by Lynch (1979), which includes illustrations of the head and hand, is adequate. Distribution and habitat.—In Ecuador, this species is known from elevations of 1710-2620 m in the southern part of the Cordillera Oriental, from 1700 m in the Cordillera de Cutucu, and from 1550 m on the western slope of the Cordillera del Condor (Duellman and Lynch, 1988; Lynch, 1979). The only Peruvian record is from the upper Rio Comainas at the base of Cerro Machinaza, 1750 m, on the eastern slopes of the Cordillera del Condor, Provincia Cordoncanqui, Departamento Amazonas (Fig. 31). All specimens have been found in bromeliads in humid mon- tane forest. Remarks.—The single Peruvian specimen (USNM 525447) is a juvenile having a SVL of 11.6 mm. Despite its small size, it has the distinguishing characteristics of Eleutherodactylus proserpens—small hands, papilla on snout, and cloacal sheath. In preservative, the dorsum is grayish tan with a large, dark brown, middorsal blotch extending from the anterior edges of the orbits to the sacrum. Dark brown canthal and supratympanic stripes and transverse bars on the limbs are present; labial bars are absent. The venter is heavily flecked with brown. Eleutherodactylus quaquaversus Lynch Eleutherodactylus quaquaversus Lynch, 1974b:9. Holotype: KU 123745, adult female, from south slope Cordillera del Due, above Rio Coca, Provincia Napo, Ecuador. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum shagreen, that on venter areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic mem- brane absent; tympanic annulus absent or evident only ventrally; (3) snout subacuminate in dorsal view, rounded or weakly pointed in profile; (4) upper eyelid bearing coni- cal tubercle, slightly narrower than IOD; cranial crests ab- sent; (5) vomerine odontophores prominent, triangular; (6) males having vocal slits but lacking nuptial pads; (7) Fin- ger I shorter than II; discs broad, elliptical; (8) fingers bear- ing lateral fringes; (9) ulnar tubercles absent; (10) heel bear- ing conical tubercle; outer and inner edges of tarsus bear- ing small tubercles; (11) inner metatarsal tubercle ellipti- cal, 4-5x rounded outer metatarsal tubercle; plantar su- pernumerary tubercles few; (12) toes bearing narrow lat- eral fringes; webbing absent; Toe V much longer than III; discs as large as those on fingers; (13) dorsum pale brown to reddish brown with darker brown interorbital bar, chev- rons or spots, and narrow diagonal bars on limbs; canthal and supratympanic stripes absent; labial bars variably present; posterior surfaces of thighs reddish brown with brown reticulations; venter white, with or without brown spots; (14) SVL in males 19.6-22.5 mm, in females 24.6— 31.3 mm (Lynch and Duellman, 1980). Of the species in the Andes in northern Peru, Eleutherodactylus quaquaversus is like E. galdi in having a conical tubercle on the upper eyelid and a conical tubercle on the heel, but the predominately green E. galdi has a prominent tympanum, cranial crests, a row of tubercles along the outer edge of the tarsus and foot, and larger, trun- cate discs on the fingers. Eleuwtherodactylus quaquaversus is superficially similar to E. ockendeni and E. percnopterus, both of which have well-defined tympana and usually a W- shaped mark in the scapular region (never present in E. quaquaversus), and they lack conical tubercles on the up- per eyelids and heels and lack brown reticulations on the posterior surfaces of the thighs. Description.—The original description by Lynch (1974b) was augmented by Duellman (1978c) and Lynch and Duellman (1980). ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 63 Distribution and habitat.—Although Eleutherodactylus quaquaversus occurs at elevations as low as 200 m in the Amazonian lowlands of Peru (Duellman and Mendelson, 1995), the species is distributed primarily on the eastern slopes of the Andes, where it is known in Ecuador from elevations of 920-1740 in the Cordillera Oriental (Lynch and Duellman, 1980), 1700 m in the Cordillera de Cutucu (Duellman and Lynch, 1988), and 1500-1550 m on the west- ern slopes of the Cordillera del Condor (Almendariz, 1997; Lynch and Duellman, 1980). We now report the species from the eastern slopes of the Cordillera del Condor— upper Rio Comainas, base of Cerro Machinaza, 1750 m, Provincia Condorcanqui, Departamento Amazonas, Peru (Fig. 31). Nearly all individuals have been found on low vegetation at night. Remarks.—In life, juvenile (USNM 525444) with a SVL of 14.8 mm from the upper Rio Comainas was described as: “dorsum tan; brown bar between orbits; lips barred; two spots on scapula; legs barred; chin gray; belly green.” (Robert P. Reynolds field notes, 18 July 1994). Eleutherodactylus rhodoplichus Duellman and Wild Eleutherodactylus rhodoplichus Duellman and Wild, 1993:13. Holotype: KU 219786, adult male, from El Tambo, 2770 m, 31 km [by road] ENE Canchaque, Provincia Huancabamba, Departamento Piura, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum coarsely shagreen, bearing scattered low, round or subconical tubercles, that on venter areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus prominent, round, its length no more than 60% length of eye; (3) snout subacuminate in dorsal view, rounded in profile; canthus rostralis rounded; (4) upper eyelid bearing low, round tu- bercles, narrower than IOD; cranial crests absent; (5) vomerine odontophores absent in males in most females, small and elliptical in other females; (6) males having vo- cal slits but lacking nuptial pads; (7) Finger I shorter than II; discs broad, elliptical; (8) fingers bearing lateral fringes; (9) ulnar tubercles few, low; (10) heel bearing small, subconical tubercles; outer edge of tarsus bearing round, diffuse tubercles; inner edge of tarsus bearing fold; (11) inner metatarsal tubercle elliptical, 3x conical outer meta- tarsal tubercle; plantar supernumerary tubercles numer- ous, low, round; (12) toes bearing lateral fringes; webbing absent; Toe V much longer than III; discs slightly smaller than those on fingers; (13) dorsum brown with fine, irregu- lar darker brown markings; venter cream to tan with dark brown flecks; (14) SVL in males 21.8—28.9 mm, in females 30.1-34.2 mm. Of the other species in the Eleutherodactylus unistrigatus Group in the Andes of northern Peru, only E. anemerus, incomptus, and percnopterus lack vomerine odontophores. Eleutherodactylus anemerus differs by lacking dorsal mark- ings and having a tubercle on the tip of the snout; both E. incomptus and E. percnopterus lack the small tubercle on the heel that is present in E. rhodoplichus (Fig. 10), and E. incomptus also differs by being much smaller and by lack- ing tubercles on the upper eyelid. Description.—The description by Duellman and Wild (1993) is adequate. Distribution and habitat.—This species is known only from elevations of 2770-3050 m in humid montane forest on the western slopes and crest of the Cordillera de Huancabamba (Fig. 31). Individuals were found at night on low vegetation and under rocks by day. Eleutherodactylus rhodostichus new species Holotype.—KU 212264, an adult male, from the west slope of Abra Tangarana, 1080 m, 7 km [by road] NE San Juan de Pacaysapa, Provincia Lamas, Departamento San Martin, Peru, one of a series collected on 5 February 1989 by William E. Duellman and Rainer Schulte. Paratypes.—KU 212265-67 collected with the holo- type. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum finely shagreen with scattered, small tubercles, that on venter areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane evident, and tym- panic annulus distinct, round, its diameter about 40% length of eye; (3) snout long, acuminate in dorsal view, acutely rounded above and inclined posteroventrally in profile; (4) upper eyelid narrower than IOD, with low tu- bercles posterolaterally; cranial crests absent; (5) vomer- ine odontophores low, ovoid; (6) males having vocal slits, lacking nuptial pads; (7) Finger I shorter than II; discs on outer fingers rounded, less than twice width of digit proxi- mal to pad; (8) fingers bearing distinct lateral fringes; (9) ulnar tubercles present; (10) heel lacking tubercles; outer and inner edges of tarsus bearing small tubercles; (11) in- ner metatarsal tubercle elliptical, about 4 subconical outer metatarsal tubercle; supernumerary plantar tubercles nu- merous; (12) toes bearing narrow lateral fringes; webbing absent; Yoe V much longer than III; discs nearly as large as those on outer fingers; (13) dorsum tan with brown mark- ings; venter tan with fine dark brown flecks; posterior sur- faces of thighs tan; (14) SVL in an adult male 19.3 mm and a subadult female 19.7 mm. By having a snout that is acuminate in dorsal view and inclined posteroventrally in profile, Eleutherodactylus rhodostichus differs from all other members of the genus in the region, except E. acuminatus and E. schultei, both of which lack a pattern on the dorsum. Furthermore, E. acuminatus differs by lacking a tympanic membrane, and 64 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS E. schultei differs by having a small tubercle on the heel. Red streaks on the dorsum of the body in living individu- als also distinguish E. rhodostichus from other species in the region. Description.—( = 1 male, 1 female; proportions are for the male, followed by those of the female). Head no- ticeably wider than body; HW 38.3, 38.7% SVL ; HL 36.3, 39.6% SVL; snout long, acuminate in dorsal view, acutely rounded above and inclined posteroventrally in profile; E-N 76.0, 79.2% length of eye; nostrils slightly protuber- ant, directed laterally; canthus rostralis weakly angular, nearly straight; loreal region weakly concave sloping abruptly to lips; lips not flared; upper eyelid with indis- tinct tubercles posterolaterally; upper eyelid width 85.0, 66.7% IOD; cranial crests absent; supratympanic fold weak, barely obscuring upper edge of tympanic annulus; side of head slightly inclined ventrolaterally; tympanic annulus thin; tympanic membrane not pustular or thickened; tym- panic annulus round; diameter of tympanic annulus 40.0, 41.7% length of eye; postrictal tubercles small, subconical, posteroventral to tympanic annulus; skin on head smooth. Choanae large, nearly round, not concealed by palatal shelf of maxillary arch; vomerine odontophores, low, oblique, posteromedian to choanae, oval in outline, each about size of choana, separated medially by distance greater than width of odontophore, bearing 1-0, 3-3 teeth; tongue longer than wide, its posterior border shallowly notched, poste- rior half not adherent to floor of mouth. Male with vocal slits extending posterolaterally from midlateral base of tongue; vocal sac single, median, subgular; nuptial pads absent. Dorsum of head, body, and limbs smooth with scat- tered, small, subconical tubercles, more pronounced in fe- male than in male; dorsolateral folds absent; flanks tuber- culate, especially in female; cloacal sheath and enlarged tubercles in cloacal region absent; skin on throat weakly areolate, on belly coarsely areolate; discoidal fold evident posteriorly. Ulnar tubercles few, low, round; thenar tubercle ovoid, about 2 size of weakly bifid palmar tubercle; su- pernumerary palmar tubercles few, minute; subarticular tubercles prominent, subconical; fingers bearing distinct lateral fringes; Finger I shorter than II; disc on thumb barely expanded; disk on Finger II slightly larger; discs on Fin- gers III-IV broadly rounded, nearly twice width of digits; all fingers having ventral pads defined by circumferential grooves. Upper surfaces of hind limbs smooth with scat- tered small tubercles; heel lacking tubercle; outer edge of tarsus bearing three or four subconical tubercles; inner edge of tarsus with two or three low tubercles; inner metatarsal tubercle flat, elliptical, about 4x subconical outer metatar- sal tubercle; supernumerary plantar tubercles low, diffuse; subarticular tubercles round, subconical; toes bearing nar- row lateral fringes; webbing absent; discs on toes slightly smaller than those on fingers; tip of Toe V extending to middle of distal subarticular tubercle on Toe IV; tip of Toe III not extending to that tubercle; when hind limbs flexed perpendicular to axis of body, heels broadly overlapping; shank 53.8, 54.3% SVL. Coloration in preservative: Dorsum tan with sparse brown markings—KU 212264: small spots laterally in scapular and sacral regions, narrow interorbital bar, weakly defined labial bars and transverse bars on hind limbs; KU 212265: short lines laterally in scapular and sacral regions, narrow interorbital bar, diagonal marks on top of snout, weakly defined labial bars and transverse bars on hind limbs. Small dark spots on flanks; posterior surfaces of thighs tan; venter pale tan with minute dark brown flecks laterally on throat and belly. Coloration in life: KU 212264: Dorsum green with red and tan marks; anterior and posterior surfaces of thighs pale green; heels and elbows tan (Fig. 10); vocal sac yel- low; belly cream; ventrals surfaces of limbs pale green; iris reddish brown. Measurements of holotype: SVL 19.3, tibia length 10.4, foot length 9.0, head width 7.4, head length 7.0, IOD 2.0, upper eyelid width 1.7, E-N 1.9, eye 2.5, tympanum 1.0. Distribution and habitat——The species is known only from one locality at 1080 m on the road from Abra Pardo Miguel to Moyobamba on the east slope of the northern part of the Cordillera Central in northern Peru (Fig. 32). Two adults and two juveniles were in terrestrial bromeli- ads in lower humid montane forest by day. Etymology.—The specific name is derived from the Greek rhodon— meaning red and the Greek stichos meaning line; the name is applied in reference to the red linear mark- ings on the dorsum of the body. Remarks.—Two juveniles (KU 212266-67 ) have SVLs of 15.8 and 16.9 mm, respectively. Both are colored like the female paratype, except that both have a middorsal, brown, triangular mark (apex anterior) just anterior to the level of the sacrum. Eleutherodactylus rufioculis new species Holotype.—KU 212313, a subadult female, from the east slope of Abra Pardo de Miguel, 2180 m, Provincia Rioja, Departamento San Martin, Peru, obtained on 31 January 1988 by Michael E. Morrison. Paratypes.—KU 212312, collected with the holotype. Referred specimens.—USNM 525465, subadult fe- male, USNM 525464, 525467-68, and 525473 from the up- per Rio Comainas, base of Cerro Machinaza, 1750 m, Provincia Condorcanqui, Departamento Amazonas, Peru; USNM 525474-75 from Alfonso Ugarte, 1138 m, upper Rio Comainas, Provincia Condorcanqui, Departamento Amazonas, Peru. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 65 < 1000 m 1000-2000 m 2000-3000 m 3000-4000 m GES > 4000 m 50 —— = Kilometers ©. rhodostichus =. rufioculis ©. schultei ©. seredipitus ). sternothylax ©. versicolor . wilenst Fig. 32. Localities of known occurrence of seven species in the Eleutherodactylus unistrigatus group in the Andes of southern Ecuador and northern Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum smooth, that on venter areolate; discoidal fold present; dorsolateral folds absent; (2) tympanic membrane smooth, and tympanic annulus visible beneath skin, its length about 30% length of eye; (3) snout moderately long, rounded in dorsal view and in profile; (4) upper eyelid lacking tubercles, narrower than IOD; cranial crests absent; (5) vomerine odontophores elongately ovoid; (6) males lacking vocal slits and nuptial pads; (7) Finger I shorter than II; discs on outer fingers expanded, about twice width of digit proximal to pad; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tu- bercles; (11) inner metatarsal tubercle oval, 3x outer meta- tarsal tubercle; supernumerary plantar tubercles minute, diffuse; (12) toes bearing narrow lateral fringes; webbing absent; Toe V longer than III; discs slightly smaller than those on outer fingers; (13) dorsum tan; venter cream, suf- fused with brown; throat brown with cream flecks; groin brown with cream spots; posterior surfaces of thighs brown with cream flecks; (14) SVLin 1 male 18.1 mm, in 1 female 20.6 mm. Eleutherodactylus rufioculis is like E. colodactylus and E. versicolor in lacking vocal slits in males. Of these, E. colodactylus differs by completely lacking a tympanic mem- brane and by having much shorter fingers with round discs, and E. versicolor differs by having a prominent tym- panum, diagonal bars on the flanks, and dark reticulation on the belly. The flanks are dark brown with pale yellow or white spots in only two species in the region—E. rufioculis and E. muscosus, which differs from E. rufioculis by being larger and having tubercles on the upper eyelid and a conical tubercle on the heel. From the sympatric E. nephophilus, which also has a red iris, E. rufioculis can be distinguished by having a smooth dorsum and no differ- entiated tympanum, and no tubercles on the eyelid, heel, or tarsus. Description.—( = 1 male, 1 female; proportions are for the male, followed by those of the female). Head as wide as body; HW 36.5, 37.4% SVL; HL 39.8, 40.1% SVL; snout moderately long, rounded in dorsal view and in pro- file; E-N 135, 115% length of eye; nostrils barely protuber- ant, directed laterally; canthus rostralis rounded, straight; loreal region barely concave; lips rounded; upper eyelid smooth; upper eyelid width 77.7, 70.3% IOD in female; cranial crests absent; supratympanic fold weak, not ob- scuring upper edge of tympanum; tympanic membrane smooth; tympanic annulus distinctly visible beneath skin; length of tympanic annulus 35%, 26% length of eye; postrictal tubercles low, diffuse; side of head nearly verti- cal. Choanae ovoid, not obscured by palatal shelf of max- illary arch; vomerine odontophores posteromedial to choa- nae, elongately ovoid in outline, each more than twice size of choana, separated medially by distance about equal to width of odontophore, each bearing 2-4 teeth in single row; tongue more than twice as long as wide, its posterior bor- der barely notched; posterior half not adherent to floor of mouth; vocal slits and vocal sac absent. Dorsum smooth; dorsolateral folds absent; flanks smooth; cloacal sheath and enlarged tubercles in cloacal region absent; skin on throat and belly weakly areolate; discoidal fold weak, evident only posteriorly. Upper sur- faces of arms smooth; ulnar tubercles absent; thenar tu- bercle low, oval, smaller than bifid palmar tubercle, su- pernumerary palmar tubercles minute, diffuse; subarticular tubercles round; fingers lacking lateral fringes; Finger I shorter than II; disc on thumb not enlarged; discs on Fingers II and IV twice width of digits, that on Finger II 1.5x width of digit; all fingers bearing ventral pads de- fined by circumferential grooves; nuptial pads absent in male. Upper surfaces of hind limbs smooth; tubercles on heel and tarsus absent; inner metatarsal tubercle flat, oval, 34x of outer metatarsal tubercle; supernumerary plantar tubercles minute, diffuse; subarticular tubercles round, non conical; toes bearing narrow lateral fringes; webbing ab- sent; discs on toes slightly smaller than those on fingers; tip of Toe V extending to middle of distal subarticular tu- 66 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS bercle on Toe IV; tip of Toe II] not extending to that tu- bercle; when hind limbs flexed perpendicular to axis of body, heels overlapping by about 4% length of shank; shank 56.4, 55.8% SVL. Coloration in preservative: Dorsum reddish brown with faint darker brown interorbital bar, and flecks later- ally in scapular region; indistinct darker brown transverse bars on limbs; flanks pale brown with large cream spots posteriorly; anterior surfaces of thighs creamy tan with vertical brown marks; posterior surfaces of thighs brown with cream flecks; venter creamy tan, heavily suffused with brown flecks. Male with pale tan on snout anterior to in- terorbital bar and on elbows and on heels. Coloration in life: Dorsum olive (with two pairs of dull red spots in female; grayish-white snout and tan elbows and heels in male) (Fig. 10); groin and anterior surfaces and thighs mottled yellow and dull red; venter yellow with brown mottling; iris red (WED field notes, 31 January 1988). Measurements of holotype: SVL 20.6, tibia length 11.5, foot length 9.2, head width 7.7, head length 8.2, IOD 2.7, upper eyelid width 1.9, E—-N 2.2, eye 2.6, tympanum 0.7. Distribution and habitat.—The species is known from the type locality at 2180 m on the eastern slopes of the northern part of the Cordillera Central and from 1138 m and 1750 m on the eastern slopes of the Cordillera del Condor (Fig. 32). All individuals were on low vegetation (< 1 m) in humid montane forest at night. Etymology.—The specific name is derived from the Latin rufus, meaning red, and the Latin oculis, meaning eye; the name is used in reference to the red iris. Remarks.—The small series from the Cordillera del Condor consists of one subadult female having a SVL of 20.2 mm and six juveniles having SVLs of 13.4-17.3 (x = 15.1) mm. The color patterns of these specimens are like that of the type series, except that two juveniles have dark W-shaped marks in the scapular region and transverse bars in the sacral region. Another juvenile has dark brown trans- verse marks in the scapular and sacral regions and a broad, pale bar immediately anterior to the interorbital bar. Eleutherodactylus schultei Duellman Eleutherodactylus schultei Duellman, 1990a:348. Holotype: KU 212222, adult male, from 5 km N Levanto, 2850 m, Departamento Amazonas, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum shagreen, bearing low, tubercles on forelimbs and in tympanic region, that on venter areolate; discoidal fold absent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus distinct, round, its length slightly less than 2 length of eye; (3) snout subacuminate in dorsal view, inclined posteroventrally in profile; canthus rostralis rounded; (4) upper eyelid bearing low tubercles, narrower than IOD; cranial crests absent; (5) vomerine odontophores present or absent, oblique; (6) males having vocal slits but lacking nuptial pads; (7) Finger I shorter than II; discs broad, rounded; (8) fingers bearing lateral fringes; (9) ul- nar tubercles present, low; (10) heel and outer edge of tar- sus bearing many, low tubercles; inner edge of tarsus lack- ing fold; (11) inner metatarsal tubercle ovoid, 2x subconical outer metatarsal tubercle; plantar supernumerary tubercles numerous; (12) toes bearing lateral fringes; webbing ab- sent; Toe V much longer than III; discs equal to those on fingers; (13) dorsum tan, reddish brown, or green, bordered or not by narrow dark brown line from snout to point above vent; venter white; (14) SVL in males 23.5—26.6 mm, in fe- males 28.4-34.0 mm. Eleutherodactylus schultei is one of three species in the region in which the snout is acuminate or subacuminate in dorsal view and inclined posteroventrally in profile (Fig. 10). Of these, E. acuminatus, which like E. schultei, lacks a pattern on the dorsum of the body, differs by lacking a tympanic annulus; E. rhodostichus has dark markings on the dorsum of the body and lacks a tubercle on the heel. Description.—The description by Duellman (1990) is adequate. Distribution and habitat.—This species is known from only two localities at elevations of 2400 and 2850 m in hu- mid montane forest in the northern part of the Cordillera Central in Departamento Amazonas, Peru (Fig. 32). The second locality (5 km [by road] NW of Mendoza) is about 42 km [airline] east of the type locality. The frogs were found in large terrestrial and arboreal bromeliads by day. Eleutherodactylus serendipitus new species Holotype.—KU 181279, adult male, from 8 km [by road] NNE Balzapata, 1850 m, Provincia Bongara, Departamento Amazonas, Peru, obtained on 3 March 1979 by William E. Duellman. Paratype.—KU 181280, an adult male, collected with the holotype. Referred specimens.—LSUMZ 39360, subadult fe- male, and LSUMZ 39363 and 39376, juveniles, from 12 km [by trail] E La Peca, 1700 m, western slope of the Cordil- lera Colan, Provincia Bagua, Departamento Amazonas, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum finely tuberculate, that on venter areolate; discoi- dal fold barely evident posteriorly; dorsolateral folds ab- sent; (2) tympanic membrane smooth, and tympanic an- nulus distinct, its length about 40% length of eye; (3) snout moderately long, subacuminate in dorsal view and bluntly round in profile; (4) upper eyelid lacking tubercles, nar- rower than IOD; cranial crests absent; (5) vomerine ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 67 odontophores oblique; (6) males having vocal slits and nuptial pads; (7) Finger I shorter than II; discs on outer fingers expanded, nearly truncate, more than twice width of digit proximal to pad; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tar- sus lacking tubercles; weak inner tarsal fold on distal third of tarsus (11) inner metatarsal tubercle elevated, elongately elliptical, about 8x outer subconical metatarsal tubercle; supernumerary plantar tubercles low, subconical; (12) toes lacking lateral fringes; webbing absent; Toe V much longer than III; discs slightly smaller than those on outer fingers; (13) dorsum tan with diffuse brown markings; venter cream with brown flecks; groin cream; posterior surfaces of thighs brown; (14) SVL in 2 males 20.4-21.2 (x= 20.8). Eleutherodactylus serendipitus has uniformly brown flanks and hidden surfaces of the thighs; three other spe- cies in the Eleutherodactylus unistrigatus group in the Andes of northern Peru are similarly colored. Of these, E. ockendeni and E. pecki differ by having tubercles on the upper eyelid and a tubercle on the heel. Furthermore, in the former the width of the upper eyelid equals the IOD, and in the latter the venter is densely flecked with brown. Eleutherodactylus percnopterus differs by having a more rounded snout in profile and diagonal (instead of transverse) bars on the hind limbs, and by lacking vomerine odontophores. Description.—( = 2 males). Head about as wide as body; HW 34.9-37.3 (x = 36.1)% SVL; HL 37.7-38.7 (x = 38.2)% SVL; snout moderately long, subacuminate in dor- sal view, bluntly rounded in profile; E-N 76.0-78.6 (x = 77.3)% length of eye; nostrils distinctly protuberant, di- rected dorsolaterally; canthus rostralis rounded, straight; loreal region barely concave; lips rounded; upper eyelid smooth; upper eyelid width 70.0—71.4 (X= 70.7)% IOD; cra- nial crests absent; supratympanic fold weak, barely obscur- ing upper edge of tympanum; tympanic membrane smooth; tympanic annulus prominent; length of tympanic annulus 39.3-44.0 (x = 41.7)% length of eye; postrictal tu- bercles low, diffuse; side of head nearly vertical. Choanae ovoid, not obscured by palatal shelf of maxillary arch; vomerine odontophores posteromedial to choanae, small, elliptical, separated medially by distance more than width of odontophore, each bearing two teeth in single row (ab- sent in KU 181280); tongue nearly twice as long as wide, its posterior border not notched; posterior half not adher- ent to floor of mouth; vocal slits and median, subgular vocal sac absent. Dorsum of snout smooth; other dorsal surfaces smooth with scattered, small tubercles, especially prominent on flanks; dorsolateral folds absent; cloacal sheath and en- larged tubercles in cloacal region absent; skin on throat and chest smooth, that on belly areolate; discoidal fold barely evident posteriorly. Upper surfaces of arms smooth with scattered, minute tubercles; ulnar tubercles absent (indistinct ulnar fold in KU 181279); thenar tubercle ovoid, about same size as bifid palmar tubercle, supernumerary palmar tubercles minute; subarticular tubercles round; fin- gers lacking lateral fringes; first finger shorter than sec- ond; disc on thumb barely enlarged; discs on Fingers III and IV more than twice width of digits, that on Finger II 1.5x width of digit; all fingers bearing ventral pads de- fined by circumferential grooves; nuptial pads absent. Upper surfaces of hind limbs smooth; tubercles on heel and outer edge of tarsus absent; low, indistinct inner tar- sal fold on distal end of tarsus; inner metatarsal tubercle elevated, elongately elliptical, 8x subconical outer meta- tarsal tubercle; supernumerary plantar tubercles distinct, in single row on proximal segments; subarticular tubercles round, nonconical; toes lacking lateral fringes; webbing absent; discs on toes slightly smaller than those on fingers; tip of Toe V extending to middle of distal subarticular tu- bercle on Toe IV; tip of Toe III extending to middle of penultimate subarticular tubercle on Toe IV; when hind limbs flexed perpendicular to axis of body, heels overlap- ping by about 4 length of shank; shank 55.2-55.4 (55.3) % WAL. Coloration in preservative: Dorsum tan with diffuse brown marks—broad interorbital bar, canthal stripe, single labial bar, supratympanic stripe, diagonal mark on either side in scapular region (Fig. 10); chevron (KU 181279) or transverse mark (KU 181280) in sacral region, flanks pale brown anteriorly, cream with minute brown flecks poste- riorly; posterior surfaces of thigh pale brown; venter cream with brown flecks. Coloration in life: At night, pale gray; by day, brown with darker brown markings; venter gray and throat dull yellow, both heavily flecked with gray; iris dull bronze with median, horizontal red-brown streak (WED field notes, 3 March 1979). Measurements of holotype: SVL 20.4, tibia length 11.3, foot length 9.5, head width 7.6, head length 7.9, IOD 2.9, upper eyelid width 2.0; E-N 1.9, eye 2.5, tympanum 1.1. Distribution and habitat——The species is known from the type locality at 1850 m on the upper western slopes of the northern part of the Cordillera Central, which is drained by tributaries of the Rio Chiriaco that flows north- ward into the Rio Maranon, and from 1700 m of the west- ern slopes of the Cordillera Colan (Fig. 32). At the type locality, two males were calling at night from leaves of low herbaceous plants in highly disturbed humid upper mon- tane forest. In the Cordillera Colan, a subadult female was on a bush near a stream, and two juveniles were in leaf litter in humid montane forest. Etymology.—The specific name is the Latinized En- glish word seredipitous, meaning characterized by 68 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS seredipity, which comes from the Persian fairy tale, The Three Princes of Serendip, who were known for finding valu- able things that were not sought for. The collection of am- phibians made at the type locality was seredipitous be- cause a stop was made there only after we were unable to proceed on a road closed by an accident. Remarks.—Of the specimens from the Cordillera Colan, the subadult female has a SVL of 21.7 mm, and the juveniles have SVLs of 13.8 and 14.6 mm. The female (LSUMZ 39360) has narrow dorsolateral dark streaks pos- terior to the sacrum and a short, median interorbital mark. The type locality of this species also is the type local- ity of Hyla aperomea Duellman, Phyllomedusa duellmani Cannatella, and Scinax oreites Duellman and Wiens. Eleutherodactylus sternothylax Duellman and Wild Eleutherodactylus sternothylax Duellman and Wild, 1993:17. Holotype: KU 219793, adult male, from 16 km [by road] ENE Canchaque, 1840 m, Provincia Huancabamba, Departamento Piura, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum shagreen, bearing few, low, round tubercles pos- teriorly and laterally, that on venter areolate; discoidal fold evident; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus distinct, round, its length about % length of eye; (3) snout subacuminate in dorsal view, acutely rounded in profile; canthus rostralis acutely rounded; (4) upper eyelid bearing low, round tubercles, narrower than IOD; cranial crests absent; (5) vomerine odontophores prominent, oval; (6) males having vocal slits but lacking nuptial pads; (7) Finger I shorter than II; discs broad, truncate; (8) fingers bearing lateral fringes; (9) ul- nar tubercles absent; (10) heel and tarsus lacking tubercles; inner edge of tarsus lacking fold; (11) inner metatarsal tu- bercle elliptical, 34x round outer metatarsal tubercle; plan- tar supernumerary tubercles diffuse; (12) toes bearing lat- eral fringes; webbing absent; Toe V much longer than II]; discs smaller than those on fingers; (13) dorsum tan with dark brown markings; venter cream with minute brown flecks; (14) SVL in males 18.3-29.1 mm, in females 28.3— 36.7 mm. The presence of an acuminate snout and absence of tu- bercles on the upper eyelids and heels distinguishes Eleutherodactylus sternothylax from all other members of the Eleutherodactylus unistrigatus Group in the region, except E. acuminatus and E. anemerus. The former differs by lack- ing a tympanic membrane and dark markings on the dor- sum and flanks, whereas the latter differs by having a prominent tubercle on the tip of the snout, as well as lack- ing dark markings. Description.—The description by Duellman and Wild (1993) is adequate. Distribution and habitat.—This species is known only from elevations of 1735-1840 m in humid montane forest on the western slopes of the Cordillera de Huancabamba (Fig. 32). Eleutherodactylus versicolor Lynch Eleutherodactylus versicolor Lynch, 1979:45. Holotype: KU 119858, adult male, from 13.5 km [by road] E Loja, 2800 m, Provincia Loja, Ecuador. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum shagreen with fine tubercles, that on venter ar- eolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus prominent, length of tympanum about 40-50% length of eye; (3) snout subacuminate in dorsal view, rounded in profile; canthus rostralis angular; (4) upper eyelid bearing tubercles or not, narrower than IOD; cranial crests absent; (5) vomerine odontophores prominent, oblique; (6) males lacking vocal slits and nuptial pads; (7) Finger I shorter than II; discs broad, elliptical; (8) fingers bearing weak lateral fringes or none; (9) ulnar tubercles low, diffuse; (10) heel lacking tu- bercles; outer edge of tarsus bearing row of small, conical tubercles; inner edge of tarsus bearing low fold distally; (11) inner metatarsal tubercle ovoid, 3-4x conical outer metatarsal tubercle; plantar supernumerary tubercles nu- merous, conical; (12) toes lacking lateral fringes; webbing absent; Toe V much longer than III; discs slightly smaller than those on fingers; (13) dorsum brown with darker brown markings—chevrons, interorbital bar, canthal and supratympanic stripes; flanks usually with vertical bars; venter cream with brown reticulations; (14) SVL in males 19.3—25.2 mm, in females 22.7-29.8 mm (Lynch, 1979). The only other species in the region that can be con- fused with Eleutherodactylus versicolor is E. ardalonychus, which also has diagonal bars on the flanks and brown re- ticulations on the venter, but the reticulations are much finer than those in E. versicolor. Furthermore, E. ardalonychus differs by having a smooth dorsum, no tympanic mem- brane and tympanic annulus visible only ventrally, and rounded, instead of elliptical, discs on the outer fingers. Description.—The description by Lynch (1979) requires slight modification. Specimens herein assigned to this spe- cies are like topotypic material except that tubercles are present on the upper eyelids—1/2 in one specimen, 2/3 in one, and many in four. Distribution and habitat.—This species has been re- ported from an elevation of 3100 m in subparamo north of San Lucas, Provincia Loja, Ecuador and from 2500-2800 m in subparamo and cloud forest at Abra de Zamora, Ec- uador (Lynch, 1979). The specimens reported herein are from somewhat lower elevations on the eastern slopes of the Cordillera del Condor and represent the first records ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 69 of the species from Peru (Fig. 32). The following localities are in humid montane forest in Departamento Amazonas: upper Rio Comainas, base of Cerro Machinaza, 1750 m (USNM 525466, 525477), Alfonso Ugarte, upper Rio Comainas, 1138 m (USNM 525469, 525471—72); Puesto Vigilancia Comainas, upper Rio Comainas, 665 m (USNM 525441). All of the Peruvian specimens were collected at night on vegetation up to 2 m above the ground. Remarks.—Three of the specimens from the Cordillera del Condor are females having SVLs of 26.0, 26.2, and 27.3 mm; two are males having SVLs of 21.8 and 24.7 mm, and one is a juvenile with a SVL of 17.5 mm. Five individuals have dark, irregular, chevron-shaped marks on the dor- sum; one male (USNM 525477) has pale cream dorsolat- eral stripes extending from the posterior edge of the eye- lid to a point above the insertion of the hind limbs. Eleutherodactylus wiensi Duellman and Wild Eleutherodactylus wiensi Duellman and Wild, 1993:22. Holotype: KU 219795, adult male, from 12.7 km [by road] ENE Canchaque, 1600 m, Provincia Huancabamba, Departamento Piura, Peru. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having (1) skin on dorsum smooth with scattered, small, conical tubercles, that on venter areolate; discoidal fold evident; dorsolat- eral folds weak; (2) tympanic membrane absent and tym- panic annulus evident only ventrally, its length about 30% length of eye; (3) snout acutely rounded in dorsal view, rounded in profile; canthus rostralis rounded; (4) upper eyelid bearing tubercles on outer edge or not, narrower than IOD; cranial crests absent; (5) vomerine odontophores prominent, oblique; (6) males having vocal slits but lack- ing nuptial pads; (7) Finger I shorter than II; discs broad, truncate; (8) fingers bearing lateral fringes; (9) ulnar tu- bercles few, diffuse; (10) heel bearing single, conical tu- bercle; outer edge of tarsus bearing few, low, round tu- bercles; inner edge of tarsus bearing low fold distally; (11) inner metatarsal tubercle ovoid, 4x subconical outer meta- tarsal tubercle; plantar supernumerary tubercles minute; (12) toes bearing lateral fringes; webbing basal; Toe V much longer than III; discs equal to or slightly smaller than those on fingers; (13) dorsum grayish tan with irregular brown markings; venter cream with brown spots or reticulations; (14) SVL in males 27.8-33.0 mm, in female 37.0 mm. This is one of four species in the Eleuwtherodactylus unistrigatus Group in the region that lacks a tympanic mem- brane but has a tympanic annulus visible ventrally (Fig. 10). Of these, E. acuminatus, which lacks dorsal markings, and E. ardalonychus, which has dark reticulations on the belly, differ from E. wiensi by having a tubercle on the heel; E. rhodoplichus differs by lacking vomerine odontophores and having tubercles on the upper eyelid. Description.—The description by Duellman and Wild (1993) is adequate. Distribution and habitat.—This species is known from only two localities at elevations of 1600 and 1735 m in hu- mid montane forest on the western slope of the Cordillera de Huancabamba, Departamento Piura, Peru (Fig. 32). In- dividuals were perched on low vegetation at night. BIOGEOGRAPHY Because of the complex topography and diverse envi- ronments in the Andes of northern Peru and southern Ec- uador, together with the intervening Huancabamba De- pression, this region has many endemics, and it is the north- ern or southern limits of distributions of many taxa, such as toads of the Bufo spinulosus and veraguensis groups (Duellman and Schulte, 1992) and lizards of the genus Pholidobolus (Montanucci, 1973; Reeder, 1996). Aside from frogs of the genus Eleutherodactylus, many species of other groups of anurans are known only from this region— centrolenids (Cadle and McDiarmid, 1990; Duellman and Schulte, 1993), Gastrotheca (Duellman, 1987; Duellman and Trueb, 1988; Trueb and Duellman, 1978,), and leptodactylids of the genera Ischnocnema (Duellman, 1990b; Lynch, 1974a), Phrynopus (Cannatella, 1984, Lynch, 1975a), and Phyllonastes (Duellman (1991b; Lynch, 1976), as well as lizards of the genus Stenocercus (Cadle, 1991). The dry valleys even are barriers to the distributions of montane- forest birds (Parker et al., 1985). The following discussion refers to patterns of distri- bution of Eleutherodactylus only in the Andes of northern Peru and southern Ecuador, and in the intervening Huancabamba Depression. Most of the species of Eleutherodactylus in the region inhabit humid montane for- ests, and many are restricted to that habitat or extend up- ward into very humid montane forest (including subparamo and paramo), where species richness is lower (Table 4). With few exceptions, only members of the Eleutherodactylus conspicillatus Group occur in the dry habi- tats in the Huancabamba Depression, where they usually are found along streams. Species in this group are wide- spread geographically and altitudinally in western South America (Lynch and Duellman, 1997). Two members of this group (E. karcharias and E. metabates) are known only from dry environments below 1000 m, and E. lymani in- habits these same environments, as well as montane habi- tats. In the region of the Huancabamba Depression, three members of the group (E. avicuporum, condor, and cuneirostris) are known only from humid montane forest. Three other species in the group (E. citriogaster, lanthanites, and peruvianus) primarily occupy humid tropical forest in the Amazon Basin, but ascend to humid montane forest 70 ScrENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Table 4. Geographic and latitudinal distribution of Eleutherodactylus in the Andes of northern Peru and southern Ecuador. Numbers in columns are elevations in meters. Species followed by an asterisk (*) also occur in the upper Amazon Basin. Cordillera Cordillera Cordillera Cordillera Cordillera Cordillera Cordillera Cordillera Occidental, Oriental de del de Central, Occidental de Huan- Species Ecuador Ecuador Cutucu Condor Colan Peru Peru cabamba Eleutherodactylus conspicillatus Group E. avicuporum — = = = 1700-2030 = a = E. citriogaster = = = == = 600-800 == = E. condor = = 1975 1500-1750 = = — — E. cuneirostris _ — — — 1700 = —= _— E. karcharias — = = = = 1000 — = E. lanthanites* = 1000-1490 = = = 1630 = = E. lymani' 610-3000 _ _ _ — — — 1120-1850 E. metabates* — — — = — —_ = = E. peruvianus* = 1410-1910 1700-1975 665-1750 = = = = E. w-nigrum 800-3200 1100-2220 — — — — — _— Eleutherodactylus nigrovittatus Group E. araiodactylus = = = = = 3370 = = Eleutherodactylus orestes Group E. atrabracus — — — = 2960-3330 = = = E. melanogaster = = — = = 3300-3470 ca a E. orestes — 2720-3120 = = = = oo = E. pataikos — — — _ = 3470 = = E. pinguis — = — _ = = 3050-3760 = E. vidua — 2710-3100 = = = = -= = Eleutherodactylus unistrigatus Group E. acuminatus* — — = 830 = 950 = = E. altamazonicus* — == = 1500-1600 = — — = E. anemerus — — — — _— — — 2770 E. ardalonychus = = — = = 680-1200 = = E, atratus a 2195-2850 = = = = — = E. balionotus — 2800 _— _ — = = = E. baryecuus — 2195-1990 a i — — a — E. bearsei — _ -— — — 500-730 = = E. bromeliaceus — 1710-2620 1700 1500-1600 = 2180 = = E. cajamarcensis 2660-3000 = = = = = 2200 3050-3100 E. ceuthospilus — — — — = = 1500 1735-2870 E. colodactylus — 2195-3140 = = = = = 2745-3110 E. cryophilus 3080-3200 2835-3355 = = = = = = E. cryptomelas — 2470-3100 = — — — — 2770-2820 E. exoristus — — = 665-1550 a = = = E. galdi — 1000-1740 1700-1975 1500-1550 1700 = = — E. ganonotus — = 1700 = — = = = E. incomptus = 1270-1910 = 1300 = = = = E. infraguttatus — — _ _ = 2000-2180 = = E. lirellus -~ — -- — — 470-1200 —_ — E. muscosus — — _ — — 1800 = = E. nephophilus _ — a _— — 1080-2180 = = E. nigrogriseus = 1180-2835 1700 1150 = = = = E. ockendeni* — 1000-1100 — 900 = = = = E. pecki — a 1700 1138-1550 = an = == E. percnopterus = — _ 1138-1750 = 1830 — = E. percultus — 2850 — = = = = = E. percnopterus —_ _— _— 1138-1750 = 1830 — a E. petrobardus — _ _ — — —_ 2500 _ E. phoxocephalus 1800-3100 = — — _ _ 1800 1850-2770 E. prolatus —_ 1140-1490 1700 = — = = = E. proserpens = 1710-2620 1700 1550 = = = = E. pycnodermis = 2650-3000 = _ — = _ = E. quaquaversus — 920-1740 1700 1500-1550 — = = = E. rhodoplichus _— — — — — — _ 2770-3050 E. rhodostichus — _ — — — 1080 = — ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU Tf Table 4 continued . versicolor — 2500-3100 — . wiensi — — —_— Cordillera Cordillera Cordillera Occidental, Oriental de Species Ecuador Ecuador Cutucu E. riveti 2620-3420 2720-3280 — E. ruidus 2317 oe — E. rufioculis _ — = E. schultei — — — E. serendipitus — — E. spinosus — 1700-2835 — E. sternothylax — — - E. trachyblepharis — 950-1250 — E. ventrimarmoratus* _ 1000-1740 1700 E iE Cordillera Cordillera Cordillera Cordillera Cordillera del de Central, Occidental de Huan- Condor Colan Peru Peru cabamba 1138-1750 — 2180 — —_ = — 2400-2850 _— = —_— 1700 1850 = = 1550 — — —_— — -- — —_ —_ 1735-1840 900-1600. —_ — _— — 665-1750 — — — a — — — = 1600-1735 ‘Also at elevations of 500-1000 m in the Rio Maranon Valley Only at elevation of 525 m in the Rio Maranon Valley *Also at elevation of 725 m in the Rio Maranon Valley on the eastern slopes of the Andes. Most species in this group are terrestrial and capable of lengthy leaps, as evi- denced by their long hind limbs. The Eleutherodactylus nigrovittatus Group consists of five species, one of which (E. nigrovittatus) has a broad dis- tribution in the upper Amazon Basin in Ecuador and north- ern Peru. The other four species inhabit montane forests and subparamo in the Andes of Colombia, Ecuador, and northern Peru. Species in this group are small, terrestrial frogs that hop about in leaf litter. According to Lynch et al. (1997), the lowland E. nigrovittatus is the sister group to E. latens + E. manipus in the Colombian Andes, and these three species are the sister group to E. elassodiscus in the Ecua- dorian Andes; they postulated dispersal of E. nigrovittatus as a means of explaining the geographical and cladistic relationships. The seven species making up the Eleutherodactylus orestes Group are small, terrestrial frogs that make short hops; this mode of locomotion is correlated with the short hind limbs. Members of this group primarily have allo- patric distributions in humid or very humid montane for- ests or subparamo and paramo. The distributions of E. orestes and E. vidua are mostly allopatric, but the two spe- cies occur sympatrically in the southern part of their ranges; E. melanogaster and E. pataikos occur sympatrically in the Cordillera Central in Peru. Members of the large Eleutherodactylus unistrigatus Group reach their greatest diversification in humid mon- tane forests, but many species occur in humid tropical for- ests, and a few extend upward above treeline. Although some species are encountered on the ground by day, these inhabitants of forests generally are arboreal, or at least perch on leaves and limbs of bushes at night. Only a few clades have been identified within this large group. Of the species known from northern Ecuador, only E. galdi has been placed in a clade (Lynch, 1996; Lynch and Rueda- Almonacid, 1997); the four species in this clade have allo- patric distributions in the Andes from northern Colombia to northern Peru. Because phylogenetic relationships have not been as- certained among the Eleutherodactylus in the Andes of northern Peru and their extralimital congeners, only a de- scriptive analysis of their distributions is possible. Even this is hampered by the absence of collections from many areas. Nonetheless, some patterns seem to be apparent. More species (26) inhabit the southern part of the Cordil- lera Oriental in Ecuador than any other mountain system under consideration here’; this is followed by the Cordil- lera del Condor (19) and the northern part of the Cordil- lera Central in Peru (16). An intermediate number (11) is known from the Cordillera de Cutucti and Cordillera de Huancabamba, whereas many fewer species are known from the southern part of the Cordillera Occidental in Ec- uador (7) and the Cordillera Occidental in Peru (5). These low numbers are indicative of the scarcity of suitable habi- tat for most species of Eleutherodactylus. The relatively low number of species recorded from the Cordillera de Cutuct and Cordillera Colan probably reflect the fact that no her- petologists have collected in those cordilleras; in fact, our existing knowledge of the anuran faunas in those cordille- ras comes from collections made by ornithologists. Endemism in the various cordilleras varies from 0% in the Cordillera del Condor to 75% in the Cordillera Cen- tral in northern Peru (Table 5). Endemism is highest in species that are distributed at elevations above 2000 m. ‘Seventy-two species are known from the Cordillera Occidental in Co- lombia (J. D. Lynch, pers. comm.). 72 Table 5. Numerical comparisons of Eleutherodactylus in the Andes of southern Ecuador and northern Peru. Number Number Percent Region Species Endemic Endemic Cordillera Occidental, Ecuador 7 | 14 Cordillera Oriental, Ecuador 26 4 27 Cordillera de Cutucu 11 1 9 Cordillera del Condor 19 0 0 Cordillera Colan 5 3 60 Cordillera Central, Peru 16 12 75 Cordillera Occidental, Peru 5 2 40 Cordillera de Huancabamba 1] 4 36 The apparent absence of endemism in the Cordillera del Condor probably is related to the absence of collections from high elevations in that cordillera. On the other hand, the apparent high endemism in the Cordillera Central in Peru is owing to several species that are known only from the eastern slopes of that cordillera; once the Cordillera Colan becomes better known, it is assumed that several species now known in the Cordillera Central will show up in the Cordillera Colan. Each of the eight cordilleras shares species with at least two other cordilleras (Fig. 33). Generally, the degree of commonality of species is related to distance, as well as elevations of intermediate regions. The two cordilleras just east of the Cordillera Oriental in Ecuador, the Cordillera de Cutucu and Cordillera del Condor share more species than do any other cordilleras. However, some of these spe- cies, as well as those shared with the Cordillera Central in Peru (e.g., Eleutherodactylus acuminatus, lanthanttes, peruvianus, and ventrimarmoratus), are primarily lowland species that ascend the slopes of the Andes. However, four species (E. bromeliaceus, galdi, nigrogriseus, and proserpens) that are shared by the Cordillera Oriental in Ecuador, Cor- dillera de Cutucu, and Cordillera del Condor are not known to occur at elevations of less than 1000 m. Occupation of more than one cordillera is less com- mon in the west, where drier environments occur at the lower elevations. However, Eleutherodactylus cajamarcensis occurs in three cordilleras and is unknown at elevations of less than 2200 m. Assuming that each species has changed little, if any, physiologically and ecologically during its existence, interpretation of present patterns of distribution ScrenTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Cordillera i i de Cordillera Cordillera ; Occidental, Oriental, Cutucu Ecuador Ecuador 11 U Cordillera del Condor 8) Cordillera Cordillera de Huan- Colan cabamba 5 11 Cordillera Occidental, Peru Cordillera Central, 5 Peru 16 Fig. 33. Diagrammatic representation of eight cordilleras in southern Ecuador and northern Peru. Numbers in boxes are the number of species of Eleutherodactylus in each cordillera; numbers in circles indicate number of species shared by cordilleras at the ends of arrows. must invoke past climatic change to account for present disjunctions. If the predictions of Pleistocene climatic de- pression in the Andes by Dollfus (1976), Hastenrath (1967), and Sauer (1971) are anywhere near correct, regions of humid montane forest now isolated on the various cordil- leras would have been connected, thereby permitting dis- persal among the cordilleras; subsequent elevation (in in- terglacial phases) would have resulted in isolation of popu- lations and possibly speciation. This climatic-speciation model championed by Simpson (1979) was refined in theory by Lynch and Duellman (1997) for Eleutherodactylus in western Ecuador. Cladograms of various lineages of Eleutherodactylus in the Andes of Colombia follow this pat- tern of allopatric speciation within the same elevational limits (Lynch et al., 1997). LITERATURE CITED Almendariz, A. 1997. Amphibian and reptile species recorded in the north- ern and western Cordillera del Condor. Pp. 199-201 in T. S. Schulenberg and K. Awbrey (eds.), The Cordillera del Condor Re- gion of Ecuador and Peru: a biological assessment. RAP Working Papers No. 7. Washington, D. C.: Conservation International. Andersson, L. G. 1945. Batrachians from east Ecuador collected 1937. 1938 by Wm. Clarke-Macintyre and Rolf Blomberg. Arkiv for Zoologi, 37A (2):1-88. , Aubodin, J., A. V. Borrello, G. Cecioni, R. Charrier, P. Chotin, J. Frutos, R. Thiele, and J.-C. Vicente. 1973. Esquisse paléogéographique et structurale des Andes Méridionales. Revue de Geographie Physique et de Geologie Dynamamique 15:11-—72. Bokermann, W. C. A. 1958. A preoccupied name of a neotropical frog, genus Eleutherodactylus. Herpetologica 14:95. Boulenger, G. A. 1908. Descriptions of new batrachians and reptiles dis- covered by Mr. M. G. Palmer in south-western Colombia. Annals & ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU 73 Magazine of Natural History, Ser, 8, 12:515—-522. Boulenger, G. A. 1912. 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Copeia 1978:498-503. van der Hammen, T. 1974. The Pleistocene changes in vegetation and climate in tropical South America. Journal of Biogeography 1:3-26. van der Hammen, T., and A. M. Cleef. 1986. Development of the high Andean paramo flora and vegetion. Pp. 153-201 in F. Vuilleumier and M. Monasterio (eds.), High Altitude Tropical Biogeography. New York, London: Oxford Univ. Press. Zeil, W. 1979. The Andes: A Geological Review. Berlin: Gebriider Borntraeger. ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU v5) APPENDIX 1 SPECIMENS EXAMINED The specimens from the Andes of northern Peru and comparative material from elsewhere in Peru and from Ecuador are documented below, alphabetically by species. Localities are arranged alphabetically within departamentos or provinces, which also are arranged al- phabetically within countries. Within a locality, specimens are listed chronologically by catalogue number following museum abbreviations, which are arranged alphabetically. Eleutherodactylus acuminatus (17 specimens) Ecuapor: Prov. Morona-Santiago: San José, 830 m, KU 146976. Prov. Napo: Limoncocha, KU 183523; Rio Yasuni, KU 175105. Prov. Sucumbios: Puerto Libre, 570 m, KU 123255-60; 15 km ENE Umbaqui, 720 m, KU 123254. Prov. Pastaza: Canelos, 530 m, KU 119473. Peru: Depto. Loreto: San Jacinto, 175-190 m, KU 221995. Depto. San Martin: Rio Cainarachi, 33 km NE Tarapoto 330 m, KU 209466; Rio Shilcayo, near Tarapoto, 500 m, KU 209467; 15.4 km SW Zapatero, 950 m, KU 217308-10. Eleutherodactylus anemerus (1 specimen) Peru: Depto. Piura: El Tambo, 31 km [by road] ENE Canchaque, 2770 m, KU 219789 (holotype). Eleutherodactylus araiodactylus (1 specimen) Peru: Depto. Amazonas: 24 km [by road] SW Leimebamba, 3370 m, UF 40764 (holotype). Eleutherodactylus ardalonychus (4 specimens) Peru: Depto. San Martin: Abra Tangarana, 7 km NE San Juan de Pacaysapa, 1080 m, KU 212299; Rio Cerranayacu, 76 km [by road] NW Rioja, 1200 m, KU 212301 (holotype), 212310; 8 km [by road] NE Tarapoto, 680 m, KU 212300. Eleutherodactylus atrabracus (2 specimens) Peru: Depto. Amazonas: Cordillera Colan, E La Peca, 2963-3330 m, LSU 49144 (holotype), 45090. Eleutherodactylus avicuporum (13 specimens) Peru: Depto. Amazonas: 12 km [by trail] E La Peca, 1700 m, KU 288628, LSUMZ 39359, 39361, 39365 (holotype), 39366-68, 39370-72, 39374-75, 45089. Eleutherodactylus bearsei (11 specimens) Peru: Depto. San Martin: Cataratas Ahuashiyacu, 14 km [by road] NE Tarapoto, 730 m, KU 212268 (holotype), 212269-74, 217314-15; 30 km [by road] SW Zapatero, 500 m, KU 212275-76. Eleutherodactylus bromeliaceus (9 specimens) Ecuapor: Prov. Morona-Santiago: 8.8 km [by road] WSW Plan de Milagro, 2370 m, KU 202316-17; Rio Piuntza, Cordillera del Condor, 1830 m, KU 146974; between Logrono and Yaupi, 1700, ANSP 29254, 29272, 29278. Prov. Zamora-Chinchipe: 22 km [by road] W Zamora, 1730 m, KU 141772. Peru: Depto. San Martin: East slope Abra Pardo de Miguel, 2180 m, KU 212213-14. Eleutherodactylus cajamarcensis (147 specimens) Ecuapor: Prov. Azuay: Luz Maria, 1880 m, KU 221706-09. Prov. Loja: 5 km NE Cariamanga, 1870 m, KU 141896~-909; 12 km [by road] NE Catacocha, 2060 m, KU 141890-95; 6.8 km [by road] E Loja, 2640 m, KU 21785152; 9 km [by road] E Loja, 2660 m, KU 119949-50; 12 km [by road] S Loja, KU 2250 m, KU 165190-92; 6 km [by road] N San Lucas, 2850- 2900 m, KU 16519399, 120007-15; 7-8 km [by road] N San Lucas, 2940- 3000 m, KU 119951-120006; 13 km [by road] E Veracruz, 2250 m, KU 141860-89. Peru: Depto. Cajamarca: San Andres de Cutervo, KU 221706-09. Depto. Piura: between Canchaque and Huancabamba, 3100 m, KU 135495, 135502; 25.5 km [by road] SW Huancabamba, 3010 m, KU 181250-61; 26 km SW [by road] Huancabamba, 3050 m, KU 196508; 29.3 km [by road] SW Huancabamba, 3100 m, KU 18124449. Eleutherodactylus ceuthospilus (53 specimens) Peru: Depto. Cajamarca: 12 km [by road] W Lamas, 1500 m, KU 212215-18. Piura: 15-16 km [by road] ENE Canchaque, 1735-1840 m, KU 181270, 181272-78, 196492-98, 219775 (holotype), 219776-85, LSUMZ 32321-31, MHNSM 15387-97; 29 mi (46 km) [by road] E Canchaque, 9200 ft (2840 m), UF 34103. Eleutherodactylus citriogaster (22 specimens) Peru: Depto. San Martin: Cataratas Ahuashiyacu, 14 km [by road] NE Tarapoto, 730 m, KU 212277 (holotype), 212278-85, 217316, 217339-— 42, MHNSM 6080-82; Rio Cainarachi, 33 km [by road] NE Tarapoto, 330 m, KU 209384; 8 km [by road] NE Tarapoto, 680 m, KU 212286; 18 km [by road] NE Tarapoto, KU 209483; 28 km [by road] NE Tarapoto, KU 212289-90. Eleutherodactylus colodactylus (131) Ecuapor: Prov. Loja: 14 km [by road] E Loja, 2770 m, KU 142160-61; 15 km [by road] E Loja, 2710 m, KU 142162-64. Prov. Zamora-Chichipe: Abra de Zamora, 2800 m, KU 142151—54, 142156—-59, 165219-21. Peru: Depto. Piura: 23 mi (37 km) [by road] E Canchaque, 9300 ft (2870 m), UF 52077-89, 52091-138, 112513-33; 31 km [by road]SW Huancabamba, 3080 m, KU 181262-64; 33 km [by road] SW Huancabamba, 3050 m, KU 196443-46; Summit of Cordillera de Huancabamba (on road between Canchaque and Huancabamba), 3100 m, KU 135494, 135496-501. Eleutherodactylus condor (48 specimens) Ecuapor: Prov. Morona-Santiago: “Camp 3,” Cordillera de Cutucu, 1975 m, ANSP 29235, 29255—56, 29258; Rio Piuntza, Cordillera del Condor, KU 146991, 146992 (holotype), 146993-147033. Peru: Depto. Amazonas: upper Rio Comainas, base of Cerra Machinaza, 1750, USNM 525437. Eleutherodactylus crytomelas (6 specimens) Ecuapor: Prov. Loja: 15 km [by road] E Loja, 2710 m, KU 141992-93; 8~9 km [by road] N San Lucas, 3000-3100 m, KU 120995—96. Peru: Depto. Piura: El Tambo, 31 km [by road] ENE Canchaque, 2770 m, KU 181269, MHNSM 15398. Eleutherodactylus cuneirostris (1 specimen) Peru: Depto. Amazonas: 12 km [by trail] E La Peca, 1700 m, LSUMZ 39369 (holotype). Eleutherodactylus exoristus (16 specimens) Ecuapor: Prov. Morona-Santiago: Rio Piuntza, Cordillera del Condor, 1830 m, KU 147047-50, 147051 (holotype), 147052-58. Peru: Depto. Amazonas: Alfonso Ugarte, upper Rio Comainas, 1138 m, USNM 525448, 525462, 525470; Puesto Vigilancia Rio Comainas, 665 m, USNM 525442. Eleutherodactylus galdi (15 specimens) Ecuapor: Prov. Morona-Santiago: “Camp 2,” 1700 m, Cordillera de Cutucu, ANSP 29245, 29247; “Camp 3,” 1975 m, Cordillera de Cutucu, ANSP 29248; Rio Piuntza, Cordillera del Condor, KU 146977-85. Prov. Napo: Rio Azuela, 1740 m, KU 143416, 165422. Peru: Depto. Amazonas: 12 km [by trail] E La Peca, Cordillera Colan, LSUMZ 39362. Eleutherodactylus incomptus (63 specimens) Ecuapor: Prov. Napo: Rio Azuela, 1740 m, KU 143497; Rio Salado, + 1 km upstream from Rio Coca, 1410 m, KU 146169, 165844-58, 177298— 319; 2 km [by road] SSW Rio Reventador, 1490 m, KU 165932; 16.5 km 76 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS [by road] NNE Santa Rosa, 1700 m, KU 143455, 143461—64, 143484 (holo- type), 14348596. Prov. Pastaza: 9.5 km [by road] NW Mera, 1270 m, KU 179000-03. Peru: Depto. Cajamarca: Santa Rosa de la Yunga, KU 21731920. Eleutherodactylus infraguttatus (6 specimens) Peru: Depto. San Martin: East slope Abra Pardo de Miguel, 2180 m, KU 212297 (holotype), 212298, 212314-16; 14 km [by road] W Venceremos, 2000 m, KU 217317. Eleutherodactylus lanthanites (50 specimens) Ecuapbor: Prov. Sucumbios: Santa Cecilia, 340 m, KU 123852-77, 126215-16, 146144 (holotype), 146145-60. Peru: Depto. Loreto: janction Rio Yanomono and Rio Amazonas, KU 220446, 220898; San Jacinto, 175-190 m, KU 222000-01. Depto. San Martin: Venceremos, 1630 m, KU 212225. Eleutherodactylus lirellus (27 specimens) Peru: Depto. San Martin: Abra Tangarana, 7 km [by road] NW San Juan de Pacaysapa, 1080 m, KU 212240 (holotype), 212241, 212251—-59; Ponga de Shilcayo, 4 km [by road] NNW Tarapoto, 470 m, KU 212260- 63; Rio Cerranayacu, 76 km [by road] NW Rioja, 1200 m, KU 212226-32, 212234, 212236-39. Eleutherodactylus lymani (34 specimens) Ecuapor: Prov. Azuay: 55.9 km [by road] E Pasaje, 1000 m, KU 152009. Prov. Loja: Loja, 2150 m, KU 119502; 2 km [by road] E Loja, 2210 m, 3.9 km [by road] E Loja, 2460 m, KU 202418; KU 119504-12; 7 km [by road] E Loja, 2500 m, KU 119503; 9 km [by road] E Loja, KU 202416; 7.6 km [by road] S Loja, 2210 m, KU 141962-64; 9 km [by road] S Loja, 2230 m, KU 165539-40; 12.2 km [by road] S Loja, 2275 m, KU 141292; 14.4 km [by road] S Loja, 2260 m, KU 202417; 17 km [by road] NE Macara, 1240 m, KU 141965. Peru: Depto. Cajamarca: 28 km [by road] N Santa Cruz, 725 m, KU 196464, 196509, LSUMZ 19553. Depto. Piura: Canchaque, 1120 m, MHNSM 11172; 15 km [by road] E Canchaque, 1850 m, KU 196465-69, 181265-67. Eleutherodactylus melanogaster (5 specimens) Peru: Depto. Amazonas: north slope Abra Barro Negro, 28 km [by road] SSW Leimebamba, 3470 m, KU 212321 (holotype), 212322-23, 218513; 25.5 km [by road] SSW Leimebamba, 3300 m, KU 181281. Eleutherodactylus metabates (2 specimens) Peru:Depto. Amazonas: 20 km [by road] SW Chiriaco, 525 m, KU 196504 (holotype), LSUMZ 32460. Eleutherodactylus muscosus (4 specimens) Peru: Depto. San Martin: east slope Abra Pardo de Miguel, 1800 m, KU 200479-81, 2094782 (holotype). Eleutherodactylus nephophilus (8 specimens) Peru: Depto. San Martin: Abra Tangarana, 7 km [by road] E San Juan de Pacaysapa, 1080 m, KU 212311; east slope Abra Pardo de Miguel, 2180 m, KU 212305, 212306 (holotype), 212307—-09, 212317; 14 km [by road] W Venceremos, 2000 m, KU 212311. Eleutherodactylus ockendeni (71 specimens) Peru: Depto. Amazonas: 4 km [by road] SW Chiriaco, 725 m, KU 196470. Depto. Cuzco: Atalaya, 650 m, KU 154797-99, 154805. Depto. Huanuco: Finca Panguana, Rio Llullapichis, KU 154761-78, 154804, 17183148; south slope Serrania de Sira, 690-1280 m, KU 154779-85. Depto. Loreto: junction Rio Sucusari and Rio Napo, KU 220356-57, 220449; San Jacinto, 175-190 m, KU 222021; Teniente Lopez, 200 m, KU 222022; 1.5 km N Teniente Lopez, 310-340 m, KU 222023. Depto. Madre de Dios: Cocha Cashu, 400 m, KU 154788-96; Manu, 365 m, KU 154786, 154866. Depto. San Martin: Rio Cainarachi, 33 km [by road] NE Tarapoto, 330 m, KU 209468-71. Eleutherodactylus pataikos (1 specimen) Peru: Depto. Amazonas: north slope Abra Barro Negro, 28 km [by road] SSW Leimebamba, 3470 m, KU 212320 (holotype). Eleutherodactylus pecki (9 specimens) Ecuapor: Prov. Morona-Santiago: Rio Piuntza, 1550 m, Cordillera del Condor, KU 147040 (holotype), 14704142; trail between Logrono and Yaupi, 1700 m, ANSP 29271, 29273-77. Peru: Depto. Amazonas: Alfonso Ugarte, Rio Comainas, 1138 m, USNM 525476. Eleutherodactylus percnopterus (24 specimens) Peru: Depto. Amazonas: Alfonso Ugarte, Rio Comainas, 1138 m, USNM 52544546, 525449-63; 20 km [by road] SW Chiriaco, 525 m, KU 196506; 33 km [by road] SE Ingenio, 1830 m, KU 196505, LSUMZ 32462— 63; 5 km [by road] NW Mendoza, 2400 m, KU 209472; upper Rio Comainas, base of Cerro Machinaza, 1750 m, USNM 525443; Santa Rosa de la Yunga, 1300 m, KU 217318 (holotype). Eleutherodactylus peruvianus (79 specimens) Ecuapor: Prov. Morona-Santiago: “Camp 2,” Cordillera de Cutucu, ANSP 29234, 2923744, 29249-50, 29259, 29264; “Camp 3,” Cordillera de Cutucu, ANSP 29286; Rio Pintza, Cordillera del Condor, 1830 m, KU 147034—38; trail between Logrono and Yaupi, ANSP 29236. Peru: Depto. Amazonas: Puesto Vigilancia, Rio Comainas, 665 m, USNM 525440; upper Rio Comainas, base of Cerro Machinaza, 1750 m, USNM 525438-39. Depto. Ayacucho: Estero Ruana in Tambo-Valle de Apurimac trail, 1800 m, KU 196473; Huanuachayocc on Tambo-Valle de Apurimac trail, 1650 m, KU 196472, 196474. Depto. Huanuco: Finca Panguana, Rio Llullapichis, KU 154835-47, 154858-62, 1718667-91; South slope Serrania sw Sira, 690 m, KU 154848-52, 154863-68. Depto. Loreto: Quebrada Oran, 5 km N Rio Amazonas, 85 km NE Iquitos, KU 206094; Quebrada Vainilla, 10 km SSW mouth of Rio Napo, KU 206093; Teniente Lopez, 200 m, KU 222030; 1.5 km N Teniente Lopez, 310-340 m, KU 222024-29. Depto. Pasco: Santa Cruz, 9 km [by road] SSE Oxapampa, 2050 m, KU 206095-97. Depto. San Martin: Abra Tangarana, 7 km [by road] NE San Juan de Pacaysapa, 1080 m, KU 212291; Rio Cainarachi, 33 km [by road] NE Tarapoto, 330 m, KU 208475-77; 20 km [by road] NE Tarapoto, KU 209473-74; 40 km [by road] NE Tarapoto, KU 217312; 15.4 km [by road] SW Zapatero, 950 m, KU 217313. Eleutherodactylus petrobardus (7 specimens) Peru: Depto. Cajamarca: 2 km [by road] W Huambos, 2500 m, KU 212292 (holotype), 212293-96, MHNSM 6196-97. Eleutherodactylus phoxocephalus (36 specimens) Ecuapor: Prov. Azuay: 10 km [by road] SW Victoria del Portete, 2700 m, KU 131281-82. Prov. Canar: 18.4 km [by road] NW El Tambo, 2860 m, KU 142118-31. Prov. Loja: Rio Zamora, 6.5 km [by road] N Loja, 2060 m, KU 142113; Saraguro, 2500 m, KU 135460-62; 3.3 km [by road] NNE Saraguro, 2400 m, KU 142117; 2 km [by road] S Saraguro, 2680 m, KU 142114-16. Prov. Zamora-Chinchipe: 15 km [by road] E Loja, 2710 m, KU 142104-12. Peru: Depto. Cajamarca: San Andres de Cutervo, KU 221710-13. Depto. Piura: 15 km [by road] ENE Canchaque, 1850 m, KU 181271; El Tambo, 31 km [by road] ENE Canchaque, 2770 m, MHNSM 15399. Eleutherodactylus pinguis (4 specimens) Peru: Depto. Cajamarca: 57 km [by road] N Cajamarca, 3760 m, UF 40766; 23 km [by road] SW Celendin, 3050 m, KU 181282, 181283 (holo- type); 33 km [by road] SW Celendin, 3200 m, KU 181284. Eleutherodactylus proserpens (10 specimens) Ecuapor: Prov. Loja: Abra de Zamora, 15 km [by road] E Loja, 2800 m, KU 202130-31. Prov. Morona-Santiago: “Camp 2,” 1700 m, Cordillera de Cutucu, ANSP 29230-33; Rio Piuntza, 1830 m, Cordillera del Condor, KU 14704446. Peru: Depto. Amazonas: Upper Rio Comainas, base of Cerro Machinaza, 1750 mm, USNM 525447. Eleutherodactylus quaquaversus (52 specimens) Ecuapor; Prov. Morona-Santiago, “Camp 2,” 1700 m, Cordillera de Cutuct, ANSP 29257, 29263, 29265-66, 29270, 29281, 29283-85; Rio ELEUTHERODACTYLUS IN THE ANDES OF NORTHERN PERU Wi Piuntza, 1830 m, Cordillera del Condor, KU 146987-90. Prov. Napo: Rio Azuela, 1740 m, KU 143448-54; 2 km [by road] SSW Rio Reventador, 1490 m, KU 165563-65; Rio Salado, + 1 km upstream from Rio Coca, 1410 m, KU 165566-85; 16.5 km [by road] NNE Santa Rosa, 1700 m, KU 14344147. Peru: Depto. Amazonas: Upper Rio Comainas, base of Cerro Machinaza, 1750 m, USNM 525444. Depto. Loreto: 1.5 km N Teniente Lopez, 310-340 m, KU 222031. Eleutherodactylus rhodoplichus (24 specimens) Peru: Depto. Piura: El Tambo, 31 km [by road] ENE Canchaque, 2770 m, KU 219786 (holotype), 219787-91, MHNSM 15400-04; 12.7 km [by road] NE El Tambo, 2820 m, KU 219792; crest of Cordillera de Huancabamba, 33 km [by road] SW Huancabamba, 3050 m, KU 196499— 503, LSUMZ 32417, 32428-33. Eleutherodactylus rhodostichus (4 specimens) Peru: Depto. San Martin: West slope Abra Tangarana, 7 km [by road] NE San Juan de Pacaysapa, 1080 m, KU 212264 (holotype), 212265-67. Eleutherodactylus rufioculis (10 specimens) Peru: Depto. Amazonas: Alfonso Ugarte, Rio Comainas, 1138 m, USNM 525474-75; Upper Rio Comainas, base of Cerro Machinaza, 1750 m, USNM 525464-65, 525467-68, 525473-74. Depto. San Martin: East slope Abra Pardo de Miguel, 2180 m, KU 212312, 212313 (holotype). Eleutherodactylus schultei (16 specimens) Peru: Depto. Amazonas: 5 km [by road]N Levanto, 2850 m, KU 212220-21, 212222 (holotype), 212223-24, 209496-97, MHNSM 6172-73; 5 km [by road] NW Mendoza, 2400 m, KU 209498-504. Eleutherodactylus serendipitus (5 specimens) Peru: Depto. Amazonas: 8 km [by road] NNE Balzapata, 1850 m, KU 181279 (holotype), 181280; 12 km [by trail] E La Peca, Cordillera Colan, LSUMZ 39360, 39363, 39377. Eleutherodactylus sternothylax (49 specimens) Peru: Depto. Piura: 15 km [by road] ENE Canchaque, 1735 m, 196479— 91, LSUMZ 32320, 32332-50, 32352, 32418-27, 32458; 16 km [by road] ENE Canchaque, 1840 m, KU 219793 (holotype), 212794, MHNSM 15405—06. Eleutherodactylus versicolor (69 specimens) Ecuapor: Prov. Loja: 13.5 km [by road] E Loja, 2800 m, KU 1198858 (holotype), 119859-71, 119911-44, 141449-60, 141465-66; 8-9 km [by road] N San Lucas, 3000-3100 m, KU 119945. Peru: Depto. Amazonas: Alfonso Ugarte, Rio Comainas, 1138 m, USNM 525469, 525471-72; Puesto Vigilancia Rio Comainas, 665 m, USNM 525441; Upper Rio Comainas, base of Cerro Machinaza, 1750 m, USNM 525466, 525477. Eleutherodactylus wiensi (7 specimens) Peru: Depto. Piura: 12.7 km [by road] ENE Canchaque, 1600 m, KU 219795 (holotype), 219706-97, MHNSM 15407-08; 15 km [by road] ENE Canchaque, 1735 m, KU 196510-11. APPENDIX 2 GAZETTEER Following is a list of localities in the Andes of northern Peru where specimens of Eleutherodactylus were collected. After each place name the departamento is given in pa- rentheses, followed by geographic coordinates, elevation, and vegetation type. Except for localities on the eastern slope of the Cordillera del Condor documented by Reynolds and Icochea (1997), coordinates were determined from maps (Mapa Fisico Politico del Peru, 1:1,000,000, 1973, and Carta Nacional del Peru, 1:100,000, 1986). Elevations were obtained from altimeter readings or from maps. When known, specific sites and habitats are given followed by names of collectors (only those responsible for field notes) and the month and years that they collected at the sites. Names of collectors are abbreviated: AM = Alfonso Miranda, ERW = Erik R. Wild, FRG = Fred G. Thompson, GKN =G. K. Noble, KLC = Kenneth L. Campbell, LSU = Louisiana State University ornithologists; RPR = Robert P Reynolds, RAM = Russell A Mittermeier, RS = Rainer Schulte, RT = Richard Thomas, WED = William E. Duellman. All localities are shown on the map (Fig. 1). Abra Barro Negro (Amazonas)—06°41' S, 77°51 W, 3500 m; very hu- mid montane forest. A pass in the Cordillera Central on the road be- tween Balsas and Leimebamba. Collections made on the north slope along the road at 25,5 km SSW of Leimebamba, 3300 m, and 28 km SSW of Leimebamba 3470 m in January 1989 by WED. At both sites there were few trees; dominant vegetation consisted of small shrubs (Baccharis), bunch grass, ferns, scattered flower herbs, and mosses 10-20 cm deep. Abra Pardo de Miguel (Amazonas-San Martin) —05 ‘46'S, 77°42’ W, 2210 m; very humid montane forest. A pass in the eastern range of the Cordil- lera Central. Collections made on the east slope at about 1800 m in Janu- ary 1981 by RS and just below the crest on the road to La Rioja, at an elevation of 2180 m in January 1989 by WED. Many trees laden with bromeliads, tree ferns, spiny bamboo (Chusquea spicata) and broad-leafed Gunnera; rocky cliffs covered with mosses and lichens. Abra Tangarana (San Martin)—06 ‘12’ S, 76°44’ W, 1120 m; humid sub- tropical forest. A pass ina low ridge of the Cordillera Central, 7 km NE of the small village San Juan de Pacaysapa on the road between Moyobamba and Tarapoto. Collections were made in a rocky ravine on the western slope by WED in February 1989. Trees in the forest held many bromeli- ads; elephant-ear plants (Xanthosoma) were abundant along the stream. Alfonso Ugarte (= Puesto Vigilancia 3) (Amazonas)—03°54' S, 78°25' W, 1138 m; humid montane forest. Military post on the upper Rio Comainas, eastern slope of Cordillera del Condor. Collections made in July-August 1994 by RPR. Alva (Amazonas)—Approximately 05°53' S, 78°56' W, 1000 m; pre- sumably thorn forest. Site on slopes of Cordillera Central above Pedro Ruiz Gallo in Rio Utcubamba Valley. Collection in May 1974 by RAM. Balzapata (Amazonas)—05 ‘46'S, 77°51' W, 1640 m; humid montane forest. A village just west of the crest of the Cordillera Central. Collec- tions made in disturbed and partly cultivated forest 8 km NNE (1850 m) on the road to La Rioja in March 1979 by WED. Cajamarca (Cajamarca)—07 °12' S, 78°30' W, 2750 m; montane dry for- est. City in Cordillera Occidental. Collection from grassland in humid montane forest at 3750 m, 57 km N of Cajamarca in April 1972 by FGT. Canchaque (Piura)—05°22' S, 79°36' W, 1120 m; tropical dry forest. A town near the western base of the Cordillera de Huancabamba. Collec- tions were made in dry forest near the town and humid montane forest at varying distances from the town on the road across the Cordillera de Huancabamba to the town of Huancabamba in April 1970 by KLC, De- cember 1974 by RT, March 1979 by WED, and January 1991 by ERW. Cataratas Ahuashiyacu (San Martin) —06°30' S, 76‘20' W, 730 m; humid subtropical forest. Waterfall and rocky stream in deep ravine 14 km NNE of Tarapoto on the road to Yurimaguas. Collections in February 1989 by WED. Celendin (Cajamarca)—06°52' S, 78°08 W, 2625 m; humid montane forest. City in the Cordillera Occidental. Collections from the east slope of Abra Comulica, 23 km SW (3050 m) and 33 km SW (3200 m) of Celendin 78 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS in March 1979 by WED. Both sites heavily disturbed, mostly grassy marsh (3050 m) and bunch-grass-Baccharis association. Chiriaco (Amazonas)—05 ‘09'S, 78°21' W, + 450 m; thorn forest. Town on the Rio Maranon a the confluence of the Rio Chiriaco. Collections made in dry forests 4 km SW and 20 km SW (525 m) on road to Bagua in December 1974 by RT. El Tambo (Piura)—05°21’S, 79°33' W, 2770 m; humid montane forest. Settlement on the western slope of the Cordillera de Huancabamba, 31 km NNE of Canchaque on the road to Huancabamba. Collections in rela- tively undisturbed forest in March 1979 by WED and January 1991 by ERW. Huambos (Cajamarca) —06 27'S, 78°57' W, +2200 m; montane dry for- est. Town on the western slopes of the Cordillera Occidental. Collections from “pre-Incan ruins” in 1916 by GKN and in scrubby forest with ter- restrial bromeliads 2 km W (2500 m) in February 1989 by WED. Huancabamba (Piura)—05‘14' S, 79°28' W, 1957 m; dry forest. City on the Rio Huancabamba at eastern base of the Cordillera de Huancabamba; area in immediate vicinity mostly cultivated. Collections from very hu- mid montane forest just east of and at the crest of the Cordillera de Huancabamba, 25-26 km SW (3010-3050 m) in March 1979 by WED and 31-33 km SW (5050-3100 m) in December 1974 by RT and in March 1979 by WED. Ingenio (Amazonas)—05°55' S, 78°55' W, 1280 m; thorn forest. Settle- ment in the Rio Utucubamba Valley. Collection from dry forest on slopes of Cordillera Central 33 km SE (1830 m) on road to Chachapoyas in De- cember 1974 by RT. Lamas (Llama) (Cajamarca) —06°31' S, 79°07' W, +2200 m; thorn forest. Village in valley on west slope of Cordillera Occidental. Collection from 12 km W (1500 m) in December 1974 by RT. La Peca (Amazonas)—05°36' S, 78°22' W, + 1400 m; thorn forest. Vil- lage near western base of Cordillera Colan. Collections from the Cordil- lera Colan—humid montane forest 12 km (by trail) E (1700 m) and very humid montane forest E (2963-3330 m) in August-September 1978 by LSU. Leimebamba (Amazonas)—06 33'S, 77°49' W, 2200 m; humid montane forest. Town in the upper Rio Utcubamba drainage in the Cordillera Cen- tral. Collection from 24 km (by road) SW, 3370 m in very humid montane forest in April 1972 by FGT. Levanto (Amazonas)—06°17' S, 77°21' W, 2650 m; humid montane for- est. Village south of Chachapoyas in the Cordillera Central. Collection from bromeliad-laden trees in highly disturbed forest 5 km N (2850 m) in January 1989 by WED. Herp aL66s. E257 D84 1999 rogs of we genus Eleutherodactyh iii i TU 3 2044 062 4 Mendoza (Amazonas)—06°18’W, 77°25' W, 1650 m; humid montane forest. Town in the Cordillera Central. Collection from pass (headwaters of Rio Chiriaco) 5 km NW (2400 m) in July 1981 by RS. Palambla (Piura)—05°23'S, 79°36' W, 1120 m; tropical dry forest. Vil- lage near the western base of the Cordillera de Huancabamba. Collec- tion in 1916 by GKN. Perico (Cajamarca) —05 ‘22'S, 78°47' W, +1000 m; thorn forest. Village in Rio Maranon Valley; collection in 1916 by GKN. Puesto Vigilancia Comainas, Rio Comainas (Amazonas)—04 ‘06'S, 78°23' W, 665 m; humid subtropical forest. Military camp on the west bank of the Rio Comainas, eastern slope of Cordillera del Condor. Collections made in July-August 1994 by RPR. Rio Cerranayacu (San Martin)—05°46' S, 77°27' W, 1200 m; humid montane forest. Tributary of the Rio Mayo on the eastern flank of the Cordillera Central. Collection from narrow valley near crossing of the Balzapata—La Rioja road in February 1989 by WED. Rio Comainas, base of Cerro Machinaza (Amazonas)—03°53' S, 78°25' W, 1750 m; humid montane forest. Site on the eastern slopes of the Cor- dillera del Condor. Collection in July-August 1994 by RPR. San Andres de Cutervo (Cajamarca)—05°14' S, 78°42’ W, + 2500 m; hu- mid montane forest. Village in the Cordillera Occidental. Collection in June and December 1992 by AM. Santa Cruz (Cajamarca)—06‘05' S, 78°51' W, + 1300 m; thorn forest. Village in the Rio Chamaya Basin. Collection from 28 km N (725 m) in December 1974 by RT. Santa Rosa de la Yunga (Cajamarca) —06 ‘05' S, 78°43’ W, 1000 ms; dry forest. Village on the southern slope of Cordillera del Condor. Collection from disturbed dry forest immediately above village in July 1989 by RS. Tarapoto (San Martin)—06°31' S, 76°23’W, 370 m; humid tropical for- est. Town near confluence of Rio Mayo and Rio Huallaga just east of Cordillera Central. Collections from various distances and elevations in February 1989 by RS and WED. Venceremos (San Martin) —05 ‘44'S, 77°31' W, 1630 m; humid montane forest. Former road camp and now small settlement on the road between Balzapata and La Rioja. Collections from immediate vicinity in February 1989 by WED and from 14 km W (2000 m on road to Balzapata) in July 1981 by RS. Zapatero (San Martin) —06°34' S, 76°30' W, 320 m; humid tropical for- est. Village at base of hills north of the Rio Huallaga. Collections from disturbed humid montane forest 15.4 km SW (950 m) in June 1989 by RS and from ravine in cutover humid tropical forest 30 km SW in February 1989 by WED. PRL DATE DUE MAY 0 4 2 No Vn) DEMCO INC 38-2931 ULY PUBLICATIONS OF THE NATURAL HISTORY MUSEUM, THE UNIVERSITY OF KANSAS 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 published in the above series were published as The University of Kansas Natural History Museum Occasional Papers until Number 180 in Decem- ber 1996. The Miscellaneous Publications of The University of Kansas Natural His- tory Museum began with Number 1 in 1946 and ended with Number 68 in Febru- ary 1996. Monographs of The University of Kansas Natural History Museum were initiated in 1970 and discontinued with Number 8 in 1992. 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